Footprint evidence of early hominin locomotor diversity at Laetoli, Tanzania.

Bipedal trackways discovered in 1978 at Laetoli site G, Tanzania and dated to 3.66 million years ago are widely accepted as the oldest unequivocal evidence of obligate bipedalism in the human lineage1-3. Another trackway discovered two years earlier at nearby site A was partially excavated and attributed to a hominin, but curious affinities with bears (ursids) marginalized its importance to the paleoanthropological community, and the location of these footprints fell into obscurity3-5. In 2019, we located, excavated and cleaned the site A trackway, producing a digital archive using 3D photogrammetry and laser scanning. Here we compare the footprints at this site with those of American black bears, chimpanzees and humans, and we show that they resemble those of hominins more than ursids. In fact, the narrow step width corroborates the original interpretation of a small, cross-stepping bipedal hominin. However, the inferred foot proportions, gait parameters and 3D morphologies of footprints at site A are readily distinguished from those at site G, indicating that a minimum of two hominin taxa with different feet and gaits coexisted at Laetoli.

Adaptations for bipedal walking: Musculoskeletal structure and three-dimensional joint mechanics of humans and bipedal chimpanzees (Pan troglodytes).

Humans are unique among apes and other primates in the musculoskeletal design of their lower back, pelvis, and lower limbs. Here, we describe the three-dimensional ground reaction forces and lower/hindlimb joint mechanics of human and bipedal chimpanzees walking over a full stride and test whether: 1) the estimated limb joint work and power during the stance phase, especially the single-support period, is lower in humans than bipedal chimpanzees, 2) the limb joint work and power required for limb swing is lower in humans than in bipedal chimpanzees, and 3) the estimated total mechanical power during walking, accounting for the storage of passive elastic strain energy in humans, is lower in humans than in bipedal chimpanzees. Humans and bipedal chimpanzees were compared at matched dimensionless and dimensional velocities. Our results indicate that humans walk with significantly less work and power output in the first double-support period and the single-support period of stance, but markedly exceed chimpanzees in the second double-support period (i.e., push-off). Humans generate less work and power in limb swing, although the species difference in limb swing power was not statistically significant. We estimated that total mechanical positive 'muscle fiber' work and power were 46.9% and 35.8% lower, respectively, in humans than in bipedal chimpanzees at matched dimensionless speeds. This is due in part to mechanisms for the storage and release of elastic energy at the ankle and hip in humans. Furthermore, these results indicate distinct 'heel strike' and 'lateral balance' mechanics in humans and bipedal chimpanzees and suggest a greater dissipation of mechanical energy through soft tissue deformations in humans. Together, our results document important differences between human and bipedal chimpanzee walking mechanics over a full stride, permitting a more comprehensive understanding of the mechanics and energetics of chimpanzee bipedalism and the evolution of hominin walking.

The loss of the 'pelvic step' in human evolution.

Human bipedalism entails relatively short strides compared with facultatively bipedal primates. Unique non-sagittal-plane motions associated with bipedalism may account for part of this discrepancy. Pelvic rotation anteriorly translates the hip, contributing to bipedal stride length (i.e. the 'pelvic step'). Facultative bipedalism in non-human primates entails much larger pelvic rotation than in humans, suggesting that a larger pelvic step may contribute to their relatively longer strides. We collected data on the pelvic step in bipedal chimpanzees and over a wide speed range of human walking. At matched dimensionless speeds, humans have 26.7% shorter dimensionless strides, and a pelvic step 5.4 times smaller than bipedal chimpanzees. Differences in pelvic rotation explain 31.8% of the difference in dimensionless stride length between the two species. We suggest that relative stride lengths and the pelvic step have been significantly reduced throughout the course of hominin evolution.

Comparison of pediatric brain-dead donors to donation after circulatory death donors in the United States.

