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A Nearly Complete Foot from Dikika, Ethiopia and Its Implications for the Ontogeny and Function of Australopithecus Afarensis

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A Nearly Complete Foot from Dikika, Ethiopia and Its Implications for the Ontogeny and Function of Australopithecus Afarensis Dartmouth College Dartmouth Digital Commons Dartmouth Scholarship Faculty Work 7-4-2018 A Nearly Complete Foot From Dikika, Ethiopia and its Implications for the Ontogeny and Function of Australopithecus Afarensis Jeremy M. DeSilva Dartmouth College Corey M. Gill Boston University Thomas C. Prang New York University Miriam A. Bredella Harvard Medical School Zeresenay Alemseged University of Chicago Follow this and additional works at: https://digitalcommons.dartmouth.edu/facoa Part of the Anthropology Commons Dartmouth Digital Commons Citation DeSilva, Jeremy M.; Gill, Corey M.; Prang, Thomas C.; Bredella, Miriam A.; and Alemseged, Zeresenay, "A Nearly Complete Foot From Dikika, Ethiopia and its Implications for the Ontogeny and Function of Australopithecus Afarensis" (2018). Dartmouth Scholarship. 2838. https://digitalcommons.dartmouth.edu/facoa/2838 This Article is brought to you for free and open access by the Faculty Work at Dartmouth Digital Commons. It has been accepted for inclusion in Dartmouth Scholarship by an authorized administrator of Dartmouth Digital Commons. For more information, please contact [email protected]. SCIENCE ADVANCES | RESEARCH ARTICLE ANTHROPOLOGY Copyright © 2018 The Authors, some rights reserved; A nearly complete foot from Dikika, Ethiopia and its exclusive licensee American Association implications for the ontogeny and function of for the Advancement of Science. No claim to Australopithecus afarensis original U.S. Government Jeremy M. DeSilva1*, Corey M. Gill2,3,4, Thomas C. Prang5,6, Works. Distributed 3 7 under a Creative Miriam A. Bredella , Zeresenay Alemseged * Commons Attribution NonCommercial The functional and evolutionary implications of primitive retentions in early hominin feet have been under debate License 4.0 (CC BY-NC). since the discovery of Australopithecus afarensis. Ontogeny can provide insight into adult phenotypes, but juvenile early hominin foot fossils are exceptionally rare. We analyze a nearly complete, 3.32-million-year-old juvenile foot of A. afarensis (DIK-1-1f). We show that juvenile A. afarensis individuals already had many of the bipedal features found in adult specimens. However, they also had medial cuneiform traits associated with increased hallucal mo- bility and a more gracile calcaneal tuber, which is unexpected on the basis of known adult morphologies. Selection for traits functionally associated with juvenile pedal grasping may provide a new perspective on their retention in Downloaded from the more terrestrial adult A. afarensis. INTRODUCTION RESULTS Hominin foot fossils are exceptionally rare, especially those of juve- DIK-1-1f measures only 54.6 mm in total proximodistal length niles. DIK-1-1f is a left foot found during excavation by D. Geraads from the calcaneal tuber to the broken metatarsal shafts. The ankle http://advances.sciencemag.org/ on 21 January 2002 at Dikika, Ethiopia (Fig. 1 and figs. S1 and S2) is human-like in having a moderately wedged talar trochlea with medial about a meter away from the skull of the DIK-1-1 skeleton (1). It and lateral rims of equal height resulting in a human-like talar axis was almost entirely embedded in matrix, with only the broken sur- angle (fig. S3), thereby positioning the tibia orthogonally over the foot, face of the tibial shaft and the broken diaphyses of the metatarsals as in A. afarensis adults (4). Humans and A. afarensis differ from apes observable externally. The partially prepared foot was briefly dis- in having a relatively large calcaneal tuber adaptive for dissipating cussed and illustrated in the preliminary announcement (1). Addi- loads during heel strike (5). Here, we find that apes have gracile calca- tional preparatory work by Z.A. and C. Kiarie continued through neal tubers as juveniles that become relatively more gracile in adults. 2013, exposing most of the elements, cemented together in an articu- In humans, relative calcaneal tuber cross-sectional area is high and lated foot. DIK-1-1f preserves all of the tarsals, the bases of all five does not change ontogenetically, indicating that humans are born metatarsals, and partial shafts of all but the first metatarsal. No pedal with a calcaneus adapted for the rigors of heel-striking bipedalism phalanges or metatarsal heads were recovered. There is some post- (Fig. 2). DIK-1-1f lacks the tuber enlargement seen in young humans on September 18, 2018 depositional shifting of the bones, although they remain in a near- and is more similar to juvenile great apes. Yet, adult calcanei (A.L. 333-8, anatomical position. The shafts of the metatarsals are sheared A.L. 333-37, and A.L. 333-55) are human-like in measures of relative cleanly by a break that runs obliquely from the preserved epiphysis cross-sectional area (6), suggesting that A. afarensis was characterized