Sunday, December 30, 2012

Indirect effects of parasites in invasions

Recent research emerging from the Global Invasions Network increases our understanding of the subtle but important role of parasites in invaded communities through density mediated and trait mediated indirect effects. The extent to which eco-evolutionary dynamics will shape the adaptive landscape of both native and invasive species in these communities as yet remains unknown.

One of the most dramatic and long-lasting effects of humans on the natural environment is the global spread of non-native species. As is well known, the astounding demographic success of non-native biological invaders has had dramatic effects on ecological communities and is considered a major driver of global losses of biodiversity, second only to degradation and destruction of habitat due to human development according to the International Union for the Conservation of Nature (IUCN). As biological invasions play a central role in the global environmental crisis, the global spread of species has significant consequences for contemporary ecological processes and can also fundamentally alter evolutionary trajectories, not only for native communities, but for the invaders themselves. Considering biological invasions under an eco-evolutionary framework is critical, as it can provide important insights providing a more complete understanding of the causes and consequences of biological invasions in this globalized ‘brave new world’ we’re heading toward that will have, effectively, a single continent: Neo-Pangea.

Recognizing the need to understand the ecological and evolutionary implications of biological invasions on a global scale, an international consortium of scientists convenes under the banner of the Global Invasions Network (GIN), a Research Coordination Network (RCN) funded by the NSF (PI’s: Ruth Hufbauer, Associate Professor at Colorado State and Mark Torchin, staff scientist at the Smithsonian Tropical Research Institute, in Panama). The 2011 GIN meeting in Panama City, Panama, generated some exciting new ideas about the effects of parasites in invasions. Over the past year, I’ve been involved in a working group that emerged out of this GIN meeting, chaired by Alison Dunn and Sarah Perkins, to consider various ways – some rather non-obvious – in which parasites can affect biological invasions.

Parasites are known to strongly influence the success of biological invasions across taxa, since parasite infection (or the absence thereof) can affect both native and invading species. The most obvious impact that parasites have on shaping the ecology and evolution of invaded systems is through direct effects on their hosts, but parasites also have widespread density and trait mediated indirect effects on other species, whether competitors, resources, consumers, or other parasites that interact with a given host that mediate the impact and success of invasion. These indirect effects include both density-mediated effects, resulting from decreases in host survival and reproduction, and trait-mediated indirect effects that result from changes in host life history traits, physiology and behavior. In a review recently published in Functional Ecology (Dunn et al. 2012), we considered the importance of these density- and trait-mediated indirect effects of parasites on the success of biological invasions, and discussed the extent to which these effects scale up to the community and ecosystem levels through the process of invasion. The extent to which parasites affect the eco-evolutionary dynamics of native and invasive species is as yet unknown, but provides exciting opportunities for empirical and theoretical research by identifying the mechanisms by which infectious processes shape the adaptive landscape in invaded systems and the traits that are likely under strong selection. Here I will draw on several examples in the review that illustrate how understanding the indirect indirections of parasites in invaded systems clearly generates natural experiments for evolutionary biology.

Parasites can affect competitive interactions between native and introduced species, which can facilitate the success of invasion by shifting the competitive balance in favor of the invasive species through density mediated indirect effects. For example, the invasive Pox virus, which is not highly virulent to the invasive grey squirrel (S. carolinensis), spills over from the grey squirrel to the native red squirrel (Sciurus vulgaris), which suffers high levels of mortality due to the virulent infection of this invasive pathogen. Theoretical models predict that parasites therefore increase the competitive replacement of red squirrels by the invader (Tompkins White & Boots 2003). Competitive interactions can also be affected by trait mediated indirect effects, affecting both the behavior and physiology of competing species that facilitates the success of invasion. The native ant, Solenopsis geminata) adopts a defensive behavior in the presence of a native phorid parasitoid, Pseudacton browni, that causes a very significant decline in their foraging rates. However, the behavior of the invasive ant, S. invicta, is less affected by the parasitoid, and can usurp available resources, increasing their competitive ability (Morrison 1999). Competitive interactions with invasive species can impose strong selection pressure on native species. Yet as these examples demonstrate, these competitive interactions are mediated by density mediated and trait mediated indirect effects of parasites. Using an eco-evolutionary approach to understand the consequences of these parasitic interactions can generate a framework for predicting how native communities will respond to the impacts of the parasite to mediate the selection strength of the competitive interaction with the invasive species.

Parasites can also increase the demographic success and ecological impact of an introduced species through trait mediated indirect effects on potential resources. For example, a native spionid polychaete (Polydora sp.) weakens the structural integrity of native whelks’ (Nucella lapillus) shells, which broadens the size range of individuals that can be preyed upon by the invasive green grab (Carcinus maenas). By amplifying the resource base of the invasive crab, polychaete infection thus increases both the ecological impact and success of the invasive crab (Fisher 2010). If predation by the invasive green crab imposes strong selection pressure on the whelk population, we can predict an evolutionary response for resistance to the polychaete, or the development of a thicker shell, in order to mediate the impact of infection which increases the ecological impact of the invasive predator. Examples from the plant literature also demonstrate that parasites of resources can increase the ecological impact of invasion through indirect effects on host physiology. For example, an invasive scale insect, Cryptococcus fagisuga, attacks the native American beech Fagus grandifolia, causing mechanical damage to the tree that facilitates infection by an emerging fungal pathogen, causing significant populating declines in the beech across the United States (Kenis et al. 2009). In a similar manner, the invasive bark beetle, scolytus multistriatus, burrows into the elm tree, Ulmus americana, transporting the invasive fungi, Ophiostoma ulm & O. novo-ulmi, which cause Dutch elm disease. This pathogen has caused significant losses (>50%) to elm trees in North America, changing both the community composition and structure of elm forests in North America. By imposing strong selection pressures on elm populations, the bark beetle and the fungal pathogen have the potential to induce an evolutionary response from the elm to resist the impact of either the beetle or the fungus.

Parasites can impose strong selection pressures on affected species, stronger yet if parasites increase the success and impact of invasive species. Ultimately, we need to integrate eco-evolutionary dynamics into investigations of invaded communities to generate a predictive framework of the causes and consequences of biological invasions. Recent investigations elucidate both the direct and indirect effects of parasites in mediating the impacts of invasion. By identifying the underlying processes by which parasites affect the process of invasion through density and trait mediated indirect effects, we may identify traits that are under strong selection, are likely to generate an eco-evolutionary response, and ultimately shape the adaptive landscape in invaded communities.

