Calcaneal external shape differs among nonhuman primates relative to locomotion. Such relationships between whole-bone calcaneal trabecular structure and locomotion, however, have yet to be studied. Here we analyze calcaneal trabecular architecture in Gorilla gorilla gorilla, Gorilla beringei beringei, and G. b. graueri to investigate general trends and fine-grained differences among gorilla taxa relative to locomotion.
Calcanei were micro-CT scanned. A three-dimensional geometric morphometric sliding semilandmark analysis was carried out and the final landmark configurations used to position 156 volumes of interest. Trabecular thickness (Tb.Th), trabecular spacing (Tb.Sp), and bone volume fraction (BV/TV) were calculated using the BoneJ plugin for ImageJ and MATLAB. Non-parametric MANOVAs were run to test for significant differences among taxa in parameter raw values and z-scores. Parameter distributions were visualized using color maps and summarized using principal components analysis.
There are no significant differences in raw BV/TV or Tb.Th among gorillas, however G. b. beringei significantly differs in z-scores for both parameters (p = <0.0271). All three taxa exhibit relatively lower BV/TV and Tb.Th in the posterior half of the calcaneus. This gradation is exacerbated in G. b. beringei. G. b. graueri significantly differs from other taxa in Tb.Sp z-scores (p < 0.001) indicating a different spacing distribution.
Relatively higher Tb.Th and BV/TV in the anterior calcaneus among gorillas likely reflects higher forces associated with body mass (transmitted through the subtalar joint) relative to forces transferred through the posterior calcaneus. The different Tb.Sp pattern in G. b. graueri may reflect proposed differences in foot positioning during locomotion.

Despite qualitative observations of wild primates pumping branches before leaping across gaps in the canopy, most studies have suggested that support compliance increases the energetic cost of arboreal leaping, thus limiting leaping performance. In this study, we quantified branch pumping behavior and tree swaying in wild primates to test the hypothesis that these behaviors improve leaping performance.
We recorded wild colobine monkeys crossing gaps in the canopy and quantitatively tracked the kinematics of both the monkey and the compliant support during behavioral sequences. We also empirically measured the compliance of a sample of locomotor supports in the monkeys\' natural habitat, allowing us to quantify the resonant properties of substrates used during leaping.
Analyses of three recordings show that adult red colobus monkeys (Piliocolobus tephrosceles) use branch compliance to their advantage by actively pumping branches before leaping, augmenting their vertical velocity at take-off. Quantitative modeling of branch resonance periods, based on empirical measurements of support compliance, suggests that monkeys specifically employed branch pumping on relatively thin branches with protracted periods of oscillation. Finally, an additional four recordings show that both red colobus and black and white colobus monkeys (Colobus guereza) utilize tree swaying to cross large gaps, augmenting horizontal velocity at take-off.
This deliberate branch manipulation to produce a mechanical effect for stronger propulsion is consistent with the framework of instrumental problem-solving. To our knowledge, this is the first study of wild primates which quantitatively shows how compliant branches can be used advantageously to augment locomotor performance.

Climbing animals theoretically should optimize the energetic costs of vertical climbing while also maintaining stability. Many modifications to climbing behaviors have been proposed as methods of satisfying these criteria, focusing on controlling the center of mass (COM) during ascent. However, the link between COM movements and metabolic energy costs has yet to be evaluated empirically. In this study, we manipulated climbing conditions across three experimental setups to elicit changes in COM position, and measured the impact of these changes upon metabolic costs across a sample of 14 humans. Metabolic energy was assessed via open flow respirometry, while COM movements were tracked both automatically and manually. Our findings demonstrate that, despite inducing variation in COM position, the energetic costs of climbing remained consistent across all three setups. Differences in energetic costs were similarly not affected by body mass; however, velocity had a significant impact upon both cost of transport and cost of locomotion, but such a relationship disappeared when accounting for metabolic costs per stride. These findings suggest that climbing has inescapable metabolic demands driven by gaining height, and that attempts to mitigate such a cost, with perhaps the exception of increasing speed, have only minimal impacts. We also demonstrate that metabolic and mechanical energy costs are largely uncorrelated. Collectively, we argue that these data refute the idea that efficient locomotion is the primary aim during climbing. Instead, adaptations towards effective climbing should focus on stability and reducing the risk of falling, as opposed to enhancing the metabolic efficiency of locomotion.

