Winds under the Rain Forest Canopy: The Aerodynamic Environment of Gliding Tree Frogs

Author(s) :Michael G. McCay Source: BIOTROPICA, 35(1):94-102. 2003. Published By: The Association for Tropical Biology & Conservation DOI: 10.1646/0006-3606(2003)035[0094:WUTRFC]2.0.CO;2 URL: http://www.bioone.org/doi/full/10.1646/0006-3606%282003%29035%5B0094%3AWUTRFC %5D2.0.CO%3B2

BioOne (www.bioone.org) is a a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOnes Terms of Use, available at www.bioone.org/page/terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder.

BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research.

BIOTROPICA 35(1): 94102

2003

Winds under the Rain Forest Canopy: The Aerodynamic Environment of Gliding Tree Frogs1
Michael G. McCay 2 University of CaliforniaBerkeley, Department of Integrative Biology, Berkeley, California 94720-3140, U.S.A.

ABSTRACT
Movement of air under the canopy of a forest affects the gliding of animals such as frogs, snakes, geckos, and squirrels; the dispersal of pollen, seeds, and spores; as well as convective transport of heat and carbon dioxide. Wind speed proles were measured under the canopy of a lowland rain forest during the morning, afternoon, and night at three sites in Costa Rica to determine the aerodynamic environment in which tree frogs maneuver while gliding. During the course of a day, average and maximum wind speeds were highest in the morning and midday, and lowest at night. Wind speeds under the canopy were highest near the top of the canopy and were lowest near the canopy oor in the morning and afternoon, and exhibited little variation with respect to height at night. Turbulence intensity (a common measure of gustiness) was constant (ca 1) for all times of day, heights in the canopy, and sites, but the absolute magnitudes of wind gust speeds were higher during the day than at night. Power spectral densities revealed that most of the variation in wind speeds occurred at frequencies that could potentially affect the gliding of tree frogs. Tree frogs (and many other gliding animals), however, glide at night and thereby avoid the higher wind speeds that occur by day. Computer simulations of the dynamic motions of frogs while gliding revealed that the night levels of wind gusts have little effect on the direction of gliding of tree frogs. Key words: Agalychnis; biomechanics; Costa Rica; gliding; La Selva; spectral analysis; tree frog; tropical wet forest; turbulence; wind speed.

TREETOPS ARE HOME TO A WIDE DIVERSITY OF ANIMALS that utilize gliding for low-energy transport within the canopy: mammals such as gliding squirrels (Hanski et al. 2000), reptiles such as Draco lizards (Mori & Hikida 1994) and gliding snakes (Socha 2002), and amphibians such as gliding tree frogs (Scott & Starrett 1974, Emerson & Koehl 1990, Roberts 1994). Because gliding is passive (i.e., does not involve active apping), once a gliding animal is airborne it is at the mercy of the airow around it. Wind gusts (caused by air turbulence) could potentially affect whether or not a gliding animal arrives at its intended target. Thus, turbulent airow within a canopy may have been a factor in the evolution of gliding within the animal groups described above. For taxa of gliding animals that evolved in environments characterized by turbulent airow, natural selection may have favored either active steering or high levels of aerodynamic stability (Maynard Smith 1952). Gliding tree frogs are among animals that have evolved the ability to glide within an arboreal can1 Received 24 February 2002; revision accepted 11 November 2002. 2 Current address: University of CaliforniaDavis, Division of Ecology and Evolution, One Shields Avenue, Davis, California 95616, U.S.A.; e-mail: mgmccay@ ucdavis.edu

opy. Gliding has evolved independently in at least two families of tree frog, Hylidae and Rhacophoridae (Emerson & Koehl 1990, Duellman 2001). Within each family are extant species spanning a full range of aerodynamic abilities from basal nongliding species to derived gliding species that descend aerially from their habitat in the rain forest canopy down to mating sites over temporary pools on the forest oor. The most derived frogs within Agalychnis, a genus from the family Hylidae (Duellman 2001), possess morphological features (enlarged, extensively webbed hands and feet) and behavioral features (gliding posture) that are associated with enhanced gliding performance (Savile 1962, Rayner 1981, Emerson & Koehl 1990). Two aspects of gliding performance thought to be profoundly affected by morphological transformations among tree frog species are aerodynamic stability and maneuverability (Emerson & Koehl 1990). Aerodynamic stability is the ability of a ier to stay on course passively in the presence of random wind gusts. Maneuverability is the ability of a ier to change course actively (i.e., ability to steer) by means of movements of its body or appendages. In a previous study (McCay 2001), I measured aerodynamic forces and torques acting on physical models of tree frogs in a wind tunnel and concluded that morphological transformations among species in the tree frog genus Agalychnis made little
94

