Suspensory locomotion differs significantly from upright quadrupedal locomotion in mammals. Nevertheless, we know little concerning joint kinematics of suspensory movement. Here, we report three-dimensional kinematic data during locomotion in brown-throated three-toed sloths (Bradypus variegatus). Individuals were recorded with four calibrated high-speed cameras while performing below-branch locomotion on a simulated branch. The elbow (range 73°-177°; mean 114°) and knee (range 107°-175°; mean 140°) were extended throughout support phase, with elbow extension increasing with speed. Both the fore- and hindlimb displayed abducted proximal limb elements (i.e., arm and thigh) and adducted distal elements (i.e., forearm and leg) during all support phase points. Comparisons of elbow and knee angles between brown-throated three-toed sloths and Linnaeus's two-toed sloths (Choloepus didactylus) showed that brown-throated three-toed sloths had significantly more extended joint positions during all support phase points. Additionally, across all kinematic measurements, brown-throated three-toed sloths showed significant differences between homologous fore- and hindlimb segments, with the knee being more extended than the elbow and the arm being more abducted than the thigh. These results are consistent with previously established morphological and behavioral differences between extant sloth genera, with three-toed sloths showing significantly longer forelimbs than hindlimbs and typically favoring locomotion on angled supports. Our findings show that, despite overall similarities in the use of below-branch quadrupedal locomotion, the two sloth lineages achieve this locomotor mode with differing kinematic strategies (e.g., degree of joint flexion). These differences may be attributed to the distinct evolutionary pathways through which obligate suspensory locomotion arose in each lineage.

The gait characteristics associated with arboreal locomotion have been frequently discussed in the context of primate evolution, wherein they present as a trio of distinctive features: a diagonal-sequence, diagonal-couplet gait pattern; a protracted arm at forelimb touchdown; and a hindlimb-biased weight support pattern. The same locomotor characteristics have been found in the woolly opossum, a fine-branch arborealist similar in ecology to some small-bodied primates. To further our understanding of the functional link between arboreality and primate-like locomotion, we present comparative data collected in the laboratory for three musteloid taxa. Musteloidea represents an ecologically diverse superfamily spanning numerous locomotor specializations that includes the highly arboreal kinkajou (Potos flavus), mixed arboreal/terrestrial red pandas (Ailurus fulgens) and primarily terrestrial coatis (Nasua narica). This study applies a combined kinetic and kinematic approach to compare the locomotor behaviors of these three musteloid taxa, representing varying degrees of arboreal specialization. We observed highly arboreal kinkajous to share many locomotor traits with primates. In contrast, red pandas (mixed terrestrial/arborealist) showed gait characteristics found in most non-primate mammals. Coatis, however, demonstrated a unique combination of locomotor traits, combining a lateral-sequence, lateral-couplet gait pattern typical of long-legged, highly terrestrial mammals, varying degrees of arm protraction, and a hindlimb-biased weight support pattern typical of most primates and woolly opossums. We conclude that the three gait characteristics traditionally used to describe arboreal walking in primates can occur independently from one another and not necessarily as a suite of interdependent characteristics, a phenomenon that has been reported for some primates.

Climbing represents a critical behavior in the context of primate evolution. However, anatomically modern human populations are considered ill-suited for climbing. This adaptation can be attributed to the evolution of striding bipedalism, redirecting anatomical traits away from efficient climbing. Although prior studies have speculated on the kinetic consequences of this anatomical reorganization, there is a lack of data on the force profiles of human climbers. This study utilized high-speed videography and force plate analysis to assess single limb forces during climbing from 44 human participants of varying climbing experience and compared these data with climbing data from eight species of non-human primates (anthropoids and strepsirrhines). Contrary to expectations, experience level had no significant effect on the magnitude of single limb forces in humans. Experienced climbers did, however, demonstrate a predictable relationship between center of mass position and peak normal forces, suggesting a better ability to modulate forces during climbing. Humans exhibited significantly higher peak propulsive forces in the hindlimb compared with the forelimb and greater hindlimb dominance overall compared with non-human primates. All species sampled demonstrated exclusively tensile forelimbs and predominantly compressive hindlimbs. Strepsirrhines exhibited a pull-push transition in normal forces, while anthropoid primates, including humans, did not. Climbing force profiles are remarkably stereotyped across humans, reflecting the universal mechanical demands of this form of locomotion. Extreme functional differentiation between forelimbs and hindlimbs in humans may help to explain the evolution of bipedalism in ancestrally climbing hominoids.