of the first metatarsal through slightly beyond the midshaft of the by an ontogenetic trajectory of calcaneal growth unique among living fifth metatarsal. There is minimal damage or surface erosion on the taxa. Similarly, while the distal calcaneus of DIK-1-1f is ape-like in bones. being elongated and dorsoplantarly narrow, an adult A. afarensis has DIK-1-1f is the oldest hominin foot to preserve all of the tarsals. distal calcanei that are dorsoplantarly wide (fig. S4). Although A. afarensis The Olduvai Hominid 8 (OH 8) foot, which is geologically much adults had a robust calcaneal tuber, whether they had a human-like younger, is similarly complete in preserving most of the foot elements, lateral plantar process has been questioned (7). The Dikika foot pro- save for the metatarsal heads and phalanges (2). However, there is vides important resolution on this issue. DIK-1-1f has a human-like a significant damage to the OH 8 calcaneal tuber, hindering the plantarly positioned, although weakly developed, apophyseal flange for evaluation of important morphological information. Furthermore, the lateral plantar process (5) and lacks the beaked medial plantar DIK-1-1f preserves the elements (tarsals and proximal metatarsals) process found in apes. that the single adult Australopithecus afarensis partial forefoot (A.L. The evolution of obligate bipedalism resulted in the sacrifice of 333-115) lacks (Fig. 1) (3). hallucal grasping, as evidenced by a more distally directed and medio- laterally flattened first tarsometatarsal joint in humans compared to apes (8). The DIK-1-1f medial cuneiform has a strongly convex distal 1Department of Anthropology, Dartmouth College, Hanover, NH 03755, USA. 2De- facet for the base of the first metatarsal. In adult humans, the medial 3 partment of Anthropology, Boston University, Boston, MA 02215, USA. Department cuneiform distal facet is flat or mildly convex, but the developing osteo- of Radiology, Massachusetts General Hospital and Harvard Medical School, Boston, MA 02114, USA. 4Department of Medicine, Icahn School of Medicine at Mount Sinai, chondral surface of the juvenile modern human medial cuneiform New York, NY 10029, USA. 5Center for the Study of Human Origins, Department of is convex, as found in the Dikika juvenile (fig. S5) (9). The curvature Anthropology, New York University, New York, NY 10003, USA. 6New York Consor- 7 and angular orientation of the hallucal facet of the Dikika medial tium in Evolutionary Anthropology, New York, NY, USA. Department of Organismal cuneiform are intermediate between gorillas and modern humans of Biology and Anatomy, University of Chicago, Chicago, IL 60637, USA. *Corresponding author. Email: [email protected] (J.M.D.); alemseged@ similar size and developmental ages (Fig. 3). In humans, the ossified uchicago.edu (Z.A.) surface of the developing facet for the first metatarsal rapidly flattens DeSilva et al., Sci. Adv. 2018; 4 : eaar7723 4 July 2018 1 of 8 SCIENCE ADVANCES | RESEARCH ARTICLE Downloaded from Fig. 1. A. afarensis juvenile foot DIK-1-1f. DIK-1-1f shown in (clockwise from top left) medial, dorsal, and lateral views. To right, dorsal view of DIK-1-1f and the adult A. afarensis foot A.L. 333-115. Scale bars, 1 cm. with age (9). In contrast, apes maintain a convex surface and exhibit the presence of a well-developed transverse arch. A transverse arch http://advances.sciencemag.org/ only moderate flattening of the ossified part of the joint surface as they in a bipedal foot requires the external torsion of the lateral meta- grow. An adult A. afarensis medial cuneiform (A.L. 333-28) has a tarsals (13), evident in DIK-1-1f by the orientation of the sectioned hallucal facet that is flatter than in an ape of similar size but more shaft of the fourth metatarsal (fig. S7). As in the human foot, the convex than that found in modern humans (Fig. 3) (8, 9). Since A.L. intermediate cuneiform is proximodistally elongated in DIK-1-1f 333-28 and DIK-1-1f are representatives of adult and juvenile A. afarensis, (fig. S8); the adult morphology is unknown but presumed to be the change in facet curvature during growth in A. afarensis was prob- similarly elongated (10). The DIK-1-1f lateral cuneiform is some- ably similar to the pattern in apes and distinct from that found in what foreshortened, while the adult form (A.L. 333-79) is more humans (fig. S5). These data suggest that multidirectional loading of human-like (fig. S8). DIK-1-1f and A.L. 333-79 may be sampling the hallucal tarsometatarsal joint in juveniles may have maintained a distribution of A. afarensis lateral cuneiform elongation greater the curved facet in A. afarensis throughout ontogeny. than that found in apes but slightly less than that found in modern The orientation of the medial cuneiform distal facet for the first humans. on September 18, 2018 metatarsal relative to the proximal navicular facet differs between hu- While it is generally accepted that A. afarensis had a rigid mid- mans and apes at all developmental ages (Fig. 3 and fig. S9). The Dikika foot, whether it was arched or not remained controversial (13–15).
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