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John H. Langdon
Human
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Bones, Cultures, and Genes
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John H. Langdon
Human Evolution
Bones, Cultures, and Genes
John H. Langdon
Human Biology Program
University of Indianapolis
Indianapolis, IN, USA
ISSN 2730-6135 ISSN 2730-6143 (electronic)

ISBN 978-3-031-14156-0 ISBN 978-3-031-14157-7 (eBook)
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To all my ancestors, remembered, forgotten, or undiscovered.
Preface
I find the study of paleoanthropology both fascinating and humbling. The

fascination begins in the imagination. What would it have been like to be alive
in the past? What did the men and women of prehistory experience? What
thoughts did they have and why were not they more inventive? Fascination
continues with each new discovery. Often the field has been compared to com-
pleting a jigsaw puzzle with 99% of the pieces missing. Almost monthly another
piece comes to light, and each one feels like a gift to be unwrapped. Where
does it fit? What does it tell us? It may add only a tiny detail, but every few years
the picture must be redrawn based on finds so unexpected that there is no place
for them in the old version.
It is humbling to acknowledge how little we know. Many books on this
topic try to give the impression that we have the answers – that we know how
humans evolved and we are just trying to fill in the details. I sincerely hope this
volume does not fall into that category. Good questions are far more important
in guiding future understanding than hypothetical answers. Old ideas deserve
to be challenged, but to do that effectively, we must be aware of the evidence
and reasoning that created them in the first place. At the end of most chapters
are a few “Important questions we cannot answer.” I encourage the reader to
add to that list.
It is also humbling to place humankind in a broad context. As a species,
Homo sapiens is both trivial and terrifying. We share the earth with about nine
million other species today, and these are a tiny part of one percent of those
that have ever lived. Recorded history and even the first villages and crops are
so recent that they do not make it into this story. Ours has only been around
perhaps 200,000 years since its origin in Africa, but in this time we have both
improved the planet for our own purposes and seriously damaged it. Whether
other species can avoid extinction depends on how well they can tolerate
human activity. Whether we continue, evolve, or become extinct remains for
future generations to determine.
vii
viii PREFACE
My purpose in writing this book has been to bring the account of human
evolution up to date with the latest contributions from many disciplines. It
explores how recent discoveries are challenging what we thought we knew, and
attempts to build a sense of anticipation for the next revelation. The greatest
leaps in our understanding occur when we reach beyond the painstaking work
of fossil hunting and archaeological excavations to see what other disciplines
can contribute. Anatomy, climatology, cultural anthropology, ecology, embry-
ology, ethology, geology, genetics, genomics, chemistry, neuroscience, physics,
and physiology all have their place in telling our story.
Indianapolis, IN, USA John H. Langdon

Acknowledgments
This effort could not have been possible without the support of many people.
I would like to thank my teachers and students from whom I have learned;
friends and colleagues, especially Shawn Hurst; and reviewers of this work in
progress. Katherine Langdon, Amy Langdon, and David Strait have been
invaluable in assembling illustrations. José María Bermúdez de Castro, Robert
Blumenschine, Peter Brown, Ron Clarke, Thure Cerling, Christopher Dean,
Eric Delson, Fernando Diez-Martin, Paul John Myburgh, and Christine Baile
and Samantha Guenther at the Cleveland Museum of Natural History gener-
ously gave permission for the reuse of images.
Most of all, I am indebted to my beloved wife, Terry, for her consistent
patience and encouragement.
ix
Contents
Part I Introduction 1
1 The Reflection in the Mirror 3
2 Background: Evolutionary Classification and Fossil Dating 31
3 A Primate Heritage 51
Part II The First Hominins 71
4 Miocene Hominoids 73
5 The Early Hominins: Australopiths103
6 The Ecological Context of Early Hominin Evolution145
7 Understanding Australopiths165
8 The Evolution of Bipedality191
Part III The First Humans 249
9 The Earliest Humans in Africa251
10 The First Technology277
11 Diet of Early Homo299
xi
xii Contents
12 Evolution of the Brain321
13 Leaving Africa359
Part IV The Middle Pleistocene 389
14 The Erectines of Asia391
15 Erectines of the West419
16 Behavior in the Middle Pleistocene461
Part V The Emergence of Modern People 495
17 Late Pleistocene Homo and the Emergence

