A Life Outside.

How do the morphologies of organisms affect their physical interactions with the environment and other organisms? My research in marine systems couples field studies of the physical habitats, life history strategies, and ecological interactions of organisms with laboratory analyses of their biomechanics. Here, I review how we pursued answers to three questions about marine organisms: (a) how benthic organisms withstand and utilize the water moving around them, (b) how the interaction between swimming and turbulent ambient water flow affects where small organisms go, and (c) how hairy appendages catch food and odors. I also discuss the importance of different types of mentors, the roadblocks for women in science when I started my career, the challenges and delights of interdisciplinary research, and my quest to understand how I see the world as a dyslexic.

Of corpses, ghosts and mirages: biomechanical consequences of morphology depend on the environment.

Organisms are subject to the laws of physics, so comparative biomechanics is a powerful approach for identifying basic principles that apply across taxa of how morphology affects performance of mechanical functions such as locomotion, feeding or resisting damage. Journal of Experimental Biology has been a leading journal for decades in publishing studies revealing such basic biomechanical principles. However, field studies of the physical environment, ecological interactions and life-history strategies of organisms reveal which aspects of their biomechanical performance are important to their success in different types of natural habitats, and thus enable us to design ecologically relevant laboratory experiments to understand biomechanical function. Because the fitness consequences of differences in morphology are affected by the biological and physical environment, biomechanics can be used to identify how physical constraints on the performance of organisms with different body plans in variable environments can affect evolution. I illustrate these points with examples from the literature that show how the biomechanical consequences of morphology depend on the ecology of the organisms. Knowledge of the temporal patterns of interactions of organisms with their physical and biological environments is essential for understanding their functional morphology as it changes during ontogeny, and it reveals constraints on their evolution.

Ecological biomechanics of damage to macroalgae.

Macroalgae provide food and habitat to a diversity of organisms in marine systems, so structural damage and breakage of thallus tissue can have important ecological consequences for the composition and dynamics of marine communities. Common sources of macroalgal damage include breakage by hydrodynamic forces imposed by ambient water currents and waves, tissue consumption by herbivores, and injuries due to epibionts. Many macroalgal species have biomechanical designs that minimize damage by these sources, such as flexibly reconfiguring into streamlined shapes in flow, having either strong or extensible tissues that are tough, and having chemical and morphological defenses against herbivores and epibionts. If damage occurs, some macroalgae have tissue properties that prevent cracks from propagating or that facilitate tissue breakage in certain places, allowing the remainder of the thallus to survive. In contrast to these mechanisms of damage control, some macroalgae use breakage to aid dispersal, while others simply complete their reproduction prior to seasonally-predictable periods of damage (e.g., storm seasons). Once damage occurs, macroalgae have a variety of biomechanical responses, including increasing tissue strength, thickening support structures, or altering thallus shape. Thus, macroalgae have myriad biomechanical strategies for preventing, controlling, and responding to structural damage that can occur throughout their lives.

Age affects the strain-rate dependence of mechanical properties of kelp tissues.

PREMISE: The resistance of macroalgae to hydrodynamic forces imposed by ambient water motion depends in part on the mechanical properties of their tissues. In wave-swept habitats, tissues are stretched (strained) at different rates as hydrodynamic forces change. Previous studies of mechanical properties of macroalgal tissues have used either a single strain rate or a small range of strain rates. Therefore, our knowledge of the mechanical properties of macroalgae is limited to a narrow fraction of the strain rates that can occur in nature. In addition, although mechanical properties of macroalgal tissues change with age, the effect of age on the strain-rate dependence of their mechanical behavior has not been documented. METHODS: Using the kelp Egregia menziesii, we measured how high strain rate (simulating wave impingement) and low strain rate (simulating wave surge) affected mechanical properties of frond tissues of various ages. RESULTS: Stiffness of tissues of all ages increased with strain rate, whereas extensibility was unaffected. Strength and toughness increased with strain rate for young tissue but were unaffected by strain rate for old tissue. CONCLUSIONS: Young tissue is weaker than old tissue and, therefore, the most susceptible to breakage from hydrodynamic forces. The increased strength of young tissue at high strain rates can help the frond resist breaking when pulled rapidly during wave impingement, when hydrodynamic forces are largest. Because breakage of young tissue can remove a frond's meristem and negatively impact the survival of the whole kelp, strain-rate dependence of the young tissue's strength can enhance kelp's survival.