Aerodynamic Environment of Gliding Tree Frogs

95

difference in determining aerodynamic stability. All species of Agalychnis studied (a non-gliding species, a species of intermediate glider morphology, and a highly derived gliding species) possessed very low aerodynamic stability, and in some cases were aerodynamically unstable. Therefore, gliding tree frogs are susceptible to being blown off course by wind gusts. Lacking aerodynamic stability, a gliding frog would quickly diverge from its original gliding trajectory if it were hit by a wind gust unless the frog actively steered itself back on course. Thus, to fully understand the consequences of morphological and behavioral transformations in the evolution of gliding in tree frogs, it is critical to understand the nature of the wind environment in the frogs habitat. Characterization of the structure of turbulent airow within forest or crop canopies is an active area of research in micrometeorology. A number of researchers have conducted extensive studies to quantify wind speeds and turbulence that exist within canopies, either by direct measurements (Allee 1926; Odum et al. 1970; Aoki et al. 1978; Amiro 1990a,b; Lee & Black 1993; Kruijt et al. 2000) or by theoretical models (Shen & Leclerc 1997). Within the canopy, turbulent airow is essential to convective transport of heat and carbon dioxide (Campbell 1977, Monteith & Unsworth 1990), transport of insect pheromones (Murlis et al. 1992), and passive dispersal of pollen, seeds, and spores (Aylor 1990, Bullock & Clarke 2000, Cain et al. 2000). The effects of atmospheric turbulence on the trajectories of animals that glide under the canopy remain to be investigated (Fitzjarrald & Moore 1995). Airow within 50 to 100 m of the earth (the surface boundary layer) is generally comprised of turbulent ow that is characterized by random wind gusts superimposed over a mean wind speed (Sutton 1953). Turbulence can be visualized (like smoke swirling from a chimney) as a chaotic collection of irregular swirls of air (eddies) that are convected through space by the overall movement of air (Stull 1988). The formation of turbulence in the surface boundary layer is driven by the strong shearing of airow due to friction from the surface of the earth (the noslip condition). Although the magnitude of turbulence varies depending on local surface properties (a smooth lake surface vs. a rough forest canopy top), turbulence is a ubiquitous feature of airow in the surface boundary layer. Although the chaotic distribution of the eddies that comprise turbulent airow cannot be precisely

determined, turbulent airow does exhibit a universal pattern of eddy size and strength (Mathieu & Scott 2000). A relatively small number of large eddies exists in turbulent ow, which break down into increasingly smaller and smaller eddies until the smallest eddies die out due to atmospheric viscosity (Tennekes & Lumley 1972). Strong winds that occur above the surface boundary layer feed kinetic energy into the largest eddies, and this kinetic energy is gradually dissipated as the largest eddies break down into smaller and smaller eddies. Kinetic energy is supplied to the largest eddies at the same rate that kinetic energy is dissipated by the smallest eddies. Thus, turbulence contains eddies of varying size scales and timescales (long-lived large eddies vs. short-lived small eddies) distributed randomly in three-dimensional space. Crowns of trees in a forest alter the airow under the canopy in several ways. Fast-moving airow skims over the top of forest canopies and avoids the resistance offered by leaves and branches that form the canopy (Monteith & Unsworth 1990). Additional turbulence is formed in this skimming airow due to the rough outer surface formed by the canopy top. Eddies contained in the turbulent airow skimming over the canopy occasionally enter the canopy; however, the resistance of the canopy to outside airows acts as a lter so that only the largest eddies actually penetrate the volume underneath the canopy (Fitzjarrald & Moore 1990). Thus, airow under the canopy is typically much slower and comprised of a higher proportion of large eddies than airow above the canopy (Fitzjarrald et al. 1990). Turbulence may affect animals that glide under the canopy in several ways. Long-acting gusts (due to larger eddies) may blow the gliding animal off course throughout the duration of a glide (Fig. 1a), but the ier may be able to adjust its initial direction of gliding to compensate. Shorter gusts (due to smaller eddies) may also blow the gliding animal off course, especially if the gust hits after the gliding animal is airborne and unable to compensate. Eddies that are of similar size to the gliding animals own body may induce extreme ow asymmetries on the ier by inducing local velocities that are opposite in direction on different regions of the body of the ier (Fig. 1b). Flow asymmetries on the gliding animal may induce unwanted rotations of its body with respect to its direction of gliding (Etkin 1972) that may prove difcult to correct, depending on the animals aerodynamic stability and maneuverability (McCay 2001). To understand the effects of turbulence on animals gliding