of Modern Humans497
18 From the Middle Paleolithic to the Modern Mind539
19 Modern Humans Disperse From Africa581
20 Distributing Modern Peoples625
21 Life History and Reproduction651
22 Evolution Today and Tomorrow683
Appendix of Anatomical Terms697
Index709
List of Figures
Fig. 1.1 The discovery of weapons signaled the transformation from ape
into human, according to the Killer Ape hypothesis. Screen image
of an early human ancestor from the feature movie 2001: A Space
Odyssey4
Fig. 1.2 Comparative brain sizes among larger-brained mammals.
Comparisons of absolute brain size (left) alone are misleading,
because larger mammals can be expected to have larger brains.
A simple ratio of brain to body mass (right) does a better job of
capturing relative brain development among species, but there are
many other factors that will influence brain size. These will be
explored at greater length in Chaps. 5 and 16. Data primarily from
(Boddy et al., 2022). Source: author 7
Fig. 1.3 A derivative version of Zallinger’s illustration. Such pictures falsely
suggest a linear and purpose-driven view of human evolution.
Source: José-Manuel Benitos <https://commons.wikimedia.org/
wiki/File:Human_evolution_scheme.svg> CC BY-SA 3.0, via
Wikimedia Commons. Source: Wikimedia Commons 12
Fig. 1.4 Appearance and range of hominin species in time. Colors
represent different radiations of hominin species. Source: author 14
Fig. 1.5 Multiregional and Out of Africa models compared. In the
Multiregional model, the continental populations are connected
through gene flow (red). The origins of this debate lay in
competing theories of the origins of human races and the
placement of known fossils in human ancestry. Source: author 22
Fig. 1.6 Phylogenetic tree of mtDNA samples from Cann et al. (1987).
Each endpoint represents an individual, coded by continent of
origin. Because there is greater diversity and earlier divergence
among the individuals from Africa, Cann concluded the last
common ancestor lived there. Obtained with permission from
Nature Springer. Source: Nature permission 25
Fig. 2.1 A hypothetical phylogeny to illustrate concepts of classification.
See text for explanation 39
xiii
xiv List of Figures
Fig. 3.1 The primate face is restructured to allow the eyes to face
anteriorly. Shown: black howler monkey, Alouatta pigra.
Photo by the author 56
Fig. 3.2 Primates commonly assume an upright trunk posture. Shown: left,
rhesus monkey, Macaca mulatta; right, Geoffroy’s spider monkey,
Ateles geoffroyi. Photos by the author 57
Fig. 3.3 Social grooming is a very important part of life in most primates.
It strengthens individual relationships and cohesion of the larger
social group. Shown: Barbary macaque, Macaca sylvanus. Source:
Berthold Werner <https://commons.wikimedia.org/wiki/
File:Gibraltar_Barbary_Macaques_BW_2015-10-26_14-07-28.
jpg> CC BY-SA 3.0, via Wikimedia Commons 58
Fig. 3.4 Left: Old World monkeys maintain a quadrupedal posture when
walking, with a narrow ribcage and limbs of roughly equal length.
Shown: rhesus macaque, Macaca mulatta. Right: Apes commonly
assume an upright posture in the trees. A broad ribcage with
laterally facing shoulders and long arms increase their reach.
Shown: dark-handed gibbon, Sources: (left) Md. Tariq Aziz
Touhid <https://commons.wikimedia.org/wiki/File:Rhesus_
Macaque_monkey_the_look.jpg> CC BY-SA 4.0, via Wikimedia
Commons; (right) Thomas Fuhrmann <https://commons.
wikimedia.org/wiki/File:Dark-handed_or_Agile_Gibbon_
(Hylobates_agilis)_Tanjung_Puting_National_Park_-_
Indonesia_1.jpg > CC BY-SA 4.0, via Wikimedia Commons 62
Fig. 3.5 Mature male orangutan, Pongo pygmaeus. Source: Arifinal0109
<https://commons.wikimedia.org/wiki/File:ORANGUTAN_
SUMATRA_MENGAMBIL_PISANG.jpg> CC BY-SA 3.0, via
Wikimedia Commons 63
Fig. 3.6 Female mountain gorilla with infant, Gorilla gorilla. Source:
Charles J Sharp <https://commons.wikimedia.org/wiki/
File:Mountain_gorilla_(Gorilla_beringei_beringei)_female_with_
baby.jpg > CC BY-SA 4.0, via Wikimedia Commons 64
Fig. 3.7 Common chimpanzee, Pan troglodytes. Source: Tu7uh <https://
commons.wikimedia.org/wiki/File:PanTroglodytesAtZooNegara.
jpg> CC BY-SA 3.0, via Wikimedia Commons 65
Fig. 3.8 Bonobos, Pan paniscus. Source: Pierre Fidenci <https://
commons.wikimedia.org/wiki/File:Pan_paniscus11.jpg> CC
BY-SA 2,5, via Wikimedia Commons 66
Fig. 4.1 Timeline for Chap. 3. Source: author 74
Fig. 4.2 A human phylogeny from 1915 including known hominin fossils
Pithecanthropus, Neanderthals, and Eoanthropus (the Piltdown
hoax), as drawn by Sir Arthur Keith. The human lineage diverges
from the great apes in the Oligocene, which is now dated to 30
million years ago. From (Keith, 1915), public domain. Source:
public domain 74
Fig. 4.3 Partial maxilla YPM 13799 with P3-M2, type specimen of
Ramapithecus punjabicus from the Siwalik Mountains of northern
India (cast). In the 1960s this was argued to be the oldest known
hominin. It is now recognized as a female Sivapithecus sivalensis
about 12.2 Ma. Photo by Katherine Langdon 76
List of Figures xv
Fig. 4.4 Gorilla, chimpanzee, and human mandibles. The large canines on
the ape’s jaws shape the front of the mouth into a wide “U” shape
with parallel sides. The human jaw is narrow in front and the
molar rows are diverging to create a parabolic arcade. Photo by
author76
Fig. 4.5 The impact of the molecular clock on hominin classification in the
1970s. Left: a phylogeny according to then-current interpretations
of the fossil record. Human diverged early and all living lineages
are distinct and visible among the middle Miocene fossils.
O = orangutan, G = gorilla, C = chimpanzee, H = humans. Right:
A phylogeny according to the molecular clock. Humans,
chimpanzees, and gorillas are equally related and descended from
a recent common ancestor about 5 Ma. Source: author 79
Fig. 4.6 GSP 15000, the face of Sivapithecus sivalensis from the Siwalik
Mountains of northern Pakistan next to a recent orangutan
cranium. The similarities overwhelmingly linked the Asian
thick-enameled hominoids to the orang clade and not to humans.
Photo by author 82
Fig. 4.7 Occurrence of Miocene hominoid fossils and the Fayum site.
Source: author 86
Fig. 4.8 TM 266–01–060-1 Sahelanthropus tchadensis from Toros-Menalla,
Chad (model). Photo by Katherine Langdon 92
Fig. 5.1 The Taung infant, type specimen of Australopithecus africanus
from Taung quarry, South Africa, about 3.0–2.6 Ma (cast).
Source: Didier Descouens <https://commons.wikimedia.org/
wiki/File:Australopithecus_africanus_-_Cast_of_taung_child.jpg>
CC BY-SA 4.0, via Wikimedia Commons 104
Fig. 5.2 Australopithecus africanus cranium, “Mrs. Ples” Sts 5 from
Sterkfontein Cave, South Africa, about 2.5 Ma. Source: José
Braga; Didier Descouens <https://commons.wikimedia.org/
wiki/File:Mrs_Ples_Global.jpg> CC BY-SA 4.0, via Wikimedia
Commons106
Fig. 5.3 Paranthropus robustus cranium SK 48, from Swartkrans Cave,
South Africa, about 2.2–1.8 Ma. Sources: left, author; right
S. Nawrocki107
Fig. 5.4 SK 53, Paranthropus robustus mandible SK 23, from Swartkrans
Cave, South Africa. Photo by author 107
Fig. 5.5 Relative tooth size in A. africanus, P. robustus, chimpanzee, and
human. From left to right, upper first incisor through lower third
molar. Australopithecus and Paranthropus are more similar to one
another than either is to a living species. Data from (Berger et al.,
2015; Grine & Strait, 1994; Irish et al., 2016; Kaifu et al., 2015).
Source: author 109
Fig. 5.6 Known Australopith sites. The distribution in Africa is almost
entirely in the East African Rift Valley and South Africa. Source:
author111
xvi List of Figures
Fig. 5.7 The East African Rift Valley. Plio-Pleistocene fossil sites are
concentrated between the fault lines (in red) in Tanzania, Kenya,
and Ethiopia 114
Fig. 5.8 Olduvai Gorge, northern Tanzania. Source: Elitre <https://
commons.wikimedia.org/wiki/File:Olduvai_2012_05_31_2823_
(7522639084).jpg> CC BY-SA 2.0, via Wikimedia Commons 115
Fig. 5.9 OH 5 Zinjanthropus boisei OH 5 cranium from Olduvai Gorge,
Tanzania about 1.8 Ma (cast). Photo by Katherine Langdon 116
Fig. 5.10 Cranium of A. anamensis MRD-VP-1/1 from Woranso-Mille,
Ethiopia, about 3.8 Ma. Photo courtesy of the Cleveland
Museum of Natural History 118
Fig. 5.11 The canine honing mechanism in a chimpanzee (cast). The upper
canine rubs and sharpens against the lower premolar while the
lower canine hones against the upper canine. Photo by Katherine
Langdon120
Fig. 5.12 Tooth area in early hominins. The lower first premolar (LP3)
increases in size as the honing mechanism is lost. The molars
increase in area to a smaller extent. Data from (Berger
et al., 2015; Suwa et al., 2009; Ward et al., 2020) 120
Fig. 5.13 A. bahrelghazali KT 12/H1 partial mandible from Koro Toro,
Bahr-el-ghazal, Chad, about 3.6 Ma. Figure from (Brunet et al.,
1995). Obtained with permission from Nature Springer. 121
Fig. 5.14 A. deyieremeda BRT-VP-3.1from Woranso-Mille, Afar Region,
Ethiopia, about 3.5–3.3 Ma. Top row: views of maxilla; bottom
row: mandible. Figure from (Haile-Selassie et al., 2015). Obtained
with permission from Nature Springer. 122
Fig. 5.15 Maxillary tooth areas. A. garhi shows an unusual combination of
large postcanine dentition and unreduced anterior teeth. Data
from (Asfaw et al., 1999; Berger et al., 2015; Leakey & Leakey,
1978; Wood, 1991) 123
Fig. 5.16 The skeleton of Stw 573, A. prometheus, partially excavated, from
Sterkfontein Cave, South Africa, possibly 3.67 Ma. Source: Wits
University <https://commons.wikimedia.org/wiki/File:Littlle_
Foot-2.jpg> CC BY-SA 4.0, via Wikimedia Commons 123
Fig. 5.17 The cranium of MH 1, A. sediba, from Malapa Cave, South Africa,
about 1.98 Ma. Source: Brett Eloff <https://commons.
wikimedia.org/wiki/File:Australopithecus_sediba.JPG> courtesy
of Prof. Berger and Wits University GNU Free Documentation
License via Wilimedia Commons 124
Fig. 5.18 Kenyanthropus platyops, KNM-WT 40000 from West Turkana,
Kenya, about 3.5 Ma (cast). Photo by Katherine Langdon 125
Fig. 5.19 Features that distinguish the robust cranium of P. robustus (SK 48)
on the right from the gracile cranium of A. africanus (Sts 5) on
the left. A. Anterior. Photos (left) S. Nawrocki; (right) author.
B. Inferior view. Photos by author. C. Mandibles. Photos by author 126
List of Figures xvii
Fig. 5.20 Anterior and posterior tooth row lengths in australopiths (=sum of
mean MD lengths). Ardipithecus does not exhibit the postcanine
expansion of the australopiths. Paranthropus has relatively small
incisors and enlarged molars. Data sources: (Asfaw et al., 1999;
Berger et al., 2015; Brunet et al., 1995; Grine & Strait, 1994;
Haile-Selassie et al., 2015; Irish et al., 2016; Kaifu et al., 2015;
Leakey & Leakey, 1978; Schuman & Brace, 1954; Suwa et al.,
2009; Ward et al., 2020) 127
Fig. 5.21 Posterior view of KNM-ER 23000 P. boisei showing the
characteristic bell shape of the braincase. Specimen from East
Turkana, Kenya, about 1.9 Ma (cast). Photo by Katherine Langdon 128
Fig. 5.22 KNM-ER 406 P. boisei cranium from East Turkana, Kenya, about
1.7 Ma (cast). Photo by Katherine Langdon 128
Fig. 5.23 P. boisei mandible from Peninj, Tanzania, about 1.5–1.3 Ma (cast).
Photo by Katherine Langdon. 129
Fig. 5.24 KNM-WT 17000 P. aethiopicus cranium from West Turkana,
Kenya, about 2.5 Ma (cast). Photo by Katherine Langdon 130
Fig. 5.25 LH 4 mandible from Laetoli, type specimen of A. afarensis, about
Fig. 6.1 There are many forms that a savanna can present. Here are three
views of the East African grasslands taken on the same day at
Nairobi National Park, Kenya. Photos by author 148
Fig. 6.2 Comparison of C3 and C4 photosynthesis pathways. C4 pathways
require an extra step of capturing CO2 in one cell and transporting
it to another. The efficiency of this transport system results in the
incorporation of more 13C. Source: Wang et al. (2012) <https://
www.researchgate.net/figure/A-schematic-diagram-of-C3-and-
C4-photosynthesis_fig11_257881531> CC BY-SA 2.0, via
Fig. 7.1 The palate of A. afarensis AL 200 “Lucy” displaying the
megadontia characteristic of australopiths (cast). Photo by
Katherine Langdon 167
Fig. 7.2 Total postcanine (P3-M3) tooth area in australopithecines (sum
mean MD length x BL breadth). Light gray is the upper tooth
row; black is the lower tooth row. Data sources: (Asfaw et al.,
1999; Berger et al., 2015; Brunet et al., 1995; Grine & Strait,
1994; Haile-Selassie et al., 2015; Irish et al., 2016; Kaifu et al.,
2015; Leakey & Leakey, 1978; Schuman & Brace, 1954; Suwa
et al., 2009; Ward et al., 2020; Wood, 1991). Source: author 168
Fig. 7.3 KNM-ER 406 P. boisei showing the extreme development of
chewing muscles. (a) The shaded area represents the origin of
temporalis. The wide zygomatic arches support the attachment of
masseter muscle and allow the passage of temporalis between the
arch and the braincase. (b) Temporalis muscle has expanded its
attachment area (shaded area). Where the right and left muscles
meet, the sagittal crest is created to provide additional bone
surface. Photos by Katherine Langdon 170
xviii List of Figures
Fig. 7.4 Microwear pattern on Sts 56 A. africanus molar. The density pits
and scratches is indicative of the coarseness of food eaten recently.
This image also shows a high degree of anisotropy (random
direction to the scratches) consistent with a frugivorous diet.
Image by Frank L. Williams and Christopher Schmidt 171
Fig. 7.5 13
C signature of diet in early hominins from dental enamel. Ar.
ramidus has the smallest proportion of 13C. The ratio of 13C: 12C
in South African australopiths is intermediate between those for
browsers and grazers, but it is fairly stable through time. P. boisei
has a significantly higher component of C3 plants in the diet.
Data from (Lee-Thorp et al., 2012). Source: author 173
Fig. 7.6 The deep robust mandible of P. boisei (cast of SL 7A-125 from
Omo Shungura Formation deposits, Ethiopia) supported
hypermegadont molars and powerful chewing muscles. Photo by
Fig. 7.7 Estimates of EQ using different body size estimates. Data from
Table 7.2. Source: author 179
Fig. 7.8 Tooth crown formation time in days as an estimate for maturation
rate (mesiolingual cusp of the upper first molar). Enamel deposition
ceases upon eruption of the tooth. The thick enamel of the
australopiths, and especially Paranthropus, was laid down at a faster
rate than in living species and the teeth erupted at a younger age.
Data from (Smith et al., 2015). Source: author 181
Fig. 8.1 Footprints of A. afarensis at Laetoli, Tanzania, about 3.6 Ma.
Source: Fidelis T Masao and colleagues. https://commons.
wikimedia.org/wiki/File:Test-pit_L8_at_Laetoli_Site_S.jpg. CC
BY-SA 4.0, via Wikimedia Commons 192
Fig. 8.2 Selected prints from trackways “A” (above) and “G” below at
Laetoli. The G trackway is assumed to have been made by
A. afarensis and the A trackway by a different, unknown bipedal
hominin. The differences in the shape of the foot and placement
of feet within the gait pattern indicate significant diversity. Source:
(McNutt et al., 2021). https://commons.wikimedia.org/wiki/
File:Laetoli_Footprints_Site_A_(2).jpg. CC BY-SA 4.0, via
Fig. 8.3 AL 288 A reconstruction of the A. afarensis skeleton “Lucy,”
from Hadar, Afar Region, Ethiopia, about 3.2 Ma. Also see
Fig. 8.6. Photo courtesy of the Cleveland Museum of Natural
History195
Fig. 8.4 The skeleton of A. prometheus Sts 573 “Little Foot” from
Sterkfontein Cave, South Africa. Source: Paul John Myburgh,
courtesy of Paul John Myburgh and Ronald Clarke 196
Fig. 8.5 Sts 14 A. africanus partial skeleton from Sterkfontein, South
Africa, about 2.5 Ma. Photo by S. Nawrocki 197
Fig. 8.6 The two known skeletons of A. sediba, MH1 juvenile (left) and
MH 2 adult (right), from Malapa Cave, South Africa, about
1.98 Ma compared with the skeleton of Lucy (AL 288 A. afarensis).
Source: Profberger https://commons.wikimedia.org/wiki/
File:Australopithecus_sediba_and_Lucy.jpg CC BY-SA 3.0,
via Wikimedia Commons 198
List of Figures xix
Fig. 8.7 OH 8 partial foot of Paranthropus boisei from Olduvai Gorge,

