A lizard of the Draco genus suns itself in the Western Ghats of India.

The patagium, a skin membrane supported by six extended ribs, acts as wings for controlled gliding. Berkeley scientists have determined that larger Draco species are still capable of gliding, despite unfavorable surface-to-mass ratios, by evolving larger patagia.

Easy Glider
Berkeley scientists learn what it takes to y without wings
PHOTO BY JAYANTH SHARMA, WILDLIFEIMAGES.COM

here is an artor rather, a knack to ying, says Douglas Adams, the British writer and radio personality famous for his Hitchhikers Guide to the Galaxy. The knack lies in learning how to throw yourself at the ground and miss. Curiously, Adams denition leaves out one element usually thought of as essential to yingwings. While his intentions were clearly humorous, Adams was perhaps more correct than he realized: from squirrels and lizards to some tropical ant species, a surprising number of wingless creatures do have a knack for avoiding the ground. They glide. Today, researchers at UC Berkeley are studying gliding organisms and accumulating clues as to how and why nature favored

this adaptation. Not until recently, with the advent of high-speed, high-resolution video cameras, was it possible to observe the movement of individual body parts on an airborne animal or insect. Armed with these innovative techniques, researchers are now looking at the specic morphologies and movements of gliders. Such research is elucidating the biomechanics of aerial behavior in real time, in turn providing insight into the evolutionary patterns behind gliding ight. LEAPIN LIZARDS A glider is like a plane without an engine, or a bird without the ability to ap its wings. But like powered yers, good gliders can move a signicant distance horizontally

without losing much height. To do so, they maximize the surface area of their bodies and adopt specic shapes, or morphologies, to generate lift that counteracts the downward pull of gravity. Researchers at Berkeley are studying how body morphology contributes to gliding ight, and how such morphologies might have evolved in the rst place. To study the evolution of ying in a lineage, scientists typically look at the phylogeny of that groupthat is, they try to nd its common ancestor. Next, they determine the groups functional morphology, how it has made ight possible, and how it has evolved over time. Finally, they scour the fossil record for empirical evidence to support their ndings and to formulate a theory of why ight

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evolved in that particular lineage. While this conventional evolutionary approach can be useful, when it comes to Draco lizards, a genus of gliding arboreal lizards from Southeast Asia, it cant be fully applied. Ranging in mass from three to thirtyve grams (from the weight of, say, a pencil to a plum), Dracos are the only living lizards to have evolved elongated ribs; these ribs support a wing-like ap of skin, the patagium, which provides the surface area needed to glide. Unfortunately for researchers interested in the evolution of these traits, there is no fossil record for this group (or any extant lizards with a similar morphology), making it difcult to chronicle how their ability to glide evolved. Fortunately, by focusing instead on how variation in Draco body morphology affects gliding behavior, Berkeley professors of integrative biology Jim McGuire and Robert Dudley have been able to ll in some details about how ight might have evolved within this group. When McGuire began studying Draco in 1995, he was intrigued by the relationship between its body size and gliding performance. His interest was piqued in part by the groups incredible diversication and coexistence: multiple species of Draco were found in the same habitat. It made sense that to coexist, different species must use different types of limited resources in order to prevent competition. Since the only major difference across the various species was body size, McGuire wondered whether size played a role in the type of resources each species consumed, and whether this had anything to do with the lizards gliding abilities. At that point, he teamed up with Dudley, and for the next several years they measured the gliding performance of twenty-nine species of remote Malaysian Draco ranging across the full size distribution. They knew from the outset that Dracos scale isometricallythat is, as their body mass increases, their proportions stay the same. Thus, their body mass increases at a greater rate than their surface area. Since surface area is critical to generating lift, one would expect the big lizard species to be relatively poor gliders compared to the smaller ones. To conduct gliding trials, McGuire and
A Draco melanopogon lizard glides down from a 6-meter pole as part of a flight experiment. Berkeley scientists can use data on the wind speed and animals velocity to determine how the lizard directs its downward flight.

Dudley erected two tall poles 9.3 meters aparta six meter takeoff pole and a four to ve meter landing pole. They positioned the lizards on the takeoff pole and then prodded them, either by agitating them with a bamboo rod or tapping the pole, so theyd glide toward the landing pole. A video camera recorded each trial, and by taking into account the position of the camera relative to the poles, McGuire and Dudley were able to calculate the approximate height lost over a standard horizontal glide distance and the angle of each glide. They also used a digital anemometer to measure wind speed and a compass to determine wind direction. Together, these measurements allowed them to determine the velocity of each glide adjusted for wind conditions.