The effect of external flow on the feeding currents of sessile microorganisms.

Microscopic sessile suspension feeders live attached to surfaces and, by consuming bacteria-sized prey and by being consumed, they form an important part of aquatic ecosystems. Their environmental impact is mediated by their feeding rate, which depends on a self-generated feeding current. The feeding rate has been hypothesized to be limited by recirculating eddies that cause the organisms to feed from water that is depleted of food particles. However, those results considered organisms in still water, while ambient flow is often present in their natural habitats. We show, using a point-force model, that even very slow ambient flow, with speed several orders of magnitude less than that of the self-generated feeding current, is sufficient to disrupt the eddies around perpendicular suspension feeders, providing a constant supply of food-rich water. However, the feeding rate decreases in external flow at a range of non-perpendicular orientations due to the formation of recirculation structures not seen in still water. We quantify the feeding flow and observe such recirculation experimentally for the suspension feeder Vorticella convallaria in external flows typical of streams and rivers.

Selective factors in the evolution of multicellularity in choanoflagellates.

Choanoflagellates, unicellular eukaryotes that can form multicellular colonies by cell division and that share a common ancestor with animals, are used as a model system to study functional consequences of being unicellular versus colonial. This review examines performance differences between unicellular and multicellular choanoflagellates in swimming, feeding, and avoiding predation, to provide insights about possible selective advantages of being multicellular for the protozoan ancestors of animals. Each choanoflagellate cell propels water by beating a single flagellum and captures bacterial prey on a collar of microvilli around the flagellum. Formation of multicellular colonies does not improve the swimming performance, but the flux of prey-bearing water to the collars of some of the cells in colonies of certain configurations can be greater than for single cells. Colony geometry appears to affect whether cells in colonies catch more prey per cell per time than do unicellular choanoflagellates. Although multicellular choanoflagellates show chemokinetic behavior in response to oxygen, only the unicellular dispersal stage (fast swimmers without collars) use pH signals to aggregate in locations where bacterial prey might be abundant. Colonies produce larger hydrodynamic signals than do single cells, and raptorial protozoan predators capture colonies while ignoring single cells. In contrast, ciliate predators entrain both single cells and colonies in their feeding currents, but reject larger colonies, whereas passive heliozoan predators show no preference. Thus, the ability of choanoflagellate cells to differentiate into different morphotypes, including multicellular forms, in response to variable aquatic environments might have provided a selective advantage to the ancestors of animals.

Does Formation of Multicellular Colonies by Choanoflagellates Affect Their Susceptibility to Capture by Passive Protozoan Predators?

Microbial eukaryotes, critical links in aquatic food webs, are unicellular, but some, such as choanoflagellates, form multicellular colonies. Are there consequences to predator avoidance of being unicellular vs. forming larger colonies? Choanoflagellates share a common ancestor with animals and are used as model organisms to study the evolution of multicellularity. Escape in size from protozoan predators is suggested as a selective factor favoring evolution of multicellularity. Heterotrophic protozoans are categorized as suspension feeders, motile raptors, or passive predators that eat swimming prey which bump into them. We focused on passive predation and measured the mechanisms responsible for the susceptibility of unicellular vs. multicellular choanoflagellates, Salpingoeca helianthica, to capture by passive heliozoan predators, Actinosphaerium nucleofilum, which trap prey on axopodia radiating from the cell body. Microvideography showed that unicellular and colonial choanoflagellates entered the predator's capture zone at similar frequencies, but a greater proportion of colonies contacted axopodia. However, more colonies than single cells were lost during transport by axopodia to the cell body. Thus, feeding efficiency (proportion of prey entering the capture zone that were engulfed in phagosomes) was the same for unicellular and multicellular prey, suggesting that colony formation is not an effective defense against such passive predators.

Effects of cell morphology and attachment to a surface on the hydrodynamic performance of unicellular choanoflagellates.