Tree sloths evolved below-branch locomotion making them one of few mammalian taxa beyond primates for which suspension is nearly obligatory. Suspension requires strong limb flexor muscles that provide both propulsion and braking/support, and available locomotor kinetics data indicate that these roles differ between fore- and hindlimb pairs. Muscle structure in the pelvic limb is hypothesized to be a key anatomical correlate of function in braking/support during suspensory walking and propulsion and/or support during vertical climbing. This expectation was tested by quantifying architecture properties in the hindlimb limb musculature of brown-throated three-toed sloths (Bradypus variegatus: N = 7) to distinguish the roles of the flexor/extensor functional muscle groups at each joint. Measurements of muscle moment arm (rm ), mass, belly length, fascicle length, pennation angle, and physiological cross-sectional area (PCSA) were taken from n = 45 muscles. Overall, most muscles studied show properties for contractile excursion and fast joint rotational velocity. However, the flexor musculature is more massive (p = 0.048) and has larger PCSA (p = 0.003) than the extensors, especially at the knee joint and digits where well-developed and strong flexors are capable of applying large joint torque. Moreover, selected hip flexors/extensors and knee flexors have modified long rm that can amplify applied joint torque in muscles with otherwise long, parallel fascicles, and one muscle (m. iliopsoas) was capable of moderately high power in B. variegatus. The architectural properties observed in the hip flexors and extensors match well with roles in suspensory braking and vertical propulsion, respectively, whereas strong knee flexors and digital flexors appear to be the main muscles providing suspensory support in the pelvic limb. With aid in support by the forelimbs and the use of adaptive slow locomotion and slow muscle fiber recruitment patterns, structure-function in the tensile limb systems of sloths appears to collectively represent an additional mechanism for energy conservation.

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.

Living sloths exhibit numerous anatomical specializations towards inverted quadrupedalism, however, previous studies have noted a more varied locomotor repertoire than previously anticipated. In this study, we present spatiotemporal gait characteristics and triaxial kinetic data from the brown-throated three-toed sloth (Bradypus variegatus) across three locomotor modes: terrestrial quadrupedal "crawling", suspensory walking, and vertical climbing. Compared to quadrupedal crawling and suspensory walking, B. variegatus adopted longer contact times and stride durations, larger duty factors, and greater speed during vertical climbing. Net fore-aft impulses were significantly greater during vertical climbing in both limb pairs than in quadrupedal crawling and suspensory walking. Functionally, during quadrupedal crawling and vertical climbing, both limb pairs served propulsive roles, while differentiation between a propulsive forelimb and braking hindlimb was observed during suspension. Net tangential forces differentiated vertical climbing kinetics from the other modes of locomotion, with the introduction of bidirectional pulling and pushing forces in the forelimb and hindlimb, respectively. The net mediolateral impulses were similar in vertical climbing and quadrupedal crawling as both limb pairs directed forces in one direction, whereas during suspensory walking, the laterally dominant forelimb was opposed by the medially dominant hindlimb. In total, this study provides novel data on the diverse locomotor dynamics in a slow-moving arboreal tetrapod and posits new testable hypotheses about the neuroplasticity and ease of transitioning between locomotor behaviors. The strikingly similar kinetic profiles of quadrupedal crawling and suspensory walking compared to vertical climbing suggest shared neuromuscular and mechanical demands between these mirrored locomotor modes.

The biomechanical demands of arboreal locomotion are generally thought to necessitate specialized kinetic and kinematic gait characteristics. While such data has been widely collected across arboreal quadrupeds, no study has yet explored how arboreal substrates influence the locomotor behavior of birds. Parrots - an ancient arboreal lineage that exhibit numerous anatomical specializations towards life in the trees - represent an ideal model group within which to examine this relationship. Here, we quantify limb loading patterns within the rosy-faced lovebird (Agapornis roseicollis) across a range of experimental conditions to define under which circumstances arboreal gaits are triggered, and how, during arboreal walking, gait patterns change across substrates of varying diameter. In so doing, we address longstanding questions as to how the challenges associated with arboreality affect gait parameters. Arboreal locomotion was associated with the adoption of a sidling gait, which was employed exclusively on the small- and medium-poles but not terrestrially. When sidling, the hindlimbs are decoupled into a distinct leading limb (which imparts exclusively braking forces) and trailing limb (which generates only propulsive forces). Sidling was also associated with relatively low pitching forces, even on the smallest substrate. Indeed, these forces were significantly lower than mediolateral forces experienced during striding on terrestrial and large-diameter substrates. We propose that the adoption of sidling gaits is a consequence of avian foot morphology and represents a novel form of arboreal locomotion where inversion/eversion is impossible. Such movement mechanics is likely widespread among avian taxa and may also typify patterns of arboreal locomotion in humans.