The EEB & flow

Should we still be testing neutral theory? If so, how?

2012-01-30T15:09:00.000-08:00

Siepielski, A.M. , Hung, K-L., Bein, E.E.B, McPeek, M.A. 2010. Experimental evidence for neutral community dynamics governing an insect assemblage. Ecology. 91. 847-857

For many ecologists, neutral theory was a (good/bad, you choose)idea that dominated ecology for the last decade but failed to provide the burdenof empirical proof necessary for its acceptance. Even its creator Stephen Hubbell recently suggested that the controversialhypothesis is no longer a plausible description of community structure, goingas far to say that it is “good starting point”, a “valuable null model”, and a“useful baseline” (in Etienne et al 2011).

But ideas, when they’re shared, are no longer the soleproperty of their creators. Other researchers continue to study neutral theory,and despite the apparent consensus that neutral theory is not an importantexplanation of community structure and dynamics, papers testing neutral theory continueto be published. This leads to an important question: do we still want to testfor neutral dynamics? And if we do, how should we approach it, given what wehave learned from the past decade of strawman arguments and using pattern-basedevidence for processes (e.g. looking at species-area relationships and speciesabundance distributions)? What empirical evidence would provide strong supportfor the predictions of neutral theory?

Damselfly larvae
(http://www.uta.edu/biology/robinson/odonate_research.htm)

In “Experimentalevidence for neutral community dynamics governing an insect assemblage”,Siepielski et al. (2010) attempt to provide a more rigorous test of neutraltheory using two Enallagma (damselfly) larvae. Siepielski et al. focus onchanges in demographic rates (growth, mortality) in response to changes in speciesrelative and total abundances. In particular, they predicted that if nichedifferences drive coexistence, increasing a species’ relative abundance shoulddrive lower growth rates and higher mortality, since that species is above itsequilibrium; lowered relative abundances should result in higher growth ratesand lowered mortality since the species is below its equilibrium density. As aresult, species should return to their equilibrial abundances. Raising thetotal abundances but leaving the relative abundances untouched should havesimilar demographic responses across species and have no effect on the relativeabundances. In contrast, neutral theory predicts that if all species are equal,their demographic rates depend on the density of the entire group (totalabundance) and not on each individual species’ relative abundance. Thereforethe response of demographic rates to changes in species relative abundances,while the total abundance is held constant, should provide support to eitherneutral or niche theory.

For two Enallagma sp. larvae Siepielski et al. used cages in the littoral zone of lakes, with cagesreceiving different treatments of relative abundance and/or total abundancemanipulation. The result of these manipulations were that replicates withincreased total abundances and constant relative abundances had loweredper-capita growth rates, while replicates with manipulated relative abundancesand constant total abundances showed no change in demographic rates. Bothspecies had similar mortality rates across the experimental treatments,although their growth rates differed slightly. From these results, Siepielskiet al. concluded that these species are ecologically equivalent.

One of the reasons work (such as this) from Mark McPeek’s lab is interesting is because he is an outlier: someone whose workis deeply rooted in a natural system, and yet who also argues that ecologicalequivalency seems plausible, and attempts to support that argument. Regardlessof whether the Enallagma species are in fact ecologically equivalent, this paper providesan example of how coexistence theory can be more rigorously tested than simplyobserving species co-ocurrences and concluding species coexistence. Further, itprovides some interesting discussion about whether ecological equivalency ispossible within functional groups, with niche differences occurring betweenfunctional groups (see Leibold and McPeek 2006, and from MacNaughton and Wolf 1970 forsimilar suggestions). Future work might focus on questions such as how tocapture the effects of small niche differences, which, if balanced against verysimilar fitnesses could explain stable coexistence. In addition, it might bevaluable to look at how resources fluctuate and how much overlap there is inresource requirements among species, when looking at how growth and mortalitychange with species densities.

With Adam Siepielski, Mark McPeek also published thepaper “On the evidence for species coexistence: a critique of the coexistence program” about theapparently lowered standards for tests of niche-based species coexistence comparedto those of neutral theory. What is certainly true is that experimental tests ofcoexistence theory are often less rigorous than necessary to support any coexistencetheory, and should strive to take a more rigorous approach. If nothing else,this will allow criticism of particular theories to focus on the ideasthemselves, rather than on how those ideas were tested.

Trends in ecology, 2011

2012-01-10T15:09:00.000-08:00

What were the topics of research that dominated ecology in 2011, and where is ecology likely to head in 2012?

A brief answer can be found by looking at the most common keywords found in ecology papers published during 2011*. "Abundance" proved the most common keyword. Interestingly, "climate change" and "global warming" appeared less common as keywords compared to last year. In contrast, words tying research to places ("Great Barrier Reef") and systems ("rainforest") seemed more common. Although it's hard to draw any specific conclusions from this kind of thing, it's notable that many of the most common words are related to community ecology, lending credence to Marc Cadotte's assertion that community ecology is flourishing as a discipline.

*Although hardly rigorous, I analyzed the keywords from 4000 randomly selected ecology papers published in 2011 found using a Web of Science search. The most common 150 terms are represented in the word cloud, where text size represents the frequency with which a word appears on the list.

Carnival of Evolution 43!

2012-01-03T06:27:00.000-08:00

The history of human thought is an epic adventure of exploration and discovery. Since the beginning of time, humans have been curious about order and chaos in nature and our place in the world. By understanding the natural world around us, we understand ourselves better. But how we attempt to answer these fundamental questions has evolved over time. This evolving history, looks something like this:

146,000 BCE
Targ: "Hey Lerb, why big cat have long teeth?"
Lerb: "I dunno Targ, but cousin Seb went for look. He gone"
Targ: " Cat lucky, seem good for people eating. I go for closer look."

523 BCE
Anaximander: "Thales, my teacher, how is it that animals take their form?"
Thales: "Anaximander, all matter is an aggregation formed from a single substance, water, and qualities are obtained through need"
Anaximander: "Ah yes, water, I will now think about how air can be the primordial substance."

1849 CE
Thomas Thomson: "I do say, the flora of northern India is peculiar in the sheer number of forms of life that populate this region. I do wonder though, what the cause is for such brilliant numbers of species?"
Joseph Hooker: "My dear Thomas, the flora of northern India is brilliant indeed! These forms find their origins in those very places where they live. Of course Lamarck believes that the crises endured by the tissues of organisms, themselves pass on the incentive to produce offspring better equipped to endure such crises. However, in my correspondence with Charles Darwin, he confirms that variation is an inherent aspect of life and gives rise to the diversity we see."

2012 CE
You: "Man, I wish I new more about evolution. Hey, what is this Carnival of Evolution? OMG, this is totally sick."
You no longer need to ponder the mysteries of life, travel the globe making observations, or running complex experiments to test hypotheses; everything you want to know about evolution today can be found by reading the monthly installments of the Carnival of Evolution!

The first installment of 2012 (or is this the last of 2011?) offers a great smattering of many different aspects of current evolutionary understanding. These 26 posts cover many of the major areas of research that define current evolutionary biology.

Most evolutionary research aims to understand how the amazing diversity of life came to be. Core to this is studying both paleontological record and patterns among modern organisms. Early explosions of diversity have always captured scientists imaginations, and Larry Moran at Sandwalk (and fellow Torontonian) explains that recent evidence is casting doubt that the Cambrian explosion was actually an explosion, at least according to genetic evidence. Much later on, ray-finned fish became extremely successful and are now the dominate form of fish on Earth. Their success and resulting diversification is likely better explained by rapid morphological changes to head shape and not fin evolution, according to Lucas Brouwers at Thoughtomics. Nothing in the paleontological record excites the imagination more than dinosaurs. Recent work has developed a detailed understanding of the ecology and evolution of these amazing creatures. Marc Vincent at Love in the Time of Chasmosaurs describes research that indicates that head crests and feathers on many dinosaurs were likely to product of sexual selection. While, according to David Orr, also at Love in the Time of Chasmosaurs, the big toe claw on both hind feet of Deinonychus evolved to pin down small prey, and not to slice open large prey (thank you Jurassic Park). In one of the best, 'huh, I didn't know that' posts, Fins to Feet shows that Mosasaurs -giant predatory marine reptiles found during the time of the dinosaurs, are likely closely related to monitor lizards and not part of the ancestral lineage that includes dinosaurs.

Studying and explaining patterns among modern day critters is the evolutionary biologists' bread and butter, and studies of organisms seem to constantly shed light on new ways in which evolution has shaped life. The interesting story of the oil beetle and how it has evolved to hitch rides on other insects is presented by Anne Buchanan The Mermaid's Tale. As relayed by Jeremy Yoder at Denim and Tweed, birds that lay eggs in the nests of other species (nest parasites) have been associated with the same hosts for millions of years. Flower colors are commonly thought to be shaped by pollinator preference, but Zen Faulkes at NeuroDojo shows evidence that white variants of bluebells (are they still bluebells?) do not see different pollinator visitation rates. Species differences can be difficult to identify using our human senses, but Lucas Brouwers at Thoughtomics explains how echolocation has diverged between indistinguishable bat species. Lungfish are oft-cited exemplars of evolution, mainly because they are so fascinating -not only do they have lings, but they walk too. Which is why they are the subject of two posts this month (one by Matthew Cobb at Why Evolution is True and one by Carl Zimmer at the Loom), both about how they move and how they may have transitioned to walking.

Evolutionary change for many animals is often not a linear move from genes to fitness, but rather behavior has the potential to affect evolution in complex ways. In one example, Jeremy Yoder at Denim and Tweed, tells the tale of how fear of being eaten can lower fitness. In another example, Simon's Science explains research that shows female stickleback fish, which are raised by their fathers, will prefer mates from their father's species, even when experimentalists switch the species providing parental care.

To unveil the wizard, a number of posts show how evolutionary research is done and how our understanding evolves. In two posts at BEACON, researchers Tasneem Pierce and Michael DeNieu give fascinating firsthand accounts of doing research. You can sense the wonder and excitement of doing scientific research from their posts. Stan Rice at Honest Ab has a wonderful sequence of five posts relaying his dinosaur adventure -at least playing with paleontology and avoiding creationists. John Wilkins at Evolving Thoughts examines the definition of evolutionary novelty in an ongoing series (maybe his next book?), and looks at comparative versus functional definitions.

For most people, evolution is central to the ultimate questions about who we are and where we come from. True to this anthropocentric* view of evolution, there are a large number of excellent posts about human evolution and why we are the way we are. For those people who feel that the Carnival of Evolution does not provide all the answers to their questions about human evolution, Greg Laden reviews two new books on understanding human evolution. Suzanne Elvidge at Genome Engineering reports that scientists have sequenced the genome of a descendent of Genghis Khan. Why is this interesting? Well it turns out that millions of people -half a percent of the current global population, are related to Genghis Khan! The obvious question to me was how is this possible? It turns out that, according to Wikipedia, Genghis Khan had a harem of between 2000-3000 women and many of his many, many sons also had obscenely** large harems. Thus, by the time Genghis was a dirty old man, he could have had 10,000 descendents.

Often the need for evolutionary explanations comes from the question: "Why the heck do we do that?". True to this question, there are four posts that look at human behavior. In a controversial but intriguing post, Khudadad Azara at Khudadad's Knols suggests that terrorism is a macho impulse for glory and honor shaped by sexual selection. The most convincing parts are that males often do stupid things for sexual advantage, and terrorism is a stupid thing. Why the hell are yawns contagious? Well, according to Suzanne Elvidge at Genome Engineering, yawns may be evolutionary as they are most contagious among close relatives. Holly Dunsworth at The Mermaid's Tale makes the case that the uniquely human ability to throw (chimps actually aren't very good at it) is not so much an anatomical thing, but a brain thing, interesting. PZ Myers at Pharyngula asks why women menstruate and suggests that it is the evolutionary result of mother-fetus conflict.

A big part of human history, culture and belief, is our conflict with disease. This month there are several very interesting posts on evolution and human disease. Swenson at Nothing in Biology Makes Sense! discusses how reconstructing the evolutionary relationships among HIV samples dating as far back as 1959 reveals that there are deep divergences indicating that HIV has likely been in humans since the late 1800's! Carl Zimmer at the Loom relays the latest research showing that Syphilis evolved in the New World and was likely brought to Europe (Italy) from early European explorers. Ford Denison writes at This Week in Evolution that a genetic mutation increasing the risk of breast cancer in women is also associated with increased fertility. This invites the conclusion that there may be a tradeoff between longevity and fertility.

That is all for this month from the Carnival of Evolution. Everything you ever wanted to know about evolution but were afraid to ask. When you start to have new questions, luckily there will be a new edition of the Carnival in a months time.

*I realized after I wrote this sentence that it sounds negative. I do not mean the increasing pejorative 'unnatural', but rather legitimately human-focused.

**Having a harem of any size is obscene, but what adjective can you use for harems with thousands of women?

Rumors of community ecology’s death were greatly exaggerated: reflections on Lawton 1999

2011-12-26T19:10:00.001-08:00

In 1999, John Lawton, eminent British ecologist, published a lament for the state of community ecology entitled “Are there general laws in ecology?” Cited more than 600 times, Lawton’s paper forced a re-evaluation of community ecology’s value, success, and even future existence. Other scientists at the time seemed to agree, with papers starting with phrases like “Although community ecology is a struggling science…” and “Given the lack of general laws in ecology…”. Lawton appeared to be suggesting that community ecology be abandoned for the generality of macroecology or the structure of population ecology.

An important point to be made is that Lawton was simply making a particularly public expression of ecology’s growing pains. In 1999, ecology was at a crossroads between the traditional approach of in-depth system-based study, and a fairly single-minded focus on competition as an explanation for patterns (e.g., Cooper 1993 ‘The Competition Controversy in Community Ecology’ Biology and Philosophy 8: 359-384), while at the same time there were emergent approaches and explanations like neutrality, macroecology, spatial ecology, ecophylogenetics, and improved computer and molecular methods. There was also growing dissent about ecology’s philosophical approach to ecology (e.g., Peters 1991 ‘A Critique for Ecology’; Haila and Heininen 1995 ‘Ecology: A New Discipline for Disciplining’ Social Text 42: 153-171): ecologists tended to ignore the Popperian approach, which required falsification of existing hypothesis, instead tending to look for support for an existing hypothesis, or at least advocated looking for patterns without considering alternative mechanisms. Not only this, but the applications for ecology were more clear than ever – the Intergovernmental Panel for Climate Change was meeting , and the ecological consequences of human actions were perhaps more obvious they had ever been. But ecologists were failing at providing solutions –Lawton argued-correctly-that in 1999 ecologists could provide little insight into how a community might change in structure and function in response to changing climate.

Although everyone should read Lawton’s paper, a simple synthesis of his concerns would be this – that community ecology is too contingent, communities are too complex, and therefore community ecology cannot formulate any laws, cannot make predictions, cannot be generalized from one system to another. This makes community ecology suspect as a science (physics being the most common example of an “ideal” science), and certainly not very useful. Lawton suggests that population ecology, where only a few models of growth could explain the majority of species’ dynamics, or macroecology, which focuses on the most general, large-scale patterns, were a better example of how ecology should be practiced.

Community ecology, rather than dying, has experienced an incredible surge in popularity, with a large contingent represented at meetings and in journal publications. Ecology itself is also thriving, as one of the fastest growing departments in universities. So what, if anything, has changed? Has ecology addressed Lawton’s criticisms?

Two major things happened in the late 1990’s and early 2000’s, which helped ecologists see beyond this general malaise. The first was that a number of well-thought out alternative ecological mechanisms explaining community membership were published. Before the late 90’s community ecologists looked for evidence of competition in patterns of community composition, either among locales or through time following disturbance. When local competition was insufficient to explain patterns, researchers likely cited, but did not test other mechanisms. Or if they did test other mechanisms, say predation, it was as an alternative, mutually exclusive mechanism. The new publications, drawing on previous ideas and concepts formalized assembly mechanisms like neutral processes or metacommunity dynamics where uneven fitnesses in a heterogeneous landscape can affect local coexistence. More than these as solely alternative mechanisms, these allowed for a synthesis where multiple mechanisms operate simultaneously to affect coexistence. Probably the most emblematic paper of this renewed excitement is Peter Chesson’s 2000 ‘Mechanisms of maintenance of species diversity’ published in Annual Reviews of Ecology and Systematics. This paper, cited over a thousand times, offers a way forward with a framework that includes competitive and niche differences but can also account for neutral dynamics.

A second major development that rejuvenated ecology was the formation of technological and statistical tools engendering broad-scale synthetic research. Suddenly the search for general explanations – Lawton’s most piercing criticism - became more common and more successful. With the advent of on-line databases, meta-analytic procedures and centers (e.g., the National Center for Ecological Analysis and Synthesis) that foster synthetic research, ecologists routinely test hypotheses that transcend local idiosyncrasies. Often, the capstone publication on a particular hypothesis is no longer a seminal experiment, but rather a meta-analysis that is combines all the available information to assess how strongly and how often a particular mechanism affects patterns.

While these theoretical and technological developments have been essential ingredients in this ecological rejuvenation, there has also been a subtle shift the philosophical approach to what it is ecological theory can and should do. Criticism in the 1990’s (e.g., Peters 1991 ‘A Critique for Ecology’) centered on the inability of ecological theory to make accurate predictions. The concept of science common in ecology in the 1990’s was that a rigorous, precise science (i.e., with laws) results in the ability to accurately predict species composition and species abundances given a set of mechanisms. This view of ecological science has been criticized as simplistic ‘physics-envy’ (e.g., see Massimo Pigliucci’s PhD dissertation ‘Dangerous habits: examining the philosophical baggage of biological research’published by the University of Tennessee in 2003). The subtle philosophical change has been a move from law=prediction to law=understanding. This is as true for physics as it is for ecology. We don’t expect a physicist to predict precisely where a falling feather will land, but we do expect to totally understand why it landed where it did based on fundamental processes. (for more on the contrast of prediction and understanding, see Wilhelm Windelband’s nomothetic and idiographic knowledge)

While the feather example above is simplistic, it is telling. In reality a physicist can produce probability contours of where the feather is likely to land, which could be very focused on a calm day or broad on a windy one. This is exactly what ecologists do. Once they understand how differing mechanisms come together to shape diversity, they make probabilistic predictions about the outcome of a set of known mechanisms.

Ecology today is as vibrant as ever. This is not a result of finding new laws that proved Lawton incorrect. Rather, ecologists now have a more sophisticated understanding of how various mechanisms operate in concert to shape diversity. Moreover, conceptual, technological and philosophical revolutions have fundamentally changed what ecologists do and what they are trying to explain. It is a great time to be an ecologist.

Lawton, J. H. (1999). Are there general laws in ecology? Oikos, 84(2), 177-192.

By Marc Cadotte and Caroline Tucker

holiday caRd

2011-12-18T15:44:00.000-08:00

I wasn't planning on making any more "caRds", but since I received a number of requests, find below a holiday card from the EEB & flow. For optimal viewing, perhaps throw some Jingle Bells on in the background.

This code relies heavily on the rgl package in R. Be sure to run the first line of code to install this package if you don't already have it.

###holiday caRd

#install "rgl" package if necessary:
install.packages("rgl")

#load library
library(rgl)

##Data
r=1.0
h=5.0
m=h/r
x=seq(0:99)
y=x
X=x*cos(y)
Y=x*sin(y)
Z=-1*(m*x)
Z1=Z+rnorm(length(Z),Z,5)
sub<-matrix(subset(c(X,Y,Z),Z==max(Z)),ncol=3)
trunk<-matrix(c(0.54,0.84,-200,0.54,0.84,-650),byrow=TRUE,ncol=3)
sno<-c(0,rep(15,9))

##Plot card
open3d(FOV=1)
par3d(windowRect=c(100,100,600,600))
bg3d("aquamarine2")
plot3d(X,Y,Z,col="darkgreen",type="l",box=FALSE,axes=FALSE,lwd=10,zlim=c(min(Z),0),xlab="",ylab="",zlab="",top=TRUE,border=TRUE)
plot3d(X,Y,Z,col=c("yellow","red"),add=TRUE,type="s",radius=3)
points3d(sub,col="orange",size=10)
lines3d(trunk,col="brown",lwd=20)
text3d(matrix(c(0.54,-100,300),ncol=3),text="Happy Holidays",font=5,cex=1.9,color="darkred",adj=c(0.5,1))
text3d(matrix(c(0.54,0.84,-650),ncol=3),text="and",font=5,cex=1.5,color="darkred",adj=c(0.5,1))
text3d(matrix(c(0.54,75,-780),ncol=3),text="a Happy New Year",font=5,cex=1.9,color="darkred",adj=c(0.5,1))
mat<-par3d("scale")

##Play card
for(i in 1:10){
par3d(scale=mat,ignoreExtent=TRUE)
points3d(matrix(c(sample(-160:160,sno[i]),sample(-160:160,sno[i]),sample(-500:150,sno[i])),ncol=3),col="white",size=3)
play3d(spin3d(axis=c(0,0,1)),duration=1)
if(i==10){points3d(sub,col="darkorange",size=17)
plot3d(X,Y,Z,col=c("yellow","red"),type="s",radius=4,add=TRUE) } }
text3d(matrix(c(0,140,-900),ncol=3),text="From the EEB & Flow",font=1,cex=1.0,color="darkgreen",adj=c(0.5,1))

##The rgl device window allows you to use your mouse to manipulate the plot in 3-dimensions, so feel free to do so before closing it.

You can also access the script here.

What should be the basic unit of community ecology, 2011.

2011-12-01T04:10:00.001-08:00

“Why intraspecific variation matters in community ecology”Bolnick et al. 2011, Trends in Ecology and Evolution.

Intraspecific variation in gastropod
shell morphology (Goodrich 1934).

There has been a long debate in community ecology on the fundamental unit, a debate on what Tansley described as the “necessity of first determining empirically our natural units”. In early years, it involved tension between Clements' and Gleason’s view of the plant community, either as a “superorganism” or simply as a conglomeration of co-occurring species. This latter, Gleasonian view won out, signaling a move towards the species-oriented approach that dominates community ecology today. In later years, there was a push to view the individual—not the species—as the fundamental unit, championed by people like Dan Simberloff. However, though this view has had some influence, it has never been mainstream.