96

McCay

MATERIALS AND METHODS

WIND SPEED MEASUREMENTS. Wind speeds were measured under the canopy of lowland wet rain forest at the La Selva eld station of the Organization for Tropical Studies (OTS) in northeast Costa Rica. La Selva is home to three species of Agalychnis tree frog: A. callidryas (a basal non-gliding tree frog), A. saltator (a frog of intermediate gliding ability), and A. calcarifer (a derived gliding frog). Wind speeds were measured at three different localities: two treefall gaps used as mating sites by A. calcarifer and a mating site (Research Swamp) shared by A. callidryas and A. saltator (Roberts 1994). Agalychnis calcarifer sites were identied by the presence of A. calcarifer eggs hanging over pools in the fallen trees, a hallmark of this species (Duellman 2001). Wind speeds were measured during two time periods: 6 July 199916 July 1999 and 28 July 20001 August 2000. Both of these periods coincided with the peak of the Agalychnis mating season when the frogs are gliding (Duellman 2001).
FIGURE 1. Tree frogs glide through turbulent airow comprised of eddies of air of random size, spatial location, and duration. The consequences of turbulent airow on the trajectories of gliding frogs depend on the size of the eddy relative to the size of the frog. (a) Large eddies that induce gusts of longer duration (Tennekes & Lumley 1972) may blow the gliding frog off course for the entire duration of the glide, but the animal may be able to compensate by adjusting its initial direction of gliding. (b) Eddies that are of similar size to that of the frog may induce airow asymmetries on different regions of the body of the frog that in turn may induce unwanted rotations of the frogs body. The unwanted rotations may be minimized using either active control (maneuverability) or passive aerodynamic stability (McCay 2001).

under tree canopies, it is imperative to know the size, intensity, and duration of eddies in the airow under the canopy. The objectives of this study were to quantify the magnitude of wind speeds and the frequency of wind gusts within the canopy through which Agalychnis tree frogs glide. In particular, this study addressed the following questions: (1) What is average wind speed and maximum wind speed that a tree frog encounters while gliding through the canopy? (2) what is the turbulence within the canopy through which tree frogs glide? and (3) how frequent are wind gusts in the canopy? Do they occur often enough to affect a frog gliding in the canopy?

WIND SPEED DATA ACQUISITION. Wind speeds were measured using a hot-wire anemometer (Kurz 443M, Kurz Instruments, Inc., Monterey, California) suspended in a waterproof housing (except for the hot-wire probe) on a static climbing rope slung over a tree branch near the top of the canopy. Wind speed data from the anemometer were recorded by a laptop PC (Toshiba 2545CDS, Toshiba America Information Systems, Inc. with DAQCard-1200 digital acquisition card, National Instruments Corporation, Austin, Texas). The anemometer was calibrated in a wind tunnel with continuously variable speed output up to 18 m/sec. At each site, the anemometer was suspended at heights of 2.5, 5, 10, and 20 m above the forest oor and was tied off to surrounding trees with several nylon kite strings to prevent movement (swaying or twisting) while taking data. At Research Swamp, the maximum height was 18.5 m due to the local canopy structure. Wind velocity data were taken at each height in 15-minute periods at a sampling rate of 60 Hz. Measurements at each height were taken at three times of day: afternoon (13001700 h), night (20000200 h), and the next morning (0500 1100 h). Two complete data sets (wind measurements at all heights and all times of day) were recorded at Research Swamp, and one complete data set was recorded at each of the two A. calcarifer mating sites.

