Unexpected effects of global warming in novel environments: butterflies emerge later in warming urban areas.

There is now ample evidence that warming temperatures cause advances in the timing of organismal activity (i.e., phenology). Studies have shown that rising temperatures are responsible for earlier plant leafing and flowering (Miller-Rushing & Primack 2008, Wolkovich et al. 2012), pest insect emergence and abundance (Willis et al. 2008), and even local species loss and reduced diversity (Willis et al. 2008). One emerging expectation from global warming studies is that insects should emerge earlier since winters are milder and spring temperatures are warmer. This expectation should hold so long as high temperatures or other environmental stressors don’t adversely affect the insects. And the concern about shifts in emergence and insect activity is the potential for mismatches between plant flowering and the availability of pollinators (Willmer 2012) –if insects emerge too soon, they may miss the flowers.

Photo by Marc Cadotte

In a forthcoming paper in Ecology by Sarah Diamond and colleagues study 20 common butterfly species across more than 80 sites in Ohio. These sites were located in a range of places across a rural to urban gradient. Instead of finding earlier emergence in warmer places, which were typically urban areas, they found that a number of species were delayed in warmer urban areas. Even though the butterflies might emerge earlier in warmer rural habitats, they were adversely affected in urbanized areas. 

These results highlight the need to consider multiple sources of stress from different types of environmental change. Observations from a few locales or from controlled experiments may not lead to conclusions about interactive influences or warming and urbanization, and that's why this study is so important. It observes a counter-intuitive result because of the influence of multiple stressors. 

A next step should be to determine if pollinator-plant interactions are being disrupted in these urban areas. The reason why we should care so much about pollinator emergence is that they provide a key ecological service by pollinator wild, garden, and agricultural plants, as well has being an important food source to other species. A mismatch in timing and disrupt these important interactions.

References

Diamond S.E., Cayton H., Wepprich T., Jenkins C.N., Dunn R.R., Haddad N.M. & Ries L. (2014). Unexpected phenological responses of butterflies to the interaction of urbanization and geographic temperature. Ecology.

Miller-Rushing A.J. & Primack R.B. (2008). Global warming and flowering times in Thoreau's Concord: a community perspective Ecology, 89, 332-341.

Roos J., Hopkins R., Kvarnheden A. & Dixelius C. (2011). The impact of global warming on plant diseases and insect vectors in Sweden. Eur J Plant Pathol, 129, 9-19.

Willis C.G., Ruhfel B., Primack R.B., Miller-Rushing A.J. & Davis C.C. (2008). Phylogenetic patterns of species loss in Thoreau's woods are driven by climate change. Proceedings of the National Academy of Sciences, 105, 17029-17033.

Willmer P. (2012). Ecology: pollinator-plant synchrony tested by climate change. Curr. Biol., 22, R131-R132.

Wolkovich E.M., Cook B.I., Allen J.M., Crimmins T.M., Betancourt J.L., Travers S.E., Pau S., Regetz J., Davies T.J., Kraft N.J.B., Ault T.R., Bolmgren K., Mazer S.J., McCabe G.J., McGill B.J., Parmesan C., Salamin N., Schwartz M.D. & Cleland E.E. (2012). Warming experiments underpredict plant phenological responses to climate change. Nature, 485, 494-497.

Diamond, S., Cayton, H., Wepprich, T., Jenkins, C., Dunn, R., Haddad, N., & Ries, L. (2014). Unexpected phenological responses of butterflies to the interaction of urbanization and geographic temperature Ecology DOI: 10.1890/13-1848.1

Timing is everything: global warming and the timing of species interactions

While an obvious affect of climate change will be changes in the distributions or range sizes of species, more insidious and likely more consequential will be how species interactions are affected by changes in the timing of growth and reproduction. These changes in an organism's life cycle, or phenology, can create mismatches between an organism's need and resource availability or the readiness of coevolved partners -such as plants and pollinators.

In an 'Idea and Perspective' paper in Ecology Letters, Louie Yang and Volker Rudolf set out a new framework to examine the effects of phenological shifts on species interactions. They argue that one cannot understand or predict the fitness consequences of a phenology shift without knowing how interacting species' phenologies are also influenced by environmental changes. The consequences of phenological shifts are changes in fitness, and the question is: how would one go about assessing the fitness effects of phenological changes on interactions? This is where this paper really hits its stride. Yang and Rudolf set out a new conceptual framework for studying the fitness consequences of phenological shifts. They make the case that an experimental approach is required to test the three likely scenarios. The first is that there are no changes in phenology -that is, measuring the fitness levels of the two interacting species under stable conditions. Second, you induce an experimental shift in the timing of one of the species. For example, in a plant-herbivore interaction, germinate the plant earlier and when the herbivore normally has access to the plant, the plant will be older. What are the fitness changes associated with this shift? Finally, you can shift the timing of the other species relative to the first. In our example, the herbivore has access to younger plants and again are there fitness consequences?

Yang and Rudolf call the full combination of possible fitness effects, across a number of timing mismatches, 'the ontogeny-phenology landscape'. By mapping fitness changes across this ontogeny-phenology landscape, researchers can offer better predictions, on top of just changes in range size or habitat use, about the possible affects of climate change. The obvious question, and Yang and Rudolf acknowledge this, is how to extend two-species ontogeny-phenology to multi-species communities. Of course, extending two-species interactions to communities is a question that plagues most of community ecology, but I think the solution is that researchers who know their systems often have intuition about the major players, and thus those species where phenology shifts should have disproportionate effects on other species. Such species could be the place to start. Another strategy would be a food web type approach, where species are lumped into broader trophic groups and we ask how shifts in certain trophic groups affect other groups.

Regardless of how to extend this framework to multispecies assemblages, I see this paper as likely to be very influential. It gives researches a new focus and framework, where specific predictions about climate change can be made.

Yang, L., & Rudolf, V. (2010). Phenology, ontogeny and the effects of climate change on the timing of species interactions Ecology Letters, 13 (1), 1-10 DOI: 10.1111/j.1461-0248.2009.01402.x

Global warming and shifts in food web strucutre

Predicting the effects of global warming on biological systems is of critical importance for informing proactive policy decisions. Most research so far has been on trying to predict shifts in species distributions and changes in interactions within local habitats. But what many of these studies assume is that the basic biological processes and requirements of the individual species will not change -that is their biology is fixed and they simply need to find the place that best suits them. Not so, say Mary O'Connor and colleagues, in a just-released study in PLoS Biology.

O'Connor and colleagues experimentally warmed marine microcosms and tested two alternative hypotheses on food web structure: 1) that productivity increases with warming; and 2) warming increases metabolic rates, thus changing consumer-autotroph (i.e., primary producers) interactions. What they found was that warming indeed altered consumer-autotroph interactions. Warming increased base metabolic rates of consumers, as well as primary production, and the net effect was that food webs shifted towards increasing consumer control (i.e., top-down control).

What this research means is that global warming may alter food web interactions by increasing resource needs of organisms as their metabolic rates increase. This may increase the stress on communities and change diversity patterns as increased needs may shift competitive hierarchies or affect autotroph's ability to withstand consumer effects.

O'Connor, M., Piehler, M., Leech, D., Anton, A., & Bruno, J. (2009). Warming and Resource Availability Shift Food Web Structure and Metabolism PLoS Biology, 7 (8) DOI: 10.1371/journal.pbio.1000178