The basis of these debates about the basic unit is simple: do similarities matter more than differences? Recently, the argument that intraspecific differences are important and that community ecology should consider individuals has become much stronger. In “Why intraspecific variation matters in community ecology”, Bolnick et al. suggest that a species-level view of community ecology is an incomplete one, and that we should be aware of making simplifying assumptions about intraspecific variation (e.g. that it is minimal and species-level means are appropriate). Bolnick et al. state their hypothesis clearly:

“… many models of species’ interactions implicitly assume that all conspecific individuals are effectively interchangeable. In this paper we argue that this assumption is misleading and that intraspecific trait variation can substantially alter ecological dynamics.”

To that end, the paper does an excellent job of identifying the key mechanisms by which intraspecific variation might be expected to alter ecological dynamics (especially as summarized in the paper's Table 1). Some of these mechanisms might be fairly ubiquitous. For example, when there are nonlinear relationships between trait values and interaction strengths, Jansen’s Inequality means that the amount of intraspecific variation around the species mean will alter the strength of that interaction. The mechanisms discussed make a convincing argument that intraspecific variation can alter ecological interactions and evolutionary dynamics.

However, a move to individual level ecology has many practical implications*: for example, it would require that we move beyond using average species-level demographic rates, dispersal abilities, and interaction strengths, since these miss important intraspecific variation; that phylogenetic trees be built to the level of the individual, requiring additional genetic information; and that perhaps fundamental changes be made to current coexistence theory. Possibly this would mean many more hours of fieldwork, more complex theory, and much more explanatory power is required. On the other hand, it could mean breakthroughs in how we understand longstanding ecological problems like ecosystem functioning, species diversity and coexistence, or trophic web structure.

For that reason, the fact that Bolnick et al. doesn’t demonstrate very clearly the gains or breakthroughs that could result from including intraspecific differences is a bit of a disappointment. Will we find that increasingly smaller amounts of variation are explained as we divide our units increasingly smaller? Or is the key to explaining community-level interactions found at the individual scale? Most of the examples in this paper are too simplistic to be useful, and for understandable reasons of space, there is little review of the literature (though they cite a number of important papers). That’s really too bad, since there are some subfields that have focused on intraspecific differences (for example, the ecosystem functioning literature), and their findings would contribute to the question of what makes intraspecific differences so promising for community ecologists. Despite that, when the mechanisms presented in Bolnick et al. are considered in combination with papers such as Crutsinger et al. 2006, Clark et al 2010, Albert et al. 2011, and Schindler et al. 2010 (just as a few examples), there is some tantalizing evidence suggesting that intraspecific variation can and does matter.

*Although no doubt similar concerns about workload have accompanied any shift in approach throughout ecology's history. And certainly most shifts in ecological approach (spatial, phylogenetic, etc) only occur once the necessary methodological infrastructure was in place.

Google Scholar will track your citations

2011-11-17T18:36:00.001-08:00

In case you haven't noticed, Google Scholar is now offering "My citations", which tracks citations and calculates indices for your papers. Setting it up looks straightforward and fast, making it another alternative to ISI Web of Science and other services.Let the h-index one-upmanship begin...

halloween caRd

2011-10-27T09:36:00.000-07:00

To view the card, run the code below using R

###########################################################################

###required libraries
##if you do not have the "fields" and "MBA" libraries, you need this step to install them
install.packages("fields")
install.packages("MBA")
## load libraries
library(fields)
library(MBA)

###data
data<-matrix(c(44.9186,81.0824,46.9512,80.1061,49.187,80.7055,57.9268,80.1493,60.3659,81.34,69.3089,80.1941,78.4553,79.2459,91.8699,70.834,99.7967,55.9047,101.2195,30.9102,96.1381,17.3075,85.5691,7.4235,74.3902,4.2299,61.7886,1.8181,55.4878,1.0059,49.187,0.9811,45.3252,1.7532,37.1951,1.9181,20.3252,7.1666,8.9431,17.1612,4.2683,29.1507,4.2683,50.8042,9.5528,66.3761,18.4959,76.2539,29.065,80.6261,38.6179,80.8607,44.9186,81.0824),ncol=2,byrow=TRUE)
stem<-matrix(c(60.3659,81.34,57.9268,80.1493,49.187,80.7055,46.9512,80.1061,44.9186,81.0824,45.5285,83.6438,47.561,88.1794,44.1057,97.2209,48.7805,99.9952,54.878,87.6176,60.3659,81.34),ncol=2,byrow=TRUE)
eye<-matrix(c(45.935,46.4407,34.1463,44.8195,36.7886,46.9952,36.382,50.9306,32.3171,53.08,29.065,53.0672,26.0163,49.7086,28.0488,45.1892,20.3252,48.7021,20.1219,55.3942,23.5772,61.1164,30.8942,65.476,32.3171,59.7729,45.935,46.4407),ncol=2,byrow=TRUE)
eye2<-matrix(c(58.9431,46.6888,65.4472,51.2419,71.748,56.9754,74.3902,65.4504,82.7236,59.7745,84.9593,56.4368,84.7561,48.562,77.439,45.7772,78.8618,50.5072,74.3902,53.6393,69.9186,51.6532,68.6992,47.9083,70.5285,44.9627,58.9431,46.6888),ncol=2,byrow=TRUE)
mouth<-matrix(c(8.9431,44.7202,23.374,41.2338,30.0813,22.7562,38.2114,38.5363,65.0407,38.445,73.374,22.533,80.2846,40.8673,96.9512,44.673,93.4959,31.8641,84.1463,18.8352,66.2602,9.7097,61.9919,16.1889,59.3496,8.3045,45.5285,8.447,42.8862,16.1137,38.2114,9.4024,22.7642,17.4125,11.9919,31.7401,8.9431,44.7202),ncol=2,byrow=TRUE)
stars<-matrix(c(sample(seq(0,100),80),sample(seq(20,100),80,replace=TRUE)),ncol=2)

###plot
par(bg = "black")
par(mar=c(5,2,4,2))+0.1
plot(data,type="n",xlim=c(0,100),ylim=c(0,100),xaxt="n",yaxt="n",xlab="",ylab="",axes=FALSE)
symbols(x=stars[,1],y=stars[,2],circles=rep(0.001,nrow(stars)),inches=0.015,bg="white",xlim=c(0,100),ylim=c(0,100))
col<-runif(nrow(data),min=4,max=7)
datmb<-mba.surf(matrix(cbind(data,col),ncol=3),no.X=200,no.Y=200,extend=FALSE)
image(datmb$xyz,zlim=c(0,10),col=heat.colors(40),add=TRUE)
polygon(stem,col="darkgreen")
polygon(eye,col="yellow")
polygon(eye2,col="yellow")
polygon(mouth,col="yellow")
points(data,type="l")
mtext("Happy Halloween",side=3,col="green3",cex=3.5,font=4)
mtext("from the EEB and Flow",side=1,col="green3",cex=2.5,font=4)

Seed dispersal: plant height seems to be more important than seed size!

2011-10-12T07:48:00.000-07:00

I really like papers that teach me something that I didn’t know. But, I love papers that show me that what I learned is wrong. This is the case of a new paper by Fiona Thomson, Angela Moles, Tony Auld, and Richard Kingsford on seed dispersal that appears in the last issue of the Journal of Ecology. This group from Australia analyzed the effects of seed size and plant height on their dispersal abilities. They reviewed intensively the literature gathering data on 200 species from 148 studies around the world. Surprisingly to me, they found plant height was much better at predicting seed dispersal than seed size. This might not sound so surprising for many people (and after seeing the paper, kind of intuitive), but there was a lot of evidence that seed size was the best predictor of dispersal, with species with smaller seeds dispersing further than species with bigger seeds. For wind dispersed species, their results are more intuitive, but they found this pattern in a number dispersal syndromes analyzed (i.e. unassisted, wind, ballistic, ingestion, and ant dispersal). So, in your next study on seed dispersal consider adding plant height as an explanatory variable.

Thomson, F. J., A. T. Moles, T. D. Auld, and R. T. Kingsford. 2011. Seed dispersal distance is more strongly correlated with plant height than with seed mass. Journal of Ecology 99:1299-1307. DOI 10.1111/j.1365-2745.2011.01867.x

The four types of failure, or how to fail in science

2011-10-04T12:25:00.000-07:00

As scientists, we’re all wrong, at least sometimes. The question is, how are we wrong?

The arsenic bacteria saga, which we’ve discussed on this blog before, is turning out to be a very public example of failure in science. First announced by NASA press conference in December 2010, authors lead by Felisa Wolfe-Simon shared their discovery of a bacterium capable of replacing phosphorus in its DNA with arsenic, suggesting the possibility of life in phosphorus-limited conditions. This apparently momentous discovery was published in Science, and met with disbelief and severe criticism. Critics throughout the blogosphere and academic departments began to compile a comprehensive list of failings on the part of the paper—8 technical criticisms were published in Science—and as the result of the intense focus on the paper’s lead author is no longer associated with the lab group where this research was carried out. This is failure at its worst—the science was flawed and it drew immediate and intense censure. This is the kind of failure that most young scientists fear: judgment, intense criticism, career-long repercussions. But it’s also probably the least common type of failure in science.

However, it’s arguable that the saddest form of failure is the opposite of this: when a paper is right—innovative, ahead of its time—but somehow never receives the attention it deserves. There are lots of famous examples of scientific obscurity, with Gregor Mendel being the poster child for scientists who toil for years in anonymity. In ecology, for example, papers that considered species as equivalent (a la neutral theory) to explain coexistence were around in the 1950’s-1960s, but received little attention. Other papers suggesting variation in environment as a possible mechanism for plant coexistence were published prior to Chesson and Huntly's influential paper, yet essentially uncited. Most researchers can name at least one paper that foreshadows the direction the field will take many years later, yet is unacknowledged and poorly cited. There are many reasons that papers could be under recognized—they are written by scientists outside of the dominant geographical areas or social networks, or who lack the ability to champion their ideas, either in writing or in person. In some instances the intellectual climate may not be conducive to an idea that, at a later time, will take off.

If that is the saddest type of failure, then the best type of failure is when being wrong inspires an explosion of new research and new ideas. Rather than causing an implosion, as the arsenic-bacteria paper did, these wrong ideas reinvigorate their field. Great examples in ecology include Steve Hubbell’s Unified Neutral Theory of Biodiversity, which although criticized rightly for its flaws, produced a high-quality body of literature debating its merits and flaws. When Jared Diamond (1975) proposed drawing conclusions about community assembly processes based on patterns of species co-occurrence, the disagreement, led by Dan Simberloff ultimately led to the current focus on null models. Cam Webb’s hypothesis that there should be a relationship between phylogenetic patterns in communities and the importance of different processes in structuring those communities sparked a decade-long investigation into the link between phylogenetic information and community assembly. Although Webb’s hypothesis proved too simplistic, it still informs current research. This is the kind of failure on which you can build a career, particularly if you are willing to continually revisit and develop your theory as the body of evidence against it grows.

However, the most common form of failure occurs when a paper is published that is wrong, yet no one notices or worse, cares. For every paper that blows up to the proportion of the arsenic bacteria paper, or inspires years of new research, there are hundreds of papers that just fade away, poorly cited and poorly read. Is it better to fail quietly, or to take the chance at public failure, with all its risks and rewards?

The European Ecology Federation Congress, day 3

2011-10-02T19:02:00.000-07:00

*sorry for the delay in getting the last day up, I've been catching up. The first talk of the morning was by Georgina Mace -great talk, and I will have an extended post on it later. Here are the other talks. This meeting was great!

Elisa Thebault. This was a great talk. She talked about the structure and stability of mutualistc and antagonistic networks. Nested interactions means that several generalists and specialists, but specialists use the same resource as generalists and do list overlap with other specialists. She addressed two main questions. First are there differences between mutualistic and antagonistic networks? Second, do these differences have consequences for coexistence and stability? First question, herbivores seem to have less nestedness and interact with closely related plants, while pollinators are more nested but less phylogenetically structured. For the second question, with is examined using modeling, using coupled predator prey equations (with a positive effect in the mutualism model) and simulated communities. She looked at two types of stability, persistence of species and resilience. She showed some very interesting results, for mutualistic networks, connectence and diversity increase stability, while for antagonistic, the opposite. Because of diversity change in the simulation, the mutualistic networks become more nested and more connected, again the opposite for antagonistic network, which becaome less connected and nested. What happens when you put these interactions together with both mutualistc and antagonist models? The same patterns emerge with muralists being more nested and connected.

Pedro Jordano. He talked about the functional role of complex networks including different types of seed dispersers and pollinators. Can phylogenetic relationships explain patterns of interactions between the seed dispersers and plants. In degraded habitats, through hunting, only a restribected subset of species are interacting with plants. What is the minimum complexity required to maintain ecosystem function.

Jason Tylianakis. He talked about global change and ecosystem function. In an example dataset, soil resource availability and grazing intensity affected trait compositiona dn diverisyt and changed plant productivity. When resources are heterogeneous then diversity affects function, but not when resources were homogeneous. Across a gradient of land use intensitfication, networks become simpler with functional links being dominated by few species. He looked at 133 host-parasitoid interaction webs. These webs deviate from null expectation and some habitats were significantly less complex than predicted.

Daniel Stouffer. He talk about understanding species roles and importance in food webs. Different types of interactions (sub webs) have differential probabilities of being present. Certain motifs appear to differentially contribute to stability. This approach can inform species conservation if a particular species appears in different motifs that contribute to network function or stability. Certain species may be common in motifs that reduce stability. Using New Zealand river food webs, he asked three questions: is the benefit of species phylogenetically conserved -yes, certain clades add benefit. Are these benefits community specific? No, beneficial species are so in all communities (bit similar communities). How general are these results? He compared the results to webs elsewhere in the world. Similar species are similarly beneficial elsewhere.

-Here I lost my notes from Jane Memmott’s plenary talk (sorry Jane!). It was a great overview of her research in restoration. At the heart of her talk was about making restoration scientifically rigorous.

Henrique Pereira. His talk was on modeling the response of biodiversity to global change. Biodiversity indicators for global change are biassed towards North America and Europe and certain taxa. Major uncertainty in extinction rates and what are the sources of uncertainty? A big source is the differences in scenarios for land change and human population growth. Also lack of ecological knowledge. Finally there are differences between models. He proposes a countryside species area relationship (cSAR) instead of regular SAR, which assumes an uninhabited matrix. Multiplies area by the affinity of species to live in that area, and so as long as a species has an affinity greater than zero for marginal habitat, it can persist in those areas –changing our predictions about habitat loss on species persistence. The cSAR predicts much lower extinction rates compared to classical SARs. Need data to classify affinities, such as uses surveys to cluster species by where they are found. The cSAR fits real data better than SAR.

Christophe Randin. His presentation was on whether elevational limits of deciduous trees match their thermal latitudinal limits. Species often not at equilibrium with their predicted fundamental niche, may reflect dispersal limitation. Species should reach their equilibrium since climate change so much quicker. Based olots of data, he presented where the distributional limitation should be and examined the distance from that edge. Surprisingly, the latitudinal limit was less likely to be reached by. Species, thus they are lagging on mountains.

Rita Bastos. She used a Dynamic model for understanding the recovery of the Azorean bullfinch in a changing environment, a lot a land use change and invasive species. Specifically, the model is a stochastic, spatially explicit model that incorates environmental variables and projected habitat change. She was able to test different management scenarios. Certain management actions on habitats can significantly increase population sizes but not spread.

Diogo Alagador. He spoke on adjusting protected areas to account for climate range adjustments. Species will move with climate change, but reserves do not move. Planning must involve multiple potential reserves and likely assisted migration. It is difficult to extrapolate for multiple species. Persistence then is the product of suitablility and dispersal ability for a species for each time period in future projections. This can be summed across species. This was tested for seveal species across all major taxa. There is variability in persistence across species and are very sensitive to disperal pathways.

The European Ecology Federation Congress, day 2

2011-09-27T21:51:00.000-07:00

Day two of the conference, and still many great talks. I mainly stayed in the session on synthesizing community ecology, phylogenetics and macroecology. This has turned out to be a great conference and Avila is a great venue.

Carsten Rahbek. His talk is on merging the fields of macroecology to better understand patterns of diversity. Different models explain variation differentially at different scales. For example, climate models do well for wide-ranging species but not for scarce species. A model of evolution may do much better for scarce species, but not for wide-ranged species. Statistical tests confirm a correlation, but not necessarily a mechanism. One could get different conclusions if one were to compare to a null model. He advocates a spatially explicit species assembly model that integrates macroecological models with community assembly. It is scale invariant and can explain spatial and temporal variation in assemblages. In an example, he shows that, based on small scale sampling, species distribution models will over-predict richness. Need to combine macroecological models with distribution models, because acroecological models do well to predict richness but not composition while distribution models predict composition but not diversity.

Jens-Christian Svenning. He talked about paleoclimatic influences on ecological patterns and function across scales. Past climates have shown masive changes and different groups of species have evolved during these events, while other species have gone extinct. The velocity of climate change was highest in northern Europe and North temperate North America, and higher velocity results in lower endemism since it is quicker for species to migrate than diversify. Higher velocity results in lower specialization in hummingbirds. He finishes with a note about current regions undergoing fast climate change; these are not necessarily those same regions that had the most change in the past.

Adreas Prinzing. His talk was about how niche conservatism can inform our potential solutions for changing environments. Specialists are declining in changing environments and how does this apply to specialist clades of closely related species? Specialist species tend to occur in specialist genera. However, niche conservatism does not tell us everytng about species differences/similarities because closely related species clearly coexist and exhibit substantial trait differences. Species coexist within niches by key divergences.

Kenneth Kozak. He presented a way that phylogeny illuminates the origin of climate-richness relationships. Only speciation, extinction or dispersal can change richness, and many models do not ask how these processes change. He examined salamader diversity and evolutionary history using 16000 occurrence records in North America, and examined climate variables for occurrences. Diversity was highly associated with cool, moist places. Richness is strongly correlated with evolutionary time of colonization of climatic conditions. For example, evolution of warm species is recent, hence fewer species. Diversity does seem to be saturating, and so time is limiting factor, and more species can probably still emerge.

David Vietes. He gave an interesting talk on the amphibians of Madagascar, which is a diversity hotspot for amphibians. There were 132 described species n 1999 and now 263 with about 200 still needing to be described. Many are endemic to small regions of Madagascar (the whole family is endemic to Madagascar). He discussed many aspects of the distribution of these species, and looked at phylogenetic patterns. Some interesting observations include: older species pairs are further separated in space and smaller species have smaller ranges. Also, there appears to be a predictable pattern of richness hotspots, but endemism hotspots are more idiosyncratic.

Joaquin Hortal. He discussed the effect of glaciation on richness, functional diversity and phylogenetic diversity for European mammals. The hypothesis he explored was that current distributional patterns driven more by past changes since glaciation than current climate. He compare several different types of measures and it turns out that current climate is more important for explaining patterns of co-occurrence and relatedness, with more closely related species occurring together at northern locales.

Catherine Graham. She also explored patterns of richness, functional and phylogenetic diversity, but was looking at hummingbirds diversity patterns across elevation gradients in South America. She compiled an impressive dataset with several morphological traits and co-occurrence patterns. Broadly, close relatives co-occur at high elevations and more distantly at lower where competition is stronger. In local communities a mix of environmental filtering and competitive dispersion seem to be operating. At high elevations, both functional and phylogenetic diversity are high.

Rob Dunn. He gave a fantastic talk on the species on the human body and in our lives and homes. He told us about projects that involve citizen scientists from across the USA and had them sample their homes and bellybuttons. Amazingly, Dunn’s group has so far identified 1400 species in belly buttons, and many of them are unknown species –which could not be classified into known species groups. He looked at many factors like ethnicity, geography, cleanliness, but none of these explained this diversity well. A subset of these species are bellybutton specialists and dominate bellybutton floras within and among people, and are phylogenetically clustered, evidence that the bellybutton habitat is a conserved trait.

Cecil Albert. I ran to another session to see this talk on intraspecific variation in species traits. She eloquently showed that for plant assemblages, there was substantial intraspecific variation in traits. Some species showed high variation and some showed almost no variation. Importantly, she showed that this variation could substantially change our ability to explain how functional traits link to abundance and coexistence. She simulated different levels of variation and looked at the strength of the correlation between expectations from mean trait versus the actual trait that varies. The strength quickly declines for some traits as variation increases, meaning that with variable traits, the explanatory ability of using a mean trait is weak.

Sally Keith (Flash talk*). She examined the Mid-domain effect (where, because of range sizes, maximal diversity is found in the centre of a geographic landmass by random chance), and process based models to test mechanism for middomain prediction. She showed that these models seem to have limited success. Perhaps environmental gradients and species interactions could be important. But when she added interactions to the model, it then predicts humped shaped pattern predicted by the mid-domain effect.

Tamara Munkemuller (Flash talk). She examined phylogenetic relationships as a way to examine niche patterns and coexistence. She hypothesized that there should be strong filtering under stressful conditions. She examined thousands of plots across elevational gradients, and plots that were in stressful locations tended to be phylogenetically clustered, meaning certain groups of species exist there.

Susanne Fritz (Flash talk). She was looking at diversification patterns in birds. Using lineage through time plots one should expect that the rate of diversification should decline trough time, which perhaps equates to niche filling. For species in tropical Asia, she found that there is not much leveling off of diversification rate. Though interestingly, groups that have not dispersed (for example, birds of paradise) do show a plateau in diversification. Globally, diversification slows down more in more speciose regions.

Jake Alexander (Flash talk). He had a very interesting talk on elevation gradients in richness in non-native plants invading mountain habitats. Most species have narrow elevational ranges in lower and mid elevations and high ranges for those species found at high elevations. The explanation is that these non-natives are generalist and that they originate form lower elevations –where human activity dominates, and must spread up the mountain to get to the high elevations.

*Flash talks are 3 minutes long, and a great way for people to communicate new and exciting results.

The European Ecology Federation Congress, day 1

2011-09-26T22:34:00.000-07:00

I’m in the beautiful walled city of Avila, Spain for the European Ecology Congress. It is at a lovely venue and with about 800-1000 attendees, seems like just the right size. It is a young meeting, with relatively few old-timers like me, but there is an excitement, and the talks have been excellent. Each session starts with a keynote, where the person gets 25 minutes, followed by a bunch of 15 minute talks. The most interesting aspects of the sessions I went to was that they usually include several 3-minute ‘flash’ talks, which surprisingly works. I spent the day going to talks in two sessions, plus a plenary talk by Jordi Bascompte, and here are the talks I saw*:

*sorry about the abrupt, choppy nature of some of the entries, there were a lot of talks, and they go until after 7pm.