Aerodynamic Environment of Gliding Tree Frogs

97

formation about the strength and size of eddies contained in the turbulent airow underneath the rain forest canopy. WIND SPEED DATA REDUCTION. Statistics were calculated for each 15-minute wind measurement at each of three locations, three times of day, and four heights in the canopy. Summary statistics. Average wind speed was calculated by taking the average of all samples of a wind measurement. Maximum speed was dened as the largest magnitude of wind speed during each 15-minute wind record. Turbulence intensity (i), a measure of overall gustiness of the wind, was calculated using the following formula (Lee & Black 1993): i /u, where standard deviation of wind speed and u mean wind speed. Data from the entire 15-minute record of wind speed were used to calculate and u. Turbulence intensity quanties how much wind speed varies from its mean as a percentage of its mean wind speed. Because the variation is caused by eddies in the airow, turbulence intensity calculates the effect of all eddies combined on overall airow. Turbulence intensity, however, does not calculate how much variation is due to smaller eddies versus larger eddies. To determine the contributions of different-sized eddies to overall gustiness of the winds, other statistical techniques must be used. Power spectral density. Power spectral density (PSD) describes how the power (or variance) of a time series is distributed with frequency. In the case of wind speed measurements, PSD measures how much variation due to wind gusts occurs in the wind speed record at each frequency. The total of the PSD at all frequencies yields total variance for the wind speed record. Low frequency PSD values measure the variance in wind speed caused by larger eddies (longer wind gusts) and high frequency PSD values measure the variance in wind speed caused by smaller eddies (shorter wind gusts). Power spectral densities were calculated for each wind speed record using the Signal Processing Toolbox for MATLAB software (MathWorks, Inc.). MATLAB uses Welchs method to estimate PSD (Welch 1967). In Welchs (1967) method, fast Fourier transforms (FFT) are performed on detrended data in overlapping windows (subsets of wind measurements); periodograms are formed on each window by scaling the magnitude squared of each transform; and then periodograms of all overlapping windows at each frequency are averaged to yield the PSD (Stoica & Moses 1997).

FIGURE 2. Two examples (1-min segment of the 15 min measurement) of the wind speed measurements taken at a height of 20 m in the canopy over Agalychnis calcarifer site no. 1 at (a) night and during the (b) morning. The night wind speeds were consistently lower than the day wind speeds (ANOVA, P 0.05). In both cases, variation in wind speed occurred at several different frequencies. The variation, however, was of much lower absolute magnitude at night than during the day.

Figure 2 shows a representative sample of wind speed data plotted as a function of time taken at A. calcarifer site no. 1 at 20 m above the forest oor during the night (Fig. 2a) and in the morning (Fig. 2b). In both cases, the data exhibit features of turbulent ow: random uctuations of many different frequencies superimposed on a mean ow speed. Fluctuations in velocity are caused by eddies of air passing by the anemometer (Stull 1988). Large eddies induce low frequency uctuations (i.e., longer wind gusts), and small eddies induce high frequency uctuations (Mathieu & Scott 2000). Thus, wind speed data provide essential in-