In pediatrics, an increasing need for transplantable organs exists. This study aimed to describe the epidemiology of pediatric deceased donors in the United States. This retrospective observational study utilized data from the Organ Procurement and Transplantation Network (OPTN) from 2000 to 2015. Patients were stratified based on method of organ donation. Demographic variables and mechanism of death were then compared. A total of 14,481 deceased pediatric organ donors, donation after brain death (DBD) and donation after circulatory death (DCD), were included in the study, of which 8% were DCD donors. A significant difference (p<0.001) existed between the two donor groups with respect to ethnicity and mechanism of death. The annual trend of DCD and DBD donors showed an inverse relationship. During the 15-year study period the number of DBD donors decreased from 985 to 785 per year while DCD donors increased from 15 to 146 per year. As well, overall organs transplanted per year decreased from 3,475 to 3,117 over the 15-year study period. Significant differences exist between pediatric DBD donors and DCD donors, specifically with respect to ethnicity and mechanism of death. The number of pediatric DBD donors is decreasing while the number of pediatric DCD is slowly rising, making it increasingly important to be able to characterize these donors to better identify eligible DCD donors to optimize organ utilization.

The biomechanics of knuckle-walking: 3-D kinematics of the chimpanzee and macaque wrist, hand and fingers.

The origin and evolution of knuckle-walking has long been a key focus in understanding African ape, including human, origins. Yet, despite numerous studies documenting morphological characteristics potentially associated with knuckle-walking, little quantitative three-dimensional (3-D) data exist of forelimb motion during knuckle-walking. Nor do any comparative 3-D data exist for hand postures used during quadrupedalism in monkeys. This lack of data has limited the testability of proposed adaptations for knuckle-walking in African apes. This study presents the first 3-D kinematic data of the wrist, hand and metacarpophalangeal joints during knuckle-walking in chimpanzees and in macaques using digitigrade and palmigrade hand postures. These results clarify the unique characteristics of, and commonalities between, knuckle-walking and digitigrady/palmigrady in multiple planes of motion. Notably, chimpanzees utilized more wrist ulnar deviation than any macaque hand posture. Maximum extension of the chimpanzee wrist was slight (5-20 deg) and generally overlapped with macaque digitigrady. Metacarpophalangeal joint motion displayed distinct differences between digits in both species, likely related to the timing of force application. These data also reveal that maximum metacarpophalangeal extension angles during knuckle-walking (26-59 deg) were generally higher than previously considered. In macaques, maximum metacarpophalangeal extension during digitigrady and palmigrady overlapped for most digits, highlighting additional complexity in the interpretation of skeletal features that may be related to limiting metacarpophalangeal motion. Most importantly, however, these new 3-D data serve as a fundamental dataset with which evaluation of proposed musculoskeletal adaptations for knuckle-walking can be tested.

Digitization of the Nissen-Riesen chimpanzee radiological growth series.

Longitudinal morphological growth data of apes are incredibly difficult to obtain. Long life histories, combined with practical and ethical issues of obtaining such long-term data have resulted in few longitudinal data sets in chimpanzees of known chronological ages. One classic, long-term growth study of chimpanzees was that of Drs Nissen and Riesen initiated at the Yale Laboratories of Primate Biology in 1939. Through that study, whole-body radiological images were taken on a regular basis from a "normative" group of chimpanzees from birth to adulthood. Here we have digitized the known remaining radiographs from that growth study, many of which are deteriorating, and uploaded the data set to the free, online database MorphoSource. The database comprises 3,568 X-ray images of 15 of the 16 chimpanzee subjects in the normative group and 1 individual from an experimental group. Herein, we briefly review the historical context of this study and specific details of the data set.

Influences of passive intervertebral range of motion on cervical vertebral form.

OBJECTIVES: The cervical spine is the junction between the head and trunk, and it therefore facilitates head mobility and stability. The goal of this study is to test several predictions regarding cervical morphology and intervertebral ranges of motion. MATERIALS AND METHODS: Intervertebral ranges of motion for 12 primate species were collected via radiographs or taken from the literature. Morphometric data describing functionally relevant aspects of cervical vertebral morphology were obtained from museum specimens representing these species. We tested for correlations between intervertebral movement and vertebral form using phylogenetic generalized least-squares regression. RESULTS: Results demonstrate limited support for the hypothesis that range of motion (ROM) is influenced by cervical vertebral morphology. Few morphological variables correlate with ROM and no relationship is consistently significant across cervical joints. DISCUSSION: These results indicate that the relationship between vertebral morphology and joint ranges of motion is, at most, weak, providing little support the use of bony morphology to reconstruct axial mobility in fossil specimens. Future work should investigate the role of soft tissues in vertebral joint stability.

Methadone's Effect on Cardiac Repolarization: Safety in the PICU.

