The Evolution of the Primate Foot: Anatomy, Function, and Palaeontological Evidence

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The human foot is a unique and defining characteristic of our anatomy. Most primates have grasping, prehensile feet, whereas the human foot stands out as a powerful non-grasping propulsive lever that is central to our evolution as adept bipedal walkers and runners and defines our lineage.   Very few books have compiled and evaluated key research on the primate foot and provided a perspective on what we know and what we still need to know. This book serves as an essential companion to “The Evolution of the Primate Hand” volume, also in the Developments in Primatology series. This book includes chapters written by experts in the field of morphology and mechanics of the primate foot, the role of the foot in different aspects of primate locomotion (including but not limited to human bipedalism), the “hard evidence” of primate foot evolution including fossil foot bones and fossil footprints, and the relevance of our foot’s evolutionary history to modern human foot pathology.
This volume addresses three fundamental questions: 
(1) What makes the human foot so different from that of other primates? (2) How does the anatomy, biomechanics, and ecological context of the foot and foot use differ among primates and why? (3) how did foot anatomy and function change throughout primate and human evolution, and why is this evolutionary history relevant in clinical contexts today?
This co-edited volume, which relies on the insights of leading scholars in primate foot anatomy and evolution provides for the first time a comprehensive review and scholarly discussion of the primate foot from multiple perspectives.  It is accessible to readers at different levels of inquiry (e.g., undergraduate/graduate students, postdoctoral research, other scholars outside of biological anthropology).  This volume provides an all-in‐one resource for research on the comparative and functional morphology and evolution of the primate foot.

Author(s): Angel Zeininger, Kevin G. Hatala, Roshna E. Wunderlich, Daniel Schmitt
Series: Developments in Primatology: Progress and Prospects
Publisher: Springer
Year: 2022

Language: English
Pages: 526
City: Cham

Foreword
Contents
Contributors
Abbreviations
Chapter 1: Introduction
References
Chapter 2: Clinical and Evolutionary Perspectives on the Primate Foot: A Historical and Contemporary View
2.1 Introduction
References
Section I: Primate Foot Anatomy
Chapter 3: The Primate Ankle and Hindfoot
3.1 Introduction
3.2 Distal Tibia and Fibula
3.2.1 Articular Surfaces
3.2.2 Whole-Bone Morphometrics
3.3 Talus
3.3.1 Articular Surfaces
3.3.2 Talar Neck and Head
3.3.3 Posterior Tubercles
3.3.4 Whole-Bone Morphometrics
3.3.5 Internal Microarchitecture
3.4 Calcaneus
3.4.1 Length and Projections
3.4.2 Articular Surfaces
3.4.3 Whole-Bone Morphometrics
3.4.4 Internal Microarchitecture
3.5 Future Directions
3.6 Summary
References
Chapter 4: The Primate Midfoot and Human Longitudinal Arch
4.1 Introduction
4.2 Basic Anatomy of the Midfoot
4.2.1 Bony Elements
4.2.2 Supporting Ligaments
4.2.3 Motion
4.3 The Midfoot of Nonhuman Primates
4.3.1 Pedal Grasping and the Divergent Hallux
4.3.2 Foot Inversion and Eversion
4.3.3 The Weight-Bearing Midfoot
4.4 The Human Midfoot and Longitudinal Arch
4.4.1 Bony Anatomy of the Longitudinal Arch
4.4.2 Soft-Tissue Anatomy of the Longitudinal Arch
4.4.3 The Windlass Mechanism and Arch Function
4.4.4 Human Midfoot Mobility
4.5 Future Directions
4.6 Summary
References
Chapter 5: The Primate Forefoot
5.1 Introduction
5.2 Intrinsic Proportions and Functional Axes of the Forefoot
5.2.1 Pedal Length Proportions
5.2.2 Pedal Proportions Across the Digital Rays and Functional Axes of the Foot
5.3 Metatarsal Morphology, Variation, and Function
5.3.1 Metatarsal Shaft Shape
5.3.2 Metatarsal Torsion
5.3.3 Metatarsophalangeal Joints
5.4 Phalangeal Morphology, Variation, and Function
5.4.1 Proximal and Middle Phalanges
5.4.2 Distal Phalanges
5.4.3 Claws and Nails
5.5 The Hallux - Adaptations for Grasping and Leverage