Dunn, Alison M.; Torchin, Mark E.; Hatcher, Melanie J.; Kotanen, Peter M.; Blumenthal, Dana M.; Byers, James E.; Coon, Courtney A. C.; Frankel, Victor M.; Holt, Robert D.; Hufbauer, Ruth A.; Kanarek, Andrew R.; Schierenbeck, Kristina A.; Wolfe, Lorne M.; Perkins, Sarah E.. 2012. Invasions and Infections: Indirect effects of parasites in invasions. Functional Ecology. http://onlinelibrary.wiley.com/doi/10.1111/j.1365-2435.2012.02041.x/full

Monday, December 3, 2012

Niche theory and speciation

In 2011, I attend a meeting on "Niche Theory and Speciation" in Hungary. The first volume of the special issue resulting from that meeting has just been published in Evolutionary Ecology Research. This first volume includes eight empirical papers plus an introduction, from which I now quote:

"This special collection, which is split between the May and July issues, originated from the workshop ‘Niche Theory and Speciation’ (http://nichews.elte.hu). The meeting took place at Keszthely (Lake Balaton) in Hungary in August 2011, and was organized by Géza Meszéna, Åke Brännström, Ulf Dieckmann, Gabriella Magyar, Liz Pásztor, and András Szilágyi. Funding was provided by the European Science Foundation in the framework of the Research Networking Programme ‘Frontiers of Speciation Research’, and also by the International Institute for Applied Systems Analysis (Laxenburg, Austria)."

"As editors, our goal was to attract the best possible set of papers. We therefore expressly left it open to authors to decide the papers they wished to contribute. Some of the papers are written summaries of work presented at the 2011 meeting, whereas others grew out of discussions during and after the event. We also solicited some additional papers from investigators who had been invited to the original meeting but who were unable to attend. We also strove for a lively mix of empirical and theoretical papers given that progress in the field will undoubtedly require effort from both sides. In the end, all papers emphasized the connection between ecology and evolution, but these links emerged naturally rather than being forced by the editors."


And here are the papers with links to the online versions:


Géza Meszéna & Andrew P. Hendry. 2012. Introduction to Niche Theory and Speciation. 14:361-363.

Luis Fernando De León, Gregor Rolshausen, Eldredge Bermingham, Jeffrey Podos & Andrew P. Hendry. 2012. Individual specialization and the seeds of adaptive radiation in Darwin's finches. 14:365-380.

Jay J. Falk, Christine E. Parent, Deepa Agashe & Daniel I. Bolnick. 2012. Drift and selection entwined: asymmetric reproductive isolation in an experimental niche shift. 14:403-423.


Andrew D. Foote. 2012. Investigating ecological speciation in non-model organisms: a case study of killer whale ecotypes. 14:447:465.


Travis Ingram, Alan G. Hudson, Pascal Vonlanthen & Ole Seehausen. 2012. Does water depth or diet divergence predict progress toward ecological speciation in whitefish radiations? 14:487-502.


I. S. Magalhaes, B. Lundsgaard-Hansen, S. Mwaiko & Ole Seehausen. 2012. Evolutionary divergence in replicate pairs of ecotypes of Lake Victoria cichlid fish. 14:381-401.

Patrik Nosil & Paul Hohenlohe. 2012. Dimensionality of sexual isolation during reinforcement and ecological speciation in Timema cristinae stick insects. 14:467-485. 


Yuval Sapir & Rupert Mazzucco. 2012. Post-zygotic reproductive isolation among populations of Iris atropurpurea: the effect of spatial distance among crosses and the role of inbreeding and outbreeding depression in determining niche width. 14:425-445.


Jorge Soberón & David Martínez-Gordillo. 2012. Occupation of environmental and morphological space: climatic niche and skull shape in Neotoma woodrats. 14:503-517.


Sunday, December 2, 2012

Carnival of Evolution #54

Carnival of Evolution #54 is now up!  We don't have a post in this carnival, since we were so demoralized and outraged by our unjust but crushing defeat in the Halloween pumpkin-carving contest that we apparently couldn't write any further blog posts last month.  But nevertheless, #54 is an awesome carnival, with posts ranging from malaria to slime molds, eusociality to eukaryotes, pleiotropy to pedigrees.  A wild ride, as always.

The carnival theme this month is "A Walkabout Mount Improbable", which makes me think of adaptive walks on fitness landscapes.  So I'll leave you with this picture, ostensibly of a fitness landscape, and a question: do real fitness landscapes look like this?


Tuesday, November 6, 2012

Trick or Retreat

Trick or Retreat
-the Hendry Lab Halloween escape-


Uprooted from its Celtic origins of celebrating the harvest and the end of the growth season, Halloween has evolved over the centuries. For some folks, Halloween nowadays seems to be all about porches filled to the brim with rubber arthropods and absorbent wadding tangled up in fences to signal free sugar to young parents. Well-organized costume contests attract wallflowers and daredevils of all ages alike to thrive in flamboyant displays of recent horror movie themes or social media excrescences. Hours and hours are spent on carving exercises that leave behind disfigured pumpkins waiting in vain for the ghosts they were originally intended to scare away. Modern Halloween has become dominated by ominous competitions. Competitions for neighborhood reputation, party attention, and candy bars… Everybody scurries around between dollar stores and front yards to accomplish as much gory decoration as possible before the turmoil begins. Everybody? Well, not everybody. The Hendry Lab, a small group of indomitable bon vivants, decided to take it easy this year.

Over the hills and far away from all the scars and skulls of the big city, there lies a small wooden cabin northeast of Montreal. It's there the Hendry Lab tends to retreat. Busy with bonfires and hikes, they quickly forgot about all the fuzz in the city and had a great time.

The tin ghost of competition, however, had followed them up into the woods and into the cabin where it manifested in the 'Obscure Booze Festival', a boozy ballot to nominate the most obscure alcoholic beverage of the weekend.

Subset of our sample collection with some dude in the background
And thus, while the Ricciardi Lab grimly sharpened their carving tools and made plans filled with bitterness to showboat around at the yearly McGill pumpkin carving contest, the Hendry Lab explored the unsung liquors of the world. For the sake of completeness, we here report the outcomes in an ordination biplot based on people’s votes for the respective categories.
Ordination plot of the Hendry Liquor Space with category vectors overlain in red; click to enlarge
As the third day dawned over the cabin in the woods, the Hendry Lab - refreshed and brisk - set course for the big city again. Word on the street had it that old Stewart was gathering his faithful from the three wings (and the Redpath Museum) to his yearly fruit-slashing contest in order to anoint the most obedient lab of them all. Such a foreseeable circus never really appealed to the denizens of the Hendry Lab. Previous years had left them without accolade, and it wouldn't be different this year; genius is rarely recognized in its own time. What would be different this year, however, was the mark their work would leave on the souls of bystanders and judges alike – this year’s take-home message would be too scary to take home...

And so the carving began. At one extreme, the Ricciardis with their Bodum® carving set and their neatly assembled 'animaux plastique' – on the other extreme, smelling of guppies and stickleback, the Hendrys, prepared to give everything to restore Halloween’s original meaning and scare away the pagan ghosts. Well, the rest is (micro-)history. Of course, the docile Ricciardis outnumbered and displaced the ardent Hendrys and took the lead with a 'pumpkin arrangement' that might be best described as 'the potpourii of triviality'. Made from polished action figures, sponges, and cheap sparkling bogus, they presented a front-yard altar upon which creativity was laid to rest with much ado.

This kind of 'forced perspective architecture' has already been invented by bower Birds. Nice try, Ricciardis...
The Hendrys were not amused – but also they were not really paying attention. Their mission was to dispel the ghosts of the past, to save the harvest, to drive away the dark clouds that ever gather above their precious liquor cabinet. And they knew that they had to break the rules for this: Over the past centuries whenever harvest season was coming up, people have made straitlaced use of their Cucurbitas and mostly carved rectangular inlets into the fruit. No wonder harvest ghosts aren't scared anymore!

