Yesterday the BBC reported on a study by Sonia Altizer and Andrew Davis, of the University of Georgia, purporting to demonstrate that populations of the monarch butterfly (Danaus plexippus) show differences in wing size that are correlated with whether or not the populations show migratory behavior to overwintering grounds. Intrigued, I went to the Evolution website and read the paper, which is accepted but not yet copy-edited. I found what seemed to be a serious problem with the interpretation—a problem that, had the BBC reporter had some expertise in evolutionary biology—could have been caught, or at least highlighted in the news report. This underscores the recurring problem that science reporters without much formal training in science often report results without giving the proper caveats.
As you probably know (especially if you watched the NOVA program, The Incredible Journey of the Butterflies, which aired a few days ago), some populations of monarch butterflies show bizarre and wonderful migratory abilities. Individuals from the east coast overwinter in Mexico, and those from west of the Rockies migrate to Southern California. The migration is necessary because adult butterflies can’t tolerate cold, but also because their food plants aren’t available in winter.
But this is not a continuous movement of adults from summering grounds to wintering grounds. The adults do fly the entire one-way journey in the fall, but it takes them several generations (each generation lasting 6-8 weeks) to get back to their summering grounds. One of the great mysteries of monarch migration is how they’re able to return to the same summering grounds used by their great-grandparents. What mechanism guides them in the right direction? And how did natural selection produce this directionality? We don’t know the answers to these questions.
Not all populations of monarchs are migratory. Those in warmer areas, like southern Florida, Costa Rica, and Puerto Rica, have no impetus for migrating since the climate is tolerable and food plants continuously available. They stay put all year.
Based on this “dimorphism” among populations, Altizer and Davis reanalyzed old data, originally collected for a study of parasitism, to test the following prediction:
We therefore predicted that monarchs from long-distance migratory populations would have larger and more elongated forewings to increase flight surface area and reduce wingtip-induced drag.
That is, individuals from migratory populations are under strong selection to fly long distances, and thus would evolve wings more suited to this task than those populations that are more sedentary.
I won’t go into all the details of this study, but here’s what they found:
1. Wild-caught individuals from both “eastern migratory” populations (Minnesota, Georgia, and Mexican overwinterers) and “western migratory” populations (California, Utah, Nevada, Washington, and Colorado) were larger than individual from “nonmigratory” populations (Florida, Hawaii, Costa Rica, and Puerto Rico). This verified their prediction (there were also shape differences, but I won’t discuss them here).
How do we know that these are evolved genetic differences rather than purely environmentally-induced differences in body size (correlated with wing size)? After all, we know from laboratory work that insects reared in colder temperatures grow larger than genetically identical insects reared in warmer temperatures. (I’ve done this many times, for instance, with Drosophila.) To answer this, Altizer and Davis reared three groups of butterflies under constant laboratory conditions of food and temperature. They observed:
2. Under these constant conditions, individuals from eastern and western migratory populations still had bigger wings than individuals from nonmigratory populations (unfortunately, they analyzed only one nonmigratory population here: that from south Florida). From this they conclude that the differences between all populations are genetic and represent evolved adaptive differences:
Collectively, these studies suggest that the demands of long-distance flight represent an important evolutionary force operating on the physical characteristics of migratory species.
The BBC report, written by Matt Walker, echoes this conclusion in the report, titled “Supersized monarch butterflies evolved to fly far.”
These “supersized” butterflies have evolved to cope with the demands of long-distance flight.
In contrast, monarchs that live in one place all year have wings that are up to 20% smaller, report scientists in the journal Evolution. . .
Walker gives no caveats in his report. He simply blurbs the paper and gives some quotes from Altizer. The reporter made no attempt to seek out opinions or commentary from other scientists.
Is there anything wrong with that? Well, one thing: there’s another explanation for the results, not depending on migration, that neither the paper nor Walker considers. It is this: it has long been known that if you look at populations of insects from different areas of its range, those from colder locations tend to be larger (both developmentally and genetically) than those from warmer locations. In other words, they conform to Bergmann’s rule, an “ecogeographic rule” that states that the body mass of an animal is positively correlated with the latitude where it lives. In other words, populations from colder areas have bigger bodies.
The classic explanation of this “rule” involves mammals: if you’re living in a colder climate, it’s adaptive to have a larger mass, for the ratio of heat produced (proportional to the cube of a linear dimension, in other words body mass) to heat lost through radiation (proportional to the square of a linear dimension, in other words body surface area) is lower for larger animals. That is, it’s easier to stay warm if you’re bigger. Now this explanation holds only for warm-blooded animals (homeotherms), but we now know that the “rule” is also obeyed by many cold-blooded animals (poikilotherms). I’ve spent a lot of my career documenting this in Drosophila, and it’s clear that, regardless of the species, populations from colder areas evolve larger size. Why this is so in poikilotherms, who don’t produce body heat to keep warm, is an intriguing but unanswered question. But the phenomenon is real.
The apparent problem with Altizer and Davis’s result is this: all the “nonmigratory” populations live in warmer areas than do the “migratory” populations. Therefore, we expect nonmigratory individuals to be smaller than migratory individuals (i.e., have smaller wings), even if there were no difference in migration behavior. (This is aside from the fact that the authors draw sweeping conclusions about genetic differences from comparing only two migratory populations with only a single nonmigratory population.)
Now the authors don’t discuss this potential problem, which I think is serious. The reviewers of the Evolution paper should have caught it. Nor does the BBC highlight it. My verdict on the paper: it’s intriguing but nowhere near conclusive, and should have been reviewed more thoroughly.
I may be wrong in this conclusion, and perhaps the authors will point out my error. And of course further work may show that they’re correct about a correlation between migration and wing size. But in the meantime, it highlights an apparent breakdown in not only reviewing papers (which has grown more cursory with the exponentially increasing submissions reflecting both the existence of more scientists and the greater pressure on scientists to publish more), but also in the tendency of science reports to avoid looking too hard at research that produces interesting conclusions.
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Altizer, S., and A. K. Davis. 2010. Populations of monarch butterflies with different migratory behaviors show difference in wing morphology. Evolution, in press.