Tanzania, about 1.8 Ma (cast). Photo by Katherine Langdon 198
Fig. 8.8 Curvatures of the human spinal column. The curvatures help to
balance the center of mass and provide shock absorption. Upright
posture in hominins creates lumbar lordosis and decreases the
sacral angle. Image in public domain 199
Fig. 8.9 Embryonic somites develop into vertebrae and related tissues. The
differentiating factors include retinoic acid, which diffuses from
cranial to caudal; fibroblast growth factor (FGF), which diffuses
from caudal to cranial, and the products of Hox genes produced
by cells within the somites. Hox gene activation is triggered by
local ratios of retinoic acid and FGF. The activitiy of Hox genes
that are key to differentiating regions of the spine is depicted.
Changes in the control and expression of the Hox genes can
quickly change the positions of boundaries between spinal regions.
Image of spine from SMART-Servier Medical Art, part of
Laboratoires Servier© CC BY-SA 3.0, via Wikimedia Commons 202
Fig. 8.10 The position of the foramen magnum, indicated by the yellow
dots, is related to the balance of the head in an upright posture.
Australopiths (center, Sts 5 A. africanus) show an intermediate
position of the foramen between that of the chimpanzee (left)
and human (right). Photos by Katherine Langdon. Center
photo by author 203
Fig. 8.11 The major muscles of abduction of the hip in posterior view. The
lateral flare of the iliac blade brings the origin of the muscles over
the femur for greater lever action in balance. Source of skeleton
outline: LadyofHats released into public domain via Wikimedia
Commons. Source of skeleton outline: https://commons.
wikimedia.org/wiki/File:Human_skeleton_back_no-text_no-color.
svg released into public domain via Wikimedia Commons. 206
Fig. 8.12 The lateral flare of the pelvic blade places the hip abductors
directly over the joint for more effective control. This is
exaggerated in australopiths and most other fossil hominin
specimens. (a). Modern human. (b). Reconstruction of AL 288
A. afarensis from Hadar, Ethiopia. (c). Chimpanzee. Photos by
Fig. 8.13 Chimpanzee (left) and human (right) femurs aligned at the knee.
The carrying angle of the shaft of the human femur brings the
knees and feet closer to the center of mass. Photo by Katherine
Langdon208
Fig. 8.14 The anterior to posterior length of the pelvis determines lever
arms for muscles of flexion and extension. The ilium anteriorly
creates leverage for hip flexors and the ischium posteriorly
supports the hamstrings for extension. The human pelvis is
reshaped to strengthen the extensors despite the upright
orientation. (a) Chimpanzee. (b) Human. Also see Fig. 7.20 for
comparative lengths. Photos by Katherine Langdon 209
Fig. 8.15 The longitudinal arch of the human foot (top) is elevated and
strengthened during gait to provide critical leverage. This does
not occur in the chimpanzee foot (below). Source: (Elftman &
Manter, 1935), in public domain 210
xx List of Figures
Fig. 8.16 Chimpanzee and human feet compared, scaled to the same length.
Note the human great toe is elongated, rotated, immobilized and
more robust. Source: (Morton, 1922), in public domain 211
Fig. 8.17 The transverse arch. While the heads of the metatarsals (solid
outlines) rest on the ground, the bases of the same bones (dotted
outlines) are elevated at midfoot into the transverse arch. The
transverse arch of the human foot is more pronounced and gives
further support against midfoot bending. (a) Human foot. (b)
Chimpanzee foot. Source: (Morton, 1924) in public domain 212
Fig. 8.18 The toes of the human foot go into hyperextension as the heel
rises during gait. Attached ligaments and tendons tug against the
rear of the foot, pulling joints into more stable positions and
elevating the arch. Source: (Hicks, 1954) in public domain 214
Fig. 8.19 The curvature of a long bone reflects forces applied to it. In this
example of a finger, the flexor tendon is represented by the gray
band. The tendon is held in place by a retinaculum anchored
along the length of the bone (blue). When the flexor muscle
contracts, the retinaculum redirects the forces to be perpendicular
to the bone tissue (arrows). These forces are best resisted by an
architectural arch. Shown: Proximal third phalanx of the hand of
H. naledi215
Fig. 8.20 Human and chimpanzee pelvis scales to similar interacetabular
distance. The human ilium and sacrum are greatly reduced in
height to bring the sacroiliac and hip joints closer together for
more efficient weight transfer between them. Photos by Katherine
Langdon217
Fig. 8.21 The reconstructed pelvis of AL 288 A. afarensis from Hadar,
Ethiopia (model). The reduction in height exceeds that of
humans. Photo by Katherine Langdon 218
Fig. 8.22 Chimpanzee and human proximal femora. The human femur has a
larger head and neck and a more obtuse angle between the neck
and the shaft. Photo by Katherine Langdon 219
Fig. 8.23 Lateral view of calcanei of (a). chimpanzee, (b). human, (c). AL
333-8 A. afaransis from Hadar, and (d). Omo 33-74-896 P. boisei
from Omo (casts of fossils). Photo by author 221
Fig. 8.24 The calcaneal tuberosity in posterior view of (a). chimpanzee,
(b). human, and (c). Omo 33-74-896 P. boisei from Omo.
Humans have developed a medial tubercle to broaden the base
for balance and support. This feature is variable among
australopith species. Photo by author 222
Fig. 8.25 Tyrannosaurs rex and other dinosaurs achieved a bipedal posture
by balancing the trunk with a massive tail. Keeping the trunk
horizontal did not require a redesign of the pelvis. Note the
disproportion of the limbs. Source: Greg Goebel https://
commons.wikimedia.org/wiki/File:T-Rex_skeleton_%22Big_
Mike%22_at_Museum_of_the_Rockies_White_Background.jpg
Fig. 8.26 Current evidence indicates substantial diversity of body plan
among earlier hominins represented in this summary image 237
List of Figures xxi
Fig. 9.1 The geographic distribution of early fossils of genus Homo in

