Neil T. Roach

Elastic energy storage in the shoulder and the evolution of high-speed throwing in Homo

Some primates, including chimpanzees, throw objects occasionally, but only humans regularly throw projectiles with high speed and accuracy. Darwin noted that the unique throwing abilities of humans, which were made possible when bipedalism emancipated the arms, enabled foragers to hunt effectively using projectiles. However, there has been little consideration of the evolution of throwing in the years since Darwin made his observations, in part because of a lack of evidence of when, how and why hominins evolved the ability to generate high-speed throws. Here we use experimental studies of humans throwing projectiles to show that our throwing capabilities largely result from several derived anatomical features that enable elastic energy storage and release at the shoulder. These features first appear together approximately 2 million years ago in the species Homo erectus. Taking into consideration archaeological evidence suggesting that hunting activity intensified around this time, we conclude that selection for throwing as a means to hunt probably had an important role in the evolution of the genus Homo.

The Middle Stone Age of the northern Kenyan Rift: age and context of new archaeological sites from the Kapedo Tuffs

Rift Valley sites in southern Ethiopia and northern Kenya preserve the oldest fossil remains attributed to Homo sapiens and the earliest archaeological sites attributed to the Middle Stone Age (MSA). New localities from the Kapedo Tuffs augment the sparse sample of MSA sites from the northern Kenya Rift. Tephrostratigraphic correlation with dated pyroclastic deposits from the adjacent volcano Silali suggests an age range of 135-123ka for archaeological sites of the Kapedo Tuffs. Comparisons of the Kapedo Tuffs archaeological assemblages with those from the adjacent Turkana and Baringo basins show broad lithic technological similarity but reveal that stone raw material availability is a key factor in explaining typologically defined archaeological variability within this region. Spatially and temporally resolved comparisons such as this provide the best means to link the biological and behavioral variation manifest in the record of early Homo sapiens.

The effect of humeral torsion on rotational range of motion in the shoulder and throwing performance

Several recent studies have found that throwing athletes typically have lower humeral torsion (retroversion) and a greater range of external rotation at the shoulder than non‐athletes. How these two parameters are related is debated. This study uses data from a sample of both throwers and non‐throwers to test a new model that predicts torsion values from a range of motion data. The model proposes a series of predicted regressions which can help provide new insight into the factors affecting rotational range of motion at the shoulder. Humeral torsion angles were measured from computed tomography scans collected from 25 male subjects. These values are compared to predicted torsion values for the same subjects calculated from both kinematic and goniometric range‐of‐motion data. Results show that humeral torsion is negatively correlated (goniometric: r = −0.409, P = 0.047; kinematic: r = −0.442, P = 0.035) with external rotational range of motion and positively correlated (goniometric: r = 0.741, P < 0.001; kinematic: r = 0.559, P = 0.006) with internal rotational range of motion. The predicted torsion values are highly correlated (goniometric: r = 0.815, P < 0.001; kinematic: r = 0.617, P = 0.006) with actual torsion values. Deviations in the data away from predicted equations highlight significant differences between high torsion and low torsion individuals that may have significant functional consequences. The method described here may be useful for non‐invasively assessing the degree of torsion in studies of the evolution and biomechanics of the shoulder and arm, and for testing hypotheses about the etiology of repetitive stress injuries among athletes and others who throw frequently.

Fossil hominin shoulders support an African ape-like last common ancestor of humans and chimpanzees