I spent the morning in a session on biodiversity and ecosystem function under environmental change. Most of these talks we by people associated with the BACCARA project on forest biodiversity.

1) Xavier Morin, Montpellier, talked about climate change and tree diversity and productivity. Looked at SR (see glossary at end for acronyms) and FD on biomass produced. Do grassland BEF studies predict frost ones, with no opportunity for random assembly? Use forest dynamics model where species are defined by rigorous parameterization –one can examine long-term dynamics and many species combinations. Simulated 30 species monocultures and many combinations from 2-30. Strong relationship between realized richness and productivity, but a lot of variation. 93% of 30 species plots show transgressive over-yielding after 2000 years. FD predicts increase in productivity. Assess future climates from three climate change scenarios, always steeper slope with future climate, meaning diversity is more important in the future. This was a great talk.

Sibylle Stoeckli. Affects of diversity on individual tree performance. She wanted to assess the influences of tree traits (e.g., size) on the performance of neighbours. Plots planted with four species combinations, with a pool of 16 species, and treatment is different FD levels (based on 9 traits). No effect of plot diversity on tree performance. Tree height has effects on neighbours depending on whether focal tree is shade tolerant or intolerant. Diversity not important as traits of species such as growth rate. Interspecific competition lower than intraspecific.

Aitor Ameztegui. Montane-apine ecotone is diverse and are traits important for coexistence. Are interspecific differences key for coexistence, and can these tell us about biome changes. One species has advantage at low light but quickly saturated with increasing light. Silver fir had constant survivorship, while other species increased survivorship with increasing light. Fir has low plasticity, whereas Scots pine is more plasticity and should adapt to climate change.

Alfredo saldena. FD on decomposition in South American rainforest (Chile). He looked at two forest types within the Andes. In both forests strong positive relationship between FD and litter decomposition. FD is based on leaf traits.

Julia Koricheva. Forest diversity and insect communities. Boreal species (5) in southern Finland in monocultures and 2, 3 and 5 species mixtures. Looked at different types of leaf damage. For birch, increasing skelontonizing damage with diversity. During aphid outbreak, decline in density with increasing SR. They prefer birch. Leaf miner richness on birch increased with SR. In another, german, experiment (the experiment in Stoeckli's talk) where FD was manipulated. Again several types of herbivores had positive relationships with FD, again counter to expectations of more specialized herbivores declining with tree diversity.

Laura Concostrina-Zubiri. Biological soil crusts (BSC) are important for soil fertility and stability in dry ecosystems. creates soil heterogeneity. Examined the role of BSC across a grazing gradient. Measured 17 soil variables for a bunch of species. Species differ in their different soil fertilities. Less heterogeneity with higher grazing. Grazing also reduces individual species contributions to soil fertility.

Plenary talk: Jordi Bascompte. Plant-animal mutualistic networks. He talked about Global datasets to answer three questions: 1) are there regularities in network architecture; 2) Do these provide robustness to extinction; and 3) what are the contribution of species to network architecture and robustness? Networks seem to be nested such that specialist animals use most utilized plants. This means there should be a link between structure and robustness (losing an aminal should not result in plant extinction). Half of communities have interactions dictated by evolutionary history. Thus when there is extinction, it tends to be related species, nonrandom. Therefore clades are more likely to be lost. How much of the interactions that are shared can be used as a competition term in coexistence model. The higher the nestedness the lower the competition and the higher maximum diversity. Some species contribute to nestedness much more than others and therefore are much more responsible for stability and have greater probability to go extinct.

I spoke in a session on evolutionary history, ecosystem function and conservation, and (probably ignoring my talk) these were excellent.

Marten Winter. He asked whether phylogenetic studies purporting to do conservation actually did conservation and whether using PD was feasible. Assumptions, some not proven. Unsderstandability of terms like evolutionary potential, what that means for species and communities can cause confusion. Different measures can produce different patterns and he asked Don't we already conserve what we want? Or is there an added advantage to accounting for PD. There were a surprising number of papers that do make conservation recommendations.

Nicolas Mouquet. Phylogenetic constraints on BEF. Biodiversity crisis is a change for synthesis for diffect fields to come together. Positive relationships went from how much to what kind diversity. Evolution is necessary for understanding how biodiversity shapes ecosystem function. He tested these relationships with bacteria from Mediterranean and evolved in lab. There are ancestral and derived groups. Strong positive BEF relationships for both ancestral and derived taxa. For ancestors, a strong PD influence was observed. But not for derived taxa, a reshuffling of traits in the lab. Need to understand the history.

Ana Rodrigues. Species are not all the same such as mouse versus echidna. Need to be cognizant of tree structure and species distributions. Does it matter if we use PD for complimentary reserve design and compare maximizing PD vs SR vs random. SR conserved then look at PD. Little difference for mammals at global level, meaning that current reserve selection routines seem sufficient. Important for species level, but perhaps species level activities may have done a good job at conserving PD.

Sandrine Pavoine. Rockfish declines and phylogeny. Phylogenetic diversity based on period between speciation events. Sum abundances for lineages for each period. Sum period lengths times relative abundance. Calculate lineage contribution to total diversity. Which period is reponsible for abundance change. One period explained declines in rockfish and is actually quite an old period (6 million years).

Wilfried Thuiller. Preserving the tree of life and climate change. Are there winners and losers? Estimate phylogenetic consequences of climate change, if there are sensitive clusters, would one expect more loss than expected by chance. There is a phylogenetic signal in climate, kind of weak, but extremely close relatives respond similarly. Loss of PD is not much different than random. Sensitive species tend to be young. But there is a predicted loss of phylobetadiversity for all birds, mammals and plants with climate change.

Vincent Devictor. Comparing several components of biodiversity. Can SR, FD, PD serve as surrogates. Compared metrics using birds surveys in France with 22 traits. Abundance weighted measures. FD declining while SR increasing. Differential responses important for making conservation decisions.

Laure Turcoti (Flash talk 1). Comparison of SR FD PD on plant communities. SR increases with urbanization and FD and PD decrease with urbanization.

Laure Zupan (Flash talk 2). Current distribution of phylogenetic diversity. Covariation across different clades. Birds, amphibians and mammals. Mismatch between tax am amphibians high PD relative to SR, while mammals low.

Jonathan Davies. Plant extinction risk in the Cape using IUCN rankings. Genera level phylogeny for the over 700 genera in the Cape. Clustering of extinction risk on phylogeny, but plant extinction is correlated with clade size, meaning that large clades have more risk –opposite of what has been observed for mammals. The reason is that many small peripheral species with small range.

Sebastian lavergne. Dechronization of niches, i.e., travel back in time. Is there signal of niche conservatism, and for different niches for the birds for Europe. Climatic, habitat and trophic niches. Trophic niche evolves at slower rate but niches evolve in punctual way, not gradual. High clade disparity in niches since niches evolving faster.

Glossary

BEF: Biodiversity and ecosystem function

FD: Functional diversity

PD: Phylogenetic diversity

SR: Species richness

NSF funds Project Baseline

2011-09-23T10:23:00.000-07:00

NSF approved 1.2 million dollars for a unique and visionary idea: collect 12 million seeds and store them in seed banks for years to come. And while storing seeds for the future doesn't sounds so different from what other groups have already done, where Project Baseline differs is that this seed bank is not only a conservation measure--preserving natural genetic variation from plant populations for the future--but also an opportunity to track the effects of changing climate on the direction and rate of evolution in these species.

This idea was first explored in "The Resurrection Initiative: Storing Ancestral Genotypes to Capture Evolution in Action" (Franks et al 2007). By collecting and storing seeds from both within and across populations throughout the range of a species, ancestral and descendent populations can be compared in the not too distant future. The role of adaptive evolution and range shifts can be explored through this lens. Project Baseline is a great example of how much we can learn from long-term, collaborative experiments and projects (other examples include NutNet , NCEAS), and how valuable funding such projects should be considered.

Ecology needs more evolution (and vice versa)

2011-09-16T12:47:00.000-07:00

One historical weakness in community ecology is its singular focus on ecology in the absence of any consideration of the role of evolution. Ecological theory may attempt to explain and expand on mechanisms of coexistence, but this is done in ignorance of whether such a mechanism could have reasonably evolved in the first place. Evolutionary biology has equally ignored the role of ecology (for example, just-so stories invented in the absence of ecological support). Fortunately, it is becoming more common to see papers that incorporate, empirically or theoretically, evolution and community ecology.

A recent paper by Robin Snyder and Peter Adler attempts to incorporate both ecology and evolution in reference to the storage effect, a mechanism in which species coexist as a result of environmental variability and corresponding differential variation in species fecundity in response to the environment. As a simplistic example, consider a system of two annual plants for which each species has their highest recruitment at different temperatures, and temperature varies randomly between years. Each species is expected to have high recruitment in different years/environmental conditions, and this high recruitment in good years can then buffer that species’ fitnesses in years of poor conditions, provided the species have some way of “storing” fitness (such as long-lived seedbanks). The storage effect therefore predicts that environmental variability can mediate the coexistence of otherwise unequal competitors. Because the requirements of the storage effect appear so ubiquitous (environmental variation, differential species responses to the environment, some sort of buffer), it seems that the storage effect could be very common. However, there is also theory suggesting that variation in demographic rates should come at a fitness cost, since the long term mean growth rate will be lower if demographic rates vary than if they are fixed (as the result of geometric averaging). This predicts that there should be selection against flexible—rather than fixed—demographic rates, including rates that vary in response to environmental or other cues. Is it possible then for variable demographic rates, which are necessary for the storage effect, to evolve?

Snyder and Adler discuss this disconnect between community ecology and evolution, questioning whether the storage effect can be supported by both evolutionary and ecological theory. To this end, the authors explore whether, and under what conditions, the storage effect could evolve. Snyder and Adler use a simple model of competition between two annual plants, in which fecundity fluctuates due to environmental variation, and germination rate can be temporally fixed, or variable. Germination rates should be constant, despite environmental variation, due to the cost of variability. Germination rates would be expected to vary year to year only if this conferred a fitness benefit to the species. Hypothesized benefits of variable germination rates include if germination rates are positively correlated with fecundity (that is, in good years germination is higher as well), or if it allows a species to avoid competition (by having high germination when their competitor has low germination). To test this hypothesis, the authors varied the correlation between fecundity and germination, and the correlation between the two species’ germination rates. They then examined the conditions under which variable germination rates were an evolutionarily stable strategy (ESS).

Snyder and Adler’s results suggest that the storage effect is expected to evolve only under anarrow set of conditions. A variable germination rate was most likely to evolve if there was a strong correlation between fecundity and germination rate. They note that such a correlation might occur if seed production and germination depended on similar environmental cues or similar resource requirements. A variable germination rate was also a stable strategy if one species was limited in its ability to evolve, in which case the other species evolved variable germination rates. If these specific conditions didn’t hold, the storage effect was not evolutionarily stable.

These results are meaningful because they highlight how different the conclusions of community ecology, which has proposed that the storage effect could be a widespread contributor to coexistence, and evolutionary theory, which suggests that the storage effect may only occur under particular conditions, can be. This kind of reconciliation of community ecology and evolution tells us more about natural systems than either approach can on its own. It also hints that theory and conclusions we’ve drawn in community ecology in the absence of evolution may be limited and incomplete.

BES day 2: Plants, plants and way more plants

2011-09-14T18:24:00.000-07:00

From Sept 13

I attended the Journal of Ecology Centenary symposium all morning, where the talks were broad overviews of select areas in plant ecology. They were quite good; I really do feel that I was informed about recent research advances.

In the first talk of the morning, Sandra Lavorel gave a tour de force about how plant functional traits scale up to ecosystem services. She recognizes that there are trade offs in services, where one service (say agricultural value) is in direct conflict with a noter service (say species richness). She very cleverly asks whether these services are constrained by ecological trade offs or traits. It is known that functional traits affect ecosystem functions and services, and it is also known that there are strong tradeoffs in plant traits such as explained by the world wide leaf economic spectrum. Where plants have these tradeoffs they affect productivity and litter decomposition. Height for example affects productivity and other trophic levels supported. Abiotic gradients affect traits like height or leaf N, and these traits affect ecosystem function such as biomass or litter. Multiple service such as agronomic value, pollination, cultural value, richness, etc. To understand how traits relate to tradeoffs in services.

Next was Angela Moles who talked about how the study of invasions has progressed and whether there were important future directions. The have been 10,000 studies on invasions over the last 30 years and she recognizes that the fact that species evolve in their new ranges to be a critical future research need. Specifically, she asked: do exotics evolve to be more similar or different tha natives? And, can differences be predicted by environmental differences between home and away range. Most interestingly she brought up the point that if on-going change produce new species, should they then be conserved as natives? She went on to say that broad generalizations about trait differences between natives and exotics have produced largely idiosyncratic results, and so other priorities such take the forefront. She went on to say that impact on natives is actually an understudied problem, which needs to be rectified. Finally, she showed us that there is a generally positive relationship between disturbance and invasion. But invasions are favored when there is a change in disturbance rate, since natives are likely adapted to historical disturbance regime. She showed some relatively weak evidence that change in disturbance better predictor of invader richness and abundance then the amount of disturbance, but more work is needed.

Yadvinder Malhi talked about how productivity and metabolism were related to biomass in tropical forests. He sowed us that a small proportion of primary production is turned into biomass. Thus small changes in various pathways could have large consequences. In exhaustive studies in the tropics, he showed that increases in GPP (gross primary production) occurred with soils nutrient quality, and decreases with elevation, likely because of temperature effects on photosynthesis. He also showed that carbon use efficiency is lower than thought, about 30% of carbon turned to biomass. Further, higher productivity is associated with lower residence times, and he hypothesized that rapid growth leads to earlier senescence or less defences if trade offs exist. Biomass appears to be increasing with climate change, but potentially greater mortality and turnover.

Then James Bullock talked about where we are at with understanding seed dispersal. There has been a long history of not understanding long distance dispersal, LDD. The main empirical approaches have grown rapidly lately: tracking seeds, molecular methods and marking seeds or to track dispersers. But at the same time spread models have appeared and advanced. However, Bullock really supports mechanistic models for wind-dispersed species, and these models seem to really provide insights. He then compared a handful of models for invasive and scarce natives, and did mechanistic modeling with climate change. Changes in future wind speeds may result in even larger changes in spread rates. Only Ailanthus appears to have dispersal rates at or faster than the rate of climate change, most species do not appear to be able to move fast enough (except animal dispersed species). Movement on shoes major dispersal vector.

Hans Jacquemyn was the final speaker, and talked about evolution and habitat fragmentation. He studies calcareous grasslands, forests and heathlands in Belgium that have increasingly become isolated and fragmented. Observed declines in genetic diversity in populations, as they get smaller in size. More recent fragmentations have less loss of genetic diversity compared to older fragments. Reduced seed output in small populations for self-incompatible species, results in reduced population growth rates. To counteract plants can increase floral displays or increase selfing rates, which they observed. Also, he has observed changes in timing of flowering and investing more in nectar.

*Some thoughts about the BES. It is a great Society, and a great meeting. It is relatively small and it is nice to see how many members know each other. For those of us in North America, I think it is a great experience to go to one of these meetings.

**I also participated in the BESdigital workshop on communicating science in a digital era. I will have a post about this tomorrow –I've been without internet connection at both the Sheffield dorm and various airports.

British Ecological Society meeting: day 1, the Tansley Lecture by Diane Wall

2011-09-12T02:47:00.001-07:00

I am at the BES meeting in Sheffield. I will be spending most of the day in journal meetings, but I was able to attend the opening lecture. I will be able to attend more talks tomorrow.

Diana Wall gave the Tansley Lecture to open the meeting. The focus of her talk was about integrating soil biodiversity into ecosystem science. She started with a quote from Arthur Tansley about how all the aspects of an ecosystem can not be ignored, and Professor Wall argues that type soils, specifically the organisms living below ground have been largely ignored, and we do now live in an era where we can study all aspects of ecosystems. Her talk showed the ways in which soil orgasims matter and how global change may have consequences for the link between soil organisms and the ecosystem functions they provide.

This is especially important because soils are deteriorating globally, and while soils are home to an impressive diversity of organisms, so little understood about these organisms. Often we do not even know how many species are found in soils (though in many cases we are talking about hundreds or thousands of species per square meter, which means we can cannot predict how global change could affect these biota and the functions they provide.

She went through three examples to highlight the importance of soil organisms and research needs to predict the impacts of global change (here global change seems broadly defined including: temperature, precipitation, land use, and water flow). In the first example, she reviews the role of soil organisms in extreme ecosystems (Antarctic and hot desert) and how climate change may alter dynamics. Experiments show how important soil organism are for flow of nutrients and energy, and global change experiments show drastic changes in their abundance, thus we should expect large consequences as environments change, especially as moisture regimes change. In different systems, the relative importance of biotic vs abiotic drivers (e.g., the presence of plants versus moisture gradients) differs and differentially important for the ecosystem effects, and so more understanding is required for predictions.

In the second part, soil animals affect soil decomposition in moist places, thus changes in moisture affect ecosystem pathways. In the third part, she outlines the ways in which soil organisms provide ecosystem services, nutrient cycling, diseases, food, food webs, biocontrol, carbon storage, waster breakdown, etc. These services have been understudied and under appreciated.

Overall this talk was a lucid and poignant call for more work to be done on soil biota -not to know what is there necessarily, but rather to be able to link together the effects of changing environments on ecosystem function.

How is a species like a baseball player?

2011-08-25T09:51:00.000-07:00

Biomass is to runs as species is to player, and as ecologist is to Brad Pitt.

Community ecology and major league baseball have a lot to learn from each other.

Let's back up. As a community ecologist, I think about how species assemble into communities, and the consequences for ecosystems when species disappear. I'm especially interested using traits of species to address these issues. For the grassland plants that I often work with, the traits are morphological (for example, plant height and leaf thickness), physiological (leaf nitrogen concentration, photosynthetic rate), and life history (timing and mode of reproduction).

As a baseball fan, I spend a lot of time watching baseball. Actually, I'm watching my Red Sox now (multitasking as usual; I freely admit there's a lot of down time in between pitches). I care about how the team does, mostly in terms of beating the Yankees. I'm especially interested in how individual players are doing at any time; for fielders I care about their batting average and defensive skills, and for the pitchers I care about how few runs they allow and how many strikeouts they get.

So my vocation and avocation have some similarities. Both ecology and baseball have changed in the last decade or so to become more focused on 'granular' data at the individual level. In ecology this has been touted as a revolutionary shift in perspective, but is really a return to the important aspects of what roles organisms play in ecosystems, and how ecosystems are shaped by the organisms in them. This trait-based approach has shifted the collection and sharing of data on organism morphology, physiology, and life history into warp speed, to the great benefit of quantitatively-minded ecologists everywhere.

In baseball, the ability to collate and analyze data on every pitch and every play has lead to an explosion of new metrics to evaluate players. One of the simplest of these new metrics, which even the traditionalists in baseball now value, is "on base plus slugging" (OPS, see all the details here). This data-intensive approach to analyzing player performance was most famously championed by the manager of the Oakland Athletics in the late 1990's, now being played by Brad Pitt in the upcoming movie Moneyball.

There is no one ecologist in particular who can claim credit for popularizing trait-based approaches in community ecology, but for the sake of laughs let's make Owen Petchey the Brad Pitt analogue.

What can we do with this analogy? For pure nerd fun, we can think about what these two worlds can learn from each other.

What can baseball learn from community ecology?

One of the most notable trait-centric innovations in community ecology has been the use of functional diversity (FD), which represents how varied the species in a community are in terms of their functional traits. Many flavors of FD exist (one of which was authored by Owen Petchey, above), but the goal is to use one value to summarize the variation in functional traits of species in a community. A high value for a set of communities indicates greater distinctiveness among the community members, and is taken to represent greater niche complementarity.

For fun, I've taken stats from a fantastic baseball database[i] and calculated the FD of all baseball teams from 1871 to 2010. I used a select set of batting, fielding, and pitching statistics[ii], and you can see the data here. For the two teams that I pay the most attention to, I plotted their FD against wins, with World Series victories highlighted:

Given that these FD values represent how different the members of a team are, it's surprising that there is much of a pattern at all. But the negative relationship between wins and FD is strong and significant by several measures[iii]. So: the more similar a team is in terms of player statistics, the better the team does!

This pattern of less dissimilarity among players correlating with better performance at the team level has apparently been noticed before, by Stephen Jay Gould, who extrapolated this pattern also across teams to explain the gradual shrinking of differences among players over time:

"if general play has improved, with less variation among a group of consistently better payers, then disparity among teams should also decrease"

and so:

"As play improves and bell curves march towards right walls, variation must shrink at the right tail." (from "Full House", thanks to Marc for this quote!).

Interesting, but is it useful? One obvious drawback in this approach of examining variation in individual performance is that it ignores the fact that in baseball, we know that a high number of earned runs allowed is bad for a pitcher, and a low number for hits is bad for a hitter. In contrast, a high value for specific leaf area is neither good nor bad for a plant, just an indication of its nutrient acquisition strategy.

There are many exponentially more nerdy avenues to go with applying community ecology tools to baseball data, but I'll spare you from that for now!

What can community ecology learn from baseball?

One new baseball stat that gets a lot of attention during trades is 'wins above replacement'. This is such a complicated statistic to calculate that the "simple" definition is that for fielders, you add together wRAA and UZR, while for pitchers it is based off of FIP. I hope that cleared things up.

The point in the end is to say how many wins a player is worth, when compared to the average player. In ecology, the concept of 'wins above replacement' has at least two analogies.

First, community ecologists have been doing competition experiments since the dawn of time. The goal is to figure out what the effect of a species is at the community level, although fully factorial competition experiments at the community level are challenging to carry out. For example, Weigelt and colleagues showed that there can be non-additive effects of competitor plant species on a target species, but could rank the effect of competitors. This result allowed them to predict the effect of adding or removing a competitor species from a mixture, in a roughly similar way to how a general manager would want to know how a trade would change his or her team's performance.