The Hendry Lab changed all that. By deconstructing their pumpkin into elementary pumpkin-subsets and reassembling it inside-out they re-invented fear as we know it – the evil, treacly menace that forces us to question the very foundations of our existence. It is inherent to these existential foundations that we engage in a constant struggle to harmonize our inner- and outer-world relationships, to somehow achieve peace of mind. But what if inside and outside were no longer where they used to be? What if things from deep down inside us would crowd out our shiny, phony façades, pullulating in the pyic cracks of our tanned skins? Behold a glimpse into these dark times that hopefully will never rise.

Take this, harvest ghosts!

Note how the dauntless Hendry himself dares to go eye level with the daemon
We might have gone astray when it comes to old Stewart’s decoration ceremony but at least our pumpkin was actually scary, unlike some we could mention. Also, besides the all-encompassing conspiracy against us, the judges were bribed. Goodnight.

This picture shows a visiting Bridge Club from Ontario who got picked up lost on Docteur Penfield & Peel

Thursday, November 1, 2012

Carnival of Evolution #53

Carnival of Evolution #53 is up!  As Max Headroom would say, ch-ch... ch-ch-ch-check it out.  (Yes, I have just dated myself.  :->)

The theme seems to be sorting.  Luckily, there's a hat that's well-adapted to that niche!


What journal does this paper belong in?  Hmm, it's about snakes...
so perhapsssss... The Journal of Herpetology?  No?  Are you sure?
Their editors are very powerful... they could help you get published...

Wednesday, October 17, 2012

Guppies, predators, and parasites

As many people who follow the blog know, there is a lot of work being done on Trinidadian guppies. Just in case you don't know about guppies, a quick overview. Guppies are native to Venezuela and Trinidad, and in the northern mountain range of Trinidad, it so happens that there is a lovely playground for evolutionary biologists. Starting with Ben Seegher's PhD thesis and continued by very cool scientists like John Endler, David Reznick, Helen Rodd, and Anne Magurran, the guppy became a model system for rapid adaptation. Guppies in low predation areas are more colourful and differ in life history and behavioural traits, among many other things, as compared to guppies that co-habit with dangerous predators. For the purposes of our paper, we focus on male colour, a well known adaptive trait in the guppy.

Pretty guppies.
Photo by Paul Bentzen

The spatial variation in predator composition means guppies have to adapt to their environment to be able to reproduce, and thus, a lot of work has focused on the effects of predation or surveyed differences among populations in relation to the predation level. However, we know life for guppies is very complicated, and that there are other selective pressures acting on them, including parasitism. If we think of adaptation beyond a single causal force, what happens? Will it alter our perceptions of the original causal force? Will the causes interact? Could it be a stronger force than the original one? To address these questions of uni- vs. multi-factorial causes of variation in phenotype, we looked at the effects of both predation and parasitism on male guppy colour.

A pike cichlid, one of the predators
Photo by Kiyoko Gotanda

While there are many ways to consider the effects of parasitism, we chose to focus on the ectoparasite Gyrodactylus spp because previous work has shown that Gyros (as we affectionately call them) can affect a number of traits that also differ between high predation (HP) populations and low predation (LP) populations. The parasite is transferred through fish contact, and we also know that parasitism levels vary considerably in natural populations. We can see Gyros through a stereo-microscope, which means we can assess parasitism levels in the field.

Based on previous work, we know Gyros can impose selection by affecting survival, growth rate, and reproductive success. We also know Gyros can have an effect on behaviour, body size, and colouration, so it is not unreasonable for one to think parasitism can impose selection on phenotypic traits. However, relatively few of these studies are field studies, so we seek to fill that lack of information by building on previous field-surveys.

A Gyro
Photo by Joanne Cable

Specifically, the question we sought to answer was "whether or not parasitism by Gyrodactylus leaves a signature on guppy phenotypes in nature," and by phenotypes, we mean things like male colour. As stated above, we know predation is an important selective agent driving guppy colour where HP fish have less colour than LP fish. Based on previous work, we would expect parasitism to have a negative effect on guppy colour. High rates of parasitism would be expected to lead to lower levels of colour. Thus, if we jointly consider predation and parasitism, is parasitism as important in driving colour differences as predation? Can parasitism modify our perceptions of the effect of predation on guppy colour? To determine this, we surveyed 26 natural populations of guppies in Trinidad. As cool as the science was, I think everyone enjoyed the break from Canadian winters the most... We sample with butterfly nets. How cool is that?

Here guppy, guppy, guppy
Photo by Kiyoko Gotanda

If you want to know our methods, you can check us out online for the details. It involves equipment, chemicals, a digital camera, and a lot of audiobooks. Or you can ask in the comments or email me. I won't bite. I promise.

So what did we find? Well, first, we found that when we sample around the same time in two subsequent years, the parasitism levels are pretty consistent, that parasitism levels among populations varied quite a bit, and that parasitism levels were generally higher in HP sites than in LP sites.  This is consistent with previous surveys, and we speculate this could be because (1) flooding might sweep infected fish downstream from LP to HP populations, (2) HP guppies like to shoal more than LP guppies, possibly increasing transmission, (3) susceptibility might be higher in HP populations, and (4) HP sites might differ ecologically in ways conducive to parasite infection.


But what about colour? Did parasitism have an effect on male guppy colour? Well, interestingly, it appeared not to. Lab studies have shown an effect of parasitism on guppy colour, but we didn't find any direct effects of parasitism on male colour. In fact, a previous field survey didn't find an association with orange colour and parasitism either. So what gives? We have a couple of ideas on why things might be detected in the lab, but not in nature. First, our field survey was a snapshot survey, so we don't know a given guppy's infection history or what the population parasitism levels were like before or after the survey. We know that Gyro epidemics increase and decrease quite rapidly, so it's possible our measures of parasitism are not a true representation of parasitism levels. Second, lab studies can control for a lot of things such as light and resources. We can't control nature, so there might be variation in other selective agents that prevented us from detecting a parasite signature on colour patterning. Third, mortality of infected or uninfected fish can differ, potentially altering parasite and colour distributions.

So we didn't find a significant effect of parasitism on guppy colour. But what if we throw predation back into the story? Does parasitism modify our conclusions about the effect of predation on colour? When considering predation, our results were congruent with what we expected a priori: males in LP sites had more colour than HP sites in general. But we had a lot of variation. It wasn't a clear cut story. Some HP sites had more colour than some LP sites. Previous work has also found similar differences and nuances, and from this, we can conclude that predation IS an important selective agent on guppies, but it's a lot more complicated than just predation alone.

We now ask "does the predation regime story benefit from a simultaneous consideration of parasitism?" Well, we didn't find an effect of parasitism on colour, and adding a parasitism term to our models almost never affected our conclusion about the effect of predation. We still must consider the potential drawbacks of field surveys, but based on our data, we find that parasitism does not modify our interpretation of the effects of predation.However, we are not saying parasitism has no effect, but rather that the signature of an effect of parasitism on male guppy colour might be swamped by other selective agents (e.g. predation) leaving us unable to detect a significant effect of parasitism on colour.