Africa. Source: author 252
Fig. 9.2 OH 7 mandible, type specimen of H. habilis from Olduvai Gorge,
Tanzania, about t1.75 Ma (cast). Photo by Katherine Langdon 253
Fig. 9.3 OH 13 H. habilis from Olduvai Gorge, Tanzania, about 1.7 Ma
(cast). Photo by Katherine Langdon 254
Fig. 9.4 OH 16 H. habilis from Olduvai Gorge, Tanzania, about 1.7 Ma
Fig. 9.5 Comparison of individual tooth area (mean length x mean
breadth). H. habilis is very close to A. afarensis in area, but later
Homo is smaller. Data from (Berger et al., 2015; Brown & Walker,
1993; Kaifu et al., 2015; Leakey & Leakey, 1978; Wood, 1991;
Wood & Leakey, 2011). Source: author 255
Fig. 9.6 Features of the lower jaw and dentition that differentiate Homo
(right) from Australopithecus (left). Left, AL 400 A. afarensis;
right, OH 13 H. habilis (casts). Photos by Katherine Langdon 258
Fig. 9.7 Features of the cranium that differentiate Australopithecus from
Homo. Left, Sts 5, A. africanus; right, KNM-ER 3373,
H. ergaster. Sources: left, S. Nawrocki; right, Katherine Langdon 259
Fig. 9.8 Variation in Homo crania from East Turkana, Kenya, established
the coexistence of multiple species. From left to right, KNM-ER
1813 H. habilis, KNM-ER 1470 H. rudolfensis, KNM-ER 3733
H. ergaster (casts). (a). Anterior view. (b). Lateral view. Photos by
Fig. 9.9 Interpretations of early human diversity in East Africa. (a) A
standard model about 1980 and (b) a current best interpretation
of the African fossils. Source: author 261
Fig. 9.10 Cranium OH 24 cf. H. habilis from Olduvai Gorge, Tanzania,
about 1.8 Ma (cast). Photo by Katherine Langdon 261
Fig. 9.11 OH 65 habiline lower face from Olduvai Gorge, Tanzania, about
1.8 Ma, in anterior and inferior views. The fossils is contemporary
with specimens of H. habilis, but the great breadth of the anterior
mouth and palate closely resembles H. rudolfensis. Courtesy of
Robert Blumenschine and Ron Clarke 262
Fig. 9.12 KNM-ER 60000 mandible, similar to the H. rudolfensis tooth
row, but adding to the variation within early Homo. Figure from
(M. G. Leakey et al., 2012). Obtained with permission from
Nature Springer 263
Fig. 9.13 SK 847 H. gautengensis or a small-brained Australopithecus from
Swartkrans, South Africa, about 1.8–1.3 Ma. Photo by author 264
Fig. 9.14 Cranial capacities of early hominins. Differences are apparent
between Australopithecus and early habilines and between all the
early species and H. ergaster267
Fig. 9.15 Paleoclimate indicators for East Africa show local fluctuations
within long-term trends. Carbon isotope ratios from paleosols
in the Omo River basin (Shungura Formation) and the Lake
Turkana basin correlate with diminishing tree cover across a
spectrum of habitats. Source: (Cerling et al., 2011)
Copyright © 2011 Wiley Periodicals, Inc. 270
xxii List of Figures
Fig. 10.1 Early presence of humans in Africa and western Asia. Blue = early
Homo. Red = Mode 1 tools. Purple = both. Source: author 278
Fig. 10.2 Oldowan core and flake tools, each shown in three views. A. Early
chopper from Melka Kunture, Ethiopia. B. Flake chopper from
Saint-Clar-de-Rivière, Haute Garonne, France. Source Didier
Descouens. https://commons.wikimedia.org/wiki/File:Pierre_
taill%C3%A9e_Melka_Kunture_%C3%89thiopie.jpg and https://
commons.wikimedia.org/wiki/File:Galet_MHNT_
PRE.2009.0.200.1.jpg CC BY-SA 4.0, via Wikimedia Commons 283
Fig. 10.3 Spheroid hammerstones from Bed I, Olduvai Gorge, Tanzania.
Source (Diez-Martín et al., 2021), with permission courtesy of
Fernando Diez-Martin 284
Fig. 10.4 Chimpanzee hand (left) and human hand (right) compared. The
relatively long human thumb allows fingertip opposition and much
greater dexterity. Source: Denise Morgan. https://commons.
wikimedia.org/wiki/File:Chimp_and_human_hands.jpg via
Fig. 10.5 The hand of A. sediba from Malapa Cave, South Africa. Source:
Profberger. https://commons.wikimedia.org/wiki/File:Hand_
and_arm_Australopithecus_sediba_on_black.jpg CC BY-SA 3.0,
Fig. 12.1 The human brain has approximately three times the volume of the
chimpanzee brain. This figure only compares the cerebrums.
Source: Todd Preuss. https://commons.wikimedia.org/wiki/
File:Human_and_chimp_brain.png. CC BY-SA 2.5, via Wikimedia
Commons323
Fig. 12.2 Cranial capacities (cc) of fossil hominins through time. H. naledi
and H. florensis are on the lower right. The habilines, including
Dmanisi fossils, do not align with the trajectory of later species of
Homo324
Fig. 12.3 Cranial capacities (cc) of fossil hominins through time, including
only the fossils most likely to be on the direct lineage of
H. sapiens. This plot is more linear than Fig. 12.2 and suggests a
more continuous increase in genus Homo through time 327
Fig. 12.4 Expected and observed human visceral organ masses. The
additional investment in brain tissue is matched by the reduction
in the mass of the liver and intestine. Data from (Aiello &
Wheeler, 1995). Source: author 329
Fig. 12.5 The lobes of the cerebral cortex. The insula is hidden deep to
folds of the frontal, parietal and temporal lobes. Source: (Gray,
1912). https://commons.wikimedia.org/wiki/File:Lobes_of_
the_brain_NL.svg in public domain through Wikimedia Commons 338
Fig. 12.6 Decision-making in the prefrontal cortex. Emotional motivation
may compete with social and other concerns. Possible actions and
outcomes are valuated and compared to determine a course of
action. Source: author 340
Fig. 12.7 Reorganization of the human brain. The chimpanzee (left) and
human brain (right) are approximately to scale. The topography of
List of Figures xxiii
each has been averaged from many specimens to indicate to

remove individual variation. A. The human brain displays a relative
expansion of the prefrontal cortex, especially in its inferior regions.
The lateral orbital frontal cortex now contributes to the
operculum (folds of cortex obscuring the insula). The inferior
parietal cortex has also increased its relative area.
B. Reorganization of the orbital region of the brain. The
expansion of the lateral frontal orbital cortex has pushed insular
cortex posterior, to be covered by the frontal operculum. Source:
Shawn Hurst 341
Fig. 12.8 Approximate language areas of the left hemisphere. These areas
perform other funcitons, as well. Brain outline by Hankem.
https://commons.wikimedia.org/wiki/File:Human_Brain_
sketch_with_eyes_and_cerebrellum.svg released into public
domain via Wikimedia Commons 346
Fig. 12.9 The human vocal tract. The descent of the larynx, enlarging the
pharynx, is necessary for the full range of human speech sounds.
By separating the palate and epiglottis, it also makes it easier for
humans to choke on food and drink. Source: author 348
Fig. 13.1 Evidence of early Homo. While we started our story in Africa,
people had also entered Asia at an early date. Source: author 360
Fig. 13.2 Important sites in Chap. 9. Source: author 360
Fig. 13.3 Ruins of the castle and cathedral at Dmanisi. The oldest hominin
fossils outside of Africa were discovered during archaeological
excavation of this citadel. Source: Larry V. Dumlao. https://
commons.wikimedia.org/wiki/File:Ruins_of_Dmanisi_Castle.jpg.
Fig. 13.4 Mandible D2600 from Dmanisi, Republic of George, about
1.8 Ma (cast). The massive build and early date led to the naming
of a new species, Homo georgicus. Source: Gerbil. https://
commons.wikimedia.org/wiki/File:D2600.jpg. CC BY-SA 4.0,
Fig. 13.5 A digital reconstruction of the five skulls from Dmanisi, apparently
from a single population, from (Lordkipanidze et al., 2013). The
range sizes and morphology challenges our expectations of normal
species variation. Obtained with permission from Science368
Fig. 13.6 Excavations in the cave at Liang Bua on Flores, where
H. floresiensis was discovered. Source: Rosino. https://commons.
wikimedia.org/wiki/File:2016032812 2708_-_The_inside_of_
the_Liang_Bua_hobbit_cave_(homo_floresiensis)_
(26066080056).jpg. CC BY-SA 2.0, via Wikimedia Commons 370
Fig. 13.7 Wallace’s Line separates the Asian and Australian continental
plates. During much of the Pleistocene, Sumatra, Java and Borneo
were connected to the mainland. Land animals crossed the water
and colonized the more eastern islands, including Sulawesi, and
Flores, only rarely by chance events. Source: author 371
Fig. 13.8 Cranium of LB1 Homo floresiensis from Liang Bua, Flores,
Indonesia, about 76 Ka (model). Photo by Katherine Langdon 372
Fig. 14.1 Erectine sites in Asia referred to in this chapter. Source: author 392
xxiv List of Figures
Fig. 14.2 Possible explanations for the distribution of erectines. (a). African
and Asian populations diverged from a broad initial distribution of
hominins. This conventional view glosses over problems of
chronology and the question of where the species arose. (b).
African and Asian populations descended independently from the
initial migration of primitive Homo. The only skeletal evidence of
this migration, at Dmanisi, is too late and too primitive to be a
good ancestor. The model assumes they are distinct species and we
can explain similarities in morphology through parallel evolution
and a high level of population variation, particularly in Africa.
Gene flow maintains coherence of the species. (c). African
populations descended from H. erectus in Asia via a back
migration. This model is contradicted by the fact that H. ergaster
in Africa predates known H. erectus in Asia. (d). Asian populations
descended from H. ergaster in Africa via a second migration.
However, there is no independent evidence for a second
migration. Source: author 394
Fig. 14.3 Franz Weidenreich’s rendering of skull XII Homo erectus from
Zhoukoudian, China, about 750 BP. (a). Anterior view. (b).
Lateral view. (c). Superior view. (d). Posterior view. Images in
public domain via Wikimedia Commons. https://commons.
wikimedia.org/wiki/File:Sinanthropus_Skull_XII_norma_
frontalis.png. https://commons.wikimedia.org/wiki/
File:Sinanthropus_Skull_XII_lateralis_sinistra.png. https://
commons.wikimedia.org/wiki/File:Sinanthropus_Skull_XII_
norma_verticalis.png. https://commons.wikimedia.org/wiki/
File:Sinanthropus_Skull_XII_norma_occipitalis.png398
Fig. 14.4 Mandible of H. erectus ZKG D1 from Zhoukoudian (cast).
Photo by Katherine Langdon 400
Fig. 14.5 H. erectus cranium from Lantian Gongwangling, China, about
1.6 Ma. Source: Woo Ju-Kang. https://commons.wikimedia.org/
wiki/File:Gongwangling_norma_verticalis.png in public domain,
Fig. 14.6 H. erectus cranium from Hexian, China, about 412 Ka. Photo
courtesy of Peter Brown 402
Fig. 14.7 Cranial capacities through time for the lineage of Asian hominins.
Not the discontinuities at the start and end of H. erectus,
suggesting population replacements 404
Fig. 14.8 Dubois’ illustration of the first specimens of Pithecanthropus
erectus from Trinil, Java. In public domain 405
Fig. 14.9 Sangiran 2 Homo erectus cranium from Sangiran, Java (cast).
Photo by Katherine Langdon 406
Fig. 14.10 Sangiran 17 Homo erectus cranium from Java, about 1.2 Ma
(model). Photo by Katherine Langdon 406
Fig. 14.11 Mandibular fragment of Meganthropus palaeojavanicus from
Sangiran, Java. Von Koenigswald believed this was a new hominin
similar to robust australopiths. It is now believed to be an extinct
ape. Source: Senckenberg Research Institute and Natural History
Museum. https://commons.wikimedia.org/ wiki/
List of Figures xxv
File:Meganthropus_palaeojavanicus.jpg. CC BY-SA 4.0, via

