Western Tiger Swallowtail butterfly, and a very close look at butterfly wing-color

We’ve gotten a few terrific photos of butterflies this year—some posted here and here— but none of the swallowtails has cooperated by alighting within range. When I saw one that had died and fallen to the road I carefully carried it home for the chance to get a close look.

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There are at least three very similar species of swallowtail around here—the Anise, Western Tiger, and Oregon Swallowtails. Based on the red and blue markings I’m thinking this is the Western Tiger Swallowtail, Papilio rutulus.

Finer than “frog hair”—butterfly hair!

Enlarging the macro photos shows details such as hairs on the body and along the inner edges of the wings.

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These hairs, called tactile setae, are attached to nerve cells, which relay information about the hairs’ movement to the butterfly. … Adults have tactile setae on almost all of their body parts. In both adults and larvae [caterpillars], the setae play an important role in helping the butterfly sense the relative position of many body parts (e.g., where is the second segment of the thorax in relation to the third segment). This is especially important for flight, and there are several collections of specialized setae and nerves that help the adult sense wind, gravity, and the position of head, body, wings, legs, antennae, and other body parts. In monarchs, setae on the adult’s antennae sense both touch and smell. (from monarchwatch.com).

In the photo below, a ventral view of the lower wings where they meet at their lowest point, there is also a delicate fringe visible along the edges. This could have aerodynamic as well as sensory functions.

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From pointillism to nanostructures

Parts of the markings that appear as solid areas to our eye are revealed to be pointillist creations. I suspect we would need to know much more than we do about the vision of butterflies (and their predators?), in order to understand how these markings work for them.

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The odd squareness of the smallest dots of color is not some pixellation in the photo, but an accurate representation. It shows the shape of the overlapping scales which form the surface of butterfly wings. Here are some microphotographs of wing scales at various magnifications, from Wikipedia.

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And here are color microphotographs showing the same squared-off dots along with the underlying scale pattern.

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Picture source.


It’s been known for some time that the colors of butterfly wings are partly from pigments but mostly from the microstructure of the scales, scattering light to produce the colors. Blues, greens, reds and iridescence are usually structural, while blacks and browns come from pigments. (Wikipedia).

But now we know more, and the more we know the more intricate and amazing it is. Research (published this past June) has been able to identify the light-scattering shapes from the wings of several butterfly species, and they are described as “ ’mind-bendingly weird’ three-dimensional curving structures… [resembling] a network of three-bladed boomerangs”. The name for these crystalline forms is gyroids, and they were first described

in 1970 by NASA physicist Alan Schoen in his theoretical search for ultra-light, ultra-strong materials for use in space. Gyroids have what’s known as an ‘infinitely connected triply periodic minimal surface’: for a given set of boundaries, they have the smallest possible surface area. The principle can be illustrated in soap film on a wireframe (see image below). Unlike soap film, however, the planes of a gyroid’s surface never intersect. As mathematicians showed in the decades following Schoen’s discovery, gyroids also contain no straight lines, and can never be divided into symmetrical parts. (source, text and soap-bubble photo: wiredscience.com)

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Gyroid-like soap bubble. Photo from wiredscience.com

So gyroids were introduced to humans as an imagined created form, something that is a mind-boggler for non-mathematicians to envision.

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The image above is a mathematician’s representation of one of the simpler types of gyroid.

Materials scientists have learned how to make synthetic gyroids for photonic devices, such as solar cells and communication systems, that manipulate the flow of light.

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A self-assembled solar cell begins with one of two polymers forming a “gyroid” shape while the other fills in the space around it. The inner polymer is dissolved away to create a mold that is filled with a conductor of electrons. The outer polymer is then burned away, the conductor is coated with a photosensitive dye, and finally the surrounding space is filled with a conductor of positive “holes”. A solar reaction takes place at all the interfaces throughout the material, and the interlocking gyroid structure efficiently carries away the current. (Source for image and caption, Cornell Univ.)