So, in a nutshell, we found that parasites vary between populations and were relatively consistent when assessed over two years, higher infection levels are often found at HP sites as compared to LP sites, parasitism doesn't appear to have an effect on male guppy colour, and considering parasites does not alter our current conclusions about predation.

That's all she wrote!

Pretty snake
photo by Kiyoko Gotanda

Reference:  Gotanda K.M., Delaire L.C., Raeymaekers J.A.M., Pérez-Jvostov F., Dargent F., Bentzen P., Scott M.E., Fussmann G.F. & Hendry A.P. (2013). Adding parasites to the guppy-predation story: insights from field surveys. Oecologia.172(1): 155-166. DOI: 10.1007/s00442-012-2485-7

Friday, October 5, 2012

Indirect Genetic Effects and Eco-Evo Feedbacks



I’ve enjoyed reading through some of the recent entries of this blog and I’m very happy to be invited to contribute. I hope that the ideas of using indirect genetic effects as part of research into eco-evo feedbacks will be interesting.

In a new paper, out in Ecology Letters, we examined how genotype by genotype (G x G) interactions affected plant productivity and pollinator visitation.  We used three genotypes each of Solidago altissima and Solidago gigantea, planted together in monocultures and all interspecific combinations. After two years of growth in large, outdoor pots, we examined pollinator visitation rates and destructively sampled the plants to obtain biomass measurements for flowers, stems and leaves, rhizomes, and coarse roots. We found that neighbor plants exerted particularly strong influences on the belowground phenotype of focal plants. The presence of strong, genetically-based, belowground interactions among neighboring plants makes sense given the expectation of resource, rather than light, limitation. The neighbor genotype effects described above can be considered indirect genetic effects (IGEs), which are influences on the phenotype of a focal individual due to the expression of genes in an interacting individual. To be precise with terminology, we’d call the IGEs in our study interspecific indirect genetic effects (IIGEs), because the interacting individuals were members of different species. 

Earlier I mentioned that we also measured pollinator visitation. We found that genotype by genotype (G x G) interactions influenced how many pollinators visited focal plants. Although we don’t know the mechanism for sure, it may be that synchronicity (or the lack thereof) in flowering times between neighboring genotypes influenced pollinator visitation. Because we didn’t detect G x G interactions for any biomass traits, it’s likely that the mechanism goes beyond variation in productivity due intransitive competitive relationships between interacting genotypes. Whatever the mechanism, G x G interactions are a fundamental unit of the coevolutionary process, suggesting that it is possible that, over many generations, we could observe coevolutionary interactions between genotypes driving changes in the composition of associated communities.

IGEs show that evolutionary forces acting on an individual’s phenotype can come from multiple sources – changes in that individual’s genotype or changes in the genotypes of interacting individuals. Additionally, models approaches have suggested that evolution can occur much faster when phenotypes interact (see Moore et al. 1997). So, in line with this blog’s recent “Kumbaya” spirit, I’d suggest that IGEs and IIGEs could be an important concept that can help unite community genetics, niche construction, ecological genetics, and so on. Perhaps IIGEs could be the arrow that connects “communities” and “phenotypes”, or they could result in changes to a focal plant’s phenotype that affects its interactions with other species. Either way, in my view eco-evo feedbacks are complex and applying names and definitions to the interactions which make up the feedback helps to simplify the issue.

Thursday, October 4, 2012

Carnival of Evolution #52

Carnival of Evolution #52 is up!  Our contribution this time around comes from Andrew Hendry, musing about the ecosystem services perspective and its implications: Ecosystem disservices and assisted elimination.

Posts in the Carnival are, as usual, about every topic under the evolutionary sun; however, the theme of this edition of the Carnival is phylogenetic networks:


M'kay?

Thursday, September 27, 2012

The Year of the Mantis


I like the thought experiment “What would you study if you could start your career over at some previous time knowing what you now know?” The answers vary widely, but discounting the few that say “anything but biology” or “investment banking” or the like, people usually talk about some study organism that they think is really cool. For me, I have periodically thought that it would be great to go back in time and discover and describe for the first time those really bizarre animals that came to scientific attention only recently: naked mole rats or gastric brooding frogs or coelacanths and so on. At other times, I wish I could have been there to describe really cool adaptive radiations of animals – with an obvious one being ants: honey pot ants, leaf cutter ants, slave making ants, army ants, and so on. Recently, however, a trip to the island of Kyushu in Japan crystalized another possibility.

I have always thought praying mantises were really amazing creatures: from their stretched bodies that can nonetheless fly to their deadly front legs to their bizarrely mobile heads that look vaguely humanoid. (They are apparently the only insect that can look over its shoulder.) But all of my experience with them was from pictures in books or sequences in movies – like the BBC one where you think it is going to eat something and then it instead gets eaten by a chameleon. I had only ever seen one in the wild before but that was when I was in line to get into a Lollapalooza concert in eastern Washington, and so I didn’t really have much time for appreciation. This summer turned things around.

The first thing that happened was that my 6 year-old daughter, Cedar, found one in the field at the barn where my wife, Heather, keeps her horse, Delmar. They brought it home and put it in a terrarium and we began to feed it grasshoppers. It was deadly. We could put in 5 grasshoppers and within just a half an hour they would all be dead – at one point the greedy bugger had one in each arm – both alive and struggling. Most of the time, the mantis would stalk the grasshopper and then lash out at the last second with those arms of death but sometimes it would move above the grasshopper and then leap on it. It is truly an awesome predator - and hours of entertainment both parents and kids. Over the summer, its abdomen kept getting bigger and bigger and then one day it laid an egg mass, so here’s hoping we have a cute clutch of the little devils next spring. I can’t wait to release a bunch of baby grasshoppers and watch the chaos.

A Quebec mantis (actually introduced from Europe).
The next thing that happened was that I started to find mantises myself – first at Mt. St. Hilaire near Montreal where I was teaching a field course (see the blog entry here) and then also at the aforementioned barn. In each case, I would spend a long time photographing them and following them around. I never saw them catch anything in the wild but I did get a great appreciation for how their mobile head makes you feel like you are interacting with them. If you get too close, their head swivels around and looks right at you. No other insect can give you the same feeling of communication. Eerie, it is.

The Hiroa-Dai karst tableland near Fukuoka.
Then this last week I went to Japan, where I participated in a symposium on contemporary evolution at the annual meeting of the Japanese Genetics Society. The symposium was organized by Yuya Fukano and my trip – and that of fellow invitee Pierre-Olivier Cheptou – was funded by Tet Yahara with organization help from Makiko Mimura. As most of the meeting was in Japanese, we had plenty of time for excursions – and one of these was to Hirao-Dai, a karst table land near Fukuoka. Along with three Japanese students and postdocs, Pierre-Olivier and I had a great time walking around and looking at a whole series of huge arthropods: wicked looking spiders, great carpenter bees, massive hornets (which apparently kill more people in Japan than any other animal), butterflies as big as birds, katydids spanning your palm, huge grasshoppers, and yes brobdingnagian mantises. So when folks say that everything is smaller in Japan, you can correct them by saying everything but the insects. When these mantis turned their heads to look at me, I involuntarily pulled back.