5.5.1 The Hallux in Prosimians and Extinct Euprimates
5.5.2 The Hallux in Anthropoids
5.6 Future Directions
5.7 Summary
References
Chapter 6: Myology of the Primate Foot
6.1 Introduction
6.2 Extrinsic Muscles of the Foot
6.2.1 Plantar Flexors
6.2.2 Dorsiflexors
6.2.3 Evertors
6.2.4 Contribution of Different Extrinsic Foot Muscles to Lower Leg Musculature
6.3 Intrinsic Muscles of the Foot
6.3.1 First Plantar Layer
6.3.2 Second Plantar Layer
6.3.3 Third Plantar Layer
6.3.4 Fourth Plantar Layer
6.3.5 Dorsum of the Foot
6.3.6 Interspecific Differences in Intrinsic Foot Musculature
6.4 Future Directions
6.5 Summary
References
Chapter 7: The Integument and Associated Structures of the Primate Foot
7.1 Introduction
7.1.1 The Place of the Foot in Primate Evolution
7.1.2 The Main Components of the Primate Foot
7.2 Mechanics
7.2.1 The Plantar Aponeurosis
7.3 Pedal Volar Pads
7.3.1 Heel Pads
7.3.2 Effect of Body Size on Pad Geometry and Material Properties
7.3.3 The Effect of Body Size on Pad Surface Anatomy
7.3.4 Effect of Substrate on Pad Size and Distribution
7.3.5 The Effect of Limb Loading on Pad Geometry
7.4 Pedal Nails
7.5 Future Directions
References
Section II: Biomechanical and Experimental Studies
Chapter 8: Experimental Research on Foot Use and Function During Climbing by Primates
8.1 Introduction
8.2 Behavioral Studies
8.3 Spatiotemporal Studies
8.4 Kinematic Studies
8.5 Kinetic Studies
8.6 Electromyography (EMG) Studies
8.7 Studies of Energetic Costs
8.8 Future Directions
8.9 Summary
References
Chapter 9: Foot Posture During Quadrupedal Walking in Primates
9.1 Introduction
9.2 Foot Posture Definitions
9.3 Semiplantigrady
9.4 Plantigrady
9.4.1 Heel-Strike Plantigrady
9.4.2 Midfoot/Heel Contact Plantigrady
9.5 Digitigrady
9.6 Future Directions
9.7 Summary
References
Chapter 10: Primate Foot Use During Bipedal Walking
10.1 Introduction
10.2 Anatomy and Gait
10.2.1 Foot Anatomy
10.2.2 Phases of the Bipedal Gait Cycle
10.3 Human Bipedal Walking
10.3.1 First Double-Limb Support Period (0–20% of Stance)
10.3.2 Single-Limb Support Period (20–80% of Stance)
10.3.3 Second Double-Limb Support Period (80–100% of Stance)
10.4 Non-human Primate Bipedal Walking
10.4.1 First Double-Limb Support Period (0–22/25% of Stance)
10.4.2 Single-Limb Support Period (22/25–78/75% of Stance)
10.4.3 Second Double-Limb Support Period (75/78–100% of Stance)
10.5 Evolution of the Bipedal Foot
10.6 Future Directions
10.7 Summary
References
Chapter 11: Running in Addition to Walking Helped Shape the Human Foot
11.1 Introduction
11.2 Impact
11.3 Elastic Energy Storage
11.4 Propulsion
11.5 Evolutionary Evidence
11.5.1 Stage 1: Last Common Ancestor
11.5.2 Stage 2: The Earliest Hominins
11.5.3 Stage 3: Australopiths
11.5.4 Stage 4: Homo
11.6 Future Directions
11.7 Summary
References
Section III: Palaeontological Evidence
Chapter 12: The Feet of Paleogene Primates
12.1 Introduction
12.2 Comparative Morphology of Plesiadapiform Feet
12.2.1 Pedal Features Shared by Most Plesiadapiforms
12.2.2 Carpolestes simpsoni
12.3 Comparative Morphology of Adapiform and Omomyiform Feet
12.3.1 Adapiform Pedal Morphology
12.3.1.1 Diversity of Adapiform Feet
12.3.1.2 Donrussellia, Asiadapines, and the Tarsal Morphology of the Ancestral Adapiform
12.3.2 Omomyiform Pedal Morphology
12.3.2.1 Diversity of Omomyiform Feet
12.3.2.2 Foot Proportions in Archicebus achilles and Other Paleogene Primates
12.3.2.3 Pedal Prehensility in Archicebus achilles and Other Paleogene Primates
12.4 Comparative Morphology of Other Eocene and Oligocene Haplorhine Feet
12.4.1 Eosimiid Pedal Morphology
12.4.2 Amphipithecid Pedal Morphology
12.4.3 Pedal Morphology of Eocene and Oligocene Anthropoids from the Fayum Depression, Egypt
12.5 Summary Timeline and Adaptive Scenarios
12.6 Future Directions
12.7 Summary
References
Chapter 13: Miocene Ape Feet
13.1 Introduction
13.2 Miocene Hominoids
13.2.1 Africa
13.2.1.1 Proconsul and Ekembo
13.2.1.2 Afropithecus
13.2.1.3 Rangwapithecus
13.2.1.4 Nacholapithecus
13.2.1.5 Equatorius
13.2.1.6 Turkanapithecus
13.2.1.7 Small Apes
Simiolus
Dendropithecus
Kalepithecus?/Limnopithecus?
