08 February 2012

Color indicates poison in "poison dart" frogs—honestly!

A strawberry poison dart frog; apparently the San Cristobal color morph. Photo by Wilfredo Falcón.
ResearchBlogging.orgAlmost everyone knows the basic story behind the brilliant coloring of poison dart frogs. These tiny tropical rainforest amphibians secrete toxic alkaloids from their skin, and their bright colors are aposematic signals to warn away potential predators.

You'd expect species that are all sending the same message—Poison! Don't eat!—to use the same signal to do it. Local studies confirm that birds are more likely to attack poison dart frogs who look different from other poison dart frogs in a given area. Yet not all poison dart frogs have the same color pattern, or even similar color patterns. Far from it—frogs within the same species can look completely different.

One possible explanation is that frogs with different coloration are not, in fact, sending the same signal. Brighter color could indicate greater toxicity. That seems to be the case for one highly variable species, the strawberry poison dart frog Dendrobates pumilio. A paper just published as an online, open-access article in The American Naturalist demonstrates that D. pumilio's colors are "honest signals"—and those signals are directed at specific predators.

The many colors of Dendrobates pumilio. Figure 1 from Maan & Cummings (2012).
The new study's authors, Martine Maan and Molly Cummings, selected a study species that is a veritable rainbow of aposemitism, as you can see from the excerpted figure above. Different populations of Dendrobates pumilio are orange, red, green, blue, and yellow, with or without black spots. Maan and Cummings make sense of that colorful diversity in two major ways: first, by finding out whether there's a relationship between color and poison, and second, by making an educated guess about how the different color morphs look to D. pumilio's many predators.

For the first part, Maan and Cummings took an objective measure of color—reflectance spectrum of frogs' skin, measured under standardized lighting—and compared it to an objective measure of toxicity—how much discomfort mice exhibited from an injection of frog skin extract. (The mouse injection method is apparently a standard toxicity assay, and I guess it makes sense if you don't know the specific chemicals that make the frogs poisonous.) The coauthors found a strong relationship between skin brightness and toxicity—frogs with brighter coloring were more poisonous.

Objectively bright coloring isn't quite the same thing as looking bright to a predator, though. Different animals have different color vision—a frog that looks brightly colored to a frog-eating bird might not be particularly showy to a frog-eating snake, because birds and snakes have different suites of sensory cells in their eyes. So the coauthors then fed the spectral readings from the frogs into mathematical models that estimate how the frogs look to different kinds of animal vision. (This approach has been used elsewhere—for instance, to determine how well brood-parasitic cuckoo eggs blend in with their hosts'.) Maan and Cummings applied models based on the visual sensitivity of crabs, snakes, two kinds of bird vision, and frog vision.

Another strawberry poison dart frog, this time the color morph found on Aguacate. Photo by Drriss.
They found strong relationships between the frogs' toxicity and their colors as seen by birds, and as seen by other frogs. The crab vision model varied depending on what kind of material the frog would be viewed against—to a crab, the frogs were conspicuous against bark or leaf litter, but not against green leaves. Meanwhile, the snake vision model didn't perceive any particular relationship between brightness and toxicity. Those results make a lot of sense. Birds are most likely to spot prey from a distance, and make a decision to pursue it or not without getting up close. Crabs aren't likely to encounter frogs up in the foliage, but on the ground, in the leaf litter. And snakes are less likely to rely sight than on chemical senses—taste or olfaction—in evaluating a potential meal.

This study doesn't directly demonstrate the action of natural selection, and that leaves a significant question hanging: Why should Dendrobates pumilio signal its toxicity honestly? Certainly, if you're a highly toxic frog, you'd want to let predators know; but if you're less toxic than the frogs in the next population, why would you tell the world? Indeed, other species of poison dart frogs have evolved mimicry—bright colors without poison.

That suggests the honest coloration within D. pumilio is be due to more than just selection by predators. Perhaps coloration serves social functions, and then more conspicuous color morphs need to be more toxic to fend off more frequent predator attacks. Or there may be genetic constraints that link bright color and toxicity within the species, and both have evolved local differences due to genetic drift. Finding out how selection and other evolutionary forces have created this pattern would be no small project, but I think it'll make an interesting story in the end. ◼

References

Darst, C. (2006). A mechanism for diversity in warning signals: Conspicuousness versus toxicity in poison frogs Proc. Nat. Acad. Sciences USA, 103 (15), 5852-7 DOI: 10.1073/pnas.0600625103

Maan, M., & Cummings, M. (2012). Poison frog colors are honest signals of toxicity, particularly for bird predators. The American Naturalist, 179 (1) DOI: 10.1086/663197

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