Semicircular canal size constrains vestibular function in miniaturized frogs.

Miniaturization has evolved repeatedly in frogs in the moist leaf litter environments of rainforests worldwide. Miniaturized frogs are among the world's smallest vertebrates and exhibit an array of enigmatic features. One area where miniaturization has predictable consequences is the vestibular system, which acts as a gyroscope, providing sensory information about movement and orientation. We investigated the vestibular system of pumpkin toadlets, Brachycephalus (Anura: Brachycephalidae), a clade of miniaturized frogs from Brazil. The semicircular canals of miniaturized frogs are the smallest recorded for adult vertebrates, resulting in low sensitivity to angular acceleration due to insufficient displacement of endolymph. This translates into a lack of postural control during jumping in Brachycephalus and represents a physical constraint resulting from Poiseuille's law, which governs movement of fluids within tubes.

Pelvic function in anuran jumping: Interspecific differences in the kinematics and motor control of the iliosacral articulation during take-off and landing.

Although the anuran pelvis is thought to be adapted for jumping, the function of the iliosacral joint has seen little direct study. Previous work has contrasted the basal "lateral-bender" pelvis from the "rod-like" pelvis of crown taxa hypothesized to function as a sagittal hinge to align the trunk with take-off forces. We compared iliosacral movements and pelvic motor patterns during jumping in the two pelvic types. Pelvic muscle activity patterns, iliosacral anteroposterior (AP) movements and sagittal bending of the pelvis during the take-off and landing phases were quantified in lateral bender taxa Ascaphus (Leiopelmatidae) and Rhinella (Bufonidae) and the rod-like Lithobates (Ranidae). All three species exhibit sagittal extension during take-off, therefore, both pelvic types employ a sagittal hinge. However, trunk elevation occurs significantly earlier in the anuran rod-like pelvis. Motor patterns confirm that the piriformis muscles depress the urostyle while the longissimus dorsi muscles elevate the trunk during take-off. However, the coccygeoiliacus muscles also produce anterior translation of the sacrum on the ilia. A new model illustrates how AP translation facilitates trunk extension in the lateral-bender anurans that have long been thought to have limited sagittal bending. During landing, AP translation patterns are similar because impact forces slide the sacrum from its posterior to anterior limits. Sagittal flexion during landing differs among the three taxa depending on the way the species land. AP translation during landing may dampen impact forces especially in Rhinella in which pelvic function is tuned to forelimb-landing dynamics. The flexibility of the lateral-bender pelvis to function in sagittal bending and AP translation helps to explain the retention of this basal configuration in many anurans. The novel function of the rod-like pelvis may be to increase the rate of trunk elevation relative to faster rates of energy release from the hindlimbs enabling them to jump farther. J. Morphol. 277:1539-1558, 2016. © 2016 Wiley Periodicals, Inc.

Functional evolution of jumping in frogs: Interspecific differences in take-off and landing.

Ancestral frogs underwent anatomical shifts including elongation of the hindlimbs and pelvis and reduction of the tail and vertebral column that heralded the transition to jumping as a primary mode of locomotion. Jumping has been hypothesized to have evolved in a step-wise fashion with basal frogs taking-off with synchronous hindlimb extension and crash-landing on their bodies, and then their limbs move forward. Subsequently, frogs began to recycle the forelimbs forward earlier in the jump to control landing. Frogs with forelimb landing radiated into many forms, locomotor modes, habitats, and niches with controlled landing thought to improve escape behavior. While the biology of take-off behavior has seen considerable study, interspecific comparisons of take-off and landing behavior are limited. In order to understand the evolution of jumping and controlled landing in frogs, data are needed on the movements of the limbs and body across an array of taxa. Here, we present the first description and comparison of kinematics of the hindlimbs, forelimbs and body during take-off and landing in relation to ground reaction forces in four frog species spanning the frog phylogeny. The goal of this study is to understand what interspecific differences reveal about the evolution of take-off and controlled landing in frogs. We provide the first comparative description of the entire process of jumping in frogs. Statistical comparisons identify both homologous behaviors and significant differences among species that are used to map patterns of trait evolution and generate hypotheses regarding the functional evolution of take-off and landing in frogs.

Landing in basal frogs: evidence of saltational patterns in the evolution of anuran locomotion.

All frogs are assumed to jump in a similar manner by rapidly extending hindlimbs during the propulsive phase and rotating the limbs forward during flight in order to land forelimbs first. However, studies of jumping behavior are lacking in the most primitive living frogs of the family Leiopelmatidae. These semi-aquatic or terrestrial anurans retain a suite of plesiomorphic morphological features and are unique in using an asynchronous (trot-like) rather than synchronous "frog-kick" swimming gait of other frogs. We compared jumping behavior in leiopelmatids to more derived frogs and found that leiopelmatids maintain extended hindlimbs throughout flight and landing phases and do not land on adducted forelimbs. These "belly-flop" landings limit the ability for repeated jumps and are consistent with a riparian origin of jumping in frogs. The unique behavior of leiopelmatids shows that frogs evolved jumping before they perfected landing. Moreover, an inability to rapidly cycle the limbs may provide a functional explanation for the absence of synchronous swimming in leiopelmatids.

Three-dimensional launch kinematics in leaping, parachuting and gliding squirrels.

Leaping, parachuting and gliding are the primary means by which arboreal squirrels negotiate gaps in the canopy. There are notable differences among the three locomotor modes with respect to mid-air postures and aerodynamics, yet it is unclear whether variation should also be expected during the launch phase of locomotion. To address this question, launch kinematic profiles were compared in leaping (Tamias striatus), parachuting (Tamiasciurus hudsonicus) and gliding (Glaucomys volans) squirrels. Animals were filmed launching to the ground from a platform using high-speed video. Statistical comparisons among taxa indicated that only six out of 23 variables were significantly different among the three species. Two were associated with tail kinematics and were a consequence of tail morphology. Two were forelimb-related and discriminated gliding from non-gliding taxa. The remaining two variables were performance attributes, indicating significant variation among the species in take-off velocity and horizontal range. The absence of significant differences in hindlimb kinematics indicates that propulsion is essentially identical in leaping, parachuting and gliding squirrels.