Second, ecologists have shown that both niche complementarity and a 'sampling effect' are responsible for driving the positive relationship between biodiversity and ecosystem functioning. The sampling effect refers to the increasing chance of including a particularly influential species when the number of species increases. Large-scale experiments in grasslands have been carried out where plants are grown in monoculture and then many combinations, up to 60 species. The use of the monocultures allows an analysis similar in spirit to 'wins above replacement', by testing how much the presence of a particular species, versus the number of species, alters the community performance.

We could take this analogy further, and think of communities more like teams. A restoration ecologist might calculate 'wins above replacement' for all the species in a set of communities, and then create All Star communities from the top performers.

Lessons learned

A. Shockingly, there are baseball nerds, and there are ecology nerds, and there are even double-whammy basebology nerds.

B. There are quantitative approaches to analyzing individual performance in these crazily disparate realms which might be useful to each other.

C. I might need to spend more time writing papers and less time geeking out about baseball!

More analogies to consider:

Reciprocal transplants: trades?

Trophic levels: minor league system?

Nitrogen fertilization: steroids?

[i] One of the most astonishing databases around: complete downloadable stats for every player since 1871. This database is what NEON should aspire to be, except that this one was compiled completely privately by some single-minded and visionary baseball geeks!

[ii] Batting: Hits, at bats, runs batted in, stolen bases, walks, home runs

Fielding: Put outs, assists, errors, zone rating

Pitching: Earned run average, home runs allowed, walks, strike outs.

[iii] E.g. even after taking into account other more typical measures of success in offense (runs, R) and defense (runs allowed, RA), within years, there is still a negative slope for FD on wins:

lme(win ~ R + RA + FD, random = ~1|yearID, data = team)

Value Std.Err DF t-value p-value

(Intercept) 80.289 0.7411 2159 108.3 <0.001

R 0.107 0.0009 2159 116.8 <0.001

RA -0.105 0.0009 2159 -115.6 <0.001

FD -1.729 0.8083 2159 -2.1 0.0325

Day 5 in Austin

2011-08-12T12:06:00.000-07:00

The ecological community in Austin assembled for the final morning of talks today, and despite the advanced stage of the conference, the 8 o’clock talk I was at was surprisingly well-attended. There was only a morning worth of talks, but frankly, the Community Pattern and Dynamics session I attended had some of the most interesting talks I’d seen all week.

I started out in the Aquatic Terrestrial Linkages section, where Tiffany Schriever introduced me to the concept of spatial subsidies (the transfer of energy from one system to another), and described her system-temporary ponds in Ontario, Canada-in which the dual aquatic and terrestrial nature of the pond amphibian and insect life cycles couple aquatic and terrestrial systems.

Although I arrived late for Rafael D’Andrea’s talk in this session, it seemed that he did an excellent job of presenting ecological models, making his question and results both clear and interesting. He examined how tradeoff models, such as Muller Landau’s seed tolerance vs. seed fecundity model, predict far less diversity can be supported when the tradeoff changes smoothly rather than abruptly.

Nathan Sanders then explained his shift from primarily place-based research to global, macroecological studies. However, Sanders acknowledged the common criticisms of the macroecological approach, in that patterns are not necessarily evidence of mechanism, and has attempted to reach a compromise between the benefits of the two approaches. To balance place-based with global approaches, Sanders and his collaborators form a global network of researchers who are carrying out the same manipulative experiments (looking at resource limitation in ants) across different systems worldwide, and the results promise to be exciting.

In the final talk in the session, Steve Walker from the Legendre lab presented his approach to dealing with the “fourth corner problem”, that is relating species traits to environmental conditions. Rather than developing new approaches for analysis, Walker has focused on approaching this problem via data management. In particular, he has developed an R package (beta available here) called multitable in which data with different dimensions (such as a site-by-species matrix and a trait-by-species matrix) can be subscripted simultaneously, and coerced into a single data frame for use in standard R functions.

All in all it was a great week, and I’m excited to get back to work and feed off the energy of the conference. ESA gave me a chance to meet some of my favourite ecology bloggers, including Ethan White and Morgan Ernest from Jabberwocky Ecology and Jeremy Fox from the Oikos blog. It made me wonder whether there might be room next year for a more formal meeting of these online colleagues, whether in a loosely organized sense, or even as a workshop or symposium focused on the how ecologists can use (and are using) new technology—especially the internet—to communicate their science. If anyone has any comments or thoughts about whether there would be a role for something like this at ESA, I'd be happy to hear them.

See you in Portland, one year from now!

Day 4

2011-08-11T17:45:00.000-07:00

As the fourth day of ESA sessions began, it was clear that many attendees’ (including my own) energy was flagging, and 8:00 am talks were more sparsely attended. However, this didn’t mean that the presentations were of any less quality or interest. In the face of so many competing talks, I took the path of least resistance, in that I chose an interesting-sounding session, and stayed put. Today, that meant that I saw much of the Community Pattern and Dynamics IV session, and some talks in the Invasion: Invasibility, Stability, and Diversity session.

Thursday poster session

The early morning portion of Community Patterns and Dynamics provided some interesting talks focused on diversity in aquatic ecosystems, in particular ponds. Schalk et al. examined how the community structure of tadpoles related to environmental gradients in Bolivia, in particular how tradeoffs in pond permanency, predation, and canopy cover determine the tadpole species present. This talk proved that frog ovipositing behaviour can be fascinating, and provided the first example I've seen of a mule-aided sampling strategy (the mule transported supplies).

Jamie Kneitel reminded us that aquatic ecosystems, and the functions and services they provide, are under threat. One issue is human-driven increases in turbidity. To examine the impacts of turbidity, and how these impacts may differ depending on the underlying cause – turbidity may be cause by input of resource, leading to eutrophication, or directly, such as via cattle trampling in pools—Kneitel used experimental mesocosms meant to simulate vernal pools. He showed that different causes of turbidity conferred different types of changes in water chemistry, and that turbidity had different effects on vegetation and invertebrate communities.

Moving to the Invasion: Invasibility, Stability, and Diversity session, Jiaqui Tan from Lin Jiang’s lab gave a very interesting talk about the oft-observed negative relationship between invasibility and diversity. In particular he connected the suggested explanation for this pattern (the sampling effect and the niche complementarity effect) to phylogenetic patterns of relationship between species. He predicted that increasing phylogenetic relatedness would increase the sampling effect, while increasing phylogenetic diversity would increase niche complementarity. Using microcosms of aquatic bacteria, in which Serratia marcesens acted as the invader, he looked at how a factorial design of low, medium, and high phylogenetic relatedness and phylogenetic diversity effected invasibility of the bacterial communities. Perhaps surprisingly, phylogenetic diversity had only a little effect on the degree of invasibility, but phylogenetic relatedness strongly decreased invasibility. However, an explanation for these results was made more difficult by the fact that the 57 traits he measured for the bacteria showed little evidence of niche conservation.

At the same session, Karen Alofs shared some preliminary analysis of a fantastic dataset showing changes in the presence of introduced smallmouth bass over 30 years in Ontario, Canada. Ontario is a place where southern and northern range limits of many fish species occur, and ranges are limited by temperature. Smallmouth bass is an invasive species originally introduced for sport, and Karen examined how changes through time in the bass’ range related to species composition, environmental characteristics and predator presence. She found that abiotic and biotic variables were correlated with invasion probability, but made the important conclusion that community composition affects invasibility, but the reverse—invasion alters community composition—is also true.

Finally, Galen Holt from Peter Chesson’s lab gave a great talk about fitness-density covariance as a possible mechanism of coexistence, and the interaction of dispersal with it. He examined water invertebrate species in a stream, in which both symmetrical dispersal and asymmetric dispersal (i.e. stream flow) could affect the strength of this mechanism. In fact, he found that dispersal weakens fitness-density covariance since species are less likely to stay where environment is best, as is required by this mechanism.

An unidentified University of Toronto student,

letting loose after a long day of talks

ESA Austin: Day 3

2011-08-10T16:52:00.000-07:00

Another great day of talks, and surprisingly so. I went to several talks that were more or less randomly chosen, and was impressed by some of the science that graduate students and younger scientists are doing. While others see a potential decline in ecology, I see a very bright future.

M. Duffy examined how virulence susceptibility and the cost of disease resistance in daphnia and its pathogen (yeast). This talk had one of the best setups I seen, and is based on a trade off between r and resistance. Epidemics result in increased resistance, as one would expect with evolution, but equal number of lakes showed evolution for increased susceptibility because of these tradeoffs. Small epidemics should result in increased susceptibility because of a greater fitness cost from reducing r than from mortality. Large outbreaks should result in greater susceptibility as mortality is high. She also showed that environmental context could alter expectation (e.g., productivity or predation).

In the next talk N. Loeuille examined the heterogeneity of resource dispersal. Classic models assume homogeneity in resources availability for competition. But with different diffusion rates, niche competition may be decoupled from tradeoffs needed for coexistence. He used a model with differential dispersal of resources. Depending on tradeoffs, the model will produce the evolution of diverse strategies of disperal. There will be specialization on single resource if dispersal is symmetric. If dispersal rate is too high or too low, but equal, then the resources support lower diversity. If two resources disperse differentially, creates heterogeneity at different scales and will support higher consumer diversity.

I ran over to Tad Fukami’s talk. He examined phylogenetic patterns in priority effects in the assemblages of yeast that colonize flower nectar. He hypothesized that there should be a strong priority effect with close relatives, since they tend to occupy similar niches. He tested this by Introducing species in different orders and assessed relatedness effects using genetic sequences. The experiment was directed by natural history of the system, like time length of flowers, microbial population dynamics in flowers. If one species colonizes first, he showed that it can reach carrying capacity regardless of the presence of other species. If it arrives second, there are major effects on abundance, but differs between which is first species. Closely related species result in strong priority effect, but weak with distant relatives. Result robust even if you control for differential ability to deal with abiotic conditions. The potential mechanisms include differentially reducing amino acids, and different growth rates on sugars.

Kevin smith gave a great talk on extinctions. He used several large, recently assembled datasets to examine how range size correlates with extinction risk under different scenarios of habitat destruction. Randomly, you would expect that broad species have a low probability of extinction overall and endemics have a high probability. Across the datasets, widespread species are going extinct at higher probabilities then a random model. Land snails conform to random model based on species range. However, for bird and amphibian datasets, the rare species bahave as expected with high extinction risk, but the middle ranged species have higher than expected extinction risk

L. Prevost examined how the theory of Island Biogeography (IBG) explained diversity patterns in fragmented habitats in mid to high elevation habitats in Costa Rica. The short answer is not very well, there were not distance or area effects on plant diversity. Communities have low similarity, no relationship with distance, but are similar according to elevation. It seems as though species responses to heterogeneity drives the system, so she recommends that many small reserves could be valuable.

In a very interesting and stimulating talk, A. Rominger examined fluctuations in evolutionary history. He showed that there are more fluctuations than predicted by various models including random walks (which conforms to a Gaussian distribution). Gaussian is observed in small time slices, but variances change over time. Fluctuations within orders fit Gaussian very well, but different from one another. Volatility itself evolves by gamma distribution. He shows that volatility is inherited within orders, and fascinating and controversial conclusion.

John Parker testing the often assumed, but understudied assumption that exotics differ in advented populations versus native. Basically, is there an away-field advantage? Examined home and away for 1000 worst invaders, across many taxa. Looked at size, reproduction, population growth. None of these were particularly enlightening. For example they are not bigger in away sites, size at home predicts invader size 1:1. There is some variation, but no consistent trend. Fecundity, also no consistent trend, with noninvasive just as likely to increased fecundity. Abundance not consistent but slight trend to be bigger away. Survival, growth, same thing. Overall slightly better away, but not greatly. We need to reexamine our hypotheses.

Day 2 in Austin

2011-08-10T05:31:00.000-07:00

The second day of ESA got off to a good start in Austin, with a day full of more community ecology talks than one person could attend. I split my day between Community Assembly and Neutral Theory II& III (it's interesting to note that ten years in neutral theory is now included in so many eponymous sessions) and Biodiversity I, and regretted missing talks in many other sessions.

The Community Assembly and Neutral Theory II covered a diverse range of systems (from microcosms to primates), scales, and methods of study. Lin Jiang presented an experiment examining the relationship between diversity and invasibility, in particular testing whether priority effects reduce the oft-seen negative relationship between diversity and invisibility. Most manipulative experiments "assemble" communities instantaneously rather than continuously and stochastically as in natural systems, and so more realistic assembly may weaken the sampling effect and niche complementarity, which are suggested to drive the negative relationship. Using protist-based microcosms of 5 resident species and one invading species, Jiang examined how more realistic assembly of communities affected the diversity-invasibility relationship. Under these conditions, there was still evidence of a negative invasion diversity effect. His most interesting result however were that in fact the presence of a close relative had the strongest influence on the success of the invasive species, in line with other theoretical and empirical results (although not conclusive given the small number of species).

In the Community Assembly and Neutral Theory II session common topics in this session included the widely-used framework of hierarchical filters (i.e. abiotic, biotic, dispersal limitation) determining local species composition and tests of the predictions of the neutral theory (with a focus on non-SAD predictions). For example Wang et al. looked at the patterns of clade age versus abundance that were predicted by neutral theory, in comparison with empirical data from the BCI dataset. There was a clear divergence between the observed data, which included old clades with high or medium abundances compared to the neutral theory prediction that old species should have low abundances. Wang examined how relaxing the assumption of equal rates of speciation among species affected the age-abundance patterns, but concluded that different rates of speciation among species wouldn't produce the observed pattern without then failing the SAD predictions. However, one astute commenter noted that it might also be important to model the possibility of changing rates of speciation through time.

There were many other interesting talks. For example, in Biodiversity I, Matthew Leibold provided a master class on resilience to human disturbance, focusing on concept of communities and ecosystems as complex sets of coupled oscillators. Finally, Angela Brandt presented the results of a 7-year experiment in California grasslands (which no doubt represents many hours of hard work), in which she examined the relationship between invasion success, resource availability, and disturbance. In particular, she framed the question from a phylogenetic context, and discovered evidence that disturbed communities tended to be both more species-rich and phylogenetically diverse, and also less phylogenetically clustered, compared to non-disturbed communities. However, if communities received nutrient enrichment and disturbance, invasion was greater, and diversity lower, than in communities that received disturbance treatments only.

Looking forward to day 3!

***Addendum by Marc:
There were two additional talks that were particularly interesting. First was Andy Gonzalez’s talk on Evolutionary rescue, which is when a population is in demographic decline, heading towards extinction and adaptation saves it from extinction. This is particularly important in changing environments. He has previously shown this using yeast growing under salt stress. His question now was whether migration in a metapopulation with heterogeneous affects this evolutionary rescue. This is an interesting question because too little dispersal means that genetic variation or beneficial mutations do not get to other patches, and too high means that suboptimal genes are maintained in patches where they are maladapted. Not surprisingly he showed that in a constant environment, dispersal is not very important. In heterogeneous metapopulations, patches at the edge of salt tolerance thresholds increase in yield with dispersal.

The other very interesting talk was from K. Anderson on niche-based sorting in highly diverse palm assemblages. She looked at soil type and resource availability, as well as herbivore damage. She experimentally planted 13 species across a soil gradient with and without herbivore exclosures. In low nutrient soils, the palms invested more in roots while in high resources soils, there was a greater investment in above ground biomass and an increase in photosynthetic rate. Leaf toughness also increased in poor sites, meaning that they were more resistant to herbivory and plants growing in the high resource sites experience more herbivore damage. She mentioned that there were differential responses from the different species, and I am very interested to see more about this neat system.

ESA Austin: Day 1

2011-08-08T21:03:00.000-07:00

We are now through a great first day at the ESA Austin meeting, and have been having a great time both at talks and out on the town in Austin (see photos). Looking over the program, it was obvious that the day had too many good talks, and that it was impossible to see them all. Considering that I was giving a talk, I decided to spend my entire time in my session on biodiversity and ecosystem function. It may seem lazy, but there were a bunch of talks that sounded great. Here are short summaries of all the talks in the session (excluding mine of course).

The Decemberists playing at Stubbs. Fantastic show (maybe the highlight of day 1, if not for the many interesting talks)

Darwin's pub, great name, OK pub.

The first talk by Nicolas Mouquet was probably the best. It was on the relationship between species diversity and ecosystem function, asking how we can move from the question of how many species to which species. The ultimate answer, according Mouquet, comes from evolution. By understanding the evolution of specialization, one can discern the importance of niche complementarity in the additive contributions to ecosystem function. Using simulations, he showed that the relationship between richness and function is dependent on whether species are specialist or generalist and the strength of tradeoffs in resource use. He then told us about fantastic experiments that evolve bacteria on differing resources, creating specialists and generalists. Positive diversity-function relationships were higher but not stronger in assemblages of generalists, because they deal with heterogeneity better. He manipulated the amount of evolutionary history in assemblages and found that the relationship between evolutionary diversity and function was stronger with groups of specialists. This research goes beyond most others in that it explicitly links coexistence to ecosystem function.

Next was a talk by J. Passari, looking at ecosystem multifunctionality in a long term plant experiment. He examined eight different functions and examined how local, large scale and among site diversity influenced ecosystem function. He found that multifunctionality increased with increasing local diversity but less so with diversity at larger scales.

Greg Crutsinger, showed how genotypic and phenotypic differences in coyote shrub morphs resulted in differences in arthropod abundance and richness, and changes in litter communities.

M. Striebel examined how phytoplankton diversity affect function. She showed that total pigment diversity (representing photosynthesis) increased with phytoplankton diversity. Also she examine how this diversity affected zooplankton diversity and found positive relationships in oligotrophic and mesotrophic systems, but not eutrophic ones.

J. Mclaren manipulated the functional group richness in desert and arctic plant communities and examine the community and functional responses. There was some compensation by other functional groups, but only a weak overall affect on function.

J. Petermann manipulated basal resource diversity and predator richness in bromeliad aquatic communities. She measured four functions and found only weak effects, it seems as though bromeliad leaf complexity may drive some of these relationships.

E. Harvey showed how multiple extinctions in complex food webs can have important cascading effects on ecosystem function. He measured multiple functions in freshwater and marine communities, and that different extinctions had differential effects and some where non-linear.

JJ Weis gave a very interesting talk where he used a model to assess how intra- and interspecific diversity affect function. He found that high complementarity resulted when species had high genotypic variation but low genotypic breadth.

Finally, T. Hanley, who is also a student in the same lab as JJ Weis, examined how intraspecific variation affected population dynamics of daphnia and their algal prey. There wasn’t any effect of daphnia genotypes on algal or daphnia dynamics, but daphnia genotypic diversity increased through time.

What is interesting about this group of talks is the diversity of organisms, systems, scales and functions being considered. These talks are a great signal that biodiversity-ecosystem function research transcend locales and is now a broad, mature field of study.

Blogging our way through Texas

2011-08-05T03:43:00.000-07:00

We are on our way to the ESA meeting in Texas! During the meeting next week, we will provide daily updates on the EEB and Flow, recounting some of the interesting talks and happenings there. See y'all in Texas.

The empirical divide

2011-07-25T18:24:00.000-07:00

Has there been a shift in how ecology is done? In an interesting editorial in the most recent ESA Bulletin, titled “Losing the Culture of Ecology”, David Lindenmayer and Gene Likens wrote that “empirical and place-based research”, such as field studies and taxonomy, appear to be falling out of favor. They suggest that ecological modeling, meta-analysis, and data-mining (the three M’s) are more lucrative (and popular) approaches today, because these methods are faster, cheaper, and “easier” to perform, allowing more rapid publication. While they recognize the important advancements resulting from these methods, the result—they suggest—is that field-based empirical research is becoming less prevalent, to the detriment of ecology.

This is a polarizing issue, and the response of those ecologists we spoke to depended on where they position themselves on the field/theoretical divide. Those who define themselves as field ecologists tended to feel embattled in the face of long, expensive months of fieldwork, with slow returns in terms of data and publications. Some felt there is a subtle insinuation that fieldwork is less generalizable and so less valuable than techniques such as meta-analysis and ecological modeling, which by their nature tend to be theory-based and general.

On the other side, some theoretical ecologists we spoke to felt the need to defend the validity of doing “indoor” ecology, noting that theory and modeling can link pattern and process, without the confounding variation common in field experiments/observations. Although field ecologists felt that they have a more difficult time obtaining funding, theoretical ecologists noted that they often receive far less money because the assumption is that theory is “free”. Further, with the exception of very specialized funding opportunities (e.g., NCEAS), meta-analyses do not typically get funded as stand-alone projects.

It’s important to note that in its short history, ecology has frequently struggled with the balance between the field and lab. The primary criticism of field-based research at the turn of the 20th century was that it was “unscientific”, inseparable from natural history, producing lists of species names rather than furthering understanding, while labwork was considered to be too divorced from natural systems to be informative (producing so-called “armchair ecologists”). These conflicts split some of the first organismal departments in the United States (*) and tensions exist to this day. No doubt these criticisms are not unfamiliar to many modern ecologists.

There needs to be a balance between the production and consumption of data. Obviously abandoning fieldwork and using only meta-analysis, modeling, and data-mining is not sustainable, but these are important methods for modern ecology. In addition, the perceptions of bias against fieldwork may be due to a general decline in funding and greater overall competitiveness for the rewards of academic labour (jobs, grants, publishing in top journals, etc.), rather than a true decline in field ecology. As we discussed this article, it became clear that our own perceptions, and perhaps those of the broader community, have formed in the absence of empirical data. We examined the last few issues of some highly-ranked ecological journals that publish primary research (Ecology Letters, Molecular Ecology, American Naturalist), and recorded the number of papers that used empirical data, and further the number of those that collected their own data (versus using data from databases, literature, etc). Surprisingly, the vast majority of studies were based on empirical data, mostly data collected by the authors. In Molecular Ecology, 27 out of 28 papers were empirical, and 26 of these used data collected by the author(s); in Ecology Letters, 17 out of 20 papers were empirical, and 12 of these used data collected by the author(s). Even in American Naturalist, which is known for its theoretical bent, 44 out of 70 papers were empirical, and 32 used the author(s)’ own data. Overall, these journals, where competition for space is most severe, primarily publish empirical research.

