No vertebrate, living or extinct, is known to have possessed an odd number of limbs. Despite this ‘forbidden phenotype’, gaits that use odd numbers of limbs (e.g. tripedalism or pentapedalism) have evolved in both avian and mammalian lineages. Tripedal locomotion is commonly employed by parrots during climbing, who use their beaks as an additional support. However, it is unclear whether the beak functions simply as a stabilizing hook, or as a propulsive limb. Here, we present data on kinetics of tripedal climbing in six rosy-faced lovebirds ( Agapornis roseicollis ). Our findings demonstrate that parrots use cyclical tripedal gaits when climbing and the beak and hindlimbs generate comparable propulsive and tangential substrate reaction forces and power. Propulsive and tangential forces generated by the beak are of magnitudes equal to or greater than those forces generated by the forelimbs of humans and non-human primates during vertical climbing. We conclude that the feeding apparatus and neck flexors of parrots have been co-opted to function biomechanically as a propulsive third limb during vertical climbing. We hypothesize that this exaptation required substantive alterations to the neuromuscular system including enhanced force-generating capabilities of the neck flexors and modifications to locomotor central pattern generators.
Brachiation is a form of suspensory locomotion observed only in Primates. The non-human hominoids (e.g., gibbons, orangutans, chimpanzees, and gorillas) are considered specialized brachiators, yet peculiar among the living apes are anatomically modern humans (Homo sapiens), who have forgone this locomotor mode in favor of bipedal striding. Humans can, however, brachiate and seem to have retained the locomotor capabilities of their arboreal ancestors. However, the mechanics of human brachiation have not been quantified. In this study, we evaluate how closely human brachiation conforms to the expectations of simple pendular motion using triaxial accelerometry and high-speed videography. These data are compared to specialized brachiating non-human primates. We found that humans have lower energy recovery than siamangs (Symphalangus syndactylus) during brachiation and have shorter observed pendular periods than expected compared to other primates. We demonstrate that relatively long forelimb length and high grip forces, a proxy for global forelimb force-generating potential, act as the main driving factors to reduce energetic costs through effective pendular recovery. These data are the first to assess the strategies humans adopt to perform a behavior they are not anatomically specialized to execute and places them within a comparative framework amongst other brachiating primates. We show that although humans demonstrate behavioral flexibility during brachiation (e.g., differing mediolateral and vertical center of mass positional movement patterns), anatomical features are the primary driver of variation in brachiation performance.
Despite the high mechanical demands associated with climbing, the ability to ascend vertically has evolved independently in most major animal lineages. However, little is known about the kinetics, mechanical energy profiles, or spatiotemporal gait characteristics of this locomotor mode. In this study, we explore the dynamics of horizontal locomotion and vertical climbing on both flat substrates and narrow poles in five Australian green tree frogs (Ranoidea caerulea). Vertical climbing is associated with slow, deliberate movements (i.e., reduced speed and stride frequency and increased duty factors) and propulsive fore-aft impulses in both the forelimb and hindlimb. By comparison, horizontal walking was characterize by a braking forelimb and a propulsive hindlimb. In the normal plane, tree frogs mirrored other taxa in exhibiting a net pulling forelimb and a net pushing hindlimb. In terms of mechanical energy, tree frogs matched theoretical predictions of climbing dynamics (i.e., the total mechanical energetic cost of vertical climbing was predominantly driven by potential energy, with negligible kinetic contributions). Utilizing power as a means for estimating efficiency, we also demonstrate that Australian green tree frogs show total mechanical power costs only slightly above the minimum mechanical power necessary to climb, highlighting their highly effective locomotor mechanics. This study provides new data on climbing dynamics in a slow-moving arboreal tetrapod and raises new testable hypotheses about how natural selection can act upon a locomotor behavior that is notably constrained by external physical forces.
No vertebrate, living or extinct, is known to have possessed an odd number of limbs. Despite this ″forbidden phenotype″, gaits that utilize odd numbers of limbs (e.g., tripedalism or pentapedalism) have evolved in both avian and mammalian lineages. Tripedal locomotion is commonly employed by parrots during climbing, who utilize their beaks as an additional support. However, it is unclear whether the beak functions simply as a stabilizing hook, or as a propulsive limb. Here, we present data on kinetics of tripedal climbing in six rosy –faced lovebirds (Agapornis rosiecollis). Our findings demonstrate that parrots utilize cyclical tripedal gaits when climbing and the beak and hindlimbs generate comparable propulsive and tangential substrate reaction forces and power. Propulsive and tangential forces generated by the beak are of equal or greater magnitudes to those forces generated by the forelimbs of humans and non –human primates during vertical climbing. We conclude that the feeding apparatus and neck musculature of parrots has been co–opted to function biomechanically as a third limb during vertical climbing. We hypothesize that this exaptation required substantive alterations to the neuromuscular system including enhanced force–generating capabilities of the neck musculature and modifications to limb central pattern generators.
Across tetrapods, the proportional lengths of the manual and pedal phalanges are highly constrained, following a generalized blueprint of shortening in a proximodistal gradient. Despite this, several lineages of both mammals (e.g. sloths, bats and colugos) and birds (e.g. raptors, parrots and woodpeckers) have broken this pattern, shortening the proximal phalanx while elongating more distal elements. As yet, no unifying explanation for this convergence has been empirically evaluated. This study combines a comparative phylogenetic assessment of phalangeal morphology across mammals and birds with a novel bioinspired robotics approach to explicitly test functional hypotheses relating to these morphotypes. We demonstrate that shortening the proximal phalanx allows taxa to maximize forces produced at the proximal interphalangeal joint, while elongation of subsequent elements maintains total ray length—ensuring arboreal species can still enclose large-diameter supports. Within suspensory and vertically clinging mammals, we additionally observe a secondary adaptation towards maximizing grip strength: namely increasing the height of the trochleae to increase the moment arm of digital flexor muscles that cross the joint. Together, our analyses highlight that numerous tetrapod lineages independently converged upon this morphotype to maximize proximal gripping strength, an adaptation to support specialized hunting and locomotor behaviours.
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