Protocol to record and analyze primate leaping in three dimensions in the wild.

Several studies comparing primate locomotion under lab versus field conditions have shown the importance of implementing both types of studies, as each has their advantages and disadvantages. However, three-dimensional (3D) motion capture of primates has been challenging under natural conditions. In this study, we provide a detailed protocol on how to collect 3D biomechanical data on primate leaping in their natural habitat that can be widely implemented.

Pump and sway: Wild primates use compliant supports as a tool to augment leaping in the canopy.

Objectives: 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.

Materials And Methods: 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.

From such great heights: The effects of substrate height and the perception of risk on lemur locomotor mechanics.

Objectives: 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.

Materials And Methods: 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).

The Stabilizing Function of the Tail During Arboreal Quadrupedalism.

Locomotion on the narrow and compliant supports of the arboreal environment is inherently precarious. Previous studies have identified a host of morphological and behavioral specializations in arboreal animals broadly thought to promote stability when on precarious substrates. Less well-studied is the role of the tail in maintaining balance.

Asymmetrical gait kinematics of free-ranging callitrichine primates in response to changes in substrate diameter and orientation.

Arboreal environments present considerable biomechanical challenges for animals moving and foraging among substrates varying in diameter, orientation and compliance. Most studies of quadrupedal gait kinematics in primates and other arboreal mammals have focused on symmetrical walking gaits and the significance of diagonal sequence gaits. Considerably less research has examined asymmetrical gaits, despite their prevalence in small-bodied arboreal taxa.

Effects of substrate and phylogeny on quadrupedal gait in free-ranging platyrrhines.

Objectives: Primate diagonal sequence (DS) gaits are often argued to be an adaptation for moving and foraging in the fine-branch niche; however, existing data have come predominantly from laboratory studies that are limited in taxonomic breadth and fail to account for the structural and ecological variation of natural substrates. We test the extent to which substrate diameter and orientation influence gait sequence type and limb phase in free-ranging primates, as well as how phylogenetic relatedness might condition response patterns.

Materials And Methods: We filmed quadrupedal locomotion in 11 platyrrhine species at field sites in Ecuador and Costa Rica and measured the diameter and orientation of locomotor substrates using remote sensors.

The effects of natural substrate discontinuities on the quadrupedal gait kinematics of free-ranging Saimiri sciureus.

Wild primates encounter complex matrices of substrates that differ in size, orientation, height, and compliance, and often move on multiple, discontinuous substrates within a single bout of locomotion. Our current understanding of primate gait is limited by artificial laboratory settings in which primate quadrupedal gait has primarily been studied. This study analyzes wild Saimiri sciureus (common squirrel monkey) gait on discontinuous substrates to capture the realistic effects of the complex arboreal habitat on walking kinematics.

My branch is your branch: Talar morphology correlates with relative substrate size in platyrrhines at Tiputini Biodiversity Station, Ecuador.

Given that most species of primates are predominantly arboreal, maintaining the ability to move among branches of varying sizes has presumably been a common selective force in primate evolution. However, empirical evaluations of the relationships between morphological variation and characteristics of substrate geometry, such as substrate diameter relative to an animal's body mass, have been limited by the lack of quantified substrate usage in the wild. Here we use recently published quantitative data to assess the relationships between relative substrate size and talar morphology in nine New World monkey species at the Tiputini Biodiversity Station, Ecuador.

Functional and developmental influences on intraspecific variation in catarrhine vertebrae.

Objectives: We tested whether patterns of intraspecific variation in catarrhine vertebral shape are consistent with developmental or functional predictions. Intraspecific variation was compared across column regions, morphological features, and species. Transitional regions and later ossifying morphological features were predicted to exhibit increased variation.

Locomotor kinematics of tree squirrels (Sciurus carolinensis) in free-ranging and laboratory environments: Implications for primate locomotion and evolution.