Choanoflagellates, eukaryotes that are important predators on bacteria in aquatic ecosystems, are closely related to animals and are used as a model system to study the evolution of animals from protozoan ancestors. The choanoflagellate Salpingoeca rosetta has a complex life cycle with different morphotypes, some unicellular and some multicellular. Here we use computational fluid dynamics to study the hydrodynamics of swimming and feeding by different unicellular stages of S. rosetta: a swimming cell with a collar of prey-capturing microvilli surrounding a single flagellum, a thecate cell attached to a surface and a dispersal-stage cell with a slender body, long flagellum and short collar. We show that a longer flagellum increases swimming speed, longer microvilli reduce speed and cell shape only affects speed when the collar is very short. The flux of prey-carrying water into the collar capture zone is greater for swimming than sessile cells, but this advantage decreases with collar size. Stalk length has little effect on flux for sessile cells. We show that ignoring the collar, as earlier models have done, overestimates flux and greatly overestimates the benefit to feeding performance of swimming versus being attached, and of a longer stalk for attached cells.

Mechanical properties of the wave-swept kelp Egregia menziesii change with season, growth rate and herbivore wounds.

The resistance of macroalgae to damage by hydrodynamic forces depends on the mechanical properties of their tissues. Although factors such as water-flow environment, algal growth rate and damage by herbivores have been shown to influence various material properties of macroalgal tissues, the interplay of these factors as they change seasonally and affect algal mechanical performance has not been worked out. We used the perennial kelp Egregia menziesii to study how the material properties of the rachis supporting a frond changed seasonally over a 2 year period, and how those changes correlated with seasonal patterns of the environment, growth rate and herbivore load. Rachis tissue became stiffer, stronger and less extensible with age (distance from the meristem). Thus, slowly growing rachises were stiffer, stronger and tougher than rapidly growing ones. Growth rates were highest in spring and summer when upwelling and long periods of daylight occurred. Therefore, rachis tissue was most resistant to damage in the winter, when waves were large as a result of seasonal storms. Herbivory was greatest during summer, when rachis growth rates were high. Unlike other macroalgae, E. menziesii did not respond to herbivore damage by increasing rachis tissue strength, but rather by growing in width so that the cross-sectional area of the wounded rachis was increased. The relative timing of environmental factors that affect growth rates (e.g. upwelling supply of nutrients, daylight duration) and of those that can damage macroalgae (e.g. winter storms, summer herbivore outbreaks) can influence the material properties and thus the mechanical performance of macroalgae.

Escaping blood-fed malaria mosquitoes minimize tactile detection without compromising on take-off speed.

To escape after taking a blood meal, a mosquito must exert forces sufficiently high to take off when carrying a load roughly equal to its body weight, while simultaneously avoiding detection by minimizing tactile signals exerted on the host's skin. We studied this trade-off between escape speed and stealth in the malaria mosquito Anopheles coluzzii using 3D motion analysis of high-speed stereoscopic videos of mosquito take-offs and aerodynamic modeling. We found that during the push-off phase, mosquitoes enhanced take-off speed using aerodynamic forces generated by the beating wings in addition to leg-based push-off forces, whereby wing forces contributed 61% of the total push-off force. Exchanging leg-derived push-off forces for wing-derived aerodynamic forces allows the animal to reduce peak force production on the host's skin. By slowly extending their long legs throughout the push-off, mosquitoes spread push-off forces over a longer time window than insects with short legs, thereby further reducing peak leg forces. Using this specialized take-off behavior, mosquitoes are capable of reaching take-off speeds comparable to those of similarly sized fruit flies, but with weight-normalized peak leg forces that were only 27% of those of the fruit flies. By limiting peak leg forces, mosquitoes possibly reduce the chance of being detected by the host. The resulting combination of high take-off speed and low tactile signals on the host might help increase the mosquito's success in escaping from blood-hosts, which consequently also increases the chance of transmitting vector-borne diseases, such as malaria, to future hosts.

Do terrestrial hermit crabs sniff? Air flow and odorant capture by flicking antennules.

Capture of odorant molecules by olfactory organs from the surrounding fluid is the first step of smelling. Sniffing intermittently moves fluid across sensory surfaces, increasing delivery rates of molecules to chemosensory receptors and providing discrete odour samples. Aquatic malacostracan crustaceans sniff by flicking olfactory antennules bearing arrays of chemosensory hairs (aesthetascs), capturing water in the arrays during downstroke and holding the sample during return stroke. Terrestrial malacostracans also flick antennules, but how their flicking affects odour capture from air is not understood. The terrestrial hermit crab, Coenobita rugosus, uses antennules bearing shingle-shaped aesthetascs to capture odours. We used particle image velocimetry to measure fine-scale fluid flow relative to a dynamically scaled physical model of a flicking antennule, and computational simulations to calculate diffusion to aesthetascs by odorant molecules carried in that flow. Air does not flow into the aesthetasc array during flick downstrokes or recovery strokes. Odorants are captured from air flowing around the outside of the array during flick downstrokes, when aesthetascs face upstream and molecule capture rates are 21% higher than for stationary antennules. Bursts of flicking followed by pauses deliver discrete odour samples to olfactory sensors, causing intermittency in odour capture by a different mechanism than aquatic crustaceans use.