Fig. 14.12 Skull IV late H. erectus from Ngandong, Java, about 110 Ka
Fig. 14.13 Skull IX late H. erectus from Ngandong, Java, about 110 Ka.
Source: Franz Weidenreich, in public domain via Wikipedia
Commons411
Fig. 14.14 Sambungmacan 3 late H. erectus from Sambungmacan, Java,
perhaps 100 Ka (cast). Photo by Katherine Langdon 411
Fig. 15.1 Sites referred to in this chapter. (a) African sites. (b) European
sites. Source of outline map. Source: author 420
Fig. 15.2 Mauer 1, type specimen of H. heidelbergensis from near
Heidelberg, Germany, about 600 Ka. Source: (MacCurdy, 1924)
in public domain 421
Fig. 15.3 The reconstructed skull of “Eoanthropus dawsoni,” the notorious
Piltdown forgery created from parts of a human cranium and an
orangutan mandible. Source: Wellcome Collection Gallery with
permission https://commons.wikimedia.org/wiki/File:Skull_of_
the_%22Eoanthropus_Dawsoni%22_(Piltdown_Man)_Wellcome_
M0013579.jpg. via CC BY-SA 4.0, via Wikimedia Commons 421
Fig. 15.4 KNM-ER 3883 H. ergaster from East Turkana, Kenya, about
Fig. 15.5 KNM-WT 15000 cranium of the H. ergaster skeleton from
Nariokotome, Kenya, about 1.6 Ma (cast). Photo by Katherine
Langdon425
Fig. 15.6 OH 9 H. ergaster (or H. erectus) from Olduvai Gorge, Tanzania,
about 1.7 Ma (cast). Photo by Katherine Langdon 427
Fig. 15.7 Homo heidelbergensis cranium from Kabwe, Zambia, about 300
Ka. Photos by author 429
Fig. 15.8 H. heidelbergensis cranium from Bodo, Ethiopia, about 632 Ka
Fig. 15.9 Schematic section of the route in Rising Star Cave from the closest
entrance to the Dinaledi Chamber. Early hominins repeatedly
brought the bodies of deceased members through this tortuous
passage. Source: Paul H. G. M. Dirks et al. https://commons.
wikimedia.org/wiki/File:Geological_map_and_cross-section_of_
the_Rising_Star_cave_system.jpg. CC BY-SA 4.0, via Wikimedia
Commons431
Fig. 15.10 DH 1 cranial bones of the type specimen of H. naledi from the
Dinaledi Chamber of Rising Star Cave, South Africa, about 300
Ka. This includes U.W. 101–1473 cranial bones, U.W. 101–1277
maxilla, and U.W. 101–1261 mandible. Source: Berger et al.,
2015 https://commons.wikimedia.org/wiki/File:Homo_naledi_
holotype_specimen_(DH1).jpg. CC BY-SA 4.0, via Wikimedia
Commons433
Fig. 15.11 LES 1 cranium of H. naledi from the Lesedi Chamber of Rising
Star Cave, South Africa. Source: John Hawks, Marina Elliott,
Peter Schmid et al. https://commons.wikimedia.org/wiki/
File:Homo_naledi_LES1_cranium.jpg. CC BY-SA 4.0, via
xxvi List of Figures
Fig. 15.12 Part of the initial recovery of bones from the Dinaledi Chamber
arranged into a composite skeleton. Source: Lee Roger Berger
research team. https://commons.wikimedia.org/wiki/
File:Homo_naledi_skeletal_specimens.jpg. CC BY-SA 4.0, via
Fig. 15.13 Articulated bones of the right hand of H. naledi (front and back).
Note the robusticity, particularly of the thumb and the broad tufts
at the tips of all the digits. Source: Tracy L. Kivell, Andrew
S. Deane, Matthew W. Tocheri, Caley M. Orr, Peter Schmid, John
Hawks, Lee R. Berger & Steven E. Churchill. https://commons.
wikimedia.org/wiki/File:Homo_naledi_hand.jpg. CC BY-SA 4.0,
Fig. 15.14 ATE9-1: Mandible fragment from the TE9 level of the Sima del
Elefante cave site (Sierra de Atapuerca, Spain). This fossil has been
dated between 1.1 and 1.3 million years using the cosmogenic
nucleide method. This fossil has been attributed to Homo sp. With
permission, courtesy and © José María Bermúdez de Castro 439
Fig. 15.15 ATD6-15 and ATD6-69 human fossil remains (Individual 3) from
the Gran Dolina cave site, level TD6. Various dating methods
conclude that this level is approximately 0.85 million years old
(MIS 21). These fossils have been attributed to the species Homo
antecessor. With permission, courtesy and © José María Bermúdez
de Castro 440
Fig. 15.16 Atapuerca 5 cranium early H. neanderthalensis from Sima de los
Huesos, Spain, about 450–430 Ka (cast). Photo by Katherine
Langdon441
Fig. 15.17 The cranium from Petralona, Greece, an example of a robust
specimen of H. heidelbergensis, about 305–150 Ka. With
permission, courtesy and © of Eric Delson 444
Fig. 15.18 Cranium of H. heidelbergensis from Ceprano, Italy in superior and
lateral views, about 460–430 Ka. Source: UtaUtaNapishtim.
https://commons.wikimedia.org/wiki/File:Ceprano_Argil.jpg.
Fig. 15.19 Cranium from Steinheim, Germany, an example of a gracile
specimen of H. heidelbergensis, about 150 Ka (model). Photos by
Fig. 15.20 The gracile morph of H. heidelbergensis (left) has a smaller and
more lightly built face and smaller braincase. Left: Steinheim
(cast); right: Petralona. Sources: (left) photo by Katherine
Langdon; (right) photo courtesy and © of Eric Delson 446
Fig. 15.21 The gracile morph of H. heidelbergensis (right) has a smoothly
rounded occipital region, while the robust form has a strong
occipital torus and a pronounced nuchal plane. Left: Petralona;
right: Steinheim (cast). Sources (left) photo courtesy and © of
Eric Delson; (right) photo by Katherine Langdon 446
Fig. 15.22 (Above) Arago XX1 H. heidelbergensis cranium from Tautavel,
France, about 450 Ka (cast). Source: Luna04. https://commons.
wikimedia.org/wiki/File:Homme_de_Tautavel.jpg. CC BY-SA 4.0,
via Wikimedia Commons. (Below) Arago XIII and Arago II
H. heidelbergensis mandibles from Tautavel, France (casts). Source:
List of Figures xxvii
Gerbil. https://commons.wikimedia.org/wiki/File:Tautavel_
UK_2.JPG. CC BY-SA 2.0, via Wikimedia Commons. Check for
new scan 447
Fig. 15.23 Two interpretations of phylogeny in Africa and Europe. On the
left, Middle Pleistocene fossils in both continents are place is a
single species connected by occasional migration and gene flow.
(The existence of parallel lineages such as H. naledi is ignored.).
The figure on the right places the hominins into separate lineages.
Roksandic et al., 2021 propose to merge H. heidelbergensis with
Neanderthals and to name the African lineage H. bodoensis448
Fig. 16.1 The presence of Acheulean cultures (orange) and hominin species
(blue and green). Source: author 462
Fig. 16.2 Location of sites discussed in this chapter. Source: author 462
Fig. 16.3 An adult human tapeworm of the species Taenia saginata. The
head, by which it clings to the wall of the intestine, is on the right.
Source: Center for Disease Control. https://commons.wikimedia.
org/wiki/File:Taenia_saginata_adult_5260_lores.jpg. in public
domain via Wikimedia Commons 463
Fig. 16.4 Examples of Lower Paleolithic bifacial tools. A. Flint handaxe
from England. Source: https://commons.wikimedia.org/wiki/
File:ESS-4444B5_Palaeolithic_Axe_(FindID_228769).jpg. CC
BY-SA 2.0, via Wikimedia Commons. B. A finely shaped handaxe
from Israel. Source: A finely shaped handaxe from Israel. Source:
Guyassaf. https://commons.wikimedia.org/wiki/File:Hand_
ston_1.jpg in public domain via Wikimedia Commons. C. Two
more crudely made bifaces from South Korea. Source: Ismoon.
https://commons.wikimedia.org/wiki/File:Paleolithic_
Handaxe._Seoul_National_University_Museum.jpg. CC BY-SA
4.0, via Wikimedia Commons 465
Fig. 16.5 How a modest-sized handaxe might be used. Source: https://
commons.wikimedia.org/wiki/File:Lower_Palaeolithic_Bifacial_
Hand_Axe_from_Kelstern_(FindID_386979-287331).jpg. via
Fig. 16.6 Excavated handaxes in situ at Olorgesalie, Kenya, about 700 Ka.
Source: Rossignol Benoît. https://commons.wikimedia.org/
wiki/File:Olorgesailiesite1993(1).jpg. CC BY-SA 3.0, via
Fig. 16.7 Elephant bones (Palaeoloxodon) exposed at Ambrona, Spain,
about 350 Ka. Source: PePeEfe. https://commons.wikimedia.
org/wiki/File:Palaeoloxodon_antiquus_-_in_situ_fossil_bones_-_
Ambrona.JPG. CC BY-SA 2.0, via Wikimedia Commons 479
Fig. 16.8 One of the wooden spears from Schoeningen, Germany, about
300 Ka. These are the oldest wooden weapons yet discovered.
Source: P. Pfarr NLD. https://commons.wikimedia.org/wiki/
File:Sch%C3%B6ningen_Speer_VII_im_
xxviii List of Figures
Sediment_1997_%C2%A9_P._Pfarr_NLD.jpg. CC BY-SA 3.0,