It appears then, that neither grants nor publications systemically bias towards the three M’s. But is there still a cost to researchers on either side of the data producer-consumer divide? The answer is likely yes. The three M’s result in quicker publications, which means these researchers look more productive on paper, resulting in greater visibility. With more publications, they are likely to make it to the top of hiring committee lists. Conversely, unless a specific job has been advertised as a modeling position, candidates giving job talks focusing on the three M’s do not come across as knowledgeably as a very skilled field person. One of us (MWC) has seen job searches at four different institutions, and the unadvertised stipulation for many departmental faculty or committee members is that the candidate will come and establish a field program. Another common criticism of 3-M candidates is that they will not be able to secure large amounts of research funding.

Given this double-edged sword, what is the optimal strategy? The glib, easy answer is that ecologists need to become less specialized, to do both theory and empirical work, if they want a successful career. And maybe this is the solution, at least for some ecologists. But is having everyone become a generalist really the answer? Most field ecologists will tell you that they do fieldwork in part because they love being in the field and they’re good at it; most theoretical ecologists are adept at manipulating ideas and theory. Perhaps there is still a role for the specialist: after all quantitative ecology—which produces data—and theoretical ecology—which consumes it—are inseparable. They have a complementary relationship, in which field observations and data fuel new models and ideas, which in turn provides new hypotheses to be tested in the field. It’s obvious that people should be able to specialize, and that the focus should be on increasing collaboration between the two groups.

Despite the hand-wrenching, perhaps this collaboration is already happening. Many of the very best 3-M papers unite theoretically-minded with empirically-grounded ecologists. The working-group style funding by NCEAS (and its emulates) explicitly links together data producers and data consumers. These papers may be deserving of greater visibility. If collaboration is the future of ecology, why does the tension still exist between lab and field? The historical tension was not really about the laboratory vs. the field, but rather about scientific philosophy, and we think this holds true today. Ecology has tangibly moved towards hypothesis-driven research, at the expense of inductive science, which was more common in the past. The tensions between “indoor ecology” and field ecology have been conflated with changes in the philosophy of modern ecology, in the difficulties of obtaining funding and publishing as a modern ecologist, and some degree of thinking the “grass is always greener” in the other field. In fact, the empirical divide may not be as wide as is often suggested.

By Caroline Tucker and Marc Cadotte

* Robert E. Kohler. Landscapes and labscapes: Exploring the lab-field border in biology. 2002. University of Chicago Press. (This is a fascinating book about the early years of ecology, and definitely worth a read).

The reality of publishing papers

2011-06-29T09:40:00.000-07:00

This is in response to my undergrads, who ask me "Have you published any of the stuff we're working on yet?" practically every week. To which my response invariably is "not yet".

(click to make larger)

Metacommunity data and theory: the tortoise and the hare

2011-06-15T11:37:00.000-07:00

Empirical approaches to metacommunities: a review and comparison with theory
Logue et al. 2011

The recognition that community composition is a function of both local and regional-scale processes, meaning that a community cannot be understood in isolation from the network of communities with which it interacts, is the fundamental idea behind metacommunity ecology. In a relatively short period of time, metacommunity ecology has integrated concepts from spatial ecology, metapopulation ecology, and community ecology with novel ideas, and developed a strong body of theory. However, metacommunity theory has advanced much more rapidly than empirical tests of that theory. In an interesting review in TREE, Logue et al. examine whether empirical data needs to catch up with the pace of theory development, or whether theory is moving too fast to incorporate the information available from empirical data.

The types of systems used in the 34 experimental and 74 observational studies that Logue et al. found were very limited – the most common experimental approach involved setting up aquatic microcosms of unicellular organisms.* Observational studies similarly tested microorganisms, usually in aquatic systems. The organisms so beloved in the rest of community ecology (plants? vertebrates?) barely feature. Most studies focus on aquatic systems composed of multiple patches (such as microcosms, ponds, pitcher plant communities) because systems with discrete boundaries are more amenable to testing current theory. However, natural systems are rarely configured into a clear “patch” versus “matrix” dichotomy. Instead they are complex and heterogeneous, and may lack clear boundaries.

Dynamics in metacommunities are generally described using four dominant paradigms: mass-effects, species sorting, neutral perspective, or patch-dynamics. These paradigms reflect the most important processes structuring communities, that is, either dispersal between communities, environmental differences between communities, dynamics driven by the tenets of neutral theory, or extinction and colonization, respectively. Strikingly, experimental studies mostly tested for mass-effects or patch dynamics, and observational studies mostly tested for species-sorting and mass effects paradigms. The neutral paradigm was rarely tested in any type of study. Logue et al. found that many studies had difficulty designing experiments that tested for evidence of specific paradigms, because natural communities are much more complex than the simple paradigms suggest. Most studies that did test for evidence of particular paradigms found evidence for multiple paradigms or had difficulty disentangling different mechanisms.

The metacommunity theory that has developed in the last five years is among the most exciting and interesting work in ecology. However, the slower pace of experimental work means that theory has developed with little feedback. For example, Logue et al. make a strong argument that the results from these studies suggest that it is time to integrate the four-paradigm system into a single, comprehensive framework (see figure). Theory is only valuable if it’s useful - this paper is an important reminder that there is an important feedback loop between theory and data, and successful science requires input from both.

*Important disclaimer: at this very moment I'm running aquatic microcosms of microscopic protists in the lab. We all have room for improvment. :)

Navjot Sodhi, conservation for all

2011-06-13T06:21:00.000-07:00

I opened my e-mail to see the shocking and saddening news that Navjot Sodhi passed away yesterday (see here for more details). He was an absolute leader in tropical conservation biology from his base in Singapore. But more than this he made conservation biology accessible to the public and especially to those working on the front lines trying to protect biodiversity. His free edited book: 'Conservation Biology for All' set a new milestone in conservation biology and in the efforts of academics to step out of the ivory tower and reach out to broader communities.

Nature’s little blue pill

2011-05-30T11:07:00.000-07:00

Something often happens to mature community ecology studies as they get older that we don’t like to talk about much. Occasionally, when biodiversity and ecosystem functioning experiments are performed in a controlled, homogeneous setting, they can suffer from flaccid response curves. It’s perfectly normal, happens to lots of healthy microcosm communities, but it can be troubling and embarrassing nonetheless. After all, who doesn’t want a nice stiff linear response curve?

Bear with me here for a minute.

A few weeks ago, Bradley Cardinale published a study in which he tested the effects of algal biodiversity on water quality in streams. It’s a pretty classic diversity-function experiment; lots of artificial streams with different numbers of species of algae in them, and he measured productivity and nitrogen uptake. As is usually the case, the more species he put in each stream, the more these ecosystem functions increased.

But Cardinale did something else in this experiment that has never been done before, at least not on this scale. He added extra niche opportunities to some of the streams, so that they offered multiple different habitats for algae. He did this by introducing heterogeneity through flow and disturbance manipulations.

Figures from Cardinale 2011, Nature. a and b are the heterogeneous streams, d and e are the homogeneous ones.

You might be familiar with that saturating response curve that is typical of so many diversity-function experiments. It starts off with large increases in ecosystem functioning as species are added to communities, and then it levels off so that as additional species are added, they only increase ecosystem functioning by small amounts (figures d and e). The theory behind this is that there are only so many niches in an environment, and as more and more species are added some of them become redundant.

Well when Cardinale threw those extra habitats into his artificial streams, that floppy old saturating curve sprang up like a regressional jack-in-the-box (figures a and b).

What happened was the homogeneous streams became dominated by just a single species that was well adapted to that environment. The heterogeneous streams allowed different species to coexist and this let them make more efficient use of the resources in those streams.

This is a major finding for a few reasons. First, it confirms that one of the main mechanisms behind diversity-function relationships is niche partitioning. I’ve said in the past that knowledge of these mechanisms is sorely needed. Second, it links coexistence theory to ecosystem functioning, two fields that are closely related but often disconnected.

Finally, it means that biodiversity is even more valuable than we had previously thought. The natural world doesn’t contain very many homogeneous streams; it’s a complicated place. The real world is probably better represented by figures a and b than by figures d and e. So while controlled experiments have shown that some species are redundant for ecosystem functioning, there is no evidence here for any redundancy in more natural settings.

This paper also underlines the fact that these studies need to be done in nature as opposed to labs. Cardinale was able to simulate nature fairly realistically because he was using algae. That’s harder to do with more complex organisms. It’s difficult to recreate environmental heterogeneity in artificial ecosystems, and if ecosystem functioning depends on both biodiversity and heterogeneity, then it’s time to take this research outside. Manipulative field studies are a good start, but completely natural settings will probably reveal more of the true story.

So although it’s very common for artificial communities to suffer from Ecological Dysfunction, there is no reason that they can’t enjoy a healthy relationship with biodiversity like any other community. All they need is a little heterogeneity to spice things up and put that spring back in their step.

Andy Hector has written an excellent perspective on the study. I recommend reading it, particularly if you don’t want to read the entire original article.

The ecology blogosphere just got a little more crowded, and better (welcome Oikos blog)

2011-05-19T13:50:00.000-07:00

A diversity of voices is why the internet is such a powerful intellectually democratizing form of communication. Ecology blogs, long the minority in scientific blogging just received an immense boost from the new Oikos blog, obviously associated with the journal, Oikos. While some of their content is dedicated to journal business, there have been great posts on ecological research and broader intellectual topics from Jeremy Fox, aka oikosjeremy.

Welcome.

Happy 10th birthday, neutral theory!

2011-05-17T07:14:00.000-07:00

Rosindell, Hubbell, and Etienne. (2011). The unified neutral theory of biodiversity and biogeography at age ten.

I would argue that neutral theory is not only the most controversial idea, but also the most successful idea to permeate community ecology in the last ten years. A quick keyword search suggests that ~30 ecological papers related to the topic were published in the last year, including some with titles still reflecting the controversy; “Different but equal: the implausible assumption at the heart of neutral theory”. Neutral theory makes a seemingly unreasonable assumption—that species identity doesn’t matter—and yet seems to predict species-area relationships and species abundance distributions as well or better than niche theory does. This made it an infuriating challenge for many ecologists. The number and quality of papers that it inspired—both in support and opposition—are a reminder that disagreement is good for science.

It’s been a decade since the publication of “The Unified Neutral Theory of Biodiversity and Biogeography”, in which Steve Hubbell proposed a controversial model in which coexistence results from drift, dispersal and speciation, rather than ecological differences between species. To mark this anniversary, a review in TREE by James Rosindell, Stephen Hubbell, and Rampal Etienne reflects on neutral theory’s first ten years, and examine the influence neutral theory has had in many areas of community ecology. The authors also note that some of the limitations of neutral theory can be dealt with by extending the classic formulation of the model, so that unrealistic assumptions related to spatial structure, speciation rates, or the zero-sum assumption can be relaxed. The excessive interest in neutral theory’s species-abundance predictions left its other predictions unexamined, and there is still room for tests of how neutral theory informs species-time relationships, modes of speciation, and even conservation decisions.

Despite these accomplishments, the review is remarkably subdued, underlined by statements such as neutral theory is a “good starting point”, a “valuable null model”, and a “useful baseline”. However, it seems unnecessary to state, as some have, that "neutral theory is dead". Its legacy, captured in the final paragraphs, is still incredibly important: “…niches have dominated our attention and left less obvious, but still important processes forgotten… Perhaps the most important contribution of neutral theory has been to highlight the key roles of dispersal limitation, speciation and ecological drift, by showing how much can be explained by these processes alone...”

George Box said it best: “All models are wrong, but some are useful”.

Carnival three-five.

2011-05-02T12:05:00.000-07:00

The 35th installment of the Carnival of Evolution is available from Lab Rat. Want to know who said what about evolution? Go to the Carnival.

Ecological interactions and evolutionary relatedness: contrary effects of conserved niches

2011-04-29T12:50:00.000-07:00

Over the past several years a multitude of papers linking patterns of evolutionary relatedness to community structure and species coexistence. Much of this work has looked at co-occurrence patterns and looked for non-random patterns of relatedness. The key explanations of patterns has been that communities comprised of more distantly-related species is thought to be structured by competitive interactions, excluding close relatives. Alternatively, communities comprised of species that are closely related, are thought to share some key feature that allows them to persist in a particular set of environmental conditions or stress. This whole area of research is completely predicated on close relatives having more similar niche requirements then two distant relatives. This predication is seldom tested.In a recent paper in the Proceedings of the National Academy of Science, Jean Burns and Sharon Strauss examine the ecological similarity among 32 plant species and tested if evolutionary relationships offered insight into these similarities. The ecological aspects they examined were germination and early survival rates as well as interaction strengths among species. To assess how these were influenced by evolutionary relatedness, they planted each species in the presence of one of four other species varying in time since divergence from a common ancestor, creating a gradient of relatedness for each species. They found that germination and early survival decreased with increasing evolutionary distance. This surprising result means that species germinating near close relatives do better early on then if they are near distant relatives. The explanation could be that they share many of their biotic and abiotic requirements, and these conserved traits influence early success.

Conversely, when they examined interaction strengths over a longer period (measured as relative individual biomass with and without a competitor), they found that negative interactions were stronger among close relatives.

These two results reveal how evolutionary history can offer insight into ecological interactions, and that the mutually exclusive models of competitive exclusion versus environmental filtering do not capture the full and subtle influence of conserved ecologies. Evolutionarily conserved traits can explain both correlated environmental responses and competitive interactions.

Burns, J., & Strauss, S. (2011). More closely related species are more ecologically similar in an experimental test Proceedings of the National Academy of Sciences, 108 (13), 5302-5307 DOI: 10.1073/pnas.1013003108

The bellybutton, biodiversity reserve of the body

2011-04-15T17:07:00.000-07:00

Although less recognized--and less glamourous--than most biodiversity hotspots, the human bellybutton harbours it own diverse collection of species, and these species tell us something about ourselves. That's the premise behind the Belly Button Biodiversity Project, which is getting some press for its large-scale sampling of bellybutton bacteria. For interesting discussion about where the data could lead, see Rob Dunn's (one of the researchers) website. His post, including the comments, hints at how much there is to learn about the ecology of human bacteria.

Update: Rob Dunn has now published a book "The Wild Life of Our Bodies", telling more stories of our changing relationships with other species.

Carnival time.

2011-04-03T03:39:00.000-07:00

The 34th edition of the Carnival of Evolution is hosted at Quintessence of Dust. Everything from the evolution of perfection to the evolution of small importance will be found there.

White-nose syndrome and wind turbines: why biodiversity matters

2011-04-01T22:22:00.000-07:00

Linking ecosystem services to economic benefits is a vital step in connecting ecological research to policy and political action. The UN Environmental Programme’s The Economics of Ecosystems and Biodiversity (TEEB) initiative represents a concerted effort to draw attention to the economic benefits of biodiversity and cost of ecosystem degradation, and to bring together scientists, economists and policy-makers.

Accordingly, Boyles et al. (Nature, 2011) paint a troubling picture about the value of economic benefits that insectivorous bats provide to the North American economy, and the degree of extinction risk they currently face. The authors point out that bats are “among the most overlooked, yet economically important, non-domesticated animals in North America”, and their loss would cost North Americans more than 3.7 billion dollars/year. Given rapid declines in populations due to white-nose syndrome (over 1 million bats killed) and wind turbine fatalities (projected to reach up to 30,000-100,000 fatalities/year as wind turbine installations increase), the authors suggest action can't wait.

Hopefully using the universal language of money helps translate scientific knowledge into political action. After all, bats are only one group of species: imagine the true cost of current rates of biodiversity loss and ecosystem destruction, from the smallest microorganism to the largest megafauna. The total must be staggering. And so, it seems, is the scale of action required to halt this decline.

The regional community, maximum entropy, and other ideas in ecology

2011-03-18T07:20:00.000-07:00

Looking through my feed of community ecology papers this month, I couldn’t help but notice that while most tested well-established concepts–density-dependence, niche partitioning, metacommunities, competition, dispersal limitation–there was also–as I suppose is usually true–a subset of papers championing newer, less established ideas.

For example, the article “Applying a regional community concept to forest birds of eastern North America” by Robert Ricklefs, furthers the regional community concept he introduced in 2008. Ricklefs is uncomfortable with how ecologists typically define local communities – i.e as spatially and ecologically discreet entities – and the predominant focus in community ecology on local coexistence. He argues that communities make sense as entities only at a larger scale, taking into account that local communities are not isolated, but instead interact as a function of overlapping ranges and species dispersal. In this paper he applies this concept to Breeding Bird Survey data to examine the distribution and abundance of birds in eastern NA.

Partel, Szava-Kovats, and Zobel are also critical of the predominant focus on local diversity. In their paper “Dark diversity: shedding light on absent species”, they pitch the idea of “dark diversity” as a valid diversity metric. Dark diversity accounts for the number of species which belong to the species pool for a particular habitat in a region but are not actually present in a local community of that habitat type. The resulting value can be used to calculate a dimensionless ratio of local to dark diversity, suitable for comparison of diversity components in dissimilar regions.

Lastly, in “A strong test of a maximum entropy model of trait-based community assembly”, Shipley et al. further test Shipley’s model of Entropy Maximization, using it to predict the composition of communities in the South African fynbos. The model predicts community composition (species identity and relative abundances) through an assumption of random assembly (or entropy maximization) within environmental constraints on species traits.

New ideas are a constant in ecology, but they face stiff competition in an already crowded field. The possible mechanisms of local coexistence, for example, are already a long list. What determines which of these–or any–ideas become entrenched in ecology? The likelihood of a concept becoming established must be a complex function relying on a cost-benefit analysis–what does applying this idea cost compared to the gain in understanding it produces?–further adjusted by intangible variables like timing and the skill and prestige of an idea’s advocate. After all, some ideas require decades to establish properly, requiring changes in the theoretical climate or technical capabilities, for example, neutral theory or spatial ecology. Others seem to catch on immediately. Philosophers have written more cogently on how scientific ideas change and paradigms shift, but as participants in the process, we have a rather unique perspective. After all, as scientists we play an active role in driving these shifts in thought and action. You might argue that the merit of the ecological ideas that become established are as much a reflection on those who accept and institute them, as on those who propose them.

≠

2011-03-08T18:44:00.000-08:00

awesome infographic by Caroline Tucker

(click to go large)

There exists a problem in science so complicated that decades of work have yet to solve it. Its causes and consequences make some of the toughest questions in complex analysis or astrophysics look like child’s play. And yet when we consider this problem, the conclusion is immediately obvious and simple: it should not exist.

I am talking about the fact that today, in 2011, female scientists are punished solely because they are female scientists.

In theory, this problem doesn’t exist anymore. Multiple waves of feminism should have chipped away whatever glass ceilings once capped our ivory towers. Women are receiving more PhDs than men in many fields and they are earning such a high proportion of bachelor’s degrees that we may have to rethink that name.

But in the last several weeks, some disturbing realities have resurfaced in the science media. At the end of January, Nature reported that women earn fewer scholarly awards than they should, based on the proportion of their respective fields that they represent. That same week, Science published a graph showing the number of European Research Council grants awarded to women in its last funding round – 9.4%.

Stats like these are nothing new; they pop up all the time. What is new, however, is the article that followed in Proceedings of the National Academy of Sciences a few days later. It turns out that there is no longer much evidence for overt discrimination against women applying for jobs or grants in quantitative fields. Instead, disparities in available resources are causing many of the differences between women and men’s scientific careers.

Yes, there are discrepancies in publication acceptance rates and grants, but the authors attribute these to factors like women occupying more positions at teaching-intensive schools rather than research institutions. When they compared men and women with similar resources, the biases disappeared, or in some cases, favoured women. (If you don’t want to read the whole article, there’s a nice summary of it here.)

Ok great, the science community isn’t explicitly discriminating against women. But this leads us to a much more troubling conclusion; the culprits are actually deeply engrained societal expectations and constraints that likely extend well beyond the sciences, and certainly beyond the scope of this blog post, though a few of them are highlighted in this thoughtful opinion piece.

Here’s what I will say: it’s not written in our DNA. How many times have you heard lines like, “Men and women are just different, they always will be, our brains aren’t wired the same”? This kind of just so statement is rarely backed up with evidence. For a good debunking of these misconceptions, check out two new books, reviewed here.

Now it’s possible that I, as a young male grad student, do not hold the most valuable two cents on these issues. I could keep rambling about things that I don’t fully understand, but my perspective is limited, and I think maybe the most constructive thing to do at this point would be to hear about other people’s ideas and experiences in the comments section below. So I’m cutting this short and leaving it incomplete in favour of a more open forum. In particular, it occurred to me that we in the ecology and evolution community have a unique opportunity to shed light on many gender issues. The PNAS article focuses on the underrepresentation of women in math-intensive fields, but comparing mathy fields to less-mathy fields entails a lot of confounding factors. In ecology and evolution, however, we cover the whole spectrum, from the completely mathless and descriptive, to the suspender-wearing, calculator-toting quants. We generally all come from relatively similar biology backgrounds, eliminating many of those confounding factors, and it would be great to hear how you all think these issues play out in E & E. So go for it blogosphere, do your thing.

Documenting the sacrifice

2011-02-13T19:06:00.000-08:00

Whether working in remote regions or with poisonous animals, biologists often put themselves in peril in the name of discovery. The first organized expeditions of naturalists sailed to exotic lands, their bravery supplemented by their curiosity, were threatened by storms while at sea, new and deadly diseases, unfamiliar animals, and new cultures. Add plane crashes and paramilitary groups to this list and not much has changed.

In a great New York Times piece titled 'Dying for Discovery', Richard Conniff recounts several stories of naturalists, ecologists and conservation biologists killed while pursuing their passion for discovery. But just how many field biologists have died while working to understand life's secrets? This is an interesting question, and begs the further question, are they adequately memorialized?

Gary Polis (1946-2000) desert ecologist, drown with four other biologists during a storm in the Sea of Cortez.

In an attempt to tell the stories of the fallen naturalists, Conniff hosts an interactive list, called the Wall of the Dead, which lists all biologists killed in the field and that he has a record of. People are able to add names, and I've visited this list several times over the past month and it has grown substantially. I've known a few field biologists that have died -and added one to the list, and know several that survived near-death experiences, and this list is a great and important monument to their memories.

Further studies of the decline effect find decline of the decline effect

2011-02-07T10:22:00.000-08:00

“The Truth Wears Off: Is something wrong with the scientific method?”

The Decline Effect explored in an article by Jonah Lehrer in the New Yorker refers to a temporal decline in the size of an observed effect: for example, the therapeutic value of antidepressants appears to have declined threefold since the original trials. Based on the cases presented, this effect is not limited to medical and psychological studies. One example in evolutionary biology is the relationship between physical symmetry and female choice: initial studies consistently found strong selection for symmetry in mates by females, but as time passed, the evidence grew increasingly smaller.