Flexibility of crab chemosensory sensilla enables flicking antennules to sniff.

The first step in smelling is capture of odorant molecules from the surrounding fluid. We used lateral flagella of olfactory antennules of crabs Callinectes sapidus to study the physical process of odor capture by antennae bearing dense tufts of hair-like chemosensory sensilla (aesthetascs). Fluid flow around and through aesthetasc arrays on dynamically scaled models of lateral flagella of C. sapidus was measured by particle image velocimetry to determine how antennules sample the surrounding water when they flick. Models enabled separate evaluation of the effects of flicking speed, aesthetasc spacing, and antennule orientation. We found that crab antennules, like those of other malacostracan crustaceans, take a discrete water sample during each flick by having a rapid downstroke, during which water flows into the aesthetasc array, and a slow recovery stroke, when water is trapped in the array and odorants have time to diffuse to aesthetascs. However, unlike antennules of crustaceans with sparse aesthetasc arrays, crabs enhance sniffing via additional mechanisms: 1) Aesthetascs are flexible and splay as a result of the hydrodynamic drag during downstrokes, then clump together during return strokes; and 2) antennules flick with aesthetascs on the upstream side of the stalk during downstrokes, but are hidden downstream during return strokes. Aiming aesthetascs into ambient flow maintains sniffing. When gaps between aesthetascs are wide, changes in antennule speed are more effective at altering flow through the array than when gaps are narrow. Nonetheless, if crabs had fixed gap widths, their ability to take discrete samples of their odorant environment would be diminished.

Interdisciplinary Laboratory Course Facilitating Knowledge Integration, Mutualistic Teaming, and Original Discovery.

Experiencing the thrill of an original scientific discovery can be transformative to students unsure about becoming a scientist, yet few courses offer authentic research experiences. Increasingly, cutting-edge discoveries require an interdisciplinary approach not offered in current departmental-based courses. Here, we describe a one-semester, learning laboratory course on organismal biomechanics offered at our large research university that enables interdisciplinary teams of students from biology and engineering to grow intellectually, collaborate effectively, and make original discoveries. To attain this goal, we avoid traditional "cookbook" laboratories by training 20 students to use a dozen research stations. Teams of five students rotate to a new station each week where a professor, graduate student, and/or team member assists in the use of equipment, guides students through stages of critical thinking, encourages interdisciplinary collaboration, and moves them toward authentic discovery. Weekly discussion sections that involve the entire class offer exchange of discipline-specific knowledge, advice on experimental design, methods of collecting and analyzing data, a statistics primer, and best practices for writing and presenting scientific papers. The building of skills in concert with weekly guided inquiry facilitates original discovery via a final research project that can be presented at a national meeting or published in a scientific journal.

Swimming in an Unsteady World.

When animals swim in aquatic habitats, the water through which they move is usually flowing. Therefore, an important part of understanding the physics of how animals swim in nature is determining how they interact with the fluctuating turbulent water currents in their environment. We addressed this issue using microscopic larvae of invertebrates in "fouling communities" growing on docks and ships to ask how swimming affects the transport of larvae between moving water and surfaces from which they disperse and onto which they recruit. Field measurements of the motion of water over fouling communities were used to design realistic turbulent wavy flow in a laboratory wave-flume over early-stage fouling communities. Fine-scale measurements of rapidly-varying water-velocity fields were made using particle-image velocimetry, and of dye-concentration fields (analog for chemical cues from the substratum) were made using planar laser-induced fluorescence. We used individual-based models of larvae that were swimming, passively sinking, passively rising, or were passive and neutrally buoyant to determine how their trajectories were affected by their motion through the water, rotation by local shear, and transport by ambient flow. Swimmers moved up and down in the turbulent flow more than did neutrally buoyant larvae. Although more of the passive sinkers landed on substrata below them, and more passive risers on surfaces above, swimming was the best strategy for landing on surfaces if their location was not predictable (as is true for fouling communities). When larvae moved within 5 mm of surfaces below them, passive sinkers and neutrally-buoyant larvae landed on the substratum, whereas many of the swimmers were carried away, suggesting that settling larvae should stop swimming as they near a surface. Swimming and passively-rising larvae were best at escaping from a surface below them, as precompetent larvae must do to disperse away. Velocities, vorticities, and odor-concentrations encountered by larvae fluctuated rapidly, with peaks much higher than mean values. Encounters with concentrations of odor or with vorticities above threshold increased as larvae neared the substratum. Although microscopic organisms swim slowly, their locomotory behavior can affect where they are transported by the movement of ambient water as well as the signals they encounter when they move within a few centimeters of surfaces.