OBJECTIVES: Opioids are routinely used in the PICU. Methadone is an effective method of preventing and treating iatrogenic opioid withdrawal; however, it carries an Food and Drug Administration Boxed Warning due to the potential to prolong the corrected QT interval and potentially lead to life-threatening arrhythmias. Guidelines on the safe use of methadone have limited applicability to children since their cardiac intervals differ from those of adults. There is little data on the electrophysiologic effects in the pediatric population. We set out to describe the safety of methadone use in the PICU, hypothesizing that methadone does not cause a significant change in corrected QT interval from baseline. DESIGN: Retrospective cohort study. SETTING: Children's Hospital of Wisconsin, Milwaukee, WI. PATIENTS: Fifty-one patients, age less than or equal to 18 years old, initiated on methadone during PICU admission, over an 11-month period, for the prevention or treatment of opioid withdrawal. INTERVENTIONS: Retrospective data queried from the electronic health record and stored telemetry waveforms obtained from an automated real-time patient data acquisition software system (BedMasterEx; Anandic Medical Systems AG, Feuerthalen, Switzerland). MEASUREMENTS AND MAIN RESULTS: Corrected QT intervals were not significantly different at 12 hours, 96 hours, or PICU discharge (p values: 0.57, 0.54, and 0.34) when compared to baseline. The median change in corrected QT from baseline to 12 hours after the first dose of methadone was 5 ms (interquartile range, -12 to 11 ms), 0 ms to steady state (interquartile range, -18 to 18 ms), and 5 ms from baseline to 12 hours after the highest dose of methadone (interquartile range, -14 to 16 ms). The most common primary diagnosis was structural heart disease (29% of subjects) in our cohort and every subject that experienced an increase in corrected QT interval greater than or equal to 40 ms had some form of structural heart disease. CONCLUSIONS: Methadone did not significantly prolong the corrected QT interval in a population of critically ill children, suggesting that it can be safely used in this population, although patients with structural heart disease may warrant closer monitoring.

Use and Outcomes of Nasotracheal Intubation Among Patients Requiring Mechanical Ventilation Across U.S. PICUs.

OBJECTIVES: The use and outcomes of nasotracheal intubation in pediatric patients requiring mechanical ventilation have not been quantified. Our goal is to identify prevalence of use, associated factors, and outcomes of nasotracheal versus orotracheal intubation in patients requiring mechanical ventilation. DESIGN: Retrospective cohort study using deidentified data from the Virtual Pediatric Systems database. Data from PICU admissions from January 1, 2015, to December 31, 2016 were analyzed. SETTING: One hundred twenty-one PICUs located within the United States. PATIENTS: PICU admissions requiring an endotracheal tube-either nasotracheal or orotracheal-were included. Those with a tracheostomy tube present at admission were excluded from the study. INTERVENTIONS: Not applicable. MEASUREMENTS AND MAIN RESULTS: Among the 121 PICUs included in the study, 64 PICUs (53%) had zero nasotracheal intubations during the reviewed time period. There were 12,088 endotracheal intubations analyzed, and 680 of them (5.6%) were nasotracheal. Of those patients nasotracheally intubated, most were under 2 years old (88.1%), and 82.2% of them were classified as a cardiac patient. Among these young cardiac patients, the rate of unplanned extubation was 0% in the nasotracheal intubated versus 2.1% in the orotracheal intubated group (p < 0.001) CONCLUSIONS:: Nasotracheal intubation is used in a minority of U.S. PICUs and mainly among young cardiac patients. Nasotracheal intubation is associated with a lower rate of unplanned extubations in this patient population. Future prospective studies analyzing the benefits and complications of nasotracheal versus orotracheal intubation in pediatric patients requiring mechanical ventilation are indicated.

Great ape thorax and shoulder configuration-An adaptation for arboreality or knuckle-walking?