This may be a result of selective reporting – scientists focus on results that are novel and interesting, even if they are in fact simply statistical outliers, or worse, the result of unconscious human bias. This sentiment is troubling; humans – scientists or not– are proficient pattern finders, but our subconscious (or conscious) beliefs influence what we search for. Lehrer argues that replication – the process of carrying out additional, comparable but independent studies – isn’t an effective part of the scientific method. After all, if study results are biased, and replications don’t agree, how can we know what to trust?

I don’t disagree with most of the article’s points: that scientists can produce biased results, PhD not withstanding, that more effort and time should be invested in data collection and experimental methodology, that the focus on 5% statistical significance is problematic. For one, it’s not clear from the article how prevalent the decline effect is. However, I wonder whether Lehrer, similar to the scientists he’s reporting on, has selected specific, interesting data points, while ignoring the general trend of the research. In 2001, Jennions and Moller published evidence of a small negative trend in effect size over time for 200+ studies, however, they suggest this is due to a bias toward high statistical significance, which requires either large effect sizes (the early studies published), or small effect sizes in combination with large sample sizes (a scenario which takes more time).

Even if the decline effect is rampant, does it represent a failure of replicability? Lehrer states that replication is flawed because “it appears that nature often gives us different answers”. As ecologists though, we know that nature doesn’t give different answers, we ask it different questions (or the same question in different contexts). Ecology is complex and context-dependent, and replication is about investigating the general role of a mechanism that may have been studied only in a specific system, organism, or process. Additional studies will likely produce slightly or greatly different results, and optimally a comprehensive understanding of the effect results. The real danger is that scientists, the media, and journals over-emphasize the significance of initial, novel results, which haven’t (and may never be) replicated.

Is there something wrong with the scientific method (which is curiously never defined in the article)? The decline effect hardly seems like evidence that we’re all wasting our time as scientists – for one, the fact that “unfashionable” results are still publishable suggests that replicability is doing what it’s supposed to, that is, correct for unusual outcomes and produce something close to the average effect size. True, scientists are not infallible, but the strength of the scientific process today is that it doesn’t operate on the individual level: it relies on a scientific community made of peers, reviewers, editors, and co-authors, and hopefully this encourages greater accuracy in our conclusions.

Carinval #32 and still going strong

2011-02-01T09:44:00.000-08:00

Want to know what people are talking about? The 32nd Carnival of Evolution is online, hosted by Denim and Tweed. Check it out, pass it along.

Trend in ecology, 2010

2011-01-25T09:35:00.002-08:00

Sciences are always in a state of ebb and flow (sorry), and topics of study fall in or out of fashion in response to paradigms shifts, methodological advances, and to support necessary ecological applications.
For the sake of curiosity, I've compiled the top keywords from ecology publications in 2010. Obviously there are many covariates, but it should come as no surprise that the top words were "biodiversity" (667 times), "climate change" (293), and "conservation" (274); other popular keywords were "evolution" (277), "population (ecology)" (273), and the rather vague "patterns" (196).

(click image for larger view)

The evolutionary story of ecosystem function

2011-01-20T18:13:00.000-08:00

Twenty years of research has repeatedly shown that communities with greater diversity result higher functioning -namely greater production of biomass. One of the major mechanisms producing this relationship is that different species use differing resources, such that their complementary use of resources uses the total resource pool more thoroughly, thus converting more resources into biomass. Resource preference is the product of evolution and how organisms have adapted to using various resources can influence the strength of the diversity-function.

In a recent paper in Nature, Dominique Gravel and colleagues test how the evolution of specialization versus general resource use affect the strength of the diversity-function relationship. They use bacteria strains that have undergone evolution on diverse resources (generalist) versus on a singular resource (specialist). The resources in their case are different carbon substrates.

Assemblages of generalists were able to use many available resources and generally had greater productivity than specialist assemblages. Generalists also show an increasing relationship between diversity and productivity, because no generalist used all resources and they still showed some preferences. Combining multiple such generalists meant that more of the total resource pool was consumed. Specialists also resulted in the positive relationship, but a much steeper one. Because specialist use many fewer carbon substrates, additional specialists meant that new resources were tapped into. Thus increasing specialist diversity resulted in more new resources being consumed than with the generalist species.

While these results are logical, they are important for two reasons. First is that the strength of the relationship between diversity and function is mechanistically determined by the resource use efficiency of individual strains, and how many of the total substrates they can use. The mechanisms producing different relationships in previous experiments were hypothesized after the results analyzed, as opposed to being predicted. Second, recent work has shown that evolutionary history seems to be a better explanation of community function than the number of species. These results show how the history of evolution can have important consequences for function.

Gravel, D., Bell, T., Barbera, C., Bouvier, T., Pommier, T., Venail, P., & Mouquet, N. (2010). Experimental niche evolution alters the strength of the diversity–productivity relationship Nature, 469 (7328), 89-92 DOI: 10.1038/nature09592

Who is a scientist, I am a scientist: the bees of Blackawton

2011-01-11T18:37:00.000-08:00

In discussions of the larger societal implications of scientific findings, the question of who is a scientist is frequently asked. I've talked with with creationists who invoke the authority of someone who has a PhD in a scientific discipline and happens to share their belief of supernatural origins, as a scientific authority. Does the fact that I have a PhD in ecology and evolutionary biology make me scientist or is being scientist something more?

This is an important question. It goes to the core of whose authority we believe for public discussion of such issues as climate change, evolution, risks of vaccines, and so on. Regardless of how we define 'scientist', a scientist participates in science by publishing peer-reviewed research articles in scientific publications. This notion of who is a scientist has been enjoyably stretched by the publication of a paper in Biology Letters by a group of elementary school children from Blackawton, UK. In consultation with a academic scientist and under the supervision of teachers, 25 8-10 year olds devised and carried out an experiment on bee visual perception and behavior, and wrote up their results into a publishable manuscript.

The students trained bees by offering them nectar rewards in different color containers. They then allowed these trained bees to forage in multicolored arenas and they conclusively show that the bees unambiguously select the colored containers they were trained on. Bess learn and adapt their behavior based on previous experience.

Publishing a paper by a group of children may sound like a gimmick, but the study is very interesting. The commentary from the journal says it best: "The children's findings show that bees are able to alter their foraging behaviour based on previously learned colours and pattern cues in a complex scene consisting of a (local) pattern within a larger (global) pattern . As there has been little testing of bees learning colour patterns at small and large scales, the results can add considerably to our understanding of insect behaviour."

The paper is extremely enjoyable to read and will have you chuckling to yourself. Sincerity pours from the words and I was left wondering if I could have reasoned so well at that age. The children develop hypotheses using information available to them, such as watching Dave Letterman's 'Stupid Dog Tricks'. Reading this article made me realize why I love being scientist. The students note that "This experiment is important, because, as far as we know, no one in history (including adults) has done this experiment before" and because they were given the opportunity to carryout this study they "also discovered that science is cool and fun because you get to do stuff that no one has ever done before". Too true. I could not have said it better myself.

Being scientist can mean a lot of things, it can mean knowledge (which the Latin origin, Scientia means), it can mean training and acquired skills, but at its core, being a scientist means conducting research, testing hypotheses and writing publications that are deemed acceptable by other scientists. Therefore the children of Blackawton are scientists, I am a scientist.

Blackawton, P., Airzee, S., Allen, A., Baker, S., Berrow, A., Blair, C., Churchill, M., Coles, J., Cumming, R., Fraquelli, L., Hackford, C., Hinton Mellor, A., Hutchcroft, M., Ireland, B., Jewsbury, D., Littlejohns, A., Littlejohns, G., Lotto, M., McKeown, J., O'Toole, A., Richards, H., Robbins-Davey, L., Roblyn, S., Rodwell-Lynn, H., Schenck, D., Springer, J., Wishy, A., Rodwell-Lynn, T., Strudwick, D., & Lotto, R. (2010). Blackawton bees Biology Letters DOI: 10.1098/rsbl.2010.1056

The carnival is on!

2011-01-06T05:14:00.000-08:00

Check out the 31st edition of the Carnival of Evolution, hosted by The Dispersal of Darwin. Check it out, a fun read for sure!

Science 2.0 - science comes of age on the Internet

2011-01-04T08:48:00.000-08:00

by Marc Cadotte, Nicholas Mirotchnick and Caroline Tucker

The Internet is not just for lolcats and porn anymore, scientists have begun using it in constructive ways. The past few weeks’ controversy about the ability (or lack thereof) of bacteria to incorporate arsenic exemplifies how the relationship between science and the Internet is changing. If you’ve missed the debate over the recent Science paper, researchers funded by NASA’s exobiology/evolutionary biology program published experimental results suggesting that a Halomonas species could incorporate arsenic into its DNA in the absence of available phosphorus. This paper received extensive attention in the mainstream media, but also vocal criticism, which was expressed primarily through postings and comments on scientific blogs. Until recently, for scientific communication the Internet has functioned primarily as an electronic source of published journal articles. Earlier attempts to take advantage of the Internet’s potential (immediacy, accessibility, and ability to connect individuals, organizations, and ideas) in scientific discourse have been mixed (e.g. Nature Precedings versus PLoS ONE). The use of blogs as a forum for scientific debate suggests that this is changing: posters tended to be active scientists and the comments were similarly knowledgeable. In contrast to this online approach, the authors of the Science paper stated that they would only respond to peer-reviewed critiques and would not engage in discussions on the blogosphere.

The story of the arsenic-utilizing bacteria highlights an emergent tension in the transition to internet-based scientific discourse. Traditional communication in science has been primarily unidirectional, from the authors of a study to the readership of a journal. Any discourse transpired on the pages of a journal, regulated by editorial and peer review. This gatekeeping meant that this discourse was technically sound and kept personal grudges and tangential discussions to a minimum. This also meant, however, that only a few voices were heard, the discussion was slow (occurring over months) and only happened for one back and forth (journals will not devote precious page space to on-going discussions and debates).

This method of discourse is changing. Journals have experimented with online discussion or commenting features on their websites. Methods in Ecology and Evolution, for example, has a correspondence page with discussion threads for each paper they publish, and PloS ONE allows for comments to be posted to every paper they publish. While, in concept, these are positive developments for scientific communication, commenting features are seldom, if ever, used. The main obstacle to their success is that they are only available on the publishers’ websites, but scientists access articles in many different ways, from database searches to library links. Few scientists actually go to individual journal websites to access papers. This is not to say that there are not discussions about scientific papers occurring online. As highlighted by the arsenic bacterial episode, blogs are an important avenue for discussing and disseminating new ideas in science. Blogs may not, however, actually foster conversations very well. One person or a few people usually run them and there is little discussion among blogs (a comment on a blog post at blog X will not be part of the discussion of the same story at blog Y). Rather, the greatest potential to foster discourse is through virtual networks where people are linked together either through friendships or professional self-identification (e.g., as fisheries biologists), with Google Reader being a particularly powerful communication tool.

It’s exciting to think about what the future of science will look like, given the changes that we’ve already started to see. The major upside of new channels of communication is that they give us the potential to quickly reach thousands of readers, instead of the handful that usually read any given journal article. They also let us communicate in both directions, and in real time. The pitfall, of course, is that they’re free-for-alls; anyone can blog about science.

But here’s what’s unexpected: these free-for-alls have been amazingly reliable at filtering out the bad and promoting the good. Inaccuracies are pulled from Wikipedia faster than anyone had predicted, the social news site Reddit is “astonishingly” altruistic, with users eliminating offensive or erroneous comments from the site and promoting other users’ questions and problems, and the reputations of blogs are shattered if their content becomes unreliable. Social networking has revolutionized the way we consume news, with sites like Facebook and Twitter launching the best articles into viral webspace. The open-access world has evolved self-regulating mechanisms that work surprisingly well so far and if these media are to continue to grow, we will have to ensure that these mechanisms remain built-in.

Seems like an easy task, right? Apparently not. For some reason, academics are slow and conservative when it comes to adopting new media. A letter to Nature two weeks ago scolded scientists for not contributing their share to Wikipedia pages. Various facebooks for academics, like Mendeley and ResearchGATE have emerged, but last week, another Nature article complained that researchers aren’t jumping on the bandwagon. These sites are potential collaborative goldmines, but we seem to be incapable mastering what tweens can do with two thumbs.

It’s not so hard to imagine a world where anyone with a broadband connection can contribute creative ideas to science, the good ideas get automatically filtered to the top and the information is all free to anyone. In this world, children count ants (or bees!) in their backyards and upload their data to global networks. Revolutionary discoveries are published instantly on blogs and thousands of scientists get to decide if they’re valid. Every gene ever sequenced and every tree height ever measured can be readily downloaded in an Excel (or OpenOffice) spreadsheet. In this world, the report on our little arsenophilic friends might never have been published in Science, because instead of being reviewed by two referees, the thousands of readers on the blogosphere would have filtered it out, if was in fact porous.

Academics should be the first, not the last, to adopt new communication tools. We are no longer limited by the postal service, email or PDFs; the web has gone 2.0 and we should follow suit. So go forth, young researchers, and blog, edit and share. And then go tweet about it all so your eight year-old kid knows how hip you are.

Biodiversity and ecosystem functioning – without fungi?

2010-12-10T06:19:00.000-08:00

Different subfields of ecology have a propensity to remain remarkably isolated – researchers in aquatic systems independently develop hypotheses that already exist in some form in other systems, and vice versa. Population ecology and community ecology, despite their obvious relevance to each other, are rarely integrated. There is a tendency – resulting from limits on our time, experience, and possibly imagination – to stay within whatever box we’ve defined for ourselves.

Historically, it seems that biodiversity and ecosystem functioning has lost sight of the progress made in classical ecology in understanding the mechanisms behind species coexistence (and all the functional implications that follow). Studies of ecosystem functioning often vaguely reference concepts such as “niche partitioning”, which would hardly be explicit enough for most papers on coexistence. Fortunately, there are periodically attempts to unifying ecological knowledge.

One of the most important contributions to understanding coexistence is Chesson’s (2000) framework of equalizing and stabilizing effects. Unlike previous approaches to species interactions, which tended to reference these vaguely-defined “niche differences”, Chesson proposed that species interactions depended on both fitness differences (differences in absolute growth rates after niche differences are controlled for) and niche differences (ecological differences between species which cause intraspecific competition to exceed interspecific competition). He also suggested rigorous methods to quantify these concepts. This framework has been applied both to the obvious questions of species coexistence and diversity maintenance, as well as predator-prey relationships (2008) and the phylogenetic structure of communities (2010).

In a recent paper, Ian Carroll et al. apply this framework to the search for the mechanisms behind biodiversity and ecosystem functioning. They point out that the questions in studies of ecosystem functioning are directly analogous to Chesson’s concepts – selection effects result from fitness or competitive differences between species, while complementarity relates to the partitioning of resources, or niche differences between species. The added benefit is that Chesson has provided clear definitions for these concepts.

While this may not be world-altering, it’s encouraging. Anytime different areas of ecology intersect, both benefit. Of course there are difficulties – no doubt the question of how to measure niche differences and fitness differences will be contentious (as attempts to translate ecological theory into ecological methodology often are) - but the possibility that a few general ecological concepts explain diverse observations is worth pursuing.

Biodiversity and ecosystem functioning - only with fungi

2010-11-18T17:44:00.001-08:00

Once again scientists have come to an age-old conclusion: fungus is behind all of life’s great mysteries. It's responsible for curing strep throat, delicious veggie burgers, that unique musk emanating from your gym bag, the colour-morphing walls at last night’s party and now, biodiversity and ecosystem functioning.

The world of biodiversity and ecosystem functioning (BEF), like many other high-profile disciplines of science, has often been bogged down by controversy. In such situations, we often spend a disproportionate amount of time focusing on the controversy instead of actually advancing the science itself (sound familiar?)

There have been several posts about BEF on this blog in the last few months, but briefly and oversimplified, here's how it works. Ecosystem functions are things like productivity, nutrient cycling and decomposition. Ecosystems that contain many species produce higher levels of these functions than monocultures do. The controversy here surrounds the cause of this phenomenon. In the 1990s, researchers originally disagreed over whether the relationship they observed was due to complementarity (different species partitioning resources) or selection effects (the higher chance of a really productive species being included in a community with many species). The question was largely settled a few years ago; selection effects do exist, but most of the relationship is driven by complementarity. Nonetheless, many biologists who are only tangentially familiar with this area of research are unaware of the consensus and continue to believe that the issue remains unresolved. Some still dismiss the whole field of BEF because of selection effects. I guess people just love a controversy.

The result of all this is that the ecologists studying these relationships have had to spend an undue amount of time parsing their results into selection and complementarity and discussing the two phenomena. They have even come to refer to these as the “mechanisms” behind BEF. And this is where we start to have a problem. Selection and complementarity are not mechanisms - they are symptoms of mechanisms. They do not tell us what is actually causing the positive effect that biodiversity has on ecosystem functioning, only what the shape of the relationship is. In fact, very few studies have actually looked for true mechanisms that explain the effects that we have repeatedly observed.

But this week I read a new paper in Ecology Letters that actually did find a mechanism, and it wasn’t one that we expected. John Maron and his coauthors at the universities of Montana and British Columbia found that belowground fungi were causing plant productivity to increase with diversity.

In an impressively complete experiment, Maron et al. put together a classic BEF setup of many plots containing varying levels of plant diversity and then measured plant biomass. But this time they added a twist; they applied fungicide to the soil in some of these plots. The result was that in the absence of fungi, the common BEF relationship disappeared. The low diversity plots became much more productive, while productivity at high diversity only increased slightly. The authors explained their results by the fact that fungi can be both species-specific and density-dependent, so as plant diversity increases, the fungi’s negative impact on plant productivity diminishes. And for good measure, they of course also ruled out a significant selection effect in their results.

So what does this all mean? Well for one thing, it means that we now have at least one good mechanistic explanation for that black box that we’ve been calling “complementarity” for years. But perhaps more importantly, it means that the link between biodiversity and ecosystem functioning is now more real than ever. If plant species go extinct, the remaining ones will be more susceptible to fungal pathogens and productivity will decline. So let’s try to not let that happen, k?

Maron, J. L., Marler, M., Klironomos, J. N. and Cleveland, C. C. , Soil fungal pathogens and the relationship between plant diversity and productivity. Ecology Letters, DOI: 10.1111/j.1461-0248.2010.01547.x

The effects of forest fragmentation after 30 years

2010-11-05T10:24:00.000-07:00

Large-scale alteration of nature landscapes has had profound implications for biological diversity. The single biggest contributor to the current extinction crisis is the wholesale destruction of habitats. As habitats are destroyed, formerly contiguous landscapes become fragmented into smaller patches. But what exactly the effects of fragmentation are, independent of habitat destruction, is not always so clear (e.g., Simberloff 2000. What do we really know about fragmentation? Texas Journal of Science 52: S5-S22).

The biological dynamics of forest fragments project (BDFFP) in the Amazon, was started in 1979 and created 11 tropical forest patches ranging from 1 to 100 ha in size. The dynamics of these fragments have been consistently monitored and compared to plots in intact forest. This experiment represents the world's largest, longest-running fragmentation experiment and has told us more about fragmentation then any other study system. In a recent publication by William Laurance and many colleagues involved in this project, they summarize 30 years of data and show how fragmentation affects ecological patterns and processes.
Fragments turn out to be very dynamic and defined by change, compared to interior plots. They have higher tree mortality and are much more susceptible to weather events such as storms or droughts. The effects are especially pronounced at the edges of these fragments. The edge community face high mortality but also have higher tree density. Faunal communities in fragments and especially near edges are depauperate.

One interesting aspect highlighted by this 30 years of research is that the edge effects are strongly influenced by what is happening around the fragments. The fragment edge effects are sensitive to the composition of the inter-patch matrix, giving managers the opportunity to influence fragment diversity and health by managing the matrix in ways that support fragments. Because of over 30 years of perseverance of the researchers involved, this experiment give scientists, managers and policy-makers information to help manage an increasingly fragmented world and to find ways to reduce to negative impacts of habitat destruction.

Laurance, W., Camargo, J., Luizão, R., Laurance, S., Pimm, S., Bruna, E., Stouffer, P., Bruce Williamson, G., Benítez-Malvido, J., & Vasconcelos, H. (2010). The fate of Amazonian forest fragments: A 32-year investigation Biological Conservation DOI: 10.1016/j.biocon.2010.09.021

Carnival of Evolution!

2010-11-03T12:49:00.000-07:00

The 29th installment of Carnival of Evolution is here, hosted by Byte Size Biology. Check out recent evolutionary musings and rants from around the blogoshpere.

Grassland diversity increases stability across multiple functions

2010-10-17T18:03:00.000-07:00

As ecological systems are altered with cascading changes in diversity, the oft-asked question is: does diversity matter for ecosystem function? This question has been tested a multitude of times, with the results often supporting the idea that more diverse assemblages provide greater functioning (such as productivity, nutrient cycling, supporting greater pollinator abundance, etc.). Besides greater functioning, scientists have hypothesized that more diverse systems are inherently more stable. That is, the functions communities provide remain more constant over time compared with less diverse systems, which may be less reliable.

While the relationship between diversity and stability has been tested for some functions, Proulx and colleagues examined the stability of 42 variables over 7 years across 82 experimental plots planted with either 1, 2, 4, 8, 16 or 60 plant species in Jena, Germany. They examined patterns of variation (and covariation) in the functions and found that many show lower variation over time in plots with more plant species. Greater stability was found at many different trophic levels including plant biomass production, the abundance and diversity of invertebrates and the abundance of parasitic wasps -which indicate more complex food webs. They also found greater stability in gas flux, such as carbon dioxide. Despite the greater stability in these measures of above-ground functions, below ground processes, such as earthworm abundance and soil nutrients, were not less variable in high diversity plots.

How ecosystems function is of great concern; these results show that more diverse plant communities function more stably and reliably than less diverse ones. The next step for this type of research should be to address what kind of diversity matters. A greater number of species means more different kinds of species, with differing traits and functions. What aspect of such functional differences determine stability of ecosystem function?

This is an exciting paper that continues to highlight the need to understand how community diversity drives ecosystem function.