98

McCay

RESULTS
WIND SPEEDS UNDER THE CANOPY. Summary statistics and power spectral densities are presented below for A. calcarifer mating site no. 1, a site that was representative of all sites measured in this study (three-way ANOVA, P 0.05). Summary statistics. Average wind speed is shown plotted as a function of height above the forest oor (Fig. 3a). Morning winds had the highest average wind speeds and night winds had the lowest average wind speeds. Average wind speeds increased with height of the canopy for all sites during the morning and afternoon. At night, average wind speeds were low and did not change with height of the canopy. Maximum wind speed is shown plotted as a function of height above the forest oor measured at A. calcarifer mating site no. 1 (Fig. 3b). Morning and afternoon winds had the highest maximum wind speeds and night winds had the lowest maximum wind speeds. Maximum wind speeds increased with increased height of the canopy for all sites during the morning and afternoon. At night, maximum wind speeds were low and did not change with height of the canopy. Turbulence intensity is shown plotted as a function of height above the forest oor measured at A. calcarifer mating site no. 1 (Fig. 3c). Turbulence intensity was approximately equal to 1 for all times of day, heights in the canopy, and locations, indicating a consistent overall turbulence in the airow throughout the canopy, despite the fact that wind speeds increased by day and decreased by night. A turbulence intensity of 1 means that the standard deviation of the wind speed was of the same order of magnitude as the mean wind speed; however, because turbulence intensity was normalized by mean wind speed, the absolute speeds of wind gusts were faster when average wind speed was higher (as in the morning or afternoon). Spectral analysis. Figure 4 shows power spectral density (PSD) plotted as a function of frequency for night wind gusts (Fig. 4a) and for morning wind gusts (Fig. 4b). In both cases, PSD is highest at frequencies below 0.1 Hz, indicating that most of the variance in wind speed was caused by longer wind gusts (larger eddies). Overall levels of PSD calculated using night wind gusts are lower than PSD calculated using the day wind gusts because day wind gusts had higher maximum wind speeds and higher absolute variance than night wind gusts; however, the shape of the PSD curves for both wind conditions is similar, indicating a similar eddy

structure for these two cases (i.e., a similar distribution of the sizes of eddies). The shapes of PSDs calculated for all other time histories were similar to the shape of the two plots shown in Figure 4.

DISCUSSION
WIND SPEEDS UNDER THE RAIN FOREST CANOPY. Average wind speeds obtained in this study are comparable to results from previous research. During the morning at 20 m height in the canopy, average wind speed was 0.2 m/sec in this study compared to 0.5 m/sec at Pasoh (Malaya; Aoki et al. 1978), 0.33 m/sec at 16.5 m in the El Verde tower in lower montane rain forest (Odum et al. 1970), and 0.2 m/sec on Barro Colorado Island (Panama) at 23 m (Allee 1926). Wind speeds measured at all heights under the canopy at La Selva (Fig. 3) also compare well to average wind speeds measured under the canopy of an Amazonian rain forest (Kruijt et al. 2000) and under the canopy of a pine forest (Lee & Black 1993). Turbulence intensity measured under the canopy at La Selva (ca 1.0) was also comparable to that measured under the canopy of an Amazonian rain forest (ca 1.0; Kruijt et al. 2000). In addition, variation in wind speeds with respect to time of day is consistent with previous measurements. Wind speeds (Fig. 3) at all locations and heights in the canopy were highest in the morning and lowest at night. In the daytime, air conditions are less stable due to the sun heating up masses of air (Monteith & Unsworth 1990), resulting in faster winds above the canopy, and consequently under the canopy. Wind speeds at Pasoh (Malaya) were measured to be 0.5 m/sec in the middle of the day and 0.1 m/sec at night (Aoki et al. 1978). Wind speeds (Fig. 3) were highest near the top of the canopy and declined steadily as height of the canopy decreased. Leaves, branches, and trunks of the canopy understory cause increased resistance to the ow of air as the height decreases to ground level (Monteith & Unsworth 1990). On Barro Colorado Island (Panama), wind speeds also decreased as height above the ground decreased: from 0.2 m/sec at a height of 23 m up in the canopy to 0.02 m/sec at a height of 2 m (Allee 1926). WINDS IN THE CANOPY AND GLIDING TREE FROGS. Tree frogs glide at night and thus avoid the highest speeds of wind gusts that occur during the day (Fig. 3); however, wind gusts occur at a frequency that may affect tree frogs while gliding. Tree frogs take