Great apes exhibit a suite of morphological traits of the shoulder and upper thorax that have traditionally been linked to orthograde arborealism. Recently it has been proposed that these traits are instead adaptations for knuckle-walking, and more broadly, that knuckle-walking itself is an adaptation for shock absorption during terrestriality. Here we test several tenets of these hypotheses using kinematic and kinetic data from chimpanzees and macaques, and electromyographic data of shoulder muscle activity in chimpanzees. We collected 3D kinematic data to quantify motion of the acromion and trunk during quadrupedalism and vertical climbing in chimpanzees as well as ground reaction forces to investigate the presence and magnitude of impact transient forces during terrestrial locomotion in chimpanzees and macaques. We also investigated patterns of recruitment of select forelimb musculature (triceps brachii and serratus anterior) using previously collected data in chimpanzees to determine whether these muscles may function to absorb impact transient forces. We found that the acromion is significantly more elevated in vertical climbing than during knuckle-walking, while dorsoventral ranges and magnitudes of motion were similar between gaits. Ground reaction forces indicate that only a minority of strides in either chimpanzees or macaques have transient forces and, when present, these transient forces as well as loading rates are small. Electromyographic results show that activity of the triceps brachii may facilitate energy absorption while serratus anterior likely functions to support the trunk, as in other primates. Our data suggest there is little to no evidence supporting recent hypotheses that the African ape upper thorax and shoulder configuration is an adaptation for knuckle-walking, or more broadly, that knuckle-walking exists as an adaptation to absorb impact shock during terrestriality. We do however find some evidence that shoulder configuration allows greater scapular elevation in chimpanzees during arboreal behaviors (e.g., vertical climbing).

Step width and frontal plane trunk motion in bipedal chimpanzee and human walking.

Human bipedalism is characterized by mediolateral oscillations of the center of mass (CoM) between the feet. The preferred step widths and CoM oscillations used by humans likely represent a trade-off of several factors (e.g., stance and swing phase costs). However, it is difficult to assess whether human frontal plane control strategies are unique given few detailed data on frontal plane motion during facultative bipedalism in apes. Here, we collected three-dimensional kinematic and kinetic data in humans and chimpanzees to investigate the relationship between step width, mediolateral CoM motion, frontal plane trunk kinematics, and CoM power during bipedalism. Chimpanzee bipedalism entails mediolateral CoM oscillations and step widths that are (scaled to lower/hind limb length) three times larger than those of humans. Chimpanzees use a combination of linear and angular motion of the trunk and list the entire trunk, and especially thorax, over the stance side foot, generating large mediolateral shifts in the CoM, whereas humans utilize little angular motion within the trunk. Larger mediolateral CoM motions do not have a significant effect on CoM power. Similarities between bipedal chimpanzees and other bipedal non-human primates (macaques and gibbons) indicate that narrow CoM motions are unique to humans and are likely due to our adducted hips and valgus knees. Valgus knees appear early in the human fossil record (∼3.6 Ma), contemporaneous with the Laetoli footprints. However, fossils attributed to Ardipithecus ramidus (∼4.4 Ma) suggest that the earliest hominins may have lacked a hominin-like degree of knee valgus. If correct, this suggests that this species may have used wide steps, larger mediolateral CoM motions, and perhaps larger trunk motions during bipedal walking. Finally, we present a novel means to estimate mediolateral CoM motion from trackway step width, and estimate that the Laetoli G track maker used CoM motions within the human range.

Unexpected terrestrial hand posture diversity in wild mountain gorillas.

OBJECTIVES: Gorillas, along with chimpanzees and bonobos, are ubiquitously described as 'knuckle-walkers.' Consequently, knuckle-walking (KW) has been featured pre-eminently in hypotheses of the pre-bipedal locomotor behavior of hominins and in the evolution of locomotor behavior in apes. However, anecdotal and behavioral accounts suggest that mountain gorillas may utilize a more complex repertoire of hand postures, which could alter current interpretations of African ape locomotion and its role in the emergence of human bipedalism. Here we documented hand postures during terrestrial locomotion in wild mountain gorillas to investigate the frequency with which KW and other hand postures are utilized in the wild. MATERIALS AND METHODS: Multiple high-speed cameras were used to record bouts of terrestrial locomotion of 77 habituated mountain gorillas at Bwindi Impenetrable National Park (Uganda) and Volcanoes National Park (Rwanda). RESULTS: We captured high-speed video of hand contacts in 8% of the world's population of mountain gorillas. Our results reveal that nearly 40% of these gorillas used "non-KW" hand postures, and these hand postures constituted 15% of all hand contacts. Some of these "non-KW" hand postures have never been documented in gorillas, yet match hand postures previously identified in orangutans. DISCUSSION: These results highlight a previously unrecognized level of hand postural diversity in gorillas, and perhaps great apes generally. Although present at lower frequencies than KW, we suggest that the possession of multiple, versatile hand postures present in wild mountain gorillas may represent a shared feature of the African ape and human clade (or even great ape clade) rather than KW per se.