The grasping capabilities and gait kinematics characteristic of primates are often argued to be adaptations for safely moving on small terminal branches. The goal of this study was to identify whether Eastern gray squirrels (Sciurus carolinensis)-arboreal rodents that frequently move and forage on small branches, lack primate-like grasping and gait patterns, and arguably represent extant analogs of a stem primate ancestor-adjust gait kinematics to narrow and nonhorizontal branches. We studied locomotor kinematics of free-ranging and laboratory-housed squirrels moving over various substrates.

A user's guide for the quantitative analysis of substrate characteristics and locomotor kinematics in free-ranging primates.

Objectives: Laboratory studies have yielded important insights into primate locomotor mechanics. Nevertheless, laboratory studies fail to capture the range of ecological and structural variation encountered by free-ranging primates. We present techniques for collecting kinematic data on wild primates using consumer grade high-speed cameras and demonstrate novel methods for quantifying metric variation in arboreal substrates.

Developments in development: What have we learned from primate locomotor ontogeny?

The importance of locomotion to evolutionary fitness has led to extensive study of primate locomotor behavior, morphology and ecology. Most previous research has focused on adult primates, but in the last few decades, increased attention to locomotor development has provided new insights toward our broader understanding of primate adaptation and evolution. Here, we review the contributions of this body of work from three basic perspectives.

Ontogenetic changes in foot strike pattern and calcaneal loading during walking in young children.

The assumption that the morphology of the human calcaneus reflects high and cyclical impact forces at heel strike during adult human walking has never been experimentally tested. Since a walking step with a heel strike is an emergent behavior in children, an ontogenetic study provides a natural experiment to begin testing the relationship between the mechanics of heel strike and calcaneal anatomy. This study examined the ground reaction forces (GRFs) of stepping in children to determine the location of the center of pressure (COP) relative to the calcaneus and the orientation and magnitude of ground reaction forces during foot contact.

Ontogenetic changes in limb postures and their impact on effective limb length in baboons (Papio cynocephalus).

Objectives: Digitigrade hand and foot postures and extended elbows and knees are considered adaptations to running in cursorial mammals because they increase effective limb lengths (ELLs). However, the relationship between digitigrady and ELL in primates is not well understood. We documented the ontogeny of limb postures in baboons to better understand the function of digitigrady during walking.

Effects of Substrate Size and Orientation on Quadrupedal Gait Kinematics in Mouse Lemurs (Microcebus murinus).

As the smallest living primate, the mouse lemur is a suitable model for reconstructing the locomotor mechanisms by which primate ancestors might have responded to the challenges of an arboreal environment. In this study, we tested the effects of substrate diameter and orientation on quadrupedal gait kinematics in mouse lemurs (Microcebus murinus). Mouse lemurs highly preferred asymmetrical to symmetrical gaits as they moved across a flat board and poles of three diameters (2.

Human quadrupeds, primate quadrupedalism, and Uner Tan Syndrome.

Since 2005, an extensive literature documents individuals from several families afflicted with "Uner Tan Syndrome (UTS)," a condition that in its most extreme form is characterized by cerebellar hypoplasia, loss of balance and coordination, impaired cognitive abilities, and habitual quadrupedal gait on hands and feet. Some researchers have interpreted habitual use of quadrupedalism by these individuals from an evolutionary perspective, suggesting that it represents an atavistic expression of our quadrupedal primate ancestry or "devolution." In support of this idea, individuals with "UTS" are said to use diagonal sequence quadrupedalism, a type of quadrupedal gait that distinguishes primates from most other mammals.

Body size and the small branch niche: using marsupial ontogeny to model primate locomotor evolution.

Recently proposed ancestral locomotor and morphological 'stages' leading to the evolution of primates have emphasized small body size, and a transition from a clawed non-grasping stage, to a clawed, grasping stage with clawless opposable hallux, to a fully-nailed primate with grasping extremities. This evolutionary transition was presumably associated with frequent use of the small branch niche. To model elements of these evolutionary transitions, we investigate how body size, substrate size, substrate orientation and grasping morphology interact to influence quadrupedal kinematics within and between ontogenetic samples of two small-bodied marsupials, one arboreal (Petaurus breviceps) and the other mainly terrestrial (Monodelphis domestica).

A new look at the Dynamic Similarity Hypothesis: the importance of swing phase.