Aerodynamic Environment of Gliding Tree Frogs

99

ca 35 seconds to glide down from the top of the canopy (estimated from computer simulation of gliding; McCay 2001), and wind gusts that occur at this frequency (0.20.33 Hz) fall within the range of frequencies at which the maximum variation occurs in the wind speeds (Fig. 4). Wind speeds measured in this study were used as part of a computer simulation of the dynamics of gliding tree frogs during ight, taking into account the frogs aerodynamic properties, mass and inertial properties, and wind gusts (McCay 2001). Wind gusts caused the simulated frogs to veer as much as 1 m off course during the day and less than 0.25 m off course at night. Because only 5 15 percent of the tree trunks (obstacles a frog may encounter while gliding) in the vicinity of the mating sites of tree frogs were spaced less than 1 m apart (McCay 2001), wind gusts did not profoundly affect the gliding of tree frogs; however, detailed analysis of the canopy structure is necessary to more fully understand the consequences of wind gusts on the gliding of tree frogs. Because wind speeds and turbulence under forest canopies are similar for rain forests (Odum et al. 1970, Aoki et al. 1978, Kruijt et al. 2000) and pine forests (Lee & Black 1993) throughout the world, gliding animals in other localities face similar challenges with respect to wind gusts. Gliding squirrels (Hanski et al. 2000) and tree frogs (Scott & Starrett 1974, Emerson & Koehl 1990, Roberts 1994) glide at night when wind speeds are low. Reptiles, however, such as Draco lizards (Mori & Hikida 1994) and gliding snakes (Socha 2002) glide by day when wind speeds are higher. One factor that is strongly related to gliding performance is wing loading, the ratio of wing area to the weight of the animal. Animals with high wing loading (high body weight relative to wing area) are thought to be less maneuverable (Rayner

FIGURE 3. (a) Average wind speeds, (b) maximum wind speeds, and (c) turbulence intensity are plotted as a function of height above the forest oor for one location (Agalychnis calcarifer site no. 1). Data are presented for wind speeds measured in the morning (white bars), afternoon (gray bars), and at night (black bars). (a) Average wind speeds were highest in the morning and lowest at

night (pairwise ANOVAs, P 0.05). Average wind speeds increased as height above the forest oor increased for morning and afternoon (pairwise ANOVAs, P 0.05). At night, average wind speeds decreased as height in the canopy decreased from 20 to 5 m, but then increased at 2.5 m (pairwise ANOVAs, P 0.05). (b) Maximum wind speed were lowest at night (KruskalWallis, P 0.05), but did not differ between morning and afternoon (KruskalWallis, P 0.05). There were no signicant differences among different locations for average wind speed (three-way ANOVA, P 0.05), maximum wind speed (KruskalWallis, P 0.05), or turbulence intensity (KruskalWallis, P 0.05).

100

McCay

low wing loading (Rayner 1981, Mori & Hikida 1994). The wing loading of Agalychnis tree frogs was low in magnitude (50 Newtons/m2) compared to the range of gliding squirrels (30110 Newtons/ m2; Thorington & Heaney 1981), and tree frogs glide through closed canopy (Scott & Starrett 1974, Roberts 1994) in agreement with previous results. Because tree frogs possess low wing loading and low aerodynamic stability (McCay 2001), it appears that tree frogs utilize active steering to stay on course while gliding in the canopy. Much, however, remains to be determined about the physics of animal gliding. In particular, the aerodynamic mechanisms of active steering in the context of the canopy environment (spatial distribution of obstacles and wind gusts) and the behavior of the animal (active adjustment of limb postures) will prove to be a productive direction for future research.

ACKNOWLEDGMENTS
FIGURE 4. Power spectral densities (PSD) plotted as a function of frequency for typical wind data measured at (a) night and in the (b)morning. Both spectra show the same shape with respect to frequency; the highest PSDs are evident at frequencies at or below 0.1 Hz, indicating that wind gusts that caused variation in wind speeds occur at frequencies of 0.1 or below. The maximum PSDs shown for night wind data (Fig.2b) are much lower than morning wind data, indicating that there was more overall variance in morning wind speeds. I gratefully acknowledge the generous gifts bestowed upon me by my thesis advisor M. A. R. Koehl: limitless time spent with me designing experiments, sorting out the logistics of eldwork in the tropics, advising me on data analysis and interpretation, editing and reediting a mountain of thesis drafts, and generally facilitating my transformation from an engineer to a biologist; nancial support (National Science Foundation grant OCE9907120 and OCE-9217338 and Augmentation Award for Science and Engineering Research Training, Ofce of Naval Research, N00014-97-1-0726); and the courageous loan of an incredible array of equipment. I would also like to thank my two eld assistants, George Wang and Jonathon Fong, for their efforts and good humor while enduring clouds of mosquitos for the sake of one more data run and for their expertise in keeping the equipment up and running. I thank Tim Cooper and George Wang (again) for applying their technical wizardry to help build the anemometer housing and program the data acquisition software. I also thank Harry Greene, who rst encouraged me to pursue eldwork in the tropics, and all personnel at OTS and La Selva station, who greatly facilitated the process of obtaining permits, travel, and housing for research in Costa Rica. Field gear and travel expenses were nanced through the generosity of the following grants: American Society of Biomechanics Student Grant-in-Aid, Sigma Xi National Chapter Grant-in-Aid, Sigma Xi Berkeley Chapter Student Grant-in-Aid, and two UCBerkeley Student Summer Research Grants. This manuscript was greatly improved by the thoughtful comments and suggestions of two anonymous reviewers and Dr. Laurie Vitt.