Evolutionary deductions can also be made from the data. According to McGuire, the original Draco glidersthe common ancestor of those living todaywere probably either large and living in really tall trees or cliffs from where they could execute steep ballistic dives, or they were fairly small so they could take advantage of their higher surface area to body mass ratio. They could probably expand their ribs just a little bit to create more surface area for increased lift, but then were favored by selection such that their patagia got larger and larger over many generations. In the absence of a fossil record, such evolutionary hypotheses establish a framework for understanding how Draco may have started gliding. THE ANTS GO GLIDING ONE BY ONE Another striking example of a glider that may shed light on the evolution of aerial behavior is Cephalotes atratus, a species of wingless ants living in the tropical rainforest canopy. Remarkably, these ants are the rst documented case of wingless ight in any living insect. Three years ago, Robert Dudley listened as his colleague Steve Yanoviak, a University of Florida entomologist, told him hed discovered something amazing in the jungles of Panama. Hed found that wingless ants, when icked off high tree branches, glided back to the trunk of the tree from which theyd fallen rather than spiraling aimlessly to the rainforest oor. Yanoviak said hed witnessed the gliding behavior again, in the same ant species, in Peru. Since there were no documented cases of gliding ight in a living wingless insect, the discovery indeed seemed novel. In 2004, Dudley joined Yanoviak and

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If youre an ant falling from the rainforest canopy, you dont want to land on your head.
It turns out that the large Draco compensate for their extra mass by diving faster and farther: the rst part of their glide is a steep ballistic dive to generate velocity, and then they level out into a at glide. Practically speaking, though, the larger Draco are more limited than smaller ones in their access to resources within the forest. Since they need a higher initial altitude to perform a successful glide, they are likely to remain in the upper reaches of the rainforest canopy, whereas the smaller Draco may scavenge far below. This partitioning of the forest habitat may in turn explain how multiple Draco species can coexist: by living in different vertical layers of the forest, they draw on different resource pools.
PHOTO BY: JIM MCGUIRE

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PHOTO BY STEVE YANOVIAK, UNIVERSITY OF OKLAHOMA

The flight of the wingless ant Cephalotes atratus may shed some light on the evolution of gliding in insects and other animals. Slow-motion video revealed that the ants use their limbs while airborne to control their fall.

Michael Kaspari, a University of Oklahoma entomologist, in studying this phenomenon. How were the ants maneuvering themselves, they wondered? What about the ants morphology facilitated aerodynamic behavior? And what could it reveal about the evolution of insect ight? Dudley, Yanoviak, Kaspari, and others spent the next year or so using high-speed, high-resolution video cameras to document how wingless ants in the Peruvian rainforest glide. They used a rope and sit harness to pull themselves up to high, treetop branches. Sometimes they climbed with ants theyd collected elsewhere in the forest, and other times they resolved to nd a colony once they reached the upper reaches of the tree. They painted the backs of the ant bodies white (for visibility against the dark forest background), dropped them from trees, and then recorded the ants descent on video camera. The painted ants repeatedly glided back to the tree trunk. Upon further study, it turns out that these ants do what skydivers dothey use their limbs to control their posture and orientation. Their legs are slightly attened at the ends, so like skydivers who can initiate a spin by slight right and left asymmetry of hand or arm posture, the ants can initiate torque that will rotate their bodies. And its not just one ant species doing this. Dudley says, its hundreds, and theyre

not all related to each other. Nor are ants the only wingless insects gliding. Biologists are now nding that many species across all insect groups are wingless gliders. They dont have wings and never haveand yet, they glide. The evolutionary pressure to adopt gliding ight may have been as simple as accidentally falling out of a tree. Or, as Dudley puts it, If youre an ant falling from the rainforest canopy, you dont want to land on your head. You want to control your descent, try to land on something so you dont break your antennae or another body part. Put this way, gliding ight sounds like a reex, the body and limbs responding to the force of gravity by creating aerodynamic surfaces that produce lift and adjust to drag. Directed aerial descent has another possible advantage as wella falling ant may be able to return to its colony as opposed to being woefully lost on the forest oor far below. Given how common gliding behavior is throughout insect species, its natural to ask whether these gliders t somewhere on the evolutionary continuum between land-dwelling and ying insects. In other words, did todays ying insects evolve from gliding ancestors? Dudleys hypothesis is that wingless gliding ight preceded winged ightthat aerodynamic behavior in insects started without wings. Admittedly, ight without wings sounds a bit like putting the chicken before the egg, but research on

gliding by Dudley and others has denitively shown that its possible. Furthermore, the evolutionary history of insects offers some evidence that ying insects evolved from gliding ancestors. According to Dudley, the most primitive insects around todaythose which most closely resemble the earliest insects on earthare the Silversh (Thysanura) and Bristletails (Archaeognatha). Both of these groups lack wings, and the Silversh can glide, just like the Peruvian rainforest ants. This suggests that all modern winged insects may have evolved the ability to y from wingless, gliding ancestors. So far, however, scientists have not yet pinpointed the transitional morphologies that would illuminate the evolutionary steps between skydiving silversh and some of the earliest winged insects. Eventually, by understanding how gliding played a role in the evolution of insect ight, researchers hope to gain insight into the evolution of ight in vertebrates. But in the meantime, they continue to investigate the mechanics of gliding, fascinated by the many speciesinsect, lizard, or otherwisethat throw themselves determinedly at the ground, but somehow, repeatedly, miss.

ADRIENNE DAVICH is a graduate student in journalism.

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