I see you.

Hanging out.

Come to my arms.
The mantises were everywhere – or maybe it was just that one student (Ryosuke Iritani) was particularly good at spotting them. I decided that if he were a superhero he would have to be called Mantis Boy – he even looked a little bit like a mantis. (Come to think of it, mantis-like qualities would seem better suited for a villain.) In addition to finding a number of individuals of two species just hanging about apparently waiting for a hapless hopper to wander by, we found one that was munching on prey it had just caught, which brings me to a particularly macabre habit of mantises – they quite happily eat their prey alive. As I was taking pictures of this mantis and its prey, a huge carpenter bee, I noticed that the bee was still moving despite having almost all of its’ abdomen eaten away. And the mantis was quite happily tucking in while the bee continued to struggle – if a bit feebly. This isn’t a rare thing, I think, as it regularly happens with our pet mantis at home. In fact, I can still vividly remember it slowly engulfing the leg of a live and struggling grasshopper like we would eat corn on a cob. Presumably they aren’t being intentionally cruel – sometimes they first remove the head – but rather they are just indifferent to the struggling and simply start to eat whatever part is closest. But it is still pretty hideous stuff from the perspective of humans, which always at least have the courtesy to kill their food first - right?

Yum. Nice and fresh.
Humans eat many curious things, especially in Asia. On the second night after arriving in Japan, we went out to dinner and were looking at the menu. Pictured (thankfully given the lack of English) were a whole series of succulent looking meats tastefully arranged on plates. I asked what they were and was surprised to be told in a matter-of-fact fashion that they were whale. Really? Since Japan only engages in “scientific whaling,” I presume this one provided some valuable scientific insight into cetaceans – perhaps it was the N that made P less than 0.05. Seeing whale on the menu without fanfare brought back memories of Norway, where my friends (Ole and Irene Berg) served me home-cooked whale.  I felt guilty enough at the time to call Heather and ask her permission to eat it. I think she would have said no if I hadn’t been standing in their kitchen with the food on the table. The second time I ate whale was also in Norway, this time at a restaurant where I didn’t find out what it was until after having a few bites. So I told these stories to my Japanese hosts, particularly emphasizing the absurdity of calling Heather to ask permission. Then in hopes that I wouldn’t have to call Heather, I flipped the menu to see if something on the other side could divert attention away from the whale. “And what are these I asked?” After a brief consultation, the answer was “horse.” I had to laugh: if these were the only two things on the menu, Heather would probably call me to insist that I eat the whale.

On the way back to my hotel after having dinner, I met Tet Yahara and Jun Kitano, who took me for a second dinner and a fifth and sixth sake. As we walked into the new restaurant/bar, I noticed a tank of squid. Pretty cool I thought, much better than tropical fish, and then I realized what they were there for. I commented to Tet, “So I suppose people get to pick their own squid out for dinner.” Half an hour later and without warning (to me at least) a squid arrived at the table. It looked VERY fresh. It was lying there in a somewhat life-like pose and its mantle had been sliced into a series of delicate an incredibly symmetrical sections that were still in the proper position. Looking closer I noticed then squid was still moving its tentacles and had waves of color moving across it. Was I about to channel a praying mantis.

So it seems to me that this must be – at least for me – the year of the mantis, and if I were to start all over again perhaps I might make it the career of the mantis. But then I did also see mudskippers and tiger beetles, which were almost as cool. Maybe I can start over multiple times.

Japanese mudskipper.

Tiger beetle.

Saturday, September 22, 2012

Ecosystem disservices and assisted elimination

It may seem dead obvious to most of us that preserving biodiversity is the most important legacy that we can leave to future generations – but it turns out that most of us is not the same as most of everybody. Instead, it seems that a large segment of everybody thinks that biodiversity conservation somehow trades off with prosperity to the extent that preserving biodiversity will decrease human well being. This recalcitrant segment of humanity often strongly influences government policy, and so biodiversity scientists have sought convincing arguments for why biodiversity is valuable and important. One example is bioprospecting: many chemicals found in nature have proven beneficial to humans and we can only assume that more such chemicals are out there waiting to be discovered. We must therefore save biodiversity in hopes of gaining those benefits before the species that could provide them go extinct. Another big idea is ecosystem services.
Ecosystem services at the Gault Nature Reserve.
Arguments from ecosystem services embrace the notion that organisms in natural environments make crucial contributions to ecosystem functions that directly and immediately benefit humans, with particularly ubiquitous examples being clean water, productivity, nutrient cycling, and pollination. An immediately obvious limitation of a strict application of this argument is that it might suggest we should simply identify those valuable organisms and preserve them without worrying about the (expensive) rest. But maybe the rest are providing key ecosystem services that we aren’t yet aware of, or maybe they are somehow critical to the dynamics of the species providing known ecosystem services. In recognition of these possibilities, biodiversity scientists have spent considerable time establishing that biodiversity per se enhances ecosystem function – with some of the most compelling evidence coming from experiments that showed greater species diversity (of grasses for example) improves ecosystem function (higher total productivity for example).

The relationship between biodiversity and ecosystem function is not universal, linear, or (often) particularly strong. For me, the most in-your-face demonstration of ambiguity is that clean water (for example) can be found from the tropics to the Arctic despite the fact that species diversity differs by orders of magnitude. (And I can’t help but think of those images of Afganistan with lots of people walking about but not a plant or source of water in sight.) Moreover, many of the ecosystems from which we now obtain our clean water are incredibly impoverished compared to what they looked like centuries or millennia previously. So ecosystem services can be maintained – or perhaps even enhanced (clean water is perhaps hardest to come by in the tropics) – through decreased species diversity.

I suspect that weak and inconsistent relationships between biodiversity and ecosystem function are symptomatic of the reality that biodiversity also provides ecosystem disservices.  Some species are just flat-out bad for clean water, productivity, nutrient cycling, and pollination. And, of course, many species are unequivocally bad for humans in general, most obviously a number of diseases. So perhaps we have, on the one hand, a positive relationship between biodiversity and good aspects of ecosystem function (services) but also, on the other hand, a negative relationship between biodiversity and bad aspects of ecosystem function (disservices). The product of these two lines yields the optimal level of biodiversity, right? Let’s manage for that – problem solved!

Please note tongue firmly in cheek.

It would be complete nonsense to actually manage in this fashion but the absurd point highlights some difficulties of using ecosystem services as the primary argument for biodiversity conservation. In particular, focusing on ecosystem services seems to more obviously argue for the preservation of particular species and the elimination of other particular species (at least when we know which those  bad species are). This brings to mind a currently controversial topic in ecology and conservation biology: “assisted migration” or “assisted colonization.” This is the idea that – particularly under climate change – the optimal conditions for some species will be shifting from their current location to some new (perhaps more northward) location, and that species might have trouble moving to those new locations owing to limited dispersal ability or man-made or natural barriers. For these species, the argument goes, we should get off our butts and actively move individuals to the more appropriate location. The counterargument is that this tantamount to intentionally introducing species to locations where they did not previously exist and the long history of negative impacts of invasive species suggests that this is bad for biodiversity and productivity and services.