1981) but also less likely to be affected by air turbulence (Thorington & Heaney 1981). In addition, iers that possess high wing loading have higher gliding speeds than iers with low wing loading (Rayner 1981). Wing loading appears to be more strongly related to maneuverability than to resisting turbulence. In a previous study, nocturnal gliding squirrels with a high wing loading ew in open canopy, and gliding squirrels with a low wing loading ew in closed canopy where higher maneuverability is required (Jackson 1999). Similar results were obtained for bats (Norberg 1994). Draco lizards (that glide by day) glide in closed canopy and possess a

LITERATURE CITED
ALLEE, W. C. 1926. Measurement of environmental factors in the tropical rain forest of Panama. Ecology 7: 73302. AMIRO, B. D. 1990a. Comparison of turbulence statistics within three boreal forest canopies. Boundary-Layer Meteorol. 51: 99121.

Aerodynamic Environment of Gliding Tree Frogs

101

. 1990b. Drag coefcients and turbulence spectra within three boreal forest canopies. Boundary-Layer Meteorol. 52: 227246. AOKI, M., K. YABUKI, AND H. KOYAMA. 1978. Micrometeorology of Pasoh forest. Malay. Nat. J. 30: 149159. AYLOR, D. E. 1990. The role of intermittent wind in the dispersal of fungal pathogens. Annu. Rev. Phytopathol. 28: 7392. BULLOCK, J. M., AND R. T. CLARKE. 2000. Long distance seed dispersal by wind: measuring and modelling the tail of the curve. Oecologia 124: 506521. CAIN, M. L., B. G. MILLIGAN, AND A. E. STRAND. 2000. Long-distance seed dispersal in plant populations. Am. J. Bot. 87: 12171227. CAMPBELL, G. S. 1977. An introduction to environmental biophysics. SpringerVerlag, New York, New York. DUELLMAN, W. E. 2001. The hylid frogs of Middle America, vol. I. Contributions to herpetology 18. Society for the Study of Amphibians and Reptiles, Ithaca, New York. EMERSON, S. B., AND M. A. R. KOEHL. 1990. The interaction of behavioral and morphological change in the evolution of a novel locomotor type: ying frogs. Evolution 44: 19311946. ETKIN, B. 1972. Dynamics of atmospheric ight. John Wiley and Sons, New York, New York. FITZJARRALD, D. R., AND K. E. MOORE. 1990. Mechanisms of nocturnal exchange between the rain forest and atmosphere. J. Geophys. Res. 95: 16,83916,850. , AND . 1995. Physical mechanisms of heat and mass exchange between forests and the atmosphere. In M. D. Lowman and N. M. Nadkarni (Eds.). Forest canopies, pp. 4572. Academic Press, San Diego, California. , , O. M. R. CABRAL, J. SCOLAR, AND L. D. DE ABREU SA. 1990. Daytime turbulent exchange between the Amazon forest and the atmosphere. J. Geophys. Res. 95: 16,82516,838. HANSKI, I. K., P. C. STEVENS, P. IHALEMPIA, AND V. SELONEN. 2000. Home-range size, movements and nest-site use in the Siberian ying squirrel, Pteromys volans. J. Mammal. 81: 798809. JACKSON, S. M. 1999. Glide angle in the genus Petaurus and a review of gliding in mammals. Mammal. Rev. 30: 9 30. KRUIJT, B., Y. MALHI, J. LLOYD, A. D. NOBRE, A. C. MIRANDA, M. G. PERIERA, A. CULF, AND J. GRACE. 2000. Turbulence statistics above and within two Amazon rain forest canopies. Boundary-Layer Meteorol. 