Significance Knowing the direction and pace of evolutionary change is critical to understanding what selective forces shaped our ancestors. Unfortunately, the human fossil record is sparse, and little is known about the earliest members of our lineage. This unresolved ancestor complicates reconstructions of what behavioral shifts drove major speciation events. Using 3D shape measurements of the shoulder, we tested competing evolutionary models of the last common ancestor against the fossil record. We found that a sustained shift in scapular shape occurred during hominin evolution from an African ape-like ancestor to a modern human-like form, first present in our genus, Homo. These data suggest a long, gradual shift out of the trees and increased reliance on tools as our ancestors became more terrestrial. Reconstructing the behavioral shifts that drove hominin evolution requires knowledge of the timing, magnitude, and direction of anatomical changes over the past ∼6–7 million years. These reconstructions depend on assumptions regarding the morphotype of the Homo–Pan last common ancestor (LCA). However, there is little consensus for the LCA, with proposed models ranging from African ape to orangutan or generalized Miocene ape-like. The ancestral state of the shoulder is of particular interest because it is functionally associated with important behavioral shifts in hominins, such as reduced arboreality, high-speed throwing, and tool use. However, previous morphometric analyses of both living and fossil taxa have yielded contradictory results. Here, we generated a 3D morphospace of ape and human scapular shape to plot evolutionary trajectories, predict ancestral morphologies, and directly test alternative evolutionary hypotheses using the hominin fossil evidence. We show that the most parsimonious model for the evolution of hominin shoulder shape starts with an African ape-like ancestral state. We propose that the shoulder evolved gradually along a single morphocline, achieving modern human-like configuration and function within the genus Homo. These data are consistent with a slow, progressive loss of arboreality and increased tool use throughout human evolution.

Footprints reveal direct evidence of group behavior and locomotion in Homo erectus

Bipedalism is a defining feature of the human lineage. Despite evidence that walking on two feet dates back 6–7 Ma, reconstructing hominin gait evolution is complicated by a sparse fossil record and challenges in inferring biomechanical patterns from isolated and fragmentary bones. Similarly, patterns of social behavior that distinguish modern humans from other living primates likely played significant roles in our evolution, but it is exceedingly difficult to understand the social behaviors of fossil hominins directly from fossil data. Footprints preserve direct records of gait biomechanics and behavior but they have been rare in the early human fossil record. Here we present analyses of an unprecedented discovery of 1.5-million-year-old footprint assemblages, produced by 20+ Homo erectus individuals. These footprints provide the oldest direct evidence for modern human-like weight transfer and confirm the presence of an energy-saving longitudinally arched foot in H. erectus. Further, print size analyses suggest that these H. erectus individuals lived and moved in cooperative multi-male groups, offering direct evidence consistent with human-like social behaviors in H. erectus.

Pleistocene footprints show intensive use of lake margin habitats by Homo erectus groups

Reconstructing hominin paleoecology is critical for understanding our ancestors’ diets, social organizations and interactions with other animals. Most paleoecological models lack fine-scale resolution due to fossil hominin scarcity and the time-averaged accumulation of faunal assemblages. Here we present data from 481 fossil tracks from northwestern Kenya, including 97 hominin footprints attributed to Homo erectus. These tracks are found in multiple sedimentary layers spanning approximately 20 thousand years. Taphonomic experiments show that each of these trackways represents minutes to no more than a few days in the lives of the individuals moving across these paleolandscapes. The geology and associated vertebrate fauna place these tracks in a deltaic setting, near a lakeshore bordered by open grasslands. Hominin footprints are disproportionately abundant in this lake margin environment, relative to hominin skeletal fossil frequency in the same deposits. Accounting for preservation bias, this abundance of hominin footprints indicates repeated use of lakeshore habitats by Homo erectus. Clusters of very large prints moving in the same direction further suggest these hominins traversed this lakeshore in multi-male groups. Such reliance on near water environments, and possibly aquatic-linked foods, may have influenced hominin foraging behavior and migratory routes across and out of Africa.

Upper body contributions to power generation during rapid, overhand throwing in humans

High-speed and accurate throwing is a distinctive human behavior. Achieving fast projectile speeds during throwing requires a combination of elastic energy storage at the shoulder, as well as the transfer of kinetic energy from proximal body segments to distal segments. However, the biomechanical bases of these mechanisms are not completely understood. We used inverse dynamics analyses of kinematic data from 20 baseball players fitted with four different braces that inhibit specific motions to test a model of power generation at key joints during the throwing motion. We found that most of the work produced during throwing is generated at the hips, and much of this work (combined with smaller contributions from the pectoralis major) is used to load elastic elements in the shoulder and power the rapid acceleration of the projectile. Despite rapid angular velocities at the elbow and wrist, the restrictions confirm that much of the power generated to produce these distal movements comes from larger proximal segments, such as the shoulder and torso. Wrist hyperextension enhances performance only modestly. Together, our data also suggest that heavy reliance on elastic energy storage may help explain some common throwing injuries and can provide further insight into the evolution of the upper body and when our ancestors first developed the ability to produce high-speed throws.