Fig. 17.1 Important sites in this chapter. Blue circles = Neanderthal
remains. Green diamonds = archaic specimens. Purple
cross = Denisovan. Red triangles = Early modern remains.
(a). Important Neanderthal sites in Europe and the near East.
(b). Africa. (c). Asia. Source: author 498
Fig. 17.2 Denisova Cave in southern Siberia, Russia. Both Neanderthals and
Denisovans were present here on and off from about 250Ka until
the arrival of H. sapiens about 45 Ka. Source: Xenochka. https://
commons.wikimedia.org/wiki/File:Таинственная пещера.jpg. CC
Fig. 17.3 Known geographic range of Neanderthal skeletal remains. Colors
represent identified subpopulations. Cultural remains are more
extensive within the dotted boundary. Source: Nienbert, Nicholas
Perrault III. https://commons.wikimedia.org/wiki/File:Range_
of_NeanderthalsAColoured.png. CC BY-SA 3.0, via Wikimedia
Commons504
Fig. 17.4 T. H. Huxley’s illustration of the Neanderthal cranium, type
specimen from the Feldhofer site, Germany (Huxley, 1863). In
public domain 506
Fig. 17.5 Various views of a Neanderthal cranium, La Ferrassie from France
(model). Photos by Katherine Langdon 506
Fig. 17.6 Neanderthal cranium from Forbes Quarry, Gibraltar. Photo by
author, courtesy of the British Museum of Natural History 507
Fig. 17.7 The profile of the classic Neanderthal cranium is evident in these
specimens from (top to bottom) Spy, Belgium; La Chapelle, France;
and La Quina, France; Guattari Cave, Italy. Source: (MacCurdy,
1924) in public domain; photo by Katherine Langdon 507
Fig. 17.8 Upper and lower Neanderthal dentition. Note the excessive
unusual wear on the incisors that opens the pulp cavities. A. La
Ferrassie from (Boule, 1923) in public domain. B. Shanidar 2
(casts). Photos by Katherine Langdon 509
Fig. 17.9 Mandible of a Neanderthal from La Ferrassie, France. Source:
(Boule, 1923) in public domain 509
Fig. 17.10 Neanderthal skeleton from La Ferrassie. Source Thilo Parg.
https://commons.wikimedia.org/wiki/File:La_Ferrassie_1_
MdlH_1_2018-10-20.jpg. CC BY-SA 4.0, via Wikimedia
Commons510
Fig. 17.11 The first attempt to reconstruct the Neanderthal skeleton by
Marcellin Boule, 1912, based primarily on the specimen from La
Chapelle-aux-Saints, France. Note the enlarged ribcage,
robusticity of the limb bones, and bowing of the femur. However,
Boule incorporated apish features to make Neanderthals less than
human, including the slouching neck, bent hip and knee, and an
abducted first toe. Source: (Boule, 1923) in public domain 511
Fig. 17.12 Cranial capacities in the European lineage. There are parallel
increases between European and African populations (compare
to Fig. 12.3), supporting a genetic linkage between the two
populations. Note the marked increase in cranial capacity between
List of Figures xxix
the early Neanderthals at Sima de los Huesos and the later one.
The magnitude of variation is similar 516
Fig. 17.13 Partial mandible from Baishiya Cave, Xiahe County, China, at least
160 Ka. Protein analysis confirms this as a Denisovan fossil. It is
also the largest identified specimen known. Source Dongiu Zhang
CC BY-SA 4.0, via Wikimedia Commons. https://commons.
wikimedia.org/w/index.php?curid=80532434518
Fig. 17.14 The cranium from Harbin, China, probably between 309 and 138
Ka. This is the best preserved cranium from a population of
lager-brained hominins that succeeded H. erectus in the later
Middle Pleistocene. Source: Ni et al., 2021 with permission 520
Fig. 17.15 Cranium from Dali, China, about 260 Ka. Photo courtesy of
Peter Brown 521
Fig. 17.16 Partial cranium from Maba, China, more than 237 Ka. Photo
courtesy of Peter Brown 521
Fig. 17.17 Cranium from Jinniushan, China, about 200 Ka. Photo courtesy
of Peter Brown 522
Fig. 17.18 Cranium from Boskop, South Africa with an unusually large
cranial capacity (cast). The age is uncertain. Photo by Katherine
Langdon525
Fig. 17.19 Cranium from Singa, Ethiopia, 145–133 Ka (cast). There is a
pathologic lesion in the right forehead. Photo by Katherine
Langdon525
Fig. 17.20 Hominin cranium from Eliye Springs, Kenya (cast). Photo by
Fig. 17.21 LH17 Ngaloba cranium from Ndutu Beds near Laetoli, Tanzania
Fig. 17.22 Digital reconstruction of the Jebel Irhoud cranium, Morocco,
about 300 Ka. Source: Philipp Gunz. https://commons.
wikimedia.org/wiki/File:Homo_sapiens_from_Jebel_Irhoud.jpg.
Fig. 17.23 Modern H. sapiens cranium from Border Cave, South Africa,
about 100 Ka (cast). Photo by author 528
Fig. 17.24 Skuhl V early modern cranium from Skhul Cave, Mount Carmel,
Israel, about 115 Ka (cast). Photo by Katherine Langdon 529
Fig. 17.25 A. Qafzeh 10 early modern juvenile cranium from Djebel Qafzeh,
Mount Carmel, Israel, about 90 Ka (cast). B. Qafzeh 11 early
modern adult cranium from Israel (cast). Photos by Katherine
Langdon529
Fig. 18.1 Middle and Late Paleolithic cultures in Africa and Europe.
Source: author 540
Fig. 18.2 Sites mentioned in this chapter. (a). Africa. (b). Europe.
Source: author 541
Fig. 18.3 Part of the ceiling of the Polychrome Chamber at Altamira.
Paintings in the cave date as far back as 36 Ka. Source: Lissy
Burges. https://commons.wikimedia.org/wiki/File:Altamira_
Nordspanien_Pr%C3%A4historische_H%C3%B6hlenmalerei_
xxx List of Figures
Deckengem%C3%A4lde_UniDiaVerlag_Nr._41501.tif. CC BY-SA
4.0, via Wikimedia Commons 542
Fig. 18.4 The Levaloissian technique permits the shaping of a tool before it
is removed from the core. This commonly results in broad but flat
flakes with sharp edges. Further retouching allows it to be shaped
for a specific purpose. (a). Mousterian point from Beuzeville,
France. (b). Mousterian scraper from La Quina, France. Source:
Didier Descouens. https://commons.wikimedia.org/wiki/
File:Pointe_levallois_Beuzeville_MHNT_PRE_.2009.0.203.1_(3).
jpg and https://commons.wikimedia.org/wiki/File:Racloir_
MHNT_PRE_2009.0.206.2_Fond_(2).jpg. CC BY-SA 4.0, via
Fig. 18.5 First appearances of signs of modern behavior. Often described as
a “revolution”, the elements of modern cultures and technology
occurred over a long period of time. Source: author 548
Fig. 18.6 Upper Paleolithic backed blade from Brassempouy, France. The
manufacture of long slender blades required more sophisticated
flaking techniques. Source: Didier Descouens. https://commons.
wikimedia.org/wiki/File:Lame_MHNT_PRE.2009.0.214.5.jpg.
Fig. 18.7 Aterian tanged point from Algeria. Such small points were
attached to the shafts of lighter weapons. Source: Michel-georges
bernard. https://commons.wikimedia.org/wiki/
File:At%C3%A9rien_(Djelfa).JPG. CC BY-SA 3.0, via Wikimedia
Commons550
Fig. 18.8 Microlithic blades from Dolni Věstonice, Czech Republic. These
would have been hafted into a handle to make a weapon or a
serrated knife. Source Thilo Parg. https://commons.wikimedia.
org/wiki/File:Dolni_Vestonice_II_Pavlov_I_microsaws.jpg. CC
Fig. 18.9 Nassarius shell beads from Blombos Cave, South Africa. Similar
beads are the earliest direct evidence personal ornamentation. Source:
Chenshilwood. https://commons.wikimedia.org/wiki/File:BBC-
shell-beads.jpg. CC BY-SA 3.0, via Wikimedia Commons 557
Fig. 18.10 Fragments of engraved and decorated ostrich eggshells from the
Middle Stone Age at Diepkloof, South Africa. The shells may have
been used as containers, but the decorations serve no functional
purpose except personal expression and identification. Source:
www.sciencemag.org/news/2010/03/engraved-eggs-suggest-
early-symbolism. CC BY-SA 3.0, via Wikimedia Commons 559
Fig. 18.11 Symbolic notation? Engraved and notched bone from the
Aurignacian of Abri Blanchard des Roches à Sergeac, France.
Numerous artifacts with similar notches or holes suggest the
possibility of counting. Source: Don Hitchcock. https://
commons.wikimedia.org/wiki/File:Blanchard_plaque.jpg. CC
Fig. 18.12 Constructed circles of stalagmites, Bruniquel Cave, France, date to
the Neanderthal period. The circles required the relocation of tons
of stone within the darkness of the cave for an unknown purpose.
Source: Luc-Henri Fage/SSAC. https://commons.wikimedia.
List of Figures xxxi
org/wiki/File:La_structure_de_la_grotte_de_Bruniquel.jpg. CC
Fig. 18.13 Blombos Cave, South Africa. This site held numerous advanced
types of artifacts from the Still Bay Culture. Source: Vincent
Mourre/Inrap. https://commons.wikimedia.org/wiki/
File:Blombos.jpg. CC BY-SA 3.0, via Wikimedia Commons 563
Fig. 18.14 Engraved block of ochre, Blombos Cave, South Africa. This rock
has one of the earliest known engravings. Source: Chris
S. Henshilwood. https://commons.wikimedia.org/wiki/
File:Blombo.jpg. CC BY-SA 4.0, via Wikimedia Commons 564
Fig. 19.1 Introgression events identified among Late Pleistocene hominin
species. Each event left an identifiable genetic sequence in the
genome of later generations. Numerous additional hybridization
events in Europe left no trace in the modern human genome.
Source: author 582
Fig. 19.2 Caves in Mount Carmel at time of excavations in 1959. Source:
Benno Rothenberg/Meitar Collection/National Library of Israel/
The Pritzker Family National Photography Collection https://
commons.wikimedia.org/wiki/File:Mount_Carmel_(997009
157796305171.jpg. CC BY-SA 4.0, via Wikimedia Commons 582
Fig. 19.3 Archaic partial cranium from Nesher Ramla, Israel, a third lineage
in the Near East, about 140–120 Ka. Source: Yossi Zaidner.
https://commons.wikimedia.org/wiki/File:Nesher_Ramla_
Homo_fossils-_a_skull_fragment_and_a_lower_jaw.jpg.
Fig. 19.4 Frontal bone from Zuttiyeh, Israel, “Galilee Man”, about
320–300 Ka. Source:‫לי‬,. https://commons.wikimedia.org/wiki/
File:IMJ_view_20130115_192522.jpg. CC BY-SA 3.0,
Fig. 19.5 Hominin fossils from Tabun Cave, Israel. (a). Tabun 2 mandible.
(b). Tabun 1 cranium, H. neanderthalensis, about 130 Ka (casts).
Photos by Katherine Langdon 585
Fig. 19.6 Amud 1 from Israel shows classic Neanderthal characteristics,
about 70–50 Ka (cast). Photos by Katherine Langdon 586
Fig. 19.7 Sites in the Near East mentioned in this chapter. Blue
circles = Neanderthal remains. Green diamonds = archaic
specimens. Red triangles = Early modern remains 588
Fig. 19.8 Coalescence time of two haplotypes compared. (a) The last
common ancestor of all modern human mtDNA lived about the
time that modern H. sapiens appears. (b) CMAHp is a deactivated
or pseudogene that produced a membrane antigen in other
species. The coalescence time is nearly three million years ago, so
at least two variants survived from the start of Homo. Another
variant arose about the beginning of H. heidelbergensis and many
additional versions were present by the start of H. sapiens. From
(Garrigan & Hammer, 2006). Obtained with permission from
Nature Springer 591
Fig. 19.9 Sites in Europe mentioned in this chapter. Source: author 592
Fig. 19.10 Cranium Předmostí 3 from the Upper Paleolithic in the Czech
Republic shows suggestions of Neanderthal traits on a modern
skull (cast). Photo by Katherine Langdon 593
xxxii List of Figures
Fig. 19.11 The cranium and mandible from Peștera cu Oase, Romania,
represent different individuals with mixed Neanderthal and
H. sapiens ancestry, 42–37 Ka (casts). Photos by Katherine
Langdon593
Fig. 19.12 Chatelperronian bone, ivory and stone tools and ornaments from
the Neanderthal layers at Grotte du Renne, France. The
manufacture of pendants and bone points was probably inspired
by encounters with Upper Paleolithic H. sapiens entering Europe.
Source: François Caron, Francesco d’Errico, Pierre Del Moral,
Frédéric Santos, and João Zilhão. https://commons.wikimedia.
org/wiki/File:Grotte_du_Renne_ivory.png. CC BY-SA 4.0, via
Fig. 19.13 Uluzzian blades from Fumane Cave, Italy, made by early
H. sapiens migrants into Europe. In addition to the blades in this
collection, the Uluzzian has many similarities to the older
Mousterian traditions. Source: Armando Falcucci. https://
commons.wikimedia.org/wiki/File:Protoaurignacian_Fumane_
Cave_retouched_bladelets_Falcucci_PlosOne_2017.png. CC
Fig. 19.14 This ivory “Venus” figurine from the Gravettian Culture at
Kostenki, Russia, shows indications of articles of clothing.
Source: Don Hitchcock. https://commons.wikimedia.org/
wiki/File:Kostenki_I_Venus.jpg. CC BY-SA 4.0, via
Fig. 19.15 Ceramic “Venus” figurine from Dolni Věstonice, Czech Republic.
The site shows early experimentation with firing clay, long before
pottery was invented. Source Petr Novák, Wikipedia. https://
commons.wikimedia.org/wiki/File:Vestonicka_venuse_edit.jpg.
Fig. 19.16 A recent cranium from Iwo Eleru, Nigeria, very likely displays
influence from an archaic introgression, about 11 Ka. Source:
Harvati K, Stringer C, Grün R, Aubert M, Allsworth-Jones P,
et al. https://commons.wikimedia.org/wiki/File:Journal.
pone.0024024.g001-1.jpg. CC BY 4.0, via Wikimedia Commons 607
Fig. 19.17 Important sites in East Asia. Green diamonds = archaic specimens.
Purple cross = Denisovan. Red triangles = Early modern remains.
Source: author 608
Fig. 19.18 Early modern cranium from Liujiang, China, 139–111 Ka.
Source: Ryan Somma from Occoquan, USA. https://commons.
wikimedia.org/wiki/File:Liujiang_cave_skull-b._Homo_Sapiens_
68,000_Years_Old.jpg. CC BY-SA 2.0, via Wikimedia Commons 609
Fig. 19.19 Longlin 1 partial cranium of variant H. sapiens from Longlin Cave,
China, 15 Ka. Source: Darren Curnoe, Xueping Ji, Paul
S. C. Taçon, and Ge Yaozheng. https://commons.wikimedia.org/
wiki/File:Red_Deer_Cave_people_skull_anterior_and_lateral.png.
Fig. 20.1 Pthirus pubis, the human pubic louse. Source: Center for Disease
Control and Prevention. https://commons.wikimedia.org/wiki/
File:Pthirus_pubis_-_crab_louse.jpg in public domain via
List of Figures xxxiii
Fig. 20.2 Reconstructed phylogeny of human lice (red) mapped onto

