The talks I saw today were uniformly good and a number were excellent. At least half of them focused on the many implications for ecology of nature's innate variability. It appears that community ecologists have decided that now is the time to start considering the fact that the environment is not stationary, which was long a default assumption in most theoretical and empirical work. Many of the talks I saw reflected this changing approach. The other half were part of a symposium organized by Sharon Strauss that looked at coexistence among sister species. This topic, combining as it did large-scale evolutionary and biogeographic processes with local competitive interactions made for a broad range of talks and some interesting attempts to reconcile different methodologies and scales.
Our variable world
Many of the past studies on environmental variability and coexistence involve desert winter annuals. Desert winter annuals are limited by available water, and the yearly rains vary greatly in the amount and timing of onset. The hypothesis is that variable germination (via prolonged dormancy in seedbanks) may allow desert winter annuals to reduce the variance in their fitness between years. Alejandra Martinez-Berdeja presented some tidy hypothesis testing using biogeographical gradients: if variable germination is an adaptive response to variable precipitation, she hypothesized that differences in germination variability might be expected where precipitation is more or less predictable. Looking at the three North American deserts, she predicted that variable germination would be greater where rainfall was more variable (bi-seasonal) compared to winter rainfall deserts. She measured the involucres (dispersal structures determining seed release) on collected seeds and found that indeed they were larger in more variable rainfall deserts, producing greater variability in seed release. Further, in winter rainfall deserts, variability in the size of involucres was correlated with variability in rainfall at a site, again suggesting a link between germination variability and rainfall variability. Her next step will hopefully be to expand the tests look at the effect of autocorrelation in rainfall likelihood on bet-hedging, since this should increase selection for bet-hedging type adaptations.
David Vasseur gave a great talk showing how extreme environmental conditions--which we are seeing as part of the changes in mean and variance of the climate--could have particularly detrimental effects on population growth rates. Species have temperature performance curves that reflect the relationship between their fitness and the temperatures they experience. Vasseur showed that in the tropics, species tend to have much narrower temperature ranges over which they can grow and survive than species in the temperature regions, and experts agree that these narrower curves give tropical species less ability to deal with increasing temperatures. But variability is rarely considered in this equation. When variability is present, long-term species fitnesses will be subject to Jensen's inequality (nonlinear averaging) mean that shape of these performance curves is additionally important: that in some situations (concave curves) variability is particularly detrimental, and in some situations (convex curves) it may have a beneficial effect. Vasseur then used models to show that as temperature variation increases, it is increasingly likely that its effect will be negative, and high variation will produce high extinction rates. In fact, on average Vasseur predicted that temperature variation would have negative effects, a concerning conclusion.
Sisters getting along
This organized symposium was advertised as: “Whether closest relatives coexist reflects the often opposing effects of limiting similarity, mode of speciation, reproductive isolation, niche conservatism, competition and facilitation, which may be strongest in sister taxa; using new phylogenies, niche models, and experimental approaches, we explore coexistence in closest relatives in both plants and animals.” It was an interesting and useful idea – sister species (species who are each other’s most recent relative) are an important tool to understand how evolution, biogeography, and ecological interactions determine coexistence. The content of the symposium provided a number of example systems, methods, and approaches that suggested this was an important but still far from cohesive area of work. Mark McPeek spoke about the damselfly work he has done over the last many years, which shows that sister species are sympatric and ecologically identical, co-occurring happily through neutral dynamics. In contrast, Richard Glor talked about his work with Hispaniola anoles, where biogeography is an explanation for radiations, close relatives use different microhabitats and rarely compete locally and traits are divergent among close relatives. Looking at California plant species, Brian Anacker’s talk suggested something in between these extremes. A broad survey showed that 80% of sister pairs were sympatric, range overlap was modest but not uncommon, but asymmetry in range size was high. Ecological differences between sister species were not particularly clear in the handful of traits he examined, not even for reproductive traits. Sister species can and do co-occur, although not in large portions of their ranges. Having established the current state of knowledge, hopefully the symposium will stimulate greater focus on the construct of sister species as a way of understanding coexistence at multiple scales.
Finally, not being willing to miss another talk with the word “derby” in the title, I attended Daniel Atwater’s talk, “Is competition among plants like a boxing match or a demolition derby? Why competitor suppression may not matter in plant communities”. Atwater argued that there were two ways to win at plant competition – be good at suppressing your competitors, or be good at tolerating them. When in competition with a single individual, being a strong suppressor should be favoured, but in competition with multiple species, tolerance may be a better strategy. That’s because resources spent on suppressing one competitor may also benefit any other species involved in the competition. In such cases, tolerance of your competitors may provide the greatest benefit. (Apparently this scenario is like a successful (but frowned upon) strategy (sandbagging) in a demolition derby). Atwater used experimental data from blue bunch grass grown in competition with spotted knapweed to parameterize a model in which he found the optimum strategy in single versus multi-species competitions. The model agreed with his hypothesis that tolerating competitors is favoured when multiple species are competing. Although I am not clear on whether competitive strategies are easily classified as tolerant vs suppressing it was an interesting talk, and left me thinking about new questions.