Central ECMO for circulatory failure following pediatric liver transplantation.

We describe the case of a 4-year-old male with a past medical history significant for nephrotic syndrome, short-bowel syndrome and fulminant hepatic failure status post (s/p) liver transplant (LT) who developed early post-transplant allograft dysfunction (hyperbilirubinemia, coagulopathy) and septic shock requiring central extracorporeal membrane oxygenation (ECMO). He remained on ECMO for 85 hours before he was decannulated without event and later underwent repeat LT. This case highlights the potential of central ECMO to provide the circulatory output necessary to reverse distributive shock physiology in patients with sepsis and hepatic dysfunction following LT. Furthermore, this is the first documented example of central ECMO as a bridge to recovery for repeat LT.

The evolution of vertebral formulae in Hominoidea.

Primate vertebral formulae have long been investigated because of their link to locomotor behavior and overall body plan. Knowledge of the ancestral vertebral formulae in the hominoid tree of life is necessary to interpret the pattern of evolution among apes, and to critically evaluate the morphological adaptations involved in the transition to hominin bipedalism. Though many evolutionary hypotheses have been proposed based on living and fossil species, the application of quantitative phylogenetic methods for thoroughly reconstructing ancestral vertebral formulae and formally testing patterns of vertebral evolution is lacking. To estimate the most probable scenarios of hominoid vertebral evolution, we utilized an iterative ancestral state reconstruction approach to determine likely ancestral vertebral counts in apes, humans, and other anthropoid out-groups. All available ape and hominin fossil taxa with an inferred regional vertebral count were included in the analysis. Sensitivity iterations were performed both by changing the phylogenetic position of fossil taxa with a contentious placement, and by changing the inferred number of vertebrae in taxa with uncertain morphology. Our ancestral state reconstruction results generally support a short-backed hypothesis of human evolution, with a Pan-Homo last common ancestor possessing a vertebral formulae of 7:13:4:6 (cervical:thoracic:lumbar:sacral). Our results indicate that an initial reduction in lumbar vertebral count and increase in sacral count is a synapomorphy of crown hominoids (supporting an intermediate-backed hypothesis for the origins of the great ape-human clade). Further reduction in lumbar count occurs independently in orangutans and African apes. Our results highlight the complexity and homoplastic nature of vertebral count evolution, and give little support to the long-backed hypothesis of human evolution.

Chimpanzee and human midfoot motion during bipedal walking and the evolution of the longitudinal arch of the foot.

The longitudinal arch of the human foot is commonly thought to reduce midfoot joint motion to convert the foot into a rigid lever during push off in bipedal walking. In contrast, African apes have been observed to exhibit midfoot dorsiflexion following heel lift during terrestrial locomotion, presumably due to their possession of highly mobile midfoot joints. This assumed dichotomy between human and African ape midfoot mobility has recently been questioned based on indirect assessments of in vivo midfoot motion, such as plantar pressure and cadaver studies; however, direct quantitative analyses of African ape midfoot kinematics during locomotion remain scarce. Here, we used high-speed motion capture to measure three-dimensional foot kinematics in two male chimpanzees and five male humans walking bipedally at similar dimensionless speeds. We analyzed 10 steps per chimpanzee subject and five steps per human subject, and compared ranges of midfoot motion between species over stance phase, as well as within double- and single-limb support periods. Contrary to expectations, humans used a greater average range of midfoot motion than chimpanzees over the full duration of stance. This difference was driven by humans' dramatic plantarflexion and adduction of the midfoot joints during the second double-limb support period, which likely helps the foot generate power during push off. However, chimpanzees did use slightly but significantly more midfoot dorsiflexion than humans in the single limb-support period, during which heel lift begins. These results indicate that both stiffness and mobility are important to longitudinal arch function, and that the human foot evolved to utilize both during push off in bipedal walking. Thus, the presence of human-like midfoot joint morphology in fossil hominins should not be taken as indicating foot rigidity, but may signify the evolution of pedal anatomy conferring enhanced push off mechanics.

Chimpanzee ankle and foot joint kinematics: Arboreal versus terrestrial locomotion.