And when Yale evolutionary ornithologist Richard Prum got curious about exactly how butterfly wing-scales twisted light, he found gyroids. His team had to use an advanced microscopy technique with nanoscale resolution, called synchrotron small angle X-ray scattering, in order to see them, but there they were. (See note at end for citation of article in PNAS.)

The butterfly’s gyroids are made of chitin, not exactly the flashy material I would associate with iridescent wings. It’s

the tough starchy material that forms the exterior of insects and crustaceans. Chitin is usually deposited on the outer membranes of cells. The Yale team wanted to know how a cell can sculpt itself into this extraordinary form, which resembles a network of three-bladed boomerangs. They found that, essentially, the outer membranes of the butterfly wing scale cells grow and fold into the interior of the cells. The membranes then form a double gyroid—or two, mirror-image networks shaped by the outer and inner cell membranes. Double gyroids are easier to self assemble but they are not as good at scattering light as a single gyroids. Chitin is then deposited in the outer gyroid to create a single solid crystal. The cell then dies, leaving behind the crystal nanostructures on the butterfly wing.

“Like engineers, butterflies grow their optically efficient single gyroids through a series of steps that make this complex shape easier to achieve. Photonic engineers are using gyroid shapes to try to create more efficient solar cells and, by mimicking nature, may be able to produce more efficient optical devices as well,” Prum said. (Source)

In an interview about the work, Richard Prum said “We’re still trying to wrap our brains around gyroids and what they are.” The shapes seem to have evolved separately in several lineages of butterflies.

”It’s a Swiss cheese,” he adds, “with spiraling channels of air traveling through it that intersect one another. But those channels actually travel in three different dimensions through the cheese, and what you end up with is this very complicated form left behind, and that form is a gyroid.”

And while the idea of butterflies with Swiss cheese wings is slightly strange, Prum says it’s a very useful one for scientist and engineers looking for the next leap forward in electronic technology.

For example, Prum says, take the fiber-optic cables that carry phone calls under the ocean. These cables carry signals in the form of colored light, but it’s very difficult to insulate them well enough to prevent light from leaking out. Current transoceanic cables have to have booster stations built along them to keep the signal strong. But a layer of gyroids around the fiber-optic cable “would act like a perfect insulation to that fiber,” Prum says. The same tiny structures that give the Emerald-patched Cattleheart its lovely green patches could also be used to keep green light from escaping a fiber-optic cable.

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The vivid green color of the scales of this Papilionid butterfly are produced by optically efficient single gyroid photonic crystals. Caption and photo from www.physorg.com

Right now, it’s expensive and impractical to manufacture gyroids small enough to do that job. But butterflies hold the secret to growing them naturally. “If you could grow one, at exactly the right scale, as butterflies do,” says Prum, “you could make these things a lot easier.” (NPR interview, Jul 3, 2010)

This is a fine example of how curiosity can lead us to unexpected discoveries. The original question is one that could be used by certain Congressional anti-intellectuals in their periodic efforts to discredit basic research: “Imagine, all this work to find out what makes the color on butterfly wings! How ridiculous!” The research and technological developments that are thought “useful” by these folks had their origins in someone’s basic research, sparked by human curiosity. From butterfly wing-color to, perhaps, more efficient fiber-optic cables or solar energy collectors. It’s called bioengineering: investigating the functions and structures of nature, to derive principles and patterns for technological innovations. But for me it’s satisfying in itself, the revelation of these marvelous structures, underlying the evanescent beauty of a butterfly.

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Western Tiger Swallowtail butterfly on Buddleia bloom. Photo by terwilliger911, flickr.

Note: The article describing gyroids as the structure causing some colors in butterfly wings is:
Structure, function, and self-assembly of single network gyroid (I4132) photonic crystals in butterfly wing scales.
Vinodkumar Saranathan et al. Proceedings of the National Academy of Sciences. Published online before print June 14, 2010, doi: 10.1073/pnas.0909616107. The abstract is available free, but the article requires purchase or subscription to PNAS. There is a supplementary article here that contains some interesting images and very technical text. There’s even a movie you can watch showing a slice-by-slice trip through a certain sort of gyroid, or as the text says, though “the pentacontinuous volume of a level set core-shell double gyroid structure”.