13.2.2 Europe
13.2.2.1 Pierolapithecus
13.2.2.2 Hispanopithecus
13.2.2.3 Rudapithecus
13.2.2.4 Ouranopithecus
13.2.2.5 Ankarapithecus
13.2.2.6 Oreopithecus
13.2.3 Asia
13.2.3.1 Sivapithecus
13.2.3.2 Lufengpithecus
13.3 Future Directions
13.4 Summary
References
Chapter 14: The Early Hominin Foot
14.1 Introduction
14.2 Early Hominin Feet
14.2.1 Sahelanthropus tchadensis (ca. 7.0 Ma)
14.2.2 Orrorin tugenensis (5.6–6.2 Ma)
14.2.3 Ardipithecus kadabba (5.2–5.8 Ma)
14.2.4 Ardipithecus ramidus (4.3–4.5 Ma)
14.2.5 Australopithecus anamensis (3.8–4.2 Ma)
14.2.6 Australopithecus deyiremeda (3.3–3.5 Ma)
14.2.7 Burtele Foot (~3.4 Ma)
14.2.8 Australopithecus afarensis (3.0–3.7 Ma)
14.2.9 Australopithecus bahrelghazeli (3.6 Ma)
14.2.10 Kenyanthropus (Australopithecus) platyops (3.3–3.5 Ma)
14.2.11 Australopithecus prometheus (2.2–4.2 Ma?)
14.2.12 Australopithecus africanus (2.0–3.2 Ma)
14.2.13 Australopithecus sediba (1.98 Ma)
14.2.14 Australopithecus garhi (2.5 Ma)
14.2.15 Australopithecus (Paranthropus) aethiopicus (2.3–2.7 Ma)
14.2.16 Australopithecus (Paranthropus) boisei (1.3–2.3 Ma)
14.2.17 Australopithecus (Paranthropus) robustus (1.0–2.0 Ma)
14.3 Future Directions
14.4 Summary
References
Chapter 15: The Feet of Fossil Homo
15.1 Introduction
15.2 Obstacles in Understanding the Feet of the Earliest Potential Members of Homo
15.3 The Pedal Fossil Record of the Genus Homo
15.3.1 Unspecified Homo from Ledi-Geraru, Ethiopia (2.80–2.75 Ma)
15.3.2 Homo habilis (2.35–1.65 Ma) and Homo rudolfensis (2.0–1.95 Ma)
15.3.3 Homo erectus sensu lato (1.85–0.108 Ma)
15.3.4 Homo antecessor (1.0–0.936 Ma)
15.3.5 Sima de los Huesos hominins (780–427 ka)
15.3.6 Homo heidelbergensis (700–100 ka)
15.3.7 Homo naledi (335–236 ka)
15.3.8 Homo neanderthalensis (130–40 ka)
15.3.9 Homo floresiensis (100–60 ka)
15.3.10 Homo luzonensis (67–50 ka)
15.3.11 Homo sapiens (300 ka-Present)
15.4 Future Directions
15.5 Summary
References
Chapter 16: Pedal Morphology and Locomotor Behavior of the Subfossil Lemurs of Madagascar
16.1 Phylogenetic History and Relationships of the Subfossil Lemurs
16.2 Pedal Morphology and Locomotor Behaviors of the Subfossil Lemurs
16.2.1 Palaeopropithecidae
16.2.2 Archaeolemuridae
16.2.3 Megaladapidae
16.3 Below Branch Quadrupedal Locomotion
16.3.1 Definition and Ecological Relevance
16.3.2 Kinematics and Kinetics
16.3.3 Anatomy of a Foot Adapted for Suspensory Locomotion
16.4 Future Directions
16.5 Summary
References
Chapter 17: Recent Developments and Future Directions for the Study of Primate Feet
References
Appendices
Appendix A
Appendix A.1. Plesiadapiform Foot Fossil Materiala
Appendix A.2. Adapiform Foot Fossil Materiala
Appendix A.3. Omomyiform Foot Fossil Materiala
Appendix A.4. Early Haplorhine Foot Fossil Materiala
Appendix B. Early Hominin Foot Fossils
Appendix C. Pedal Skeletal Fossils Attributed to the Genus Homoa
References
Index