OBJECTIVES: Many aspects of chimpanzee ankle and midfoot joint morphology are believed to reflect adaptations for arboreal locomotion. However, terrestrial travel also constitutes a significant component of chimpanzee locomotion, complicating functional interpretations of chimpanzee and fossil hominin foot morphology. Here we tested hypotheses of foot motion and, in keeping with general assumptions, we predicted that chimpanzees would use greater ankle and midfoot joint ranges of motion during travel on arboreal supports than on the ground. METHODS: We used a high-speed motion capture system to measure three-dimensional kinematics of the ankle and midfoot joints in two male chimpanzees during three locomotor modes: terrestrial quadrupedalism on a flat runway, arboreal quadrupedalism on a horizontally oriented tree trunk, and climbing on a vertically oriented tree trunk. RESULTS: Chimpanzees used relatively high ankle joint dorsiflexion angles during all three locomotor modes, although dorsiflexion was greatest in arboreal modes. They used higher subtalar joint coronal plane ranges of motion during terrestrial and arboreal quadrupedalism than during climbing, due in part to their use of high eversion angles in the former. Finally, they used high midfoot inversion angles during arboreal locomotor modes, but used similar midfoot sagittal plane kinematics across all locomotor modes. DISCUSSION: The results indicate that chimpanzees use large ranges of motion at their various ankle and midfoot joints during both terrestrial and arboreal locomotion. Therefore, we argue that chimpanzee foot anatomy enables a versatile locomotor repertoire, and urge caution when using foot joint morphology to reconstruct arboreal behavior in fossil hominins.

Three-dimensional kinematics of the pelvis and hind limbs in chimpanzee (Pan troglodytes) and human bipedal walking.

The common chimpanzee (Pan troglodytes) is a facultative biped and our closest living relative. As such, the musculoskeletal anatomies of their pelvis and hind limbs have long provided a comparative context for studies of human and fossil hominin locomotion. Yet, how the chimpanzee pelvis and hind limb actually move during bipedal walking is still not well defined. Here, we describe the three-dimensional (3-D) kinematics of the pelvis, hip, knee and ankle during bipedal walking and compare those values to humans walking at the same dimensionless and dimensional velocities. The stride-to-stride and intraspecific variations in 3-D kinematics were calculated using the adjusted coefficient of multiple correlation. Our results indicate that humans walk with a more stable pelvis than chimpanzees, especially in tilt and rotation. Both species exhibit similar magnitudes of pelvis list, but with segment motion that is opposite in phasing. In the hind limb, chimpanzees walk with a more flexed and abducted limb posture, and substantially exceed humans in the magnitude of hip rotation during a stride. The average stride-to-stride variation in joint and segment motion was greater in chimpanzees than humans, while the intraspecific variation was similar on average. These results demonstrate substantial differences between human and chimpanzee bipedal walking, in both the sagittal and non-sagittal planes. These new 3-D kinematic data are fundamental to a comprehensive understanding of the mechanics, energetics and control of chimpanzee bipedalism.

Center of mass mechanics of chimpanzee bipedal walking.

Center of mass (CoM) oscillations were documented for 81 bipedal walking strides of three chimpanzees. Full-stride ground reaction forces were recorded as well as kinematic data to synchronize force to gait events and to determine speed. Despite being a bent-hip, bent-knee (BHBK) gait, chimpanzee walking uses pendulum-like motion with vertical oscillations of the CoM that are similar in pattern and relative magnitude to those of humans. Maximum height is achieved during single support and minimum height during double support. The mediolateral oscillations of the CoM are more pronounced relative to stature than in human walking when compared at the same Froude speed. Despite the pendular nature of chimpanzee bipedalism, energy recoveries from exchanges of kinetic and potential energies are low on average and highly variable. This variability is probably related to the poor phasic coordination of energy fluctuations in these facultatively bipedal animals. The work on the CoM per unit mass and distance (mechanical cost of transport) is higher than that in humans, but lower than that in bipedally walking monkeys and gibbons. The pronounced side sway is not passive, but constitutes 10% of the total work of lifting and accelerating the CoM. CoM oscillations of bipedally walking chimpanzees are distinctly different from those of BHBK gait of humans with a flat trajectory, but this is often described as "chimpanzee-like" walking. Human BHBK gait is a poor model for chimpanzee bipedal walking and offers limited insights for reconstructing early hominin gait evolution.