The Dynamic Similarity Hypothesis (DSH) suggests that when animals of different size walk at similar Froude numbers (equal ratios of inertial and gravitational forces) they will use similar size-corrected gaits. This application of similarity theory to animal biomechanics has contributed to fundamental insights in the mechanics and evolution of a diverse set of locomotor systems. However, despite its popularity, many mammals fail to walk with dynamically similar stride lengths, a key element of gait that determines spontaneous speed and energy costs.

Reevaluation of the lumbosacral region of Oreopithecus bambolii.

Functional interpretations of the postcranium of the late Miocene ape Oreopithecus bambolii are controversial. The claim that Oreopithecus practiced habitual terrestrial bipedalism is partly based on restored postcranial remains originally recovered from Baccinello, Tuscany (Köhler and Moyà-Solà, 1997). The lower lumbar vertebrae of BA#72 were cited as evidence that Oreopithecus exhibits features indicative of a lordotic lumbar spine, including dorsal wedging of the vertebral bodies and a caudally progressive increase in postzygapophyseal interfacet distance.

Kinematics of quadrupedal locomotion in sugar gliders (Petaurus breviceps): effects of age and substrate size.

Arboreal mammals face unique challenges to locomotor stability. This is particularly true with respect to juveniles, who must navigate substrates similar to those traversed by adults, despite a reduced body size and neuromuscular immaturity. Kinematic differences exhibited by juveniles and adults on a given arboreal substrate could therefore be due to differences in body size relative to substrate size, to differences in neuromuscular development, or to both.

Morphological correlates of tail length in the catarrhine sacrum.

Taillessness is a distinctive synapomorphy of the Hominoidea that has implications for interpretation of the locomotor behaviors and phylogenetic affinities of the clade's earliest members. However, difficulties persist in confidently identifying taillessness in the catarrhine fossil record, stemming largely from our limited knowledge of the anatomical features with which the tail is associated. Here, we compare the morphology of the sacrum, the sole bony link between the tail and the rest of the body, among extant tailless hominoids and a broad sample of extant cercopithecoids known to vary in tail length (i.

Is primate-like quadrupedalism necessary for fine-branch locomotion? A test using sugar gliders (Petaurus breviceps).

Locomotor features shared by arboreal marsupials and primates are frequently cited as a functional complex that evolved in the context of a "fine branch niche." Adaptation to a fine branch niche cannot be understood without considering that small and large arboreal mammals may differ in their biomechanical response to a given branch size. We tested the effects of substrate diameter and orientation on quadrupedal kinematics in a small arboreal marsupial (the sugar glider, Petaurus breviceps).

Understanding hind limb weight support in chimpanzees with implications for the evolution of primate locomotion.

Most quadrupedal mammals support a larger amount of body weight on their forelimbs compared with their hind limbs during locomotion, whereas most primates support more of their body weight on their hind limbs. Increased hind limb weight support is generally interpreted as an adaptation that reduces stress on primates' highly mobile forelimb joints. Thus, increased hind limb weight support was likely vital for the evolution of primate arboreality.

Fetal load and the evolution of lumbar lordosis in bipedal hominins.

As predicted by Darwin, bipedal posture and locomotion are key distinguishing features of the earliest known hominins. Hominin axial skeletons show many derived adaptations for bipedalism, including an elongated lumbar region, both in the number of vertebrae and their lengths, as well as a marked posterior concavity of wedged lumbar vertebrae, known as a lordosis. The lordosis stabilizes the upper body over the lower limbs in bipeds by positioning the trunk's centre of mass (COM) above the hips.

Morphological and functional differentiation in the lumbar spine of lorisids and galagids.

The striking contrast in positional behavior exhibited by lorisids (slow quadrupedalism/suspension) and galagids (leaping/quadrupedalism) is well reflected in their postcranial morphology, particularly in the limbs. Although they exhibit very different spinal postures and movements, vertebral adaptations have been less well explored in these taxa. This study addressed morphological and functional differentiation in the lumbar vertebrae of four species of lorisids and five species of galagids.