hominid phylogeny (blue). Splitting times of lice species and
clades are indicated. Lice transferred hosts at least twice, once
from Gorilla to hominins and a second time from early Homo
(presumably erectines) in Eurasia to later Homo. Source: author 630
Fig. 20.3 Major recent movements of populations in Africa. Today’s
Khoisan and Pygmy hunter-gatherers are remnants of older
foraging populations that once spanned the continent. The
expansions of Bantu-speaking farmers during = the last 4000 years
have encompassed nearly all of sub-Saharan Africa. Pastoralists of
East Africa occupy the drier savanna. North Africa is strongly
influenced by contacts with Eurasia in historic times. Madagascar
was settled by Indonesians crossing the Indian Ocean about
2000 years ago. Source: author 631
Fig. 20.4 Expansion of farming populations into Europe. Source: Detlef
Gronenborn, Barbara Horejs, Börner, Ober https://commons.
wikimedia.org/wiki/File:Expansion_of_farming_in_western_
Eurasia,_9600%E2%80%934000_BCE.png. CC BY-SA 4.0, via
Fig. 20.5 Expansion of Indo-European populations into Europe from the
steppes of western Asia. Source: Joshua Jonathan https://
commons.wikimedia.org/wiki/File:Indo-European_expansions.
jpg. CC BY-SA 4.0, via Wikimedia Commons 635
Fig. 20.6 Major population movement and corridors in Asia described in
the text. The initial populating of the continent by anatomically
modern humans is believed to have followed the southern
coastline about 60 Ka and subsequently spread eastward into the
islands and northward into China. A second migration by Upper
Paleolithic populations occurred across the steppes after 40 Ka
and repeatedly in historic times. Source: author 637
Fig. 20.7 Movement of populations into Australia, Polynesia, and the
Americas. Source: author 640
Fig. 21.1 Life history strategies describe the relative allocation of resources
to growth, maintenance, and reproduction. Different species may
follow different strategies. Increased investment in growth leads to
a larger organism, while increased investment in maintenance can
result in a longer lifespan. (Proportions shown are schematic, not
quantitative). Source: author 654
Fig. 21.2 Duration of life history phases of chimpanzees and human
compared. Humans experience two unique phases, childhood
and post-reproductive life. Source: author 657
Fig. 21.3 Schematic comparison of human vs. chimpanzee brain growth.
The human brain continues its velocity of increase in brain size
after birth, while the newborn chimpanzee brain is much closer
to its adult size. After (Bogin, 1999). Source: author 658
Fig. 21.4 Comparison of the reproductive lives of women in three societies
(see Table 21.3). The reproductive lifespan occupies the years
between menarche and reproductive cessation/menopause. Each
child is represented by a period of gestation (black) and lactation
xxxiv List of Figures
(green). The rest of the reproductive lifespan consists of cycles of

ovulation and menstruation. Note how cultural practices result in
very different hormonal and physiological experiences. Source:
author664
Fig. 21.5 The ovary as the final common pathway for the Reproductive
Filter. Numerous factors can affect the levels of circulating
estrogens. Each factor has an independent influence on the
probability of ovulation during a given cycle. Image source:
Sheldahl. https://commons.wikimedia.org/wiki/File:Ovary.png.
CC BY-SA 4.0, via Wikimedia Commons. Image cropped and
reversed. Source: author 668
Fig. 21.6 The rate of enamel deposition on teeth is one indication of the
rate of growth. A. Enamel is deposited more slowly and over a
longer period in humans than in African apes. B. Australopiths
have not changed the rate of deposition but have achieved a
thicker enamel coating by extending the period in which it is
created. C and D. Tooth formation in early Homo more closely
resembles the pattern of apes than of Australopiths or modern
humans. Source: (Dean et al., 2001) with permission, courtesy
of Christopher Dean 670
Fig. 22.1 The lion-man figurine from Hohlenstein-Stadel Cave, Germany.
This figurine dates to the early arrival of H. sapiens in Europe,
about 40 Ka and perhaps represents the human spirit in an
animal body 684
Fig. A1 The human and chimpanzee skulls. (a) Anterior view. (b) Lateral
view. (c) Inferior view. (d) Superior view. (e) Posterior view.
Photos by Katherine Langdon 701
Fig. A2 Human and chimpanzee mandibles. Photos by Katherine Langdon 702
Fig. A3 Human and chimpanzee dentition compared. The human
mandible has only two molars. Photo by Katherine Langdon 703
Fig. A4 The human skeleton. Public domain 704
Fig. A5 The human hand skeleton. Public domain 705
Fig. A6 The human coxal bone, lateral view. Photo by Katherine Langdon 706
Fig. A7 The human foot skeleton. Public domain 707
List of Tables
Table 1.1 Commonly recognized or recently proposed species of hominins

(see Fig. 1.1) 15
Table 1.2 Grades and species radiations in the hominin lineage 18
Table 2.1 Common tools for relative dating 42
Table 2.2 Common radiometric dating techniques 43
Table 2.3 Additional absolute dating techniques 44
Table 4.1 A discovery of diversity: List of medium and large and related
small-bodied fossil hominoids, excluding hominins (see Table
1.1), primarily from (Begun, 2007; Hartwig, 2008). It is difficult
to find consensus on names and classification of species in part
because of the speed of new discoveries and the fragmentary
representation of many taxa 83
Table 5.1 Named species of Australopithecus (Current dating from Brown
et al., 2013; Herries et al., 2013; Robinson et al., 2018; Wood
et al., 1994; Wood & Boyle, 2016) 117
Table 5.2 Summary of australopith and earlier fossil sites, arranged in
approximate chronological order 131
Table 6.1 Paleohabitats at Plio-Pleistocene fossil sites, arranged in
approximate chronological order 152
Table 7.1 Evidence for australopithecine diet 167
Table 7.2 Body mass estimates of australopithecine species 177
Table 7.3 Calculation of EQ for fossil hominins and living great apes 179
Table 8.1 Sacral angle and incidence (degrees ± SD). The sacral angle
is the relation of the body of the sacrum to the horizontal plane.
Sacral incidence is a measure that incorporates both pelvic and
sacral tilt (see Fig. 8.8). The range of values given for the fossils
indicates different reconstructions (Been et al., 2017; Tardieu
et al., 2013, 2017) 200
Table 8.2 Wedging of lumbar vertebral bodies (Latimer & Ward, 1993;
Pickering et al., 2019; Sanders, 1998; Williams et al., 2013).
Positive values for the angle between superior and inferior
surfaces indicate a greater ventral height then dorsal height;
xxxv
xxxvi List of Tables
negative values represent the opposite, contributing to a lordosis.