Proulx, R., Wirth, C., Voigt, W., Weigelt, A., Roscher, C., Attinger, S., Baade, J., Barnard, R., Buchmann, N., Buscot, F., Eisenhauer, N., Fischer, M., Gleixner, G., Halle, S., Hildebrandt, A., Kowalski, E., Kuu, A., Lange, M., Milcu, A., Niklaus, P., Oelmann, Y., Rosenkranz, S., Sabais, A., Scherber, C., Scherer-Lorenzen, M., Scheu, S., Schulze, E., Schumacher, J., Schwichtenberg, G., Soussana, J., Temperton, V., Weisser, W., Wilcke, W., & Schmid, B. (2010). Diversity Promotes Temporal Stability across Levels of Ecosystem Organization in Experimental Grasslands PLoS ONE, 5 (10) DOI: 10.1371/journal.pone.0013382

Protecting biodiversity one task at a time: have your say

2010-09-04T04:02:00.000-07:00

The fact that the Earth is in the midst of a biodiversity crisis has been repeatedly acknowledged by world governments. The greatest pronouncement was is 2002 with the '2010 Biodiversity Target' where many of the largest economies signed a pledge to halt biodiversity loss by 2010. Yet it is now 2010 and species are continuing to go extinct and habitats are continuing to be destroyed or degraded. But it shouldn't be a surprise that non-binding governmental proclamations fail to produce substantial results. Yet the reality is that we need to do something, inaction only worsens the legacy of biological deficit for future generations.

Maybe the best way forward is not more international governmental summits, but rather focusing on small scale, achievable short term goal. Guillaume Chapron started the Biodiversity 100 campaign, hosted by the Guardian (see story here), which seeks out public and professional input into the 100 immediate and achievable projects or ideas that will help protect biodiversity. The idea is to be able to go to governments and international agencies with this list and get them to make specific pledges to carry out these tasks.

There is till time to participate! If you have an idea of an action to protect biodiversity, fill out the web form. There are already a plethora of great suggestions, from protecting specific habitats to stemming population growth. This list is important because it includes the voices of the international public citizenry and that of scientists. More than that though, there will be a concrete list of tasks (ranging from very local to very global) that citizen groups can use to sustain pressure on governments.

Enhanced biodiversity-ecosystem function relationships in polluted systems

2010-07-27T07:03:00.000-07:00

*note: this text was adapted from an Editor's Choice I wrote for the Journal of Applied Ecology.

In this era of species loss and habitat degradation, understanding the link between biodiversity and functioning of species assemblages is a critically important area of research. Two decades of research has shown that communities with more species or functional types results in higher levels of ecosystem functioning, such as nutrient processing rates, carbon sequestration and productivity, among others. This research has typically used controlled experiments that standardize environmental influences and manipulate species diversity. However, a number of people have hypothesized that biodiversity may be even more important for the maintenance of ecosystem functioning during times of environmental stress or change rather than under stable, controlled conditions. It is during these times of environmental change that preserving ecological function is most important, as changes in function can have cascading effects on other trophic levels, compounding environmental stress. Therefore, explicitly testing how biodiversity affects function under environmental stress can help to inform management decisions.

Image from Wikimedia commons

In a recent paper in the Journal of Applied Ecology, Li and colleagues examine how algal biodiversity influences productivity in microcosms with differing cadmium concentrations. Cadmium (Cd) is a heavy metal used in a number of products and industrial processes, but it is toxic and Cd pollution is a concern for human populations and biological systems, especially aquatic communities. This is especially true in nations currently undergoing massive industrial expansion. In response to concerns about Cd pollution effects on aquatic productivity, Li et al. used algal assemblages from single species monocultures to eight species polycultures grown under a Cd-free control and two concentrations of Cd, and measured algal biomass.

Their results revealed that there was only a weak biodiversity-biomass relationship in the Cd-free teatment, which the authors ascribed to negative interactions offsetting positive niche partitioning. In particular, those species that were most productive in their monocultures were the most suppressed in polycultures. However, in microcosms with Cd present there were positive relationships between diversity and biomass. They attribute this to a reduction in the strength of competitive interactions and the opportunity for highly productive species to persist in the communities.

While a plethora of experiments generally find increased ecosystem function with greater diversity, Li et al.’s research indicates that the effect of biodiversity on function may be even more important in polluted systems. If this result can be duplicated in other systems, then this gives added pressure for management strategies to maintain maximal diversity as insurance against an uncertain future.

Li, J., Duan, H., Li, S., Kuang, J., Zeng, Y., & Shu, W. (2010). Cadmium pollution triggers a positive biodiversity-productivity relationship: evidence from a laboratory microcosm experiment Journal of Applied Ecology, 47 (4), 890-898 DOI: 10.1111/j.1365-2664.2010.01818.x

Reinterpreting phylogenetic patterns in communities

2010-07-22T08:01:00.000-07:00

Examining the phylogenetic structure of a community in order to understand patterns of community assembly has become an increasingly popular approach. A quick web search of “community”, “phylogenetics”, and “ecology” finds several hundred papers, most written in the last ten years.

Eco-phylogeneticists examine how patterns of evolutionary relatedness within communities may reflect the processes structuring those communities. In particular, a commonly tested hypothesis is the competition-relatedness hypothesis, which suggests that more closely-related species having more similar niches and therefore stronger competitive interactions, making coexistence between them less likely. As a result, if competition is important, communities may exhibit phylogenetic overdispersion, with species being less related on average than if drawn randomly from the regional species pool. The contrasting pattern, phylogenetic clustering, where species tend to be more closely related than expected, is often interpreted as being the result of strong environmental filtering, such that only a closely related group of species, best adapted to that environment, surviving in the community.

Evidence for the competition-relatedness hypothesis has been mixed, and since most tests of this hypothesis focus on patterns in observed data, conclusions about the underlying mechanism driving community phylogenetic patterns are rarely testable, and yet widely made.

In Mayfield and Levine (2010, Ecology Letters), the authors critique the current ecological justification for the competition-relatedness hypothesis, noting that it does not agree with a more current view of the processes driving species coexistence. As established by Chesson (2000, Annual Review of Ecology and Systematics), coexistence can involve both stabilizing forces (niche differences between species), and equalizing forces (fitness differences between species). In a simplistic example, plants using different soil types (niche differences) may coexist, while plants with similar high growth rates may exclude those species with lower growth rates (fitness differences). The final community should reflect the interplay of both these processes.

The implication of this view of species coexistence is that there is no preconceived phylogenetic pattern which should reflect competition: if species with the highest heights are competitively superior and exclude other species (coexistence driven by fitness differences), and height is a phylogenetically conserved trait, the community will appear to be phylogenetically clustered. Traditionally, a clustered pattern would not be considered to indicate the effects of competition. In fact, Mayfield and Levine show that the expected phylogenetic pattern depends entirely on whether niche and/or fitness differences are important and/or related to phylogenetic distance.

This suggest that conclusions in past studies may need to be reinterpreted. It also adds to the list of assumptions about evolutionary relatedness and ecological function which need to be tested: for example, how do niche and fitness differences tend to change through time? Do they tend to be conserved among closely related species? Does one or the other tend to dominate as a driver of coexistence in different systems? If nothing else, we need to be careful about making generalizations which don’t account for the differing evolutionary history, geographical location, and ecological setting that communities experience, when interpreting observed patterns in those communities.

Organic farming and natural enemy evenness

2010-07-07T07:31:00.000-07:00

The basic reality of agricultural activity is that it reduces biological diversity, and these reductions in diversity potentially impact ecosystem services. But do some agricultural practices impact these services less than others? In a recent paper in Nature by David Crowder and colleagues, the question of how organic versus conventional farming affects predator and herbivore pathogen diversity and how this cascades to pest suppression. They show through a meta-analysis, that organic farms tend to support greater natural enemy evenness, and they hypothesize that greater evenness of enemies should better control pest populations, resulting in larger, more productive plants.

Picture from wikipedia

This result in itself is interesting, but they also carried out an elegant enclosure experiment where they manipulate the evenness of insect predators and pathogens and measure potato plant size. They found that even communities had the lowest herbivore densities and saw the greatest increases in plant biomass. Conversely, very uneven communities, typical of conventional farms, had the largest pest populations resulting in lower plant biomass accumulation.

While, multiple farming strategies are needed for adequate agricultural production, there are strong arguments for organic farms to be a important part of agricultural practice. These results show that organic farms have cascading effects on pest predators and pathogens and show that enemy evenness, as opposed to richness, has important ecosystem service consequences. To quote myself, evenness is a critical component of biodiversity, and much research has emphasized species richness, maybe at the detriment of studying evenness.

Crowder, D., Northfield, T., Strand, M., & Snyder, W. (2010). Organic agriculture promotes evenness and natural pest control Nature, 466 (7302), 109-112 DOI: 10.1038/nature09183

New Carnival of Evolution

2010-07-06T18:20:00.000-07:00

Check out the July edition of the Carnival of Evolution written by Hannah Waters at Culturing Science.

Happy Year of Biodiversity

2010-06-12T07:15:00.000-07:00

It’s ironic that during the International Year of Biodiversity, the US is experiencing an environmental disaster on a massive scale. Unfortunately, this disaster is just another failure in environmental protection, part of a long series of failures which seem to characterize this Year of Biodiversity. Even as the political will behind the 2010 biodiversity targets seems to have waned (and most indicators suggest that declines in diversity are unchecked), evidence continues to mount for the functional value of biological diversity.

This week’s issue of Nature features a couple of pieces focusing on biodiversity through a political or economic lens. Although the economic benefits and services provided by species-level diversity has been well illustrated, in “Population diversity and the portfolio effect in an exploited species”, Schindler et al. (Nature, 465, 609-612) new evidence that at even finer divisions than the species, diversity plays an important role. In this case, they find that genetic diversity at the population level is an additional and significant contributor to ecosystem stability. Schindler et al. examine the effects of hundreds of locally-adapted populations of sockeye salmon on the valuable salmon fishery in the Bristol Bay area of Alaska. They suggest that the portfolio effect (or the robustness of biodiversity to variable conditions – like a diverse financial portfolio) can function at the population level as well as the species level. High levels of intra-specific diversity can produce temporal variation among populations in response to environmental variability, resulting in catches that are more stable year-to-year, and making fishery closures less likely, a clear economic benefit.

Populations are declining at an even faster rate than species themselves: the more we understand the importance of conserving diversity at multiple biological scales (ecosystem, species, population, even the individual?), the more complicated and onerous the task of conserving diversity becomes.

In the same issue of Nature is an editorial on the possibility of an IPCC-like panel for biodiversity. At this very moment (give or take a few time zones), government representatives from all over the world are deciding whether or not to create this panel. So far, they have a catchy name for it, the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services (IPBES), which hopefully hasn’t been written in stone. But they also have a strong recognition of the inextricable links between biodiversity, ecosystem services and human wellbeing – links that are highlighted in the Schindler et al. article. Furthermore, an explicit goal of IPBES is to address the currently tangled state of biodiversity organizations, conventions and programs by forming a unified front of sound biodiversity policy and science. The Convention on Biological Diversity had set a target of halting biodiversity loss by 2010 and we have failed spectacularly. Is IPBES the solution?

Wanted: an IPCC for biodiversity. Nature, 465, 525-525

Schindler, D.E., Hilborn, R., Chasco, B., Boatright, C.P., Quinn, T.P., Rogers, L.A. & Webster, M.S. Population diversity and the portfolio effect in an exploited species. Nature, 465, 609-612

By Nick Mirotchnick and Caroline Tucker

Another reason why a new publishing model is needed...

2010-06-09T12:26:00.000-07:00

The finances and ethics of scientific publishing are complex, and there is an inherent tension between commercial publishers and academics and their institutions. On the one hand, we as scientists are (most often) using public money to carry out research, usually in the public interest, and then we typically publish in for-profit journals that restrict public access to our publications. Authors seldom see any of the financial return from publisher profits. On the other hand, publishers provide a level of distribution and visibility for our work, which individual authors could not match. In previous posts I have discussed Open Access publications, but there is another reason to consider other publication models. Recently Nature Publishing Group notified the University of California system of an impending 400% increase in the cost for their publications. The UC administration has responded with an announced plan to boycott NPG publications. The announcement rightly points out a 400% increase is not feasible given the current plight of library budgets, especially in California, and that scientists in the UC system disproportionately contribute to publishing, reviewing and editing NPG publications and thus are the engine for NPG profits. (See a nice story about the boycott in The Chronicle of Higher Education)

This is just the latest symptom of the growing tension between publishing and academia, and is a stark reminder that other publishing models need to actively supported. Perhaps the UC system could invest in open access publishers in lieu of NPGs outrageous costs? Something has to give, and perhaps the UC boycott will remind libraries that they hold the purse strings and could be the greatest driving force for change.

Experimental test of Darwin's naturalization hypothesis

2010-06-01T17:43:00.000-07:00

Among the numerous and still informative ecological predictions made by Darwin, one posits that when species are introduced into regions where they were not formerly found, the most successful tend to not have close relatives already occupying the region. This is known as Darwin's Naturalization Hypothesis, and his logic was that among close relatives, where ecological requirements should be most similar, the struggle for existence is most severe. Thus the modern formulation is that invader success is influenced by the amount of time since two species shared a common ancestor (usually called phylogenetic distance). Tests of this hypothesis have been primarily done on large species inventories, with results from different studies either supporting or refuting it. In a new study by Lin Jiang and colleagues published in the American Naturalist, they cleverly use bacteria with known relatedness to test this hypothesis.

They used four species of bacteria: Bacillus pumilus, B. cereus, Frigoribacterium sp. and Serratia marcescens as residents in every possible 1, 2, 3 and 4-species communities and invaded them with a subspecies of S. marcescens. What they found was that the invader density was highly significantly related to phylogenetic distance, so that the invader reached its greatest density when communities contained only distantly-related species.

Though these types of laboratory experiments are simplistic (I too use these systems), they offer insights into particular mechanisms, which may otherwise be difficult to detect in noisier systems.

Jiang, L., Tan, J., & Pu, Z. (2010). An Experimental Test of Darwin’s Naturalization Hypothesis The American Naturalist, 175 (4), 415-423 DOI: 10.1086/650720

The successful launch of MEE

2010-05-25T13:55:00.000-07:00

Usually, I view the release of a new journal with some skepticism. There are so many journals and it feels like academics are over-parsing fields, isolating researchers that should be communicating. However, sometimes a journal comes along and it is obvious that there is a need and the community responds to its arrival. Such is the case with the British Ecological Society's newest journal, Methods in Ecology and Evolution, started by Rob Freckleton. The idea that a journal would be dedicated to methods papers is a great idea. This era of ecology and evolution is one that is defined by rapid advances in experimental, technological and computational tools and keeping track of these advances is difficult. Having a single journal should make finding such papers easier, but more importantly provides a home for methodological and computational ecologists and evolutionary biologists, which will hopefully spur greater communication and interaction, fostering more rapid development of tools.

Two issues have been published and they have been populated by good, entertaining articles. I especially enjoyed the one by Bob O'Hara and Johan Kotze on why you shouldn't log transform count data. As a researcher, I've done this (instead of using a GLM with proper distribution) and as an editor, I've allowed this, but it has always felt wrong somehow, and this shows that it is.

The early success of the journal is not just the product of the good papers it has already published, but also because of the savvy use of electronic communication. They Tweet on Twitter, link fans through Facebook, blog about recent advances in methods from other journals and post podcast and videocast interviews with authors. These casts give readers access to authors' own explanations of how their methods can be used.

I am excited about this new journal and hope it has a great impact on the publication of methodological papers.

Picante's coming out party

2010-05-11T06:20:00.000-07:00

This past decade has seen a rapid expansion of the use of evolutionary phylogenies in ecological studies. This expansion is largely due to the increased availability of phylogenies, but has resulted in new types of hypotheses and statistics aimed to test the phylogenetic patterns underpinning ecological communities. The main computational tool used has been phylocom, created by Cam Webb, David Ackerly and Steve Kembel, which has its own binaries to be installed on one’s computer. However, a new R package, picante has been created by Steve Kembel and colleagues which runs many of the same routines as in phylocom, but in the R framework, allowing one to tie these analyses in better with other, non-phylogenetic tests. Picante also has a number of features and tests not found in phylocom, including tests of phylobetadiversity and phylogenetic signal using Blomberg’s K.

Thanks Steve for all your hard work and for making these tests available to everyone.

Kembel, S., Cowan, P., Helmus, M., Cornwell, W., Morlon, H., Ackerly, D., Blomberg, S., & Webb, C. (2010). Picante: R tools for integrating phylogenies and ecology Bioinformatics DOI: 10.1093/bioinformatics/btq166

Niche or Neutral? Why size matters.

2010-04-27T05:59:00.000-07:00

Metacommunity dynamics (i.e. the effects of dispersal among connected communities) have become an increasingly common lens through which to explain community structure. For example, competition-colonization models explain the coexistence of superior and inferior competitors as the result of a trade-off in colonization and competitive ability. Species are either superior competitors, with high probabilities of establishing in patches, but low ability to move between patches, or superior colonizers, which have tend to lose in competitive interactions but can travel easily between patches. Under this framework, the ability of superior colonizers to reach and maintain populations in patches where their superior competitors are absent allows them to avoid extinction.

One problem with these types of models is that they rarely acknowledge the importance of ecological drift – that is, that chance events also affect species interactions. This despite the fact that we know that in “real life”, chance events likely play a major role in producing assemblages different than those we might predict based on theory. One of the strengths of the Hubbell’s neutral model is that it recognizes and embraces the importance of randomness.

A recent paper by Orrock and Watling (2010) examines how chance events can alter the predictions of the classic competition-colonization model. Orrock and Watling show that the size of communities in a metacommunity (which is assumed to correlate with the strength of ecological drift) determines whether community dynamics are niche-structured or neutral in nature. In large communities, predictions agree closely with those of the classic competition-colonization model, and niche-based interactions (i.e. competitive hierarchies) dominate. It’s in small communities that things get interesting: ecological drift becomes more important, so that differences in competitive ability between species are effectively neutralized. As a result, small communities begin to resemble neutral assemblages in which species abundances don’t relate to differences in competitive ability. An interesting consequence of this outcome is that species who are poor competitors but good colonizers have an additional refuge – simply by escaping to small communities, even if these communities contain superior competitors, they can persist in a metacommunity.

Beyond the theoretical implications of this model, the applied implications are what really matter. Habitat destruction and fragmentation are an growing problem due to human activities. Habitat patches are often smaller, and of lower quality, decreasing the size of the community each patch can support. Even if these patches are still connected and functioning as a metacommunity, species which rely on their strong competitive ability for persistence will lose this advantage as assemblages become increasingly neutral. Under this model, community diversity declines even more as habitat is lost than in the traditional competition-colonization model, and superior competitors face even greater extinction risk than previously predicted.

Since in reality, metacommunities are likely to consist of patches of different sizes, rather than all large or all small patches, the predictions here remain to be extended to more realistic metacommunities. However, Orrock and Watling have produced a useful model for understanding how ecological drift can affect diversity in a metacommunity and alter the expectations of traditional competition-colonization models.

Orrock, J.L. and Watling, J.I. (2010) Local community size mediates ecological drift and competition in metacommunities. Proc. R. Soc. B.

Teaching a quoll that cane toads are bad

2010-04-14T08:42:00.000-07:00

Often, species become endangered because of multiple stressors, with habitat destruction taking the prize as the most egregious. However, often what pushes a species into extinction is not the main driver of endangerment. For example, passenger pigeon numbers were decimated by unabated hunting, but the proximate cause of extinction was likely an inability to thrive in low densities. Yet, seldom is the case where a known single species interaction is the primary cause of engangerment and maybe extinction. The northern quoll, Dasyurus hallucatus, is an endangered marsupial predator in Australia. The current major threat to the northern quoll is the invasion of toxic can toads. Quolls, being predators of small mammals, birds, reptiles and amphibians, readily attacks cane toads, which are toxic to quolls. Quoll populations have disappeared from areas invaded by cane toads, and extinction seems almost inevitable.

Given that the spread of cane toads into the remaining quoll habitats is inevitable, research, led by Stephanie O'donnell in Richard Shine's lab at the University of Sydney and published in the Journal of Applied Ecology, is underway to train quoll's to avoid cane toads. These researchers feed a subset of captive quolls dead toads laced with thiabendazole, a chemical that induces nausea. They then fitted individuals with radio collars and released these toad-smart quolls as well as toad naive ones. Some toad-naive quolls died quickly, after attacking cane toads. Only 58% of male naive quolls survived, while 88% of toad-smart males survived. While females seemed less likely to attack toads, 84% of naive females survived and 94% of toad-smart females survived!

See the video of a toad-smart quoll deciding not to eat a cane toad, its pretty cool.

O’Donnell, S., Webb, J., & Shine, R. (2010). Conditioned taste aversion enhances the survival of an endangered predator imperilled by a toxic invader Journal of Applied Ecology DOI: 10.1111/j.1365-2664.2010.01802.x

Plant rarity: environmental or dispersal limited?

2010-04-08T18:41:00.000-07:00

In order to promote the persistence and possible spread of extremely rare plant species, ecologists need to know why a species is rare in the first place. In 1986, Deborah Rabinowitz identified seven forms of rarity, where rarity could mean several things depending on range size, habitat specificity and population sizes. When considering rarity, it often feels intuitive to look for environmental causes for these different forms of rarity. Habitat alteration is an obvious environmental change that affects abundance and distribution, but are rare species generally limited by habitat or resource availability? The alternative cause of rarity could just be that sufficient habitat exists, but that the rare species is simply unable to find or disperse to other sites. An extreme example of this would be the Devil's Hole pupfish which exists at only a single pool. It can survive elsewhere (such as in artificial tanks) but natural dispersal is impossible as its pool is in a desert.

Photo taken by Kristian Peters and available through GNU free documentation license

In a recent paper by Birgit Seifert and Markus Fischer in Biological Conservation, they examine whether an endangered plant, Armeria maritima subsp. elongata, was limited because of a lack of habitats or if it was dispersal limited. They collected seeds from eight populations and experimentally added these seeds to their original populations and to uninhabited, but apparently appropriate sites. They found that seeds germinated equally well in inhabited and uninhabited sites and seedlings had similar survivorships. They found that variation in germination rates were likely caused by originating population size and that low genetic diversity and inbreeding reduce viability.