94: 297 331. LEE, X., AND T. A. BLACK. 1993. Turbulence near the forest oor of an old-growth Douglas-r stand on a southfacing slope. For. Sci. 39: 211230. MATHIEU, J., AND J. SCOTT. 2000. An introduction to turbulent ow. Cambridge University Press, Cambridge, England. MAYNARD SMITH, J. 1952. The importance of the nervous system in the evolution of animal ight. Evolution 6: 127 129. MCCAY, M. G. 2001. The evolution of gliding in Neotropical tree frogs. Ph.D. dissertation. University of California, Berkeley, California. MONTEITH, J. L., AND M. H. UNSWORTH. 1990. Principles of environmental physics. Chapman and Hall, New York, New York. MORI, A., AND T. HIKIDA. 1994. Field observations on the social-behavior of the ying lizard, Draco volans sumatranus, in Borneo. Copeia 1994: 124130. MURLIS, J., J. S. ELKINTON, AND R. T. CARDE. 1992. Odor plumes and how insects use them. Annu. Rev. Entomol. 37: 505532. ODUM, H. T., G. DREWRY, AND J. R. KLINE. 1970. Climate at El Verde, 19631966. In H. T. Odum and R. F. Pigeon (Eds.). A tropical rain forest: a study of irradiation and ecology at El Verde, Puerto Rico, pp. 347418. Division of Technical Information, USAEC, Oak Ridge, Tennessee. NORBERG, U. M. 1994. Wing design, ight performance and habitat use in bats. In P. C. Wainwright and S. M. Reilly (Eds.). Ecological morphology: integrative organismal biology, pp. 205239. University of Chicago Press, Chicago, Illinois. RAYNER, J. M. V. 1981. Flight adaptations in vertebrates. Symp. Zool. Soc. Lond. 48: 137172. ROBERTS, W. E. 1994. Explosive breeding aggregations and parachuting in a Neotropical frog, Agalychnis saltator (Hylidae). J. Herpetol. 28: 193199. SAVILE, D. B. O. 1962. Gliding and ight in the vertebrates. Am. Zool. 2: 161166. SCOTT, N. J., JR., AND A. STARRETT. 1974. An unusual breeding aggregation of frogs, with notes on the ecology of Agalychnis spurrelli (Anura: Hylidae). Bull. South. Calif. Acad. Sci. 73: 8694. SHEN, S., AND M. Y. LECLERC. 1997. Modelling the turbulence structure in the canopy layer. Agric. For. Meteorol. 87: 325. SOCHA, J. J. 2002. Gliding ight in the paradise tree snake. Nature 418: 603604. SOLARI, G. 1993. Gust buffeting. I: Peak wind velocity and equivalent pressure. J. Struct. Engrg. 119: 365382. STOICA, P., AND R. L. MOSES. 1997. Introduction to spectral analysis. Prentice Hall, Englewood Cliffs, New Jersey. STULL, R. B. 1988. An introduction to boundary layer meteorology. Kluwer Academic, Boston, Massachusetts. SUTTON, O. G. 1953. Micrometeorology, a study of physical processes in the lowest layers of the Earths atmosphere. McGrawHill, New York, New York. TENNEKES, H., AND J. L. LUMLEY. 1972. A rst course in turbulence. MIT Press, Cambridge, Massachusetts.

102

McCay

THORINGTON, R. W., AND L. R. HEANEY. 1981. Body proportions and gliding adaptions of ying squirrels (Petauristinae). J. Mammal. 62: 101114. WELCH, R. D. 1967. The use of fast Fourier transform for the estimation of power spectra: a method based on time averaging over short, modied periodograms. IEEE Trans. Audio Electroacoust. AU- 15: 7073.