Assisted colonization buys into a hands-on approach to management. Another hands-on approach might be called “assisted elimination” (or assisted extinction or assisted extirpation). Acknowledging ecosystem disservices provided by particular species, perhaps we should actively seek out and exterminate those species. We did this for small pox and we try to do it for invasive species or pests all the time. This seems like a pretty sure-fire management strategy given that we can all surely agree on particular species that are just straight-up bad and without which the world would be a better place: HIV, TB, lyme disease, malaria, dengue, mosquitoes, rats, poison ivy (at least near my house), small yappy dogs (particularly my neighbor’s) … Hmmm, mosquitos do many things in the environment and are critical to the existence of a number of other species, such as some bats, dragonflies, and birds. So are mosquitoes or poison ivy or rats really a disservice? What is their optimal diversity or abundance? How would their service-disservice lines intersect? We are on the edge of a slippery slope here.

The point of this post isn’t to argue that we should promote management strategies based on the acknowledgement of ecosystem disservices or that we should or shouldn’t promote assisted elimination. It is instead intended to illuminate the difficulty of making arguments for biodiversity preservation solely from a consideration of ecosystem services – and also to help remember the limitations of a hands-on approach to management. We just know too little. What we really should be doing is preserving biodiversity per se – specifically because of our ignorance. And the best way to preserve biodiversity is to preserve habitats and (yes, Andy) their connectivity.

A biodiversity of scientists enhanced services and disservices at the Gault Nature Reserve.

(The ideas in this post were motivated by discussions at the just-finished DIVERSITAS EcoEvol workshop held at McGill’s Gault Nature Reserve in Quebec, Canada. Participants included myself, Andrew Gonzalez, Paul Leadley, Cornelia Krug, Makiko Mimura, Talisker, Bob Holt, Mark Kirkpatrick, Jon Bridle, Luc De Meester, Dominique Gravel, Jonathan Davies, Virginie Millien, Ulrich Mueller, Highland Park, Justin Travis, Jessica Hellmann, William Godsoe, Mark Urban, Brian McGill, and Rob Coulautti. I am quite certain that many of the other workshop participants will disagree with the ideas expressed in this post and so I hope they will provide comments that devastate what I have here suggested.)

Tuesday, September 11, 2012

The Kumbaya Model and the Jake Reset Effect


Community genetics, evolutionary metacommunities, niche construction, ecosystem engineering, ecological stoichiometry, genes to ecosystems, evolutionary ecology, ecological genetics, evolutionary quantitative genetics, ecological speciation, life history theory, evolutionary rescue, community phylogenetics, coevolutionary theory …

All of these research “fields” with their wonderfully jargony titles involve the study of interactions between ecology and evolution. Surely they can all be united into a single framework under a unified conceptual model of eco-evolutionary dynamics: the “Kumbaya Model” if you will. Adopting a motivating degree of naiveté, the working group meeting from which I just returned had set the Kumbaya Model as one of its key goals.

The previous two meetings of this working group, both funded by the Quebec Centre for Biodiversity Science, had taken place at the Gault Nature Reserve, McGill’s field station/mountain near Montreal. The working group then splintered (or adaptively diversified) into three subgroups: one considering evolutionary rescue, one considering rates and patterns of trait evolution, and one working on the Kumbaya Model. An original driving force behind the overall umbrella working group was Eric Palkovacs, previously at Duke University. In the last year, Eric moved to UC Santa Cruz and so when a meeting location for the third subgroup was debated, California was a promising candidate. Tipping the balance, my family has a vineyard and winery in Napa (http://www.hendrywines.com/) that could host us for free.

The Chardonnay and Pinot Noir harvest was in full swing.
Last Friday, ten eco-evo types (yes, Jonathan Davies, you too now fall under this appellation) descended on the Hendry Vineyard in Napa – and particularly on the house of my brother Mike, his long-to-be-suffering wife Molly, and their enthusiastic dog Jake. For three days, we held hands, sang Kumbaya, and asked of the eco-evolutionary fields “why can’t we all just get along”? When inspiration flagged, we sought more of it through walks, tours of the vineyard and winery, a lot of very fine wine, and a barbeque banquet to end all banquets (Mike and Molly crafted a feast for us that included Mike-caught coho salmon from our cabin in BC and Molly-grown Asian pears from the garden below their house.)

Jonathan used locally available fossils to time-calibrate a large fish phylogeny.
On one of the walks, we were privileged to witness and thereby recognize and appreciate the Jake Reset Effect. At this time of year, reservoirs on the vineyard have very little water in them and are just a few feet deep. With thoughts of maybe initiating some eco-evolutionary research in these reservoirs, we had walked over to have a look – and Jake had come along as a guide. As soon as we arrived, Jake followed his muse and waded into the water and around the reservoir. In doing so, he stirred up mud from the bottom in big plumes. David Post had been standing beside the reservoir and waxing poetic on the power of Daphnia as a model system for studying eco-evolutionary dynamics when he noticed Jake wading through the sediments. “Noooooo …” he wailed “Jake you are stirring up the Daphnia egg bank and destroying decades of adaptive evolution.” Although he was being a bit histrionic, his assertion could literally be true: small and rare disturbances, such as a dog walking through a reservoir, could reset or at least remix the past evolutionary history of Daphnia and shuffle its genetic variation across the decades of evolution buried in the sediments. So, in one fell swoop, it was too late and so much for Daphnia research in the Hendry Vineyard Reservoir. At least the reservoir still has stickleback and I doubt Jake can so easily reset their evolution.
  
"... and another cool thing about Daphnia is ..."
Although walking and drinking wine might seem like frivolous distractions, we actually did make great progress on the Kumbaya Model. I don’t want to spill the beans here, because you will soon (or late) be able to read it in Science or, failing that, the Proceedings of the Southwest Santa Cruz Natural History Society. We, and Eric in particular (see his previous blog entry), have railed about the fact that eco-evo review papers are more common than eco-evo studies that actually present real data, and so we promise that this new Kumbaya Model paper will be the review to end all reviews – the one review to rule them all. In the meantime, we will seek new and clever ways to invade the vineyard again and perhaps collect some real data as a part of Fanie Pelletier’s Global Eco-Evolutionary Research Consortium.

A nexus of the consortium? (Photo by Mike Hendry.)
So what could we do? An excellent way to study the role of evolution in ecological dynamics is to have two replicate populations (such as Daphnia in two ponds) where one population is allowed to evolve adaptively while the other is not. And one of the best ways to prevent directional adaptive evolution is to increase mixing between gene pools that have been selected in different environments, such as populations from different places or that exist at different times – think Daphnia eggs from different depths in the sediment (and therefore different years). As there are two reservoirs on the vineyard, I am thinking we can just have Jake regularly wade through one (adaptation interrupted) but not the other (adaptation perfected). I can see the paper title now: “Experimental Manipulation of the Jake Reset Effect Validates the Kumbaya Model of Eco-Evolutionary Dynamics.” The only question is where Jake should be in the author list.