We brake for butterflies

Butterflies everywhere in the air! so many you have to drive about 5 miles an hour, letting the current of your progress gently push them out of the way. That’s how it was one morning last week, on the paved forest road where we often walk. By 3 pm it would be 100°. Though there were still wildflowers in bloom, these butterflies seemed not to be feeding, but mostly just flying and chasing one another. Breeding season? One did land for a moment on Dan’s finger and another swooped at it aggressively, over and over.

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California Sister butterflies (Adelpha bredowii), ventral view.

As before, in a different location on this road, we saw scores of the California Sister butterfly (Adelpha bredowii) but this time none of the Lorquin’s admiral (Limenitis lorquinii) seen then. Swallowtails were present too, like sunlight in flight, but in small numbers. Unlike the others, the swallowtails never lighted for long either on vegetation or on the road, where the California Sisters clustered to get minerals from visible animal scat or from remains too small for us to see.

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California Sister butterfly, dorsal view, on the road.

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This ant was pulling along the body of a California Sister butterfly. It would move the butterfly an inch or two, then stop and scurry around looking (I thought) for a more effective place to grab on.

Swallowtail butterflies

The swallowtails never let us get close enough for a really good look or photo, and we may even have seen more than one species. Dan, whose eyes are better, says that most were a pale yellow. the others brighter. Of the three found in our area, one is a species called the Pale Swallowtail (Papilio eurymedon) that uses Ceanothus spp. for its larval host plant, and Blueblossom ceanothus (Ceanothus thyrsiflorus) is a common flowering shrub here. Very pretty too, growing to 6 feet or more in height and flowering in varying shades of blue and lilac. Most are past their peak of bloom now, beginning to fade or entirely withered; these photos are from June.

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The British biologist J. S. B. Haldane was engaged in discussion with an eminent theologian. ‘What inference,’ asked the latter, ‘might one draw about the nature of God from a study of his works?’ Haldane replied: ‘An inordinate fondness for beetles.’ Indeed, of the 1.5 million described species on the planet, 350,000 are beetles, more species than in the entire plant kingdom. So I didn’t even try to identify the mating beetles in the photo above, but Dan picked up Insects of the Pacific Northwest (by Haggard and Haggard) and found them easily: Anastrangalia laetifica, the Dimorphic Long-horned Beetle! The female’s red wingcovers are visible on the right side, beneath the male’s all-black back.

This is the Pale Swallowtail, below. [Photo by Franco Folini, from flickr]

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Different life stages of the Pale Swallowtail caterpillar are shown here, and for the Anise Swallowtail here. Caterpillars can have quite different appearances, as they pass through successive moults (stages called instars), and so the one illustrated in your field guide for a given species may not look at all like the one you find.

The other Swallowtails likely to be seen here in Southwestern Oregon are the Anise Swallowtail (Papilio zelicaon) and the Western Tiger Swallowtail (P. rutulus). Oregon’s state insect is the Oregon Swallowtail (P. oregonius, sometimes called P. bairdii) but it’s found in the dry sagebrush canyons of Eastern Oregon and Washington along with its caterpillar host plant Tarragon or Dragon’s-wort (Artemisia dracunculus). Our culinary tarragons are varieties of this same species.

Siskiyou wildflowers and butterflies

Our roadside botanizing was especially exciting today. First perhaps I should explain why we walk along forest service roads instead of hiking along trails. It has a lot to do with a single plant, although not one I would describe as a widlflower.

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Yes, it’s poison oak (Toxicodendron diversilobum), seen above early in the spring before it has reached its full diabolical potential of thickets six feet tall, stretching branches out onto trails in search of sunshine in order to grow even more monstrously large. Poison oak could be an interesting plant: it occurs in various forms from semi-vines threading up tree trunks, to a low-growing ankle-ambusher, as well as the aforementioned woody thickets. But all parts contain a chemical that is—not poisonous—but an extremely powerful allergen, an oil called urushiol. Most people are allergic to it, and I am very very allergic, so once we get off of bare ground I spend most of my time looking down and around before every step in order to find it before it finds me. (Be warned: allergies can come and go, so a history of immunity doesn’t mean you’ll always be immune.)