The Evolutionary Origin and Population History of the Grauer Gorilla

Gorillas living in western central Africa (Gorilla gorilla) are morphologically and genetically distinguishable from those living in eastern central Africa (Gorilla beringei). Genomic analyses show eastern gorillas experienced a significant reduction in population size during the Pleistocene subsequent to geographical isolation from their western counterparts. However, how these results relate more specifically to the recent biogeographical and evolutionary history of eastern gorillas remains poorly understood. Here we show that two rare morphological traits are present in the hands and feet of both eastern gorilla subspecies at strikingly high frequencies (>60% in G. b. graueri; ∼28% in G. b. beringei) in comparison with western gorillas (<1%). The intrageneric distribution of these rare traits suggests that they became common among eastern gorillas after diverging from their western relatives during the early to middle Pleistocene. The extremely high frequencies observed among grauer gorillas-which currently occupy a geographic range more than ten times the size of that of mountain gorillas-imply that grauers originated relatively recently from a small founding population of eastern gorillas. Current paleoenvironmental, geological, and biogeographical evidence supports the hypothesis that a small group of eastern gorillas likely dispersed westward from the Virungas into present-day grauer range in the highlands just north of Lake Kivu, either immediately before or directly after the Younger Dryas interval. We propose that as the lowland forests of central Africa expanded rapidly during the early Holocene, they became connected with the expanding highland forests along the Albertine Rift and enabled the descendants of this small group to widely disperse. The descendant populations significantly expanded their geographic range and population numbers relative to the gorillas of the Virunga Mountains and the Bwindi-Impenetrable Forest, ultimately resulting in the grauer gorilla subspecies recognized today. This founder-effect hypothesis offers some optimism for modern conservation efforts to save critically endangered eastern gorillas from extinction.

Evolution of the Early Hominin Hand

Over the course of early hominin evolution, two fundamental changes in hand function occurred: the loss of a locomotor role and unparalleled intensification of manipulation, tool making, and tool use. In the context of these functional changes, early hominin hand anatomy evolved a number of derived characteristics within an otherwise primitive bauplan. Here we explore the functional significance and evolutionary history of seven major anatomical changes that make our hands distinctive, including hand proportions, thumb robusticity, thumb musculature, distal tuberosities, carpal architecture, wrist mobility, and finger curvature. This chapter highlights many areas that need more research and leads to several major conclusions: the abandonment of arboreal locomotion and rise in manipulative capabilities evolved over long periods of time and in a nonlinear fashion; early hominin taxa likely varied in their locomotor repertoires and manipulative abilities, not unlike differences in behavior seen among closely related species living today; and intensification of manipulation, rather than the origin of stone tool making, was a major driver of human hand evolution. Finally, we propose a new term, hyper-opposable, to describe the derived human ability to produce extensive contact area between the thumb and other digits, and forcefully secure and precision handle objects between the thumb and other digits through pad-to-pad contact.

Humeral torsion does not dictate shoulder position, but does influence throwing speed

A debate has emerged in the last few years over the shape and position of the shoulder in early Homo. That the shoulder joint underwent changes approximately 2 million years ago is not in dispute. A number of newly discovered and relatively complete scapulae show that the orientation of the glenohumeral joint shifted caudally from the more cranial orientation seen in the apes and earlier hominins (Walker and Leakey, 1993; Larson et al., 2007; Lordkipanidze et al., 2007; Haile-Selassie et al., 2010; Green and Alemseged, 2012; Churchill et al., 2013). However, just howmodern human-like this caudally rotated shoulder complex is remains less clear. Larson (2007, 2009) has proposed that early Homo possessed a novel, transitional shoulder morphology in which the shoulder joint faced anteriorly. We have proposed that Homo erectus had an essentially modern human-like shoulder complex with a laterally oriented glenohumeral joint (Roach et al., 2013; Roach and Richmond, 2015). Why does this debate matter? These differing reconstructions of the shoulder have important functional implications for a number of crucial behavioral shifts hypothesized to occur at or near the origins of our genus (e.g., reduced climbing behavior, intensification of tool manufacture and use, endurance running, and high speed throwing). Much of this debate has hinged on the length of the clavicle. As the only bony strut attaching the shoulder complex to the torso,