Jake: for hire as a reset effect technician.
Workshop participants who can all just get along: Andrew Jones, Chris Dalton, Nash Turley, Dan Hasselman, Alison Derry, Fanie Pelletier, David Post, Eric Palkovacs, Andrew Hendry, and Jonathan Davies.

Honorary participants: Mike Hendry, Molly Hendry, and especially Jake.

Photo by Mike Hendry.

Thursday, September 6, 2012

Saunas & speciation

  This is going to be a long post, so put it off for a lazy post-lunch lull, get yourself a cup of coffee, and prop your feet up on your favorite ottoman.  You have been warned.

  I recently participated in the Helsinki Summer School on Mathematical Ecology and Evolution, which this year focused on the theory of speciation.  The school was a part of the larger series of schools, workshops and conferences organized by FROSpects, an ESF Research Networking Programme that has been doing a great job of supporting and promoting speciation research – I've attended several of their events in the past, such as the Abisko Winter School in 2011 and the Speciation 2010 conference in Austria, and they have had a big influence on my path as a graduate student.  I'll talk about the Helsinki summer school in a moment, but first, I can't resist giving you a brief touristic interlude.

  Since I've never been that far northeast in Europe before, I seized the opportunity and went for an overnight visit to St. Petersburg (only three and a half hours away by train!), which was fun but exhausting.  The high point for me was seeing the iconic Church on the Spilled Blood, which was every bit as spectacular in person as it is in photos:



  The Hermitage was amazing too, and could have taken up much, much more than the half-day I could give it.  It was heartbreaking to have to hurry through rooms filled with works by Cézanne, Matisse, Van Gogh, Corot, Caravaggio, Gauguin, Pisarro, Degas, Renoir, Seurat, Vlaminck, Picasso... I will have to go back some day.  They had great exhibits of ancient art and artifacts from around the world, too, which I enjoyed a lot, and tons of Middle Ages and Renaissance art that I mostly skipped due to lack of time.  (I did suffer the tourist hordes to glimpse the two very famous da Vincis there, though.)

  One lovely thing about the Hermitage is that unlike many stuffy, arbitrarily restrictive museums, they mostly allow you to photograph their exhibits (but no flash, of course, since it can damage the art).  The first photo below shows an example of the opulent interior of the museum, which was once a palace; the second shows a nifty Kandinsky that caught my eye; and the third shows a remarkable mask and breast collar for a horse, from around the 3rd-4th century B.C., from the Pazyryk culture of the Altai – a region in central Siberia that I had never even heard of before:




  I got to spend a little time in Helsinki, too, on the way in and out, which was great.  It has a beautiful downtown area with an open-air market, small side streets and lovely churches:


  And also quite a remarkable island fortress called Suomenlinna that you can explore, walking both on top of and inside of the walls and admiring the old cannons:




    Amusingly, the name Suomenlinna is Finnish for "Finnish castle", while the Swedish name for the fortress, Sveaborg, is Swedish for "Swedish castle".  (It was built by the Swedes in 1748, then was captured by the Russians in 1808 – but I don't know whether their name for it would translate to "Russian castle" – and then became Finnish when Finland became independent in 1917.)

  The school was actually held, not in Helsinki, but in Turku, another town in Finland to the west of Helsinki.  I didn't have much time to see Turku – in part because I managed to crack a rib early in the summer school, attempting to play soccer one fine evening, which subsequently limited my mobility – but the Turku cathedral dates back to the 13th century (although much of it was rebuilt after fires much more recently) and was quite groovy.

  Well, I'm supposed to be talking about the summer school, aren't I!  If I were Andrew, I would find a way to weave some quirk of my travel experiences together with musings about evolution to form a unified tapestry.  Suomenlinna and the evolution of defenses against predation and parasitism?  The Hermitage and the adaptive value (if any) of the human sense of aesthetics?  But I am not feeling inspired, so please pardon the abrupt transition from tourism to biology.



  The school had five main speakers: Sergei Gavrilets, Nick Barton, Sander van Doorn, Eva Kisdi, and Dan Bolnick.  First, however, Mats Gyllenberg opened the school with a talk about competitive exclusion, limiting similarity, and coexistence; this is a central question for speciation theory, of course, whether coexistence occurs with secondary contact after divergence in allopatry, or is an ongoing issue as divergence proceeds in sympatry or parapatry.  Ultimately, all life on our planet descends from one common ancestor, and yet that life has managed to subdivide the niche of "planet Earth" (which one could imagine being dominated by one extreme generalist species) into enough partitions to support the many millions of species now extant, finessing the Principle of Competitive Exclusion in a myriad ways. The upshot of Gyllenberg's talk, which I believe was based upon work by Dieckmann, Metz, and Meszéna, was that coexistence of an infinite number of species is mathematically possible – the variety of biodiversity could be continuous, in other words, rather than divided into discrete packets that we call "species" – but that this result is not robust, and collapses into a finite number of discrete species subdividing the available dimensions of niche space given any sort of perturbation from stochasticity. Paraphrasing him, the messy realities of biology prevent the coexistence of infinitely many types.

  Sergei Gavrilets gave quite a fast-paced set of lectures (despite his laptop dying at the beginning of the school).  He started with the history of population genetics and the earliest forays into mathematical modeling and theoretical speciation research.  Then he discussed fitness landscapes of various kinds, evolution on them, peak shift models and the difficulty of getting across valleys, the Bateson-Dobzhansky-Muller model of neutral mutations leading to reproductive isolation due to epistatic interactions, and movement along "nearly neutral networks" on high-dimensional "holey adaptive landscapes."  Much of this covered similar ground to his very important 2003 paper "Models of Speciation: What have we learned in 40 years?"  Then he wrapped up with an overview of models of non-random mating and sympatric speciation, touching on the importance of a cost of choosiness in inhibiting the development of assortative mating.  Little is known empirically about costs of choosiness, but this is a central consideration for speciation theory.  Empiricists, take note!

  Nick Barton walked us all through the rather intricate details of a remarkably powerful extension to population genetics that he has developed with Michael Turelli and Mark Kirkpatrick.  This method allows the effects of dominance, epistasis, and linkage disequilibrium to be accounted for much more tractably than with more traditional methods.  In many situations these effects can be ignored, because populations are often well-mixed (because selection is weak compared to recombination), and so selection acts chiefly on the marginal, additive effects of genes.  Not so in speciation theory, however, because speciation typically implies strong selection, non-random mating, and mixing of populations between divergent habitats, all of which cause linkage disequilibrium, and because reproductive isolation is often based upon epistasis, and so forth.  Indeed, he said that speciation might best be formally defined in terms of linkage disequilibrium; so clearly speciation theory must tackle the causes and consequences of that!  In later talks Dr. Barton used his method to explore speciation in the Levene model, with reference to Felsenstein's classic 1981 "Santa Rosalia" paper, and then wrapped up with a survey of models of parapatric speciation and clines.