Happily, there’s an abundance of things to see by walking along the road and making a few careful excursions. Today was a bonanza.

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There’s an audio recording of Lew Welch reading this, here.

I don’t think we saw anything that “nobody’s ever really seen”, although one must pay careful attention to Lew Welch’s language, that “really seen” part. But what we saw was marvelous. Here’s one sight:

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From a distance I thought the butterflies were gathered upon a damp patch improbably located in the middle of the hot dusty gravel road. In other such situations, I haven’t been able to approach very closely without scaring them off. I took some pictures, then moved a bit closer, closer still, and in the end I was kneeling right beside them without really disturbing them at all. And then I could see what it was that they were so attracted to.

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They were on the scat of some animal, not an uncommon territorial marker to find in the middle of these forest roads. Could be fox, raccoon, coyote. Undigested material including seeds and some woody bits (pine needles?) can be seen, and the scat is pretty dry. Unlikely to be a source of moisture. However, butterflies require minerals not found in nectar, and often get these by drinking from damp soil or applying their tongues to scat. I am curious how they get nutrients from dry materials, because their tongues are hollow tubes designed for drinking liquids.

I poured some water on a nearby area before we left in search of lilies. When we came back, all the butterflies were still on the scat.

There were two species there. One was Adelpha bredowii, California sister, shown here exploring my arm. Some photos (here, for example) show this species with blue rather than grey markings, but that may be local variation.

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What’s the “sister” about? It’s said to refer to the black and white markings (like a nun’s habit) on the other side of the wings, the dorsal side (looking down on the outspread wings and the insect’s back, from above).

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Photo source.

The other is Limenitis lorquinii, Lorquin’s admiral. There are several different butterfly species with “admiral” in their names, and the reference is not clear. Some say the names were originally “admirable” but I can find no support, just speculation. Lorquin was a Frenchman in California during the Gold Rush of 1850, who sent butterfly specimens back to France where they were described for the first time by eminent lepidopterist Jean Baptiste Alphonse Dechauffour de Boisduval.

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It is unbelievable to see these creatures in such detail. First, Limenitis lorquini.

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It is possible to see the wing-veins as the three-dimensional structures that they are. When we read that a new butterfly emerging from the chrysalis has to “pump up” its folded wrinkled wings, before they are strong enough to fly, these veins are the means. “The butterfly has to expand and dry [its wings] as soon as it emerges from the chrysalis. To do this, it uses its body as a pump and forces fluid through a series of tube-like veins. It’s a little like inflating a balloon — as the veins fill with fluid, they slowly stretch the surface of the wings.” Source.

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Adelpha bredowii, trailing its long tongue over my skin.

We went on to look at the Washington lilies described in my previous post. The blooms that were white and pink on June 24th,

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today were nearly bright pink and drooping.

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But another plant was in spectacular bloom.

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This is Philadelphus lewisii, commonly called mock orange for its fragrance. To me there was nothing citrus-y about the fragrance, but I’ve never smelled orange trees in bloom. (There are perhaps a dozen other plants also called mock orange, illustrating how treacherous common names can be.) Philadelphus lewisii is one of nearly 200 plants new to science which Lewis and Clark described. Indians used its straight stems in making arrows.

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On the drive back to the main road we saw many more, all in synchrony of bloom. It’s a shrub that can reach 12 feet, so it offers a lot of flowers! We had remarked earlier on how many butterflies were about, in the air: monarchs, tiger swallowtails, and others. Surely the Philadelphus extravaganza had something to do with the sudden abundance of butterflies, and we speculated on how insects and plants keep in step when the music of the dance—the temperature, rainfall, sunny or cloudy skies—can vary so drastically year to year. This long rainy spring was very atypical, yet after three sunny days here are the partners right in step.

Another unusual find will have to wait for my next post. It has something to do with this wild rose…

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