The wedging index is the ratio between ventral body height and
dorsal body height × 100 200
Table 8.3 Position of the foramen magnum in fossil hominins (Ahern,
2006). Distance from biporion to basion. Negative values place
the foramen anterior to biporion 204
Table 8.4 Semicircular canals, radius of curvature (Spoor et al., 1994) 205
Table 8.5 Metatarsal torsion. Positive value reflects eversion of head relative
to the base (DeSilva et al., 2019; Drapeau & Harmon, 2013) 213
Table 8.6 Limb indices of fossil hominoids and living models (Haeusler &
McHenry, 2004; Heaton et al., 2019; Johanson et al., 1987;
Johanson et al., 1982a; Lovejoy et al., 2009b; Moya-Sola &
Kohler, 1997; Napier & Napier, 1967; Schultz, 1937; S. Ward
et al., 1999). Values in parentheses have been reconstructed
from expected correlations within the skeleton 224
Table 8.7 Body mass estimated from isolated upper and lower limb bones
from Sterkfontein and Hadar (McHenry & Berger, 1998).
The numbers represent median values for unique individuals 227
Table 8.8 Summary of the appearance of indicators of bipedalism 229
Table 8.9 Selected hypotheses for the origin of bipedalism 232
Table 9.1 Early Homo and habiline fossils before 1.5 Ma 256
Table 9.2 Diagnostic Traits of Genus Homo after (Conroy & Pontzer, 2012;
Kimbel, 2009) 258
Table 9.3 Comparison of types of Early Pleistocene hominins at Koobi Fora
(Fig. 9.10)260
Table 9.4 Cranial capacities of early Homo in Africa (cc) 264
Table 10.1 Fossil and archaeological evidence for the early dispersal of
humans within and out of Africa 287
Table 12.1 Estimated encephalization quotient for hominin species, using
data from Fig. 12.2 the EQ equation of Grabowski, et al. (2016) 325
Table 12.2 Energy expenditure by great apes and humans. Data from
(Pontzer et al., 2016) 330
Table 12.3 Functional areas of the cerebrum, simplified 337
Table 13.1 Cranial capacities of Dmanisi hominins (Lordkipanidze
et al., 2013) 367
Table 14.1 Important Early and Middle Pleistocene hominin sites in China 401
Table 14.2 Cranial capacity of Middle Pleistocene hominins in Asia and Africa 403
Table 14.3 Homo erectus fossils from Java 408
Table 15.1 Erectine specimens in Africa 2.0–1.0 Ma 423
Table 15.2 Body mass estimates of early Homo species. Ruff and Walker used
a simple regression based on femoral lengths. See Chap. 7 and
discussion of Table 7.2 for other methodologies 426
Table 15.3 Encephalization quotients of early Homo species show a
substantial increase for H. ergaster. See Table 5.3 426
Table 15.4 Middle Pleistocene hominin fossils and key archaeological sites
in the Near East 437
Table 15.5 Middle Pleistocene Hominin fossils and key archaeological sites
in Europe 442
Table 15.6 Time successive populations of the Middle Pleistocene 450
List of Tables xxxvii
Table 16.1 The spread of Acheulean technologies by early appearance 467

Table 16.2 The appearance of bifacial technologies in East Asia 468
Table 16.3 Early evidence for fire by region (not a comprehensive list) 473
Table 17.1 Selected finds of Neanderthal fossils 501
Table 17.2 Parallel brain expansion in Europe and African Homo. The Sima
de los Huesos sample here represents a transition between
H. heidelbergensis and Neanderthals 508
Table 17.3 Denisovan fossil sites 517
Table 17.4 Later Pleistocene and modern H. sapiens in China. “Archaic”
refers to the conspicuous retention of primitive traits and the
failure of current species names to accommodate them 519
Table 17.5 Late Pleistocene Homo fossils in Africa 524
Table 18.1 Important MSA cultures referred to in the text 547
Table 19.1 Climatic windows of opportunity for migrations out of Africa via
the Middle East (Beyer et al., 2021) 583
Table 19.2 Neanderthal and modern sites in the Middle East 587
Table 19.3 Out-of-Africa migrations of Homo. Compare to Table 14.1 589
Table 19.4 Major Upper Paleolithic cultures in Europe 600
Table 19.5 Migration of Homo sapiens and the Upper/Late Paleolithic
cultures out of Africa, according to fossil or artifactual evidence 605
Table 19.6 The appearance of modern H. sapiens in China 609
Table 21.1 Ideal characteristics of organisms pursuing r and K strategies.
Reality is more complex (see text) 655
Table 21.2 Ratio of neonatal to adult brain size. The human brain develops
to a greater extent in a context where it can be influenced by the
environment. Data from (DeSilva, 2011) 659
Table 21.3 Reproductive histories of women from three representative
societies (May, 1978). The Colonial American is an actual figure
from vital records. Cultural infertility is a period of time when
the woman is mature but forbidden to reproduce. Unsuccessful
conceptions are assumed time lost between pregnancies. The
number of ovulations is estimated from reproductive lifespan
minus time pregnant or nursing 663
Table 21.4 Important fossil pelves providing information about obstetric shape 672
Table 22.1 Leading causes of death for 2019 (WHO, 2020) 690
Table 22.2 Examples of human gene associations with alleles undergoing
current or recent selection (Hancock & di Rienzo, 2008;
Karlsson et al., 2014; Mostafavi et al., 2017; Pennisi, 2016;
Sabeti et al., 2007) 692
PART I
Introduction
CHAPTER 1
The Reflection in the Mirror
Key Ideas in this Chapter
1. Human evolution is told as a narrative, with characters, motivations, and a

plot, because that is how the human mind is structured. Origin stories are
ubiquitous and useful in human cultures, but they are also subjective.
2. We can list numerous traits that make humans unusual; there are very few
that make us unique except in the degree of expression. There is a far smaller
distance between humans and other species than is usually assumed. We
must be wary of ascribing to our ancestors qualities of modern humans that
are not supported by objective evidence.
3. Five important traits that distinguish our species are bipedalism, endur-
ance, greater social intelligence, cooperative child-rearing, and cumula-
tive culture.
4. We have a good understanding of what the major phases of human evolu-
tion have been, but the fossil record is full of gaps and surprises. As we
gather more evidence and ask new questions, our interpretation of the
details of human evolution must be considered tentative and fluid.
5. A long-standing debate over the origin of modern populations has been
largely resolved through the use of genomic information. All modern peo-
ple are descended from a relatively recent African population, although
other recent species such as Neanderthals have made small contributions to
the human genome.
6. Theories and debates for the past, whether discredited or resolves, continue
to shape our approach to the field.
© The Author(s), under exclusive license to Springer Nature 3

Switzerland AG 2022
J. H. Langdon, Human Evolution, Springer Texts in Social Sciences,
https://doi.org/10.1007/978-3-031-14157-7_1
4 J. H. LANGDON
Origin Stories
A band of hairy primates forages in a desert. Amid prehistoric-looking animals,
they scrabble for seeds and edible plants amid sparse vegetation. They seek shel-
ter at night under overhanging rocks to escape the cold and roaming predators.
When a larger band drives them from their waterhole, they retreat, passive and
helpless in a hostile world. One morning they awaken in the presence of a strange
device planted by aliens that magically gives them the gift of insight. An adult
male discovers that an animal bone may be used as a club, and with that weapon
his tribe can obtain meat from the game animals beside them. Now stronger and
better fed, the band is ready to drive away their rivals from the water hole. The
other band screams and jumps around and makes the threats and bluffs typical of
the species; but our protagonist rises on two feet and strikes his enemy with the
club, killing him. [Opening act from the motion picture 2001: A Space Odyssey
(1968), based on the science fiction novel by Arthur C. Clarke.]
Although it is science fiction, the account above is an origin story, describing
how our species acquired intellect, bipedality, and a murderous instinct. Every
society, every culture, and every family has an origin story. It may be an oral tra-
dition, a religious statement, or a history book, and frequently is a blend of all
three. An origin story is not just a genealogy. Its purpose is less to tell a people
where they came from than that to tell them who they are. It is about values and
traditions and shared experiences. It builds an identity by telling them who they
are not. In turn, origin stories may be rewritten when identities change (Fig. 1.1).
A century ago, the origin story of the United States, as written in the text-
books and taught to children, included a cast of heroes such as Christopher
Columbus, George Washington, Daniel Boone, and Thomas Edison. It
recounted pivotal events such as the American Revolution and the Civil War
and critical documents like the Mayflower Compact and the Gettysburg
Address. Americans celebrated and reenacted this identity on Independence
Day and Thanksgiving. This “America” was white, English, and largely male.
Fig. 1.1 The discovery of weapons signaled the transformation from ape into human,
according to the Killer Ape hypothesis. Screen image of an early human ancestor from
the feature movie 2001: A Space Odyssey
1 THE REFLECTION IN THE MIRROR 5
Now our nation is redefining its identity by rewriting its origin story with new
heroes and pivotal events—Martin Luther King, Cesar Chavez, Harriet
Tubman, slavery, the Trail of Tears, and Juneteenth.
The narrative of human evolution is an origin story. It is a scientific recon-
struction of our past, but inevitably it is about our identity as human beings.
What is a human? What is human nature? How and why do we differ from
other species? Meaningful answers to these questions stray into the subjective.
The depiction of our origins in the movie 2001 built upon orthodox scientific
thinking of the 1950s. The “Killer Ape” hypothesis advanced by anthropolo-
gist Raymond Dart