The descent of ant: field-measured performance of gliding ants.

Gliding ants avoid predatory attacks and potentially mortal consequences of dislodgement from rainforest canopy substrates by directing their aerial descent towards nearby tree trunks. The ecologically relevant measure of performance for gliding ants is the ratio of net horizontal to vertical distance traveled over the course of a gliding trajectory, or glide index. To study variation in glide index, we measured three-dimensional trajectories of Cephalotes atratus ants gliding in natural rainforest habitats. We determined that righting phase duration, glide angle, and path directness all significantly influence variation in glide index. Unsuccessful landing attempts result in the ant bouncing off its target and being forced to make a second landing attempt. Our results indicate that ants are not passive gliders and that they exert active control over the aerodynamic forces they experience during their descent, despite their apparent lack of specialized control surfaces.

Ontogenetic changes in the olfactory antennules of the shore crab, Hemigrapsus oregonensis, maintain sniffing function during growth.

Malacostracan crustaceans capture odours using arrays of chemosensory hairs (aesthetascs) on antennules. Lobsters and stomatopods have sparse aesthetascs on long antennules that flick with a rapid downstroke when water flows between the aesthetascs and a slow return stroke when water is trapped within the array (sniffing). Changes in velocity only cause big differences in flow through an array in a critical range of hair size, spacing and speed. Crabs have short antennules bearing dense arrays of flexible aesthetascs that splay apart during downstroke and clump together during return. Can crabs sniff, and when during ontogeny are they big enough to sniff? Antennules of Hemigrapsus oregonensis representing an ontogenetic series from small juveniles to adults were used to design dynamically scaled physical models. Particle image velocimetry quantified fluid flow through each array and showed that even very small crabs capture a new water sample in their arrays during the downstroke and retain that sample during return stroke. Comparison with isometrically scaled antennules suggests that reduction in aesthetasc flexural stiffness during ontogeny, in addition to increase in aesthetasc number and decrease in relative size, maintain sniffing as crabs grow. Sniffing performance of intermediate-sized juveniles was worse than for smaller and larger crabs.

Interactions between benthic predators and zooplanktonic prey are affected by turbulent waves.

Predators capture prey in complex and variable environments. In the ocean, bottom-dwelling (benthic) organisms are subjected to water currents, waves, and turbulent eddies. For benthic predators that feed on small animals carried in the water (zooplankton), flow not only delivers prey, but can also shape predator-prey interactions. Benthic passive suspension feeders collect prey delivered by movement of ambient water onto capture-surfaces, whereas motile benthic predators, such as burrow-dwelling fish, dart out to catch passing zooplankton. How does the flow of ambient water affect these contrasting modes of predation by benthic zooplanktivores? We studied the effects of turbulent, wavy flow on the encounter, capture, and retention of motile zooplanktonic prey (copepods, Acartia spp.) by passive benthic suspension feeders (sea anemones, Anthopleura elegantissima). Predator-prey interactions were video-recorded in a wave-generating flume under two regimes of oscillating flow with different peak wave velocities and levels of turbulent kinetic energy ("weak" and "strong" waves). Rates of encounter (number of prey passing through a sea anemone's capture zone per time), capture (prey contacting and sticking to tentacles per time), and retention (prey retained on tentacles, without struggling free or washing off, per time) were measured at both strengths of waves. Strong waves enhanced encounter rates both for dead copepods and for actively swimming copepods, but there was so much variability in the behavior of the live prey that the effect of wave strength on encounter rates was not significant. Trapping efficiency (number of prey retained per number encountered) was the same in both flow regimes because, although fewer prey executed maneuvers to escape capture in strong waves, more of the captured prey was washed off the predators' tentacles. Although peak water velocities and turbulence of waves did not affect feeding rates of passive suspension-feeding sea anemones, increases in these aspects of flow have been shown to enhance feeding rates and efficiency of motile benthic fish that lunge out of their burrows to catch zooplankton. Faster, more turbulent flow interferes with the ability of prey to detect predators and execute escape maneuvers, and thus enhances capture rates both for passive suspension-feeding predators and for actively swimming predators. However, prey captured in the mouths of fish are not washed away by ambient flow, whereas prey captured on the tentacles of suspension feeders can be swept off before they are ingested. Therefore, the effects of flowing water on predation on zooplankton by benthic animals depend on the feeding mode of the predator.

Cooperatively generated stresslet flows supply fresh fluid to multicellular choanoflagellate colonies.

The flagellated protozoan Salpingoeca rosetta is one of the closest relatives of multicellular animals. Unicellular S. rosetta can be induced to form multicellular colonies, but colonies swim more slowly than individual cells so the advantages conferred by colony formation are uncertain. Here we use theoretical models to show that hydrodynamic cooperation between cells can increase the fluid supply to the colony, an important predictor of feeding rate. Our results suggest that hydrodynamic benefits may have been an important selective factor in the evolution of early multicellular animals.

Numerical simulations of odorant detection by biologically inspired sensor arrays.

The antennules of many marine crustaceans enable them to rapidly locate sources of odorant in turbulent environmental flows and may provide biological inspiration for engineered plume sampling systems. A substantial gap in knowledge concerns how the physical interaction between a sensing device and the chemical filaments forming a turbulent plume affects odorant detection and filters the information content of the plume. We modeled biological arrays of chemosensory hairs as infinite arrays of odorant flux-detecting cylinders and simulated the fluid flow around and odorant flux into the hair-like sensors as they intercepted a single odorant filament. As array geometry and sampling kinematics were varied, we quantified distortion of the flux time series relative to the spatial shape of the original odorant filament as well as flux metrics that may be important to both organisms and engineered systems attempting to measure plume structure and/or identify chemical composition. The most important predictor of signal distortion is the ratio of sensor diameter to odorant filament width. Achieving high peak properties (e.g. sharpness) of the flux time series and maximizing the total number of odorant molecules detected appear to be mutually exclusive design goals. Sensor arrays inspired specifically by the spiny lobster Panulirus argus and mantis shrimp Gonodactylaceus falcatus introduce little signal distortion but these species' neural systems may not be able to resolve plume structure at the level of individual filaments via temporal properties of the odorant flux. Current chemical sensors are similarly constrained. Our results suggest either that the spatial distribution of flux across the aesthetasc array is utilized by P. argus and G. falcatus, or that such high spatiotemporal resolution is unnecessary for effective plume tracking.

Using physical models to study the gliding performance of extinct animals.

Aerodynamic studies using physical models of fossil organisms can provide quantitative information about how performance of defined activities, such as gliding, depends on specific morphological features. Such analyses allow us to rule out hypotheses about the function of extinct organisms that are not physically plausible and to determine if and how specific morphological features and postures affect performance. The purpose of this article is to provide a practical guide for the design of dynamically scaled physical models to study the gliding of extinct animals using examples from our research on the theropod dinosaur, †Microraptor gui, which had flight feathers on its hind limbs as well as on its forelimbs. Analysis of the aerodynamics of †M. gui can shed light on the design of gliders with large surfaces posterior to the center of mass and provide functional information to evolutionary biologists trying to unravel the origins of flight in the dinosaurian ancestors and sister groups to birds. Measurements of lift, drag, side force, and moments in pitch, roll, and yaw on models in a wind tunnel can be used to calculate indices of gliding and parachuting performance, aerodynamic static stability, and control effectiveness in maneuvering. These indices permit the aerodynamic performance of bodies of different shape, size, stiffness, texture, and posture to be compared and thus can provide insights about the design of gliders, both biological and man-made. Our measurements of maximum lift-to-drag ratios of 2.5-3.1 for physical models of †M. gui suggest that its gliding performance was similar to that of flying squirrels and that the various leg postures that might have been used by †M. gui make little difference to that aspect of aerodynamic performance. We found that body orientation relative to the movement of air past the animal determines whether it is difficult or easy to maneuver.