These results reinforce two things. First is that conserving species may only require specific activities, such as collect and distributing seeds. Here ideas like assisted migration seem like valuable conservation strategies. Secondly, we really need to be doing these simple experiments to better understand why species are rare. If we fail to understand the causes of rarity, we may be wasting valuable resources when try to protect rare species.

Seifert, B., & Fischer, M. (2010). Experimental establishment of a declining dry-grassland flagship species in relation to seed origin and target environment Biological Conservation DOI: 10.1016/j.biocon.2010.02.028

Predicting endangered carnivores: the role of environment, space and phylogeny

2010-03-22T17:23:00.000-07:00

For conservation biology, there are several research thrusts that are of critical importance, and one of these is to find predictors of species' extinction risk. Oft-cited is the particular susceptibility of large-bodied organisms, with their large ranges and slow reproductive rates. But there should be other predictors too, especially within larger mammals. In a forthcoming paper in Global Ecology and Biogeography, Safi and Pettorelli use just a few variables to predict extinction risk in carnivores.
They quantified species extinction risk according to the IUCN risk assessments and asked how well three attributes explained variation in extinction risk. They quantified the environmental characteristics of the species' ranges (temperature, precipitation, etc.), spatial distances between species' ranges and the phylogenetic distances among species. Overall, spatial and phylogenetic distances were good predictors of threat status -generally predicting between 21-70% of variation in extinction risk, whereas the environmental variables were weaker predictors. Full models incorporating all three variables (and accounting for their covariance), were able to explain upwards of 96% of the variation in extinction risk!

Although these variables do not represent causal mechanisms of extinction risk -rather they are correlative, they do provide conservation biologists with a rapid assessment tool to evaluate extinction risk. These tools should be particularly important in cases were population data are lacking and immediate pragmatic decisions are required.

Safi, K., & Pettorelli, N. (2010). Phylogenetic, spatial and environmental components of extinction risk in carnivores Global Ecology and Biogeography DOI: 10.1111/j.1466-8238.2010.00523.x

Low impact blogging

2010-03-15T18:36:00.000-07:00

As a form of communication, blogging (and all that other stuff on the internet) is fairly environmentally friendly. Trees are not cut down to produce paper to print our posts, fuel-hungry trucks are not used to deliver these articles and stories to our many(!) readers and there is no trash to add to landfills. However, there is still the unappreciated cost associated with energy consumption for all the hours of researching, writing, and being read. The energy for all this electronic activity mainly comes from fossil fuels, meaning that my blogging has a carbon footprint.

Not anymore. No, we did not go nuclear. Rather, the ingenious people behind Mach's grun have started a great program. For writing this post about them, their 'make it green' campaign and the Arbor Day Foundation will plant a tree in Plumas National Forest in northern California. In 2007, a devastating forest fire destoyed 65,000 ha. By choosing to blog green, at least one more tree is planted. I will feel better knowing that there will be tree exhaling oxygen for our blog.

Ecology and industry: bridging the gap between economics and the environment

2010-03-09T06:11:00.000-08:00

Applied ecology is the science of minimizing human impacts and of supporting ecological systems in an economic landscape. Often though, applied ecologists work in isolation from those economic forces shaping biological landscapes, not really knowing what businesses would like to accomplish for habitat protection or sustainability. At the same businesses are seldom aware of the knowledge, tools and insight provided by ecologists. And perhaps, greater interaction could help turn ecology into a science with direct impact into how human activities proceed and how we manage the impacts of those activities.

This is the premise of a paper by Paul Armsworth and 15 other authors on the ecological research needs of business, appearing in the Journal of Applied Ecology (for an interview with Paul, by yours truly, please go to the podcast, and I should point out that I am an Editor with this journal). The authors include academics, NGOs and industrial representatives, and they've come together to analyze patterns of cooperation and to discuss ways forward.

They reviewed papers appearing in the top applied ecology journals and grant proposals to the National Environmental Research Council (NERC) in the UK to measure the degree and type of interaction between ecologists and different industries. Ten to 15 percent of publications in applied journals showed some business involvement -mostly from the traditional biological resource industries (farming, fishing and forestry). Further, 35% of NERC proposals included some business engagement, but only 1% had direct business interaction.

Further, the authors reported on a workshop where ecologists and business representatives discussed a number of topics. This included how to minimize negative biodiversity impacts and for industries, such as mining, to consider ecosystem function, and how to develop new ecologically-based economic opportunities, such as insurers managing environmental risk. While there were some challenges identified (such as differing time frames of business needs versus scientific research), the authors note the positive atmosphere and the spirit of collaboration.

The research in this paper should be emulated elsewhere. A better understanding of business needs and desires can only inform and offer opportunities for applied ecological research. Top-down governmental regulation can only take conservation and ecosystem management so far and those who are directly involved in altering and managing ecosystems must articulate goals and desires in order to successfully apply ecological principles to biodiversity protection in an economic landscape.

Armsworth, P., Armsworth, A., Compton, N., Cottle, P., Davies, I., Emmett, B., Fandrich, V., Foote, M., Gaston, K., Gardiner, P., Hess, T., Hopkins, J., Horsley, N., Leaver, N., Maynard, T., & Shannon, D. (2010). The ecological research needs of business Journal of Applied Ecology, 47 (2), 235-243 DOI: 10.1111/j.1365-2664.2010.01792.x

Competitive coexistence, it's all about individuals.

2010-03-05T10:56:00.001-08:00

Understanding how species coexist has been the raison d'etre for many ecologists over the past 100 years. The quest to understand and explain why so many species coexist together has really been a journey of shifting narratives. The major road stops on this journey have included searching for niche differences among species -from single resources to multidimensional niches, elevating the role for non-equilibrial dynamics -namely disturbances, and assessing the possibility that species actually differ little and diversity patterns follow neutral process. Along this entire journey, researchers (especially theoreticians) have reminded the larger community that that coexistence is a product of the balance between interactions among species (interspecific) and interactions among individuals within species (intraspecific). Despite this occasional reminder, ecologists have largely searched for mechanisms dictating the strength of interspecific interactions.

Image used under Flickr creative commons license, taken by Tinken

In order for two species to coexist, intraspecific competition must be stronger than interspecific -so sayeth classic models of competition. While people have consistently looked for niche differences that reduce interspecific competition, no one has really assessed the strength of intraspecific competition. Until now that is. In a recent paper in Science, Jim Clark examines intra- vs interspecific interactions from data following individual tree performances, across multiple species, for up to 18 years. This data set included annual growth and reproduction, resulting in 226,000 observations across 22,000 trees in 33 species!

His question was actually quite simple -what is the strength of intraspecific interactions relative to interspecific ones? There are two alternatives. First, that intraspecific competition is higher, meaning that among species differences only need to be small for coexistence to occur; or secondly, that intraspecific competition is lower, requiring greater species niche differences for coexistence. To answer this he looked at correlations in growth and fecundity between individuals either belonging to the same or different species, living in proximity to one another. He took a strong positive correlation as evidence for strong competition and a negative or weak correlation as evidence for resource or temporal niche partitioning. What he found was that individuals within species were much more likely to show correlated responses to fluctuating environments, than individuals among species.

This paper represents persuasive evidence that within-species competition is generally extremely high, meaning that to satisfy the inequality leading to coexistence: intra > inter, subtle niche differences can be sufficient. These findings should spur a new era of theoretical predictions and empirical tests as our collective journey to understanding coexistence continues.

Clark, J. (2010). Individuals and the Variation Needed for High Species Diversity in Forest Trees Science, 327 (5969), 1129-1132 DOI: 10.1126/science.1183506

Check out the carnival of evolution and be sure to vote for your favorite blogs

2010-03-02T06:41:00.000-08:00

Be sure to check out this month's Carnival of Evolution (number 21) posted at Mauka to Makai. The Carnival is a monthly digest of recent evolutionary musings from around the blogosphere. This month's edition includes a number of interesting posts, as well as one of our posts on what evolution offers conservation.

Also, Research Blogging has announced finalists for various blogs awards. If you are eligible, please vote, there are a lot of great blogs vying for these awards. Also, The EEB and Flow is among the finalists for best biology blog. And to the people we nominated us, thanks again for nominating our blog.

New Tool Reveals Where Ticks Eat Breakfast

2010-02-27T12:20:00.000-08:00

You have a much greater chance of getting sick from a tick bite today than you did 30 years ago. But a new tool might allow researchers to better understand why more ticks are making people sick.

“If you’re a health inspector and a bunch of people get food poisoning, the first thing you’d want to know is where they ate last. If you’re a disease ecologist and a bunch of ticks have a pathogen, the first thing you’d want to know is where the ticks ate last,” said Brian Allan, a post-doctoral researcher at the Tyson Research Station in St. Louis.

Allan led a team of researchers in developing a novel technology that probes the genetic contents of ticks’ gut. The tool can determine which wildlife species provided the tick’s last meal and which pathogens came along with that meal.

In the first study to use the new technology, Allan and his colleagues focused on several rapidly emerging diseases transmitted by the lone star tick. These include two pathogens responsible for a potentially fatal bacterial infection known as ehrlichiosis [ur-lick-ee-oh-sis]. In Missouri, over 200 cases of ehrlichiosis were documented last year.

Allan et al.'s study showed that about 80 percent of pathogen-positive ticks had fed on white-tailed deer. They also found that squirrels and rabbits were capable of infecting ticks at a higher rate than deer. However, since the lone star tick feeds on squirrels and rabbits less frequently, they account for a smaller percentage of infection.

Allan and his colleagues hope that the technique will shed light on theoretical questions in the field of ecology. They are especially interested in testing whether biodiversity is good for your health, a hypothesis known as “the dilution effect.”

Allan, B. F., L. S. Goessling, G. A. Storch, and R. E. Thach. 2010. Blood meal analysis to identify reservoir hosts for Amblyomma americanum ticks. Emerging Infectious Diseases 16: 433-440. DOI: 10.3201/eid1603.090911

How can evolution inform conservation decisions?

2010-02-22T10:37:00.000-08:00

First of all, let me apologize for the lack of blog posts over the past 2 weeks, I've been busy visiting the Olympics and reading a couple of hundred blogs, judging them for the Research Blogging awards.

The conservation of biological diversity is a major imperative for biologists. International agreements such as the Convention on Biological Diversity and intergovernmental exercises, such as the Millennium Ecosystem Assessment, call upon scientists to provide evidence on the current state of biological diversity and to evaluate solutions for reducing diversity and ecosystem function loss. Critical to these efforts have been the work of ecologists, conservation biologists and ecological economists. However, seemingly missing from the conversation about the state of biodiversity knowledge has been evolutionary biologists. Are they primarily concerned with describing historical processes and mechanisms of biological change, or do they have substantive knowledge and ideas that should be viewed as a critical component of any scheme to conserve biological diversity?

In a recent paper in Evolution, Hendry and a number of coauthors convincingly make the case that evolutionary biology is a necessary component for conservation. Evolution offer four key insights that should inform conservation and policy decisions. First, they point out that evolutionary biologists are in the business of discovering and documenting biodiversity. They are the primary drivers behind long-term, sustained biological collections, because they need to know what exists in order to better understand evolutionary history. With millions of species awaiting scientific discovery, their efforts are critical to measuring biodiversity. But not only are they discovering new species and enumerating them, they are uncovering their evolutionary relationships, which gives conservationists better information about which species to prioritize. What Vane-Wright famously called 'the agony of choice', with limited resources, we need to prioritize some species over others, and their evolutionary uniqueness ought to be a factor. More than this, evolutionary biologists have developed pragmatic tools for inventorying and sharing data on biodiversity at all levels, from genes to species, which is available for prioritization.

The second key insight is that by understanding the causes of diversification, we can better understand and predict diversity responses to environmental and climatic change. By understanding how key functional traits evolve, we can develop predictions about which species or groups of species can tolerate certain perturbations. Further, research into how and why certain evolutionary groups faced extinction can help us respond to the current extinction crisis. For example, the evolutionary correspondence between coevolved mutualists, such as plants and pollinators, can be used to assess the potential for cascading extinctions. These types of analyses can help identify those groups of related species, or those possessing some trait, which make species more susceptible to extinction.

Thirdly, evolution allows for an understanding of the potential responses to human disturbance. Evolutionary change is a critical part of ecological dynamics, and as environment change can result in reduced fitness, smaller population sizes and extinction, evolution offers an adaptive response to these negative impacts. Knowing when and how populations can evolve is crucial. Evolutionary change is a product of genetic variation, immigration, population size and stochasticity, and if the ability to evolve to environmental change is key for persistence, then these evolutionary processes are also key.

Finally, evolutionary patterns and processes have important implications for ecosystem services and economic and human well-being. Both genetic and evolutionary diversity of plant communities has been shown to affect arthropod diversity, primary productivity (including work by me) and nutrient dynamics. Thus understanding how changes in diversity affect ecosystem processes should consider evolutionary processes. Further, exotic species are often cited as one of the major threats to biodiversity, and evolutionary change in exotics has been shown to increase exotic impacts on native species.

All together, these key reasons why evolution matters for conservation, mean that developing sound management plans requires considering evolution patterns and processes. We can use evolution to our benefit only if we understand how evolution shapes current dynamics. The challenge to evolutionary biologists is the same as it was for ecologists perhaps 15 to 20 years ago, to present their understanding and conservation ideas to a broader audience and to engage policy makers. To this end, the authors highlight some recent advances in incorporating evolutionary views into existing biodiversity and conservation programmes –most notably into DIVERSITAS.

Just like ecological processes and dynamics cannot be fully understood without appreciating evolution ancestry or dynamics, developing an extensive, expansive conservation strategies must take into account evolution. I hope that this paper signals a new era of a synthesis between ecology and evolution, which produces precise, viable conservation strategies.

Hendry, A., Lohmann, L., Conti, E., Cracraft, J., Crandall, K., Faith, D., HÃ¤user, C., Joly, C., Kogure, K., Larigauderie, A., MagallÃ³n, S., Moritz, C., Tillier, S., Zardoya, R., Prieur-Richard, A., Walther, B., Yahara, T., & Donoghue, M. (2010). EVOLUTIONARY BIOLOGY IN BIODIVERSITY SCIENCE, CONSERVATION, AND POLICY: A CALL TO ACTION Evolution DOI: 10.1111/j.1558-5646.2010.00947.x

Research blogging awards; and thanks

2010-02-10T05:01:00.000-08:00

Hi all, nominations for Research Blogging's annual awards will close Feb. 11, so be sure to nominate any research blogs you think deserve consideration. The top prize is $1000, and there are several smaller, field-specific awards as well. A panel of judges (with me being a member) will create a short list of blogs for each category and registered users on Research Blogging will be able to vote for the winners.

And a thank you to whoever nominated the EEB and Flow.

Predator-human conflict: the emergence of a primordial fear?

2010-02-08T10:01:00.000-08:00

There is something terrifying and at the same time captivating about the idea of a large, wild, mysterious predator. The very idea that a large predator is near by makes us feel vulnerable. Every year, news stories about wild animal attacks appear in numerous publications and on many television shows. Human death at the fangs or claws of a wild beast is at the heart of many legendary stories and probably sown into the fabric of our being by millennia of ever present risk from large predators. This characteristic of our human experience, I think, dictates our response to animal attacks. Stories of animal attacks are usually concluded with statements about having or attempting to track down and destroy the guilty animal.

Such is the case for three recent animal attacks in Canada. In late October, 2009 in Nova Scotia, a raising 19-year old folk singer was killed by a couple of coyotes while hiking. It is difficult to find meaning in such a horrendous death, but the narrative, told by reporters, was essentially to rest assured that one of the coyotes had been killed and the other was being tracked and would be destroyed. There were two cougar attacks in early January, 2010 in British Columbia, that basically ended with the same reassurance. In the first, a boy was attacked and his pet golden retriever courageously saved his life. A police officer arrived a shot the cougar which was mauling the dog -an obviously legitimate response, and the news story again reassures us that the animal was destroyed. And don't worry the hero dog survived. In the second cougar attack, another boy was attacked, and this time his mother saved his life. But again the story narrative ended by reassuring us that the guilty cougar, and another cat for good measure, were destroyed the next day.

After reading these stories, I asked myself two things. Why is our response to destroy predators that attack? And why do we need to be reassured that this has happened? In defence of the predators, they are just doing what their instincts tell them to do, and most often their only mistake is that they selected their prey poorly. But the reality is that there are only 2-4 cougar attacks per year and only 18 fatalities over the past 100 years. Why do we fear such a low probability event? In contrast, automobile accidents are the leading cause of death in children under 12 in North America. Thousands of people die, and millions injured in car accidents every year in North America. Recently, in Toronto, were I live, 10 pedestrians were killed in 10 days, yet my heart doesn't race when I cross a street. If our fears and responses to human injury and death reflected the actual major risks, we would invoke restrictive rules regarding automobile use.

We believe that we can live with nature in our backyard. But when that close contact results in an animal attack, human fear seems to dictate an irrational response. Do we really expect predators to obey our rules? Can we punish them enough to effectively tame them? We cannot, and I hope that our approaches to dealing with human-animal conflict can better deal with animal attacks, in a way that does not subjugate large predators to whims of our fears.

The evolution of a symbiont

2010-02-03T17:57:00.000-08:00

The evolution of negative interactions seems like a logical consequence of natural selection. Organisms compete for resources or view one another as a resource, thus finding ways to more efficiently find and consume prey. However, to me, the natural selection of symbiotic or mutualistic interactions has never seemed as straight forward (expect maybe the case where one species provides protection for the other, such as in ant-plant mutualisms). A specific example is the rise of nitrogen-fixing plants, who supply nutrients to bacteria called rhizobia capable of converting atmospheric nitrogen into forms, such as ammonia, usable to the plant host. Not only has this symbiosis evolved, but has seemed to evolve in very evolutionarily distinct lineages. The question is, what are the mechanisms allowing for this?

In a recent paper, Marchetti and colleagues answer part of the question. They experimentally manipulate a pathogenic bacteria and observe it turning into a symbiont. They transferred a plasmid from the symbiotic nitrogen fixing Cupriavidus taiwanensis into Ralstonia solanacearum and infected Mimosa roots with it. Plasmid transfer among distinct bacteria species is common and referred to horizontal genetic transfer (as opposed to vertical, which is the transfer to daughter cells). The presence of the plasmid caused R. solanacearum to quickly evolve into a root-nodulating symbiont. Two regulatory genes lost function, and this caused R. solanacearum to form nodules and to impregnate Mimosa root cells.

This extremely novel experiment reveals how horizontal gene transfer can supply the impetus for rapid evolution from being a pathogen to a symbiont. More importantly it reveals that sometimes just a few steps are required for this transition and how distantly-related bacterial species can acquire symbiotic behaviors.

Marchetti, M., Capela, D., Glew, M., Cruveiller, S., Chane-Woon-Ming, B., Gris, C., Timmers, T., Poinsot, V., Gilbert, L., Heeb, P., Médigue, C., Batut, J., & Masson-Boivin, C. (2010). Experimental Evolution of a Plant Pathogen into a Legume Symbiont PLoS Biology, 8 (1) DOI: 10.1371/journal.pbio.1000280

To intervene or not to intervene: this is a real question

2010-01-27T07:04:00.000-08:00

Should land managers actively manipulate the structure and function of ecosystems within protected areas? Is intervention appropriate to protect or maintain native biodiversity and natural processes in areas such as national parks and wilderness areas? These are the questions that stem from a new paper by Richard Hobbs and others in Frontiers in Ecology and the Environment. US national parks and wilderness areas have legislative mandates to maintain ‘naturalness’, but what does this mean in the context of dynamic ecosystems with current and future changes including invasions by nonnative organisms and climate change?

Hobbs and his colleagues challenge concepts of naturalness and propose several ‘guiding principles’ for stewards of national parks and wilderness. They suggest that more useful concepts for managing protected areas relate to ecological integrity and resilience. Concepts of ecological integrity have been adopted by Parks Canada and relate to maintaining ecosystem components. Resilience concepts focus on the ability of a system to “absorb change and persist” without undergoing a “fundamental loss of character”. While maintaining ecological integrity in the face of global changes may - by definition - require protection of species, maintaining ecological resilience tends to focus more attention on ecosystem function “over preserving specific species in situ”.

Rather than protecting an area to maintain naturalness, focusing on ecological integrity and resilience acknowledges that a diversity of approaches - from non-intervention to actively managing systems - may be required. The flexibility in this view, demands that conservation planning span gradients of land uses across landscapes. Management objectives and success need to be re-evaluated in an adaptive and experimental framework, which requires careful and robust monitoring.

At The Wilderness Society and specifically here in Montana, these very questions are being wrestled with in terms of forest restoration, fire management, and climate change. Current forest conditions have been shaped by historic logging practices and fire suppression leading to altered structure and function – including increasing the severity of fires. Through active management, including removing small diameter trees and lighting prescribed fires, managers hope to restore forests and fire intensities to conditions more closely resembling those that historically occurred. Much of the research on restoration was conducted in dry forests in the American Southwest where low-severity fires occurred across large areas. However, in the Northern Rockies, many forests were shaped by a ‘mixed severity’ fire regime, where fires crept along the forest floor in some areas and torched trees in others. In many cases, these forests have not been fundamentally altered and need only the return of fire to restore their resilience. In other cases, forests are recovering from past logging practices and may benefit from thinning to restore a fire-resilient structure.

To return to the paper at hand: what is the appropriate level of intervention to maintain ecological integrity and resilience given past forest management and future climate change? If the current forest lacks integrity (novel stand structure) and resilience under a predicted climate of warmer, drier conditions, what is the appropriate level of management? While The Wilderness Society continues to work with diverse partners to answer these questions, one thing is clear: whatever actions take place, they need to be conducted with humility in an experimental framework that includes sufficient ecological monitoring. For the ‘experiment’ to be most helpful, we should maintain adequate hands-off “controls” along with the “treatments” to allow us to gauge the effects of intervention.

Richard J Hobbs, David N Cole, Laurie Yung, Erika S Zavaleta, Gregory H Aplet, F Stuart Chapin III, Peter B Landres, David J Parsons, Nathan L Stephenson, Peter S White, David M Graber, Eric S Higgs, Constance I Millar, John M Randall, Kathy A Tonnessen, Stephen Woodley (2009) Guiding concepts for park and wilderness stewardship in an era of global environmental change. Frontiers in Ecology and the Environment e-View.
doi: 10.1890/090089
http://www.esajournals.org/doi/abs/10.1890/090089