Monday, January 30, 2012

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

For many ecologists, neutral theory was a (good/bad, you choose) idea that dominated ecology for the last decade but failed to provide the burden of empirical proof necessary for its acceptance. Even its creator Stephen Hubbell  recently suggested that the controversial hypothesis is no longer a plausible description of community structure, going as 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 sole property of their creators. Other researchers continue to study neutral theory, and despite the apparent consensus that neutral theory is not an important explanation of community structure and dynamics, papers testing neutral theory continue to be published. This leads to an important question: do we still want to test for neutral dynamics? And if we do, how should we approach it, given what we have learned from the past decade of strawman arguments and using pattern-based evidence for processes (e.g. looking at species-area relationships and species abundance distributions)? What empirical evidence would provide strong support for the predictions of neutral theory?

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

For two Enallagma sp. larvae Siepielski et al. used cages in the littoral zone of lakes, with cages receiving different treatments of relative abundance and/or total abundance manipulation. The result of these manipulations were that replicates with increased total abundances and constant relative abundances had lowered per-capita growth rates, while replicates with manipulated relative abundances and constant total abundances showed no change in demographic rates. Both species had similar mortality rates across the experimental treatments, although their growth rates differed slightly. From these results, Siepielski et 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 work is deeply rooted in a natural system, and yet who also argues that ecological equivalency seems plausible, and attempts to support that argument. Regardless of whether the Enallagma species are in fact ecologically equivalent, this paper provides an example of how coexistence theory can be more rigorously tested than simply observing species co-ocurrences and concluding species coexistence. Further, it provides some interesting discussion about whether ecological equivalency is possible within functional groups, with niche differences occurring between functional groups (see Leibold and McPeek 2006, and from MacNaughton and Wolf 1970 for similar suggestions). Future work might focus on questions such as how to capture the effects of small niche differences, which, if balanced against very similar fitnesses could explain stable coexistence. In addition, it might be valuable to look at how resources fluctuate and how much overlap there is in resource requirements among species, when looking at how growth and mortality change with species densities.

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

Tuesday, January 10, 2012

Trends in ecology, 2011

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.

Tuesday, January 3, 2012

Carnival of Evolution 43!

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?