  Sander van Doorn talked about the role of sexual selection in speciation.  Dr. van Doorn is one of those speakers who is so eloquent that he makes very complex ideas seem simple... until, a few moments after he stops speaking, the illusion of perfect understanding that he had cast over you melts away.  Sexual selection is a multifarious topic.  There are cue traits and preference traits, ornaments and honest signals and nuptial gifts and parental investment, male-male competition and female choice, good-genes models and magic-trait models and Fisherian runaway sexual selection models. There is the problem of coexistence of ecologically identical species after speciation due to sexual selection, and the problem of the origin of variation in mating preferences.  He surveyed a wide range of models touching upon these and other issues, and I'm not going to try to summarize it all except to say that it was very interesting and I look forward to a review paper from him covering the same ground!

  Eva Kisdi gave a very clear and gentle introduction to adaptive dynamics, starting with the Levene model and the fine-tuning problem, then introducing the tools of adaptive dynamics from pairwise invasibility plots to the criteria for classifying singular points.  One pithy statement that she made stuck with me: natural selection is usually called "the survival of the fittest", but when an evolutionary branching point is present, it instead selects for increasing diversity!  Perhaps that nicely encapsulates why I find speciation theory fascinating.  She walked us through a very cool proof of a method of determining the constraints on trade-off functions that will produce evolutionary branching in any given model – even models for which analytical results cannot be obtained.  In later lectures, she showed how to do adaptive dynamics for sexual populations, and then applied that method to a magic trait model.

  Finally, Dan Bolnick grounded the school with talks (and a workgroup that I participated in) on the interactions between theory and data.  He started with a summary of what he thinks are the big questions in speciation research today, and summarized contributions from theorists and empiricists to those questions.  I found this to be a very helpful clarification of the "big picture": why we were all gathered together in a classroom for a week to talk about speciation.  He then took a step back and talked about how exactly theory and data can interact, what theory typically does for empirical work, what empirical work typically uses theory for, and the limitations inherent in those interactions.  He proposed that there are four types of interaction.  The first is that theory and empirical work can be "in the same room", sharing a biological context but not actually meeting or speaking to each other.  Alternatively, work can be data-driven, with novel empirical findings driving new theory to explain what was observed, or theory-driven, with interesting theoretical findings pushing empirical investigation of questions that had not previously been examined.  Most intimately, there can be a reciprocal interaction in which each prompts new work in the other, in a feedback loop between theoretical refinements and new experiments.  He capped things off with a tale of Tribolium, an insect that has been leading him and a postdoc in his lab on a wild goose chase through both theory and experiment, with a mysterious cliff-hanger ending that has yet to be resolved.

  Other speakers touched upon the interaction between theory and data too:
  • Dr. Gavrilets, for example, opined that the maturity of any scientific field can be assessed by its level of mathematical sophistication, and that evolutionary biology is therefore the most mature subfield of biology – a provocative claim! – and also quoted Haldane as saying that one should mistrust any verbal argument when an algebraic argument would instead be possible, and that skepticism is warranted whenever not enough facts are known to permit an algebraic argument from being formulated.

  • Dr. Barton emphasized that he thinks there is a need to relate theory more to things that we can actually measure, and for empiricists to then actually measure those things.  For example, adaptive dynamics predicts that populations will evolve towards stationary points and then often experience disruptive selection; this is an extremely important prediction, but it has not been tested.  Can it be tested?  If not, can the predictions of adaptive dynamics be framed somehow in a more testable way?

  • Dr. van Doorn pointed out that in reality, a species can respond in many different ways to selection pressures: evolution of dominance, concentration of genetic variance at just a few loci (evolution of polymorphism), broadening their resource utilization curve, evolving sexual dimorphism, evolving plasticity... but because of the kind of models we build, with very little genetic flexibility or realism, we have little to say about when these different outcomes should be expected instead of speciation.  Can we make better models?  How can empirical data help us to do so?

  • Dr. Kisdi, in her discussion of trade-offs, emphasized that so little is known empirically about the shape – or even the existence! – of trade-off functions in nature that theorists have more or less free rein to speculate in this area.  In the workgroup that I participated in with Dr. Bolnick, we returned to this issue in more depth, and one of my fellow students, Pavel Payne, gave a brief presentation about what little is known about trade-offs in nature.  How can we know so little about something so central?  What can we can do improve this state of affairs?  And are there ways that theorists could help what what must ultimately be an empirical project in this area?
  For me, these discussions of the interaction between theory and data were the most thought-provoking part of the summer school, because they spoke to my developing identity as an evolutionary modeler.  What does it mean to be a modeler?  If your models are untestable, are you still doing biology, or are you doing pure mathematics?  Is evolutionary modeling an undertaking whose primary aim is to inform and explain empirical work, or is it a field that stands on its own, and that does not necessarily need to be useful to empiricists to be worthwhile?  Alternatively, if Dr. Gavrilets is right that it is the mathematical models that show the maturity of the field, then is empirical work in biology perhaps an undertaking whose primary aim should be to supply data, parameter values, and scenarios as fodder for modelers?  Do theory and data have a mutualistic relationship, or a commensal relationship, or a competitive relationship – or is one perhaps even a parasite of the other?  Is the "reciprocal interaction" between theory and data that Dr. Bolnick described the best way for the two to interact, or are other ways – even just being "in the same room" – just as good, or perhaps even better?  There were as many answers to these sorts of questions as there were participants at the school, and that made for very rich and interesting discussions, to me.

  I got to go with my friend in Helsinki to a traditional public wood-burning sauna (awesome!), and then several times I used the sauna at the convention center where the summer school was held, too.  So I can definitively state that saunas are not only good in theory; I have confirmed this experimentally, and I have replicated my results several times in independent trials.  This, I think, is the best of all possible outcomes of the interaction between theory and data.  May all your research endeavors go so well.

   And perhaps that's where I can find a thread to tie together my touristy travelogue with the summer school.  Ultimately, for me, it is the real world that I wish to understand and explain.  Reality is more beautiful than any mathematical model, to me, and mathematics is just one tool (albeit both a beautiful and useful one) to help us understand reality.  No model would ever predict the Church on Spilled Blood.  Ergo, tourist snaps of St. Petersburg.  And on that note, let's shift back to the tourism to wrap things up!



  At the end of the school, we spent the final evening in Naantali, a charming little tourist trap on the coast.  For some reason, Naantali is associated with the Moomintrolls, a set of children's book characters, and there was Moomin-kitsch everywhere.  I grew up reading the Moomin-books, though, so it was fun for me (and I now regret not buying something related to the Hattifatteners, my favorite part of those books).  We had a really yummy dinner at a local restaurant and enjoyed the views of the Baltic and the amazing Archipelago Sea off the coast of Finland.  Here's a Moomin-flag for the Moomin World at Naantali, and then a shot of Dan Bolnick looking for (and finding!) stickleback in the waters of the Baltic, with the beginnings of the archipelago behind him:



  The summer school was organized by Eva Kisdi and Tadeas Priklopil, who did a great job of making the event run smoothly.  Thanks to Eva and Tadeas, to the lecturers who gave so much of their valuable time, to my fellow students for making it fun, and to FROSpects and the other organizations that made this school possible!


Participants at the summer school.  (I'm the guy in the lower
right who is so white that he looks like he'd glow in the dark.)


  Those who want more travel details and photos will eventually be able to find them on my photography website, cloudphotographic.com, but they won't be there for a couple of months, probably, and by then you will all have forgotten.