An accident during arboreal locomotion can lead to risky falls, but it remains unclear that the extent to which primates, as adept arborealists, change their locomotion in response to the perceived risk of moving on high supports in the tree canopy. By using more stable forms of locomotion on higher substrates, primates might avoid potentially fatal consequences.
Using high-speed cameras, we recorded the quadrupedal locomotion of four wild lemur species-Eulemur rubriventer, Eulemur rufifrons, Hapalemur aureus, and Lemur catta (N = 113 total strides). We quantified the height, diameter, and angular orientation of locomotor supports using remote sensors and tested the influence of support parameters on gait kinematics, specifically predicting that in response to increasing substrate height, lemurs would decrease speed and stride frequency, but increase stride length and the mean number of supporting limbs.
Lemurs did not adjust stride frequency on substrates of varying height. Adjustments to speed, stride length, and the mean number of supporting limbs in response to varying height often ran counter to predictions. Only E. rubriventer decreased speed and increased the mean number of supporting limbs on higher substrates.
Results suggest that quadrupedal walking is a relatively safe form of locomotion for lemurs, requiring subtle changes in gait to increase stability on higher-that is, potentially riskier-substrates. Continued investigation of the impact of height on locomotion will be important to determine how animals assess risk in their environment and how they choose to use this information to move more safely.

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.

Differences in talar articular morphology relative to locomotion have recently been found within Pan and Gorilla. Whole-bone talar morphology within, and shared variation among, Pan and Gorilla (sub)species, however, has yet to be investigated. Here we separately analyze talar external shape within Pan (P. t. troglodytes, P. t. schweinfurthii, P. t. verus, P. paniscus) and Gorilla (G. g. gorilla, G. b. beringei, G. b. graueri) relative to degree of arboreality and body size. Pan and Gorilla are additionally analyzed together to determine if consistent shape differences exist within the genera.
Talar external shape was quantified using a weighted spherical harmonic analysis. Shape variation both within and among Pan and Gorilla was described using principal component analyses. Root mean square distances were calculated between taxon averages, and resampling statistics conducted to test for pairwise differences.
P. t. verus (most arboreal Pan) talar shape significantly differs from other Pan taxa (p < 0.05 for pairwise comparisons) driven by more asymmetrical trochlear rims and a medially-set talar head. P. t. troglodytes, P. t. schweinfurthii, and P. paniscus do not significantly differ (p > 0.05 for pairwise comparisons). All gorilla taxa exhibit significantly different talar morphologies (p < 0.007 for pairwise comparisons). The more terrestrial subspecies of G. beringei and P. troglodytes exhibit a superoinferiorly taller talar head/neck complex.
P. t. verus exhibits talar morphologies that have been previously related to more frequent arboreality. The adaptations in the more terrestrial G. beringei and P. troglodytes subspecies may serve to facilitate load transmission.

Across vertebrate diversity, limb bone morphology is typically expected to reflect differences in the habitats and functional tasks that species utilize. Arboreal vertebrates are often recognized to have longer limbs than terrestrial relatives, a feature thought to help extend the reach of limbs across gaps between branches. Among terrestrial vertebrates, longer limbs can experience greater bending moments that might expose bones to a greater risk of failure. However, changes in habitat or behavior can impose changes in the forces that bones experience. If locomotion imposed lower loads in trees than on the ground, such a release from loading demands might have produced conditions under which potential constraints on the evolution of long limbs were removed, making it easier for them to evolve in arboreal species. We tested for such environmental differences in limb bone loading using the green iguana (Iguana iguana), a species that readily walks over ground and climbs trees. We implanted strain gauges on the humerus and femur, and then compared loads between treatments modeling substrate conditions of arboreal habitats. For hindlimbs, inclined substrate angles were most correlated with strain increases, whereas the forelimbs had a similar pattern but of lesser magnitude. Unlike some other habitat transitions, these results do not support biomechanical release as a mechanism likely to have facilitated limb elongation. Instead, limb bone adaptations in arboreal habitats were likely driven by selective pressures other than responses to skeletal loading.

Across vertebrate diversity, limb bone morphology is typically expected to reflect differences in the habitats and functional tasks that species utilize. Arboreal vertebrates are often recognized to have longer limbs than terrestrial relatives, a feature thought to help extend the reach of limbs across gaps between branches. Among terrestrial vertebrates, longer limbs can experience greater bending moments that might expose bones to a greater risk of failure. However, changes in habitat or behavior can impose changes in the forces that bones experience. If locomotion imposed lower loads in trees than on the ground, such a release from loading demands might have produced conditions under which potential constraints on the evolution of long limbs were removed, making it easier for them to evolve in arboreal species. We tested for such environmental differences in limb bone loading using the green iguana (Iguana iguana), a species that readily walks over ground and climbs trees. We implanted strain gauges on the humerus and femur, and then compared loads between treatments modeling substrate conditions of arboreal habitats. For hindlimbs, inclined substrate angles were most correlated with strain increases, whereas the forelimbs had a similar pattern but of lesser magnitude. Unlike some other habitat transitions, these results do not support biomechanical release as a mechanism likely to have facilitated limb elongation. Instead, limb bone adaptations in arboreal habitats were likely driven by selective pressures other than responses to skeletal loading.

At what inclination does climbing begin? In this paper, we investigate the transition from walking to climbing in two species of parrot (Agapornis roseicollis and Nymphicus hollandicus) that are known to incorporate both their tail and their craniocervical system into the gait cycle during vertical climbing. Locomotor behaviors ranging in inclination were observed at angles between 0° and 90° for A. roseicollis, and 45°-85° degrees for N. hollandicus. Use of the tail in both species was observed at 45° inclination, and was joined at higher inclinations (> 65°) by use of the craniocervical system. Additionally, as inclination approached (but remained below) 90°, locomotor speeds were reduced while gaits were characterized by higher duty factors and lower stride frequency. These gait changes are consistent with those thought to increase stability. At 90°, A. roseicollis significantly increased its stride length, resulting in higher overall locomotor speed. Collectively these data demonstrate that the transition between horizontal walking and vertical climbing is gradual, incrementally altering several components of gait as inclinations increase. Such data underscore the need for further investigation into how exactly \"climbing\" is defined and the specific locomotor characteristics that differentiate this behavior from level walking.

Regionalization of the vertebral column can help animals adapt to different kinds of locomotion, including arboreal locomotion. Although functional axial regionalization has been described in both chameleons and arboreal mammals, no morphological basis for this functional regionalization in chameleons has been proposed. However, recent studies have described regionalization in the presacral vertebral column of other extant squamates. To investigate possible morphological regionalization in the vertebral column of chameleons, we took morphometric measurements from the presacral vertebrae of 28 chameleon species representing all extant chameleon genera, both fully arboreal and ground-dwelling, and performed comparative analyses. Our results support chameleons exhibiting three or four presacral morphological regions that correspond closely to those in other sauropsids, but we detected evolutionary shifts in vertebral traits occurring in only arboreal chameleons. Specifically, the anterior dorsal region in arboreal chameleons has more vertically oriented zygapophyseal joints, predicting decreased mediolateral flexibility. This shift is functionally significant because stiffening of the anterior thoracic vertebral column has been proposed to help bridge gaps between supports in primates. Thus, specialization of existing morphological regions in the vertebral column of chameleons may have played an important role in the evolution of extreme arboreal locomotion, paralleling the adaptations of arboreal primates.