Effects of ingesting large prey on the kinematics of rectilinear locomotion in Boa constrictor.

Large and stout snakes commonly consume large prey and use rectilinear crawling; yet, whether body wall distention after feeding impairs rectilinear locomotion is poorly understood. After eating large prey (30-37% body mass), all Boa constrictor tested could perform rectilinear locomotion in the region with the food bolus despite a greatly increased distance between the ribs and the ventral skin that likely lengthens muscles relevant to propulsion. Unexpectedly, out of 11 kinematic variables, only two changed significantly (P<0.05) after feeding: cyclic changes in snake height increased by more than 1.5 times and the longitudinal movements of the ventral skin relative to the skeleton decreased by more than 25%. Additionally, cyclic changes in snake width suggest that the ribs are active and mobile during rectilinear locomotion, particularly in fed snakes, but also in unfed snakes. These kinematic changes suggest that rectilinear actuators reorient more vertically and undergo smaller longitudinal excursions following large prey ingestion, both of which likely act to reduce elongation of these muscles that may otherwise experience substantial strain.

Using natricine snakes to test how prey type and size affect predatory behaviors and performance.

Introduction: Predation is a complex process for which behavior, morphology, and size of both predator and prey can affect the success and effectiveness of the predator. For predators such as snakes that swallow prey whole, gape ultimately limits prey size, but the behaviors used to select, capture, and consume prey and attributes of the prey can also affect maximal prey size. For example, swallowing live, struggling prey is difficult, but using coiling or envenomation to restrain or kill prey has evolved repeatedly in snakes. Methods: To test the potential benefits of these behaviors, we manipulated the type and size of prey, and determined how stereotyped predatory behavior was in a snake species (Liodytes rigida) that uses both coiling and envenomation to restrain and immobilize its formidable prey of crayfish. We also studied a close relative (Liodytes pygaea) that eats fish and salamanders to gain insights into the evolution of these traits. Results: For L. rigida, envenomation of hard-shell crayfish via their soft underside was very stereotyped (100% of feedings). Envenomation of soft-shell crayfish was less frequent (59% of feedings) but became more likely both with increased relative prey size and increased time after molt (hardness). L. rigida coiled more for hard-shell than soft-shell crayfish (77% vs. 30%). The probability of coiling was unaffected by prey size, but it increased with increased time after molt for the soft-shell crayfish. Liodytes rigida waited to swallow crayfish until they were completely immobile in 75% and 37% of the feedings with hard- and soft-shelled crayfish, respectively. Even with large prey L. pygaea never used coiling or envenomation, whereas previous studies of L. alleni, the sister species of L. rigida, observed non-lethal coiling without envenomation when eating hard-shell crayfish. Discussion: Our findings for the Liodytes clade of three species suggest that coiling evolved ancestral to the crayfish specialists (L. alleni; L. rigida), and envenomation by L. rigida subsequently evolved as an additional means of subduing formidable prey. The proximate benefits observed for coiling and envenomation in L. rigida support the evolutionary scenario that both traits evolved to enhance the feeding performance for more formidable prey.

Scaling Relationships of Maximal Gape in Two Species of Large Invasive Snakes, Brown Treesnakes and Burmese Pythons, and Implications for Maximal Prey Size.

Snakes are a phylogenetically diverse (> 3500 species) clade of gape-limited predators that consume diverse prey and have considerable ontogenetic and interspecific variation in size, but empirical data on maximal gape are very limited. To test how overall size predicts gape, we quantified the scaling relationships between maximal gape, overall size, and several cranial dimensions for a wide range of sizes (mass 8-64,100 g) for two large, invasive snake species: Burmese pythons (Python molorus bivittatus) and brown treesnakes (Boiga irregularis). Although skull size scaled with negative allometry relative to overall size, isometry and positive allometry commonly occurred for other measurements. For similar snout-vent lengths (SVL), the maximal gape areas of Burmese pythons were approximately 4-6 times greater than those of brown treesnakes, mainly as a result of having a significantly larger relative contribution to gape by the intermandibular soft tissues (43% vs. 17%). In both snake species and for all types of prey, the scaling relationships predicted that relative prey mass (RPM) at maximal gape decreased precipitously with increased overall snake size. For a given SVL or mass, the predicted maximal values of RPM of the Burmese pythons exceeded those of brown treesnakes for all prey types, and predicted values of RPM were usually least for chickens, greatest for limbed reptiles and intermediate for mammals. The pythons we studied are noteworthy for having large overall size and gape that is large even after correcting for overall size, both of which could facilitate some large individuals (SVL = 5 m) exploiting very large vertebrate prey (e.g., deer > 50 kg). Although brown treesnakes had longer quadrate bones, Burmese pythons had larger absolute and larger relative gape as a combined result of larger overall size, larger relative head size, and most importantly, greater stretch of the soft tissues.

Lasso locomotion expands the climbing repertoire of snakes.

The diverse ways and environments in which animals move are correlated with morphology1, but morphology is not sufficient to predict how animals move because behavioral innovations can create new capacities. We document a new mode of snake locomotion - 'lasso locomotion' - that allows the brown treesnake (Boiga irregularis) to ascend much larger smooth cylinders than any previously known behavior. This lasso locomotion may facilitate exploiting resources that might otherwise be unobtainable and contribute to the success and impact of this highly invasive species. VIDEO ABSTRACT.

What Defines Different Modes of Snake Locomotion?

Animals move in diverse ways, as indicated in part by the wide variety of gaits and modes that have been described for vertebrate locomotion. Much variation in the gaits of limbed animals is associated with changing speed, whereas different modes of snake locomotion are often associated with moving on different surfaces. For several decades different types of snake locomotion have been categorized as one of four major modes: rectilinear, lateral undulation, sidewinding, and concertina. Recent empirical work shows that the scheme of four modes of snake locomotion is overly conservative. For example, during aquatic lateral undulation, the timing between muscle activity and lateral bending changes along the length of the snake, which is unlike terrestrial lateral undulation. The motor pattern used to prevent sagging while bridging gaps also suggests that arboreal lateral undulation on narrow surfaces or with a few discrete points of support has a different motor pattern than terrestrial lateral undulation when the entire length of the snake is supported. In all types of concertina locomotion, the distance from the head to the tail changes substantially as snakes alternately flex and then extend different portions of their body. However, snakes climbing cylinders with concertina exert forces medially to attain a purchase on the branch, whereas tunnels require pushing laterally to form an anchoring region. Furthermore, different motor patterns are used for these two types of concertina movement. Some snakes climb vertical cylinders with helical wrapping completely around the cylinder, whereas all other forms of concertina bend regions of the body alternately to the left and right. Current data support rectilinear locomotion and sidewinding as being distinct modes, whereas lateral undulation and concertina are best used for defining categories of gaits with some unifying similarities. Partly as a result of different motor patterns, I propose recognizing five and four distinct types of lateral undulation and concertina, respectively, resulting in a total of 11 distinct gaits previously recognized as only four.

Experimental modification of morphology reveals the effects of the zygosphene-zygantrum joint on the range of motion of snake vertebrae.

Variation in joint shape and soft tissue can alter range of motion (ROM) and create trade-offs between stability and flexibility. The shape of the distinctive zygosphene-zygantrum joint of snake vertebrae has been hypothesized to prevent axial torsion (twisting), but its function has never been tested experimentally. We used experimental manipulation of morphology to determine the role of the zygosphene-zygantrum articulation by micro-computed tomography (µCT) scanning and 3D printing two mid-body vertebrae with unaltered shape and with the zygosphene digitally removed for four species of phylogenetically diverse snakes. We recorded the angular ROM while manipulating the models in yaw (lateral bending), pitch (dorsoventral bending) and roll (axial torsion). Removing the zygosphene typically increased yaw and dorsal pitch ROM. In the normal vertebrae, roll was <2.5 deg for all combinations of pitch and yaw. Roll increased in altered vertebrae but only for combinations of high yaw and ventral pitch that were near or beyond the limits of normal vertebra ROM. In the prairie rattlesnake and brown tree snake, roll in the altered vertebrae was always limited by bony processes other than the zygosphene, whereas in the altered vertebrae of the corn snake and boa constrictor, roll ROM was unconstrained when the pre- and post-zygapophyses no longer overlapped. The zygosphene acts as a bony limit for yaw and dorsal pitch, indirectly preventing roll by precluding most pitch and yaw combinations where roll could occur and potentially allowing greater forces to be applied across the vertebral column than would be possible with only soft-tissue constraints.

Visual Contrast and Intensity Affect Perch Choice of Brown Tree Snakes (Boiga irregularis) and Boa Constrictors (Boa constrictor).

Habitat structure can affect animal movement both by affecting the mechanical demands of locomotion and by influencing where animals choose to go. Arboreal habitats facilitate studying path choice by animals because variation in branch structure has known mechanical consequences, and different branches create discrete choices. Recent laboratory studies have found that arboreal snakes can use vision to select shapes and locations of destinations that mechanically facilitate bridging gaps. However, the extent to which the appearance of objects unrelated to biomechanical demands affects the choice of destinations remains poorly understood for most animal taxa including snakes. Hence, we manipulated the intensity (black, gray, or white), contrast, structure, and locations of destinations to test for their combined effects on perch choice during gap bridging of brown tree snakes and boa constrictors. For a white background and a given perch structure and location, both species had significant preferences for darker perches. The preference for darker destinations was strong enough to override or reduce some preferences for biomechanically advantageous destinations such as those having secondary branches or being located closer or along a straighter trajectory. These results provide a striking example of how visual cues unrelated to the physical structure of surfaces, such as contrast and intensity, can bias choice and, in some cases, supersede a preference for mechanically beneficial surfaces. Because these two species are so phylogenetically distant, some of their similar preferences suggest a sensory bias that may be widespread in snakes. The manipulation of surface color may facilitate management of invasive species, such as the brown tree snakes, by enhancing the efficiency of traps or making certain objects less attractive to them.

Crawling without wiggling: muscular mechanisms and kinematics of rectilinear locomotion in boa constrictors.

A central issue for understanding locomotion of vertebrates is how muscle activity and movements of their segmented axial structures are coordinated, and snakes have a longitudinal uniformity of body segments and diverse locomotor behaviors that are well suited for studying the neural control of rhythmic axial movements. Unlike all other major modes of snake locomotion, rectilinear locomotion does not involve axial bending, and the mechanisms of propulsion and modulating speed are not well understood. We integrated electromyograms and kinematics of boa constrictors to test Lissmann's decades-old hypotheses of activity of the costocutaneous superior (CCS) and inferior (CCI) muscles and the intrinsic cutaneous interscutalis (IS) muscle during rectilinear locomotion. The CCI was active during static contact with the ground as it shortened and pulled the axial skeleton forward relative to both the ventral skin and the ground during the propulsive phase. The CCS was active during sliding contact with the ground as it shortened and pulled the skin forward relative to both the skeleton and the ground during the recovery phase. The IS shortened the ventral skin, and subsequent isometric activity kept the skin stiff and shortened during most of static contact while overlapping extensively with CCI activity. The concentric activity of the CCI and CCS supported Lissmann's predictions. Contrary to Lissmann, the IS had prolonged isometric activity, and the time when it shortened was not consistent with providing propulsive force. Decoupling propulsion from axial bending in rectilinear locomotion may have facilitated economical locomotion of early snakes in subterranean tunnels.

Three-dimensional trajectories affect the epaxial muscle activity of arboreal snakes crossing gaps.

The need for long-axis support is widespread among non-aquatic vertebrates and may be particularly acute for arboreal snakes when many vertebrae span sizable gaps between branches with diverse orientations. Hence, we used brown tree snakes (Boiga irregularis) bridging gaps to test how three-dimensional trajectories affected muscle activity and whether these motor patterns differed from those for the locomotion of terrestrial snakes and movements of other vertebrates. We used five trajectories: pitch angles of 90, 0 and -90 deg (downward) when yaw=0 deg, and 90 deg yaw angles to the left and right when pitch=0 deg. We recorded movement and electromyograms from the three largest epaxial muscles, which from dorsal to ventral are the semispinalis-spinalis (SSP), longissimus dorsi (LD) and iliocostalis (IL). Overall, the SSP had extensive bilateral activity, which resembled the motor pattern during the dorsiflexion of sidewinding snakes. Unlike any previously described terrestrial snake locomotion, bilateral activity of the LD and IL was also common during gap bridging. The largest amounts of muscle activity usually occurred for horizontal gaps, and muscle activity decreased markedly as soon as the snake's head touched the far edge of the gap. Snakes had the least amount of muscle activity for pitch=-90 deg. While turning sideways, muscles on the convex side had less activity when turning compared with the concave side. Hence, the orientation relative to gravity profoundly affected muscle activity during gap bridging, and these complex three-dimensional movements involved several previously undescribed variants of axial motor pattern.

Why arboreal snakes should not be cylindrical: body shape, incline and surface roughness have interactive effects on locomotion.

Depending on animal size, shape, body plan and behaviour, variation in surface structure can affect the speed and ease of locomotion. The slope of branches and the roughness of bark both vary considerably, but their combined effects on the locomotion of arboreal animals are poorly understood. We used artificial branches with five inclines and five peg heights (≤40 mm) to test for interactive effects on the locomotion of three snake species with different body shapes. Unlike boa constrictors (Boa constrictor), corn snakes (Pantherophis guttatus) and brown tree snakes (Boiga irregularis) can both form ventrolateral keels, which are most pronounced in B. irregularis. Increasing peg height up to 10 mm elicited more of the lateral undulatory behaviour (sliding contact without gripping) rather than the concertina behaviour (periodic static gripping) and increased the speed of lateral undulation. Increased incline: (1) elicited more concertina locomotion, (2) decreased speed and (3) increased the threshold peg height that elicited lateral undulation. Boiga irregularis was the fastest species, and it used lateral undulation on the most surfaces, including a vertical cylinder with pegs only 1 mm high. Overall, B. constrictor was the slowest and used the most concertina locomotion, but this species climbed steep, smooth surfaces faster than P. guttatus. Our results illustrate how morphology and two different aspects of habitat structure can have interactive effects on organismal performance and behaviour. Notably, a sharper keel facilitated exploiting shorter protrusions to prevent slipping and provide propulsion, which became increasingly important as surface steepness increased.

The effects of slope and branch structure on the locomotion of a specialized arboreal colubrid snake (Boiga irregularis).

The surfaces in arboreal habitats have variable diameters, slopes, and branching structure that pose functional challenges for animal locomotion. Nevertheless, many lineages of snakes have independently evolved arboreality. We tested the effects of arboreal habitat structure on the locomotion of a highly arboreal species, the brown tree snake (Boiga irregularis), moving on seven diameters (0.6-21 cm) of cylinders oriented at three slopes (0°, 45°, 90°) and with or without pegs. Intermediate diameters of horizontal cylinders maximized speed, and some of the large-diameter cylinders without pegs were impassable when they were inclined. With increased slope the snakes were slower, and they changed from using lateral undulation with sliding contact and balancing to concertina locomotion with periodic static gripping. The presence of pegs increased the speeds of the brown tree snakes and resulted in them only using lateral undulation. Surface diameter, slope, and the occurrence of pegs also had widespread significant effects on the kinematics of the brown tree snakes. Overall, compared to anatomically less specialized corn snakes, brown tree snakes use more lateral undulation, are usually much faster, and are able to move on a wider variety of surfaces. Unlike some of the trade-offs found previously between two less specialized species of snakes with different stoutness when they used modes of arboreal locomotion that involved either balancing or gripping, the slender-bodied brown tree snakes excel at both. Hence, this species may not only be a "jack of all trades" but also a master of many.

Gripping during climbing of arboreal snakes may be safe but not economical.

On the steep surfaces that are common in arboreal environments, many types of animals without claws or adhesive structures must use muscular force to generate sufficient normal force to prevent slipping and climb successfully. Unlike many limbed arboreal animals that have discrete gripping regions on the feet, the elongate bodies of snakes allow for considerable modulation of both the size and orientation of the gripping region. We quantified the gripping forces of snakes climbing a vertical cylinder to determine the extent to which their force production favoured economy or safety. Our sample included four boid species and one colubrid. Nearly all of the gripping forces that we observed for each snake exceeded our estimate of the minimum required, and snakes commonly produced more than three times the normal force required to support their body weight. This suggests that a large safety factor to avoid slipping and falling is more important than locomotor economy.

Lungfish axial muscle function and the vertebrate water to land transition.

The role of axial form and function during the vertebrate water to land transition is poorly understood, in part because patterns of axial movement lack morphological correlates. The few studies available from elongate, semi-aquatic vertebrates suggest that moving on land may be powered simply from modifications of generalized swimming axial motor patterns and kinematics. Lungfish are an ideal group to study the role of axial function in terrestrial locomotion as they are the sister taxon to tetrapods and regularly move on land. Here we use electromyography and high-speed video to test whether lungfish moving on land use axial muscles similar to undulatory swimming or demonstrate novelty. We compared terrestrial lungfish data to data from lungfish swimming in different viscosities as well as to salamander locomotion. The terrestrial locomotion of lungfish involved substantial activity in the trunk muscles but almost no tail activity. Unlike other elongate vertebrates, lungfish moved on land with a standing wave pattern of axial muscle activity that closely resembled the pattern observed in terrestrially locomoting salamanders. The similarity in axial motor pattern in salamanders and lungfish suggests that some aspects of neuromuscular control for the axial movements involved in terrestrial locomotion were present before derived appendicular structures.

Incline and peg spacing have interactive effects on the arboreal locomotor performance and kinematics of brown tree snakes, Boiga irregularis.

Many animals move using lateral undulations, but snakes are noteworthy for using this type of locomotion in an unusual diversity of environments, including trees in which both the spacing and orientation of branches vary considerably. Despite branches providing discrete locations for snakes to generate propulsive forces during lateral undulation, the consequences of branch spacing for the locomotion of snakes are poorly understood. Hence, we determined maximal speeds and kinematics of an arboreal snake (Boiga irregularis) crawling on horizontal and vertical cylinders with pegs that simulated different spacing between secondary branches. Peg spacing, perch orientation and their two-way interaction term had widespread, significant effects on both performance and kinematics. For the horizontal surfaces, maximal locomotor speed occurred with intermediate peg spacing, and it was nearly twice as fast as for both the smallest and largest peg spacings. By contrast, the locomotor speed of snakes on the vertical surfaces was unaffected by peg spacing, and was uniformly slower than that for the horizontal surfaces. For both perch orientations, the number of pegs touched by the snake decreased as peg spacing increased, and while touching only one peg the snakes crawled with apparent ease and steady speed. The snakes crawled vertically with only one peg as quickly as they did using 2-10 pegs. Pegs on a horizontal cylinder are probably important both for propulsion of snakes and prevention of long-axis rolling, whereas pegs protruding from vertical cylinders and those protruding from horizontal planar surfaces are probably used almost exclusively for propulsion.

Three-dimensional locations of destinations have species-dependent effects on the choice of paths and the gap-bridging performance of arboreal snakes.

Challenges for moving in arboreal environments include sizable gaps and variable three-dimensional locations between the discrete destinations created by branches. Different three-dimensional orientations of the body change the mechanical demands of crossing gaps, especially for elongate animals, but these effects on both the maximal gap distance crossed (Gapmax ) and the choice of destinations are poorly understood despite many animals frequently encountering these circumstances. We tested and compared the effects of three-dimensional gap location on the bridging ability of three species of snakes with similar length but moderately stout (Boa constrictor), intermediate (Pantherophis guttatus) or slender (Boiga irregularis) shapes, and we tested for bias in choice of destinations for the latter two species. The effects of both gap location and species on Gapmax were highly significant, and the rank order of species from greatest to least Gapmax was often from the most slender to the stoutest species. Although the effects of yaw (horizontal) angles were statistically significant, they were small compared to those of pitch (vertical) angles, which created nearly twofold variation in the values of Gapmax within each species. The greatest values of Gapmax occurred when snakes went straight down. Within a horizontal plane, P. guttatus and B. irregularis had similar preferences for choosing gaps with smaller yaw angles. However, when pitch angle was varied, P. guttatus preferred lower destinations, whereas B. irregularis preferred higher destinations. Thus, despite quite uniform effects of gap location on bridging abilities and torques acting on the animals, some preferences for locations were species specific. J. Exp. Zool. 319A:124-137, 2013. © 2012 Wiley Periodicals, Inc.

Surface shape affects the three-dimensional exploratory movements of nocturnal arboreal snakes.

Movement and searching behaviors at diverse spatial scales are important for understanding how animals interact with their environment. Although the shapes of branches and the voids in arboreal habitats seem likely to affect searching behaviors, their influence is poorly understood. To gain insights into how both environmental structure and the attributes of an animal may affect movement and searching, we compared the three-dimensional exploratory movements of snakes in the dark on two simulated arboreal surfaces (disc and horizontal cylinder). Most of the exploratory movements of snakes in the dark were a small fraction of the distances they could reach while bridging gaps in the light. The snakes extended farther away from the edge of the supporting surface at the ends of the cylinder than from the sides of the cylinder or from any direction from the surface of the disc. The exploratory movements were not random, and the surface shape and three-dimensional directions had significant interactive effects on how the movements were structured in time. Thus, the physical capacity for reaching did not limit the area that was explored, but the shape of the supporting surface and the orientation relative to gravity did create biased searching patterns.

Arboreal habitat structure affects locomotor speed and perch choice of white-footed mice (Peromyscus leucopus).

Arboreal habitats pose several challenges for locomotion resulting from narrow cylindrical surfaces, steep slopes, and secondary branches that can form obstructions. We used laboratory trials to test whether different diameters, slopes, or complexity of branches affected maximum speeds and perch choice of the semi-arboreal white-footed mouse (Peromyscus leucopus). We tested locomotor performance of mice running horizontally and up and down 45° slopes for cylindrical artificial branches with five diameters ranging from 10 to 116 mm and on a subset of diameters for cylinders that were horizontal and had pegs (e.g., secondary branches) every 10 or 20 cm. Slope, diameter, and presence of pegs on top of cylinders had significant and interactive effects on locomotor performance. On horizontal cylinders the speed of mice increased with increased diameter among the three smallest diameters, but changed little with further increases in diameter, whereas for sloped surfaces the extreme diameters had lower speeds than an intermediate diameter. For a given diameter, the speeds of mice were usually faster when running horizontally rather than running uphill or downhill. The presence of pegs greatly decreased running speed compared to unobstructed surfaces, but the magnitude of this effect decreased as diameter increased. The difficulties of maintaining balance and avoiding toppling may have caused much of the decrease in speed and associated increased amounts of pausing. Only 1 of 11 choice tests detected a significant bias of mice favoring the perch that maximized locomotor performance.

The effects of three-dimensional gap orientation on bridging performance and behavior of brown tree snakes (Boiga irregularis).

Traversing gaps with different orientations within arboreal environments has ecological relevance and mechanical consequences for animals. For example, the orientation of the animal while crossing gaps determines whether the torques acting on the body tend to cause it to pitch or roll from the supporting perch or fail as a result of localized bending. The elongate bodies of snakes seem well suited for crossing gaps, but a long unsupported portion of the body can create large torques that make gap bridging demanding. We tested whether the three-dimensional orientation of substrates across a gap affected the performance and behavior of an arboreal snake (Boiga irregularis). The snakes crossed gaps 65% larger for vertical than for horizontal trajectories and 13% greater for straight trajectories than for those with a 90 deg turn within the horizontal plane. Our results suggest that failure due to the inability to keep the body rigid at the edge of the gap may be the primary constraint on performance for gaps with a large horizontal component. In addition, the decreased performance when the destination perch was oriented at an angle to the long axis of the initial perch was probably a result of the inability of snakes to maintain balance due to the large rolling torque. For some very large gaps the snakes enhanced their performance by using rapid lunges to cross otherwise impassable gaps. Perhaps such dynamic movements preceded the aerial behavior observed in other species of arboreal snakes.

Perch diameter and branching patterns have interactive effects on the locomotion and path choice of anole lizards.

Natural branches vary conspicuously in their diameter, density and orientation, but how these latter two factors affect animal locomotion is poorly understood. Thus, for three species of arboreal anole lizards found on different size branches and with different limb lengths, we tested sprinting performance on cylinders with five diameters (5-100 mm) and five patterns of pegs, which simulated different branch orientations and spacing. We also tested whether the lizards preferred surfaces that enhanced their performance. The overall responses to different surfaces were similar among the three species, although the magnitude of the effects differed. All species were faster on cylinders with larger diameter and no pegs along the top. The short-limbed species was the slowest on all surfaces. Much of the variation in performance resulted from variable amounts of pausing among different surfaces and species. Lizards preferred to run along the top of cylinders, but pegs along the top of the narrow cylinders interfered with this. Pegs on top of the 100-mm diameter cylinder, however, had little effect on speed as the lizards ran quite a straight path alongside pegs without bumping into them. All three species usually chose surfaces with greater diameters and fewer pegs, but very large diameters with pegs were preferred to much smaller diameter cylinders without pegs. Our results suggest that preferring larger diameters in natural vegetation has a direct benefit for speed and an added benefit of allowing detouring around branches with little adverse effect on speed.

Perch size and structure have species-dependent effects on the arboreal locomotion of rat snakes and boa constrictors.

Arboreal habitats create diverse challenges for animal locomotion, but the numerical and phylogenetic diversity of snakes that climb trees suggest that their overall body plan is well suited for this task. Snakes have considerable diversity of axial anatomy, but the functional consequences of this diversity for arboreal locomotion are poorly understood because of the lack of comparative data. We simulated diverse arboreal surfaces to test whether environmental structure had different effects on the locomotion of snakes belonging to two distantly related species with differences in axial musculature and stoutness. On most cylindrical surfaces lacking pegs, both species used concertina locomotion, which always involved periodic stopping and gripping but was kinematically distinct in the two species. On horizontal cylinders that were a small fraction of body diameter, the boa constrictors used a balancing form of lateral undulation that was not observed for rat snakes. For all snakes the presence of pegs elicited lateral undulation and enhanced speed. For both species maximal speeds decreased with increased incline and were greatest on cylinders with intermediate diameters that approximated the diameter of the snakes. The frictional resistances that we studied had small effects compared with those of cylinder diameter, incline and the presence of pegs. The stouter and more muscular boa constrictors were usually faster than the rat snakes when using the gripping gait, whereas rat snakes were faster when using lateral undulation on the surfaces with pegs. Thus, variation in environmental structure had several highly significant effects on locomotor mode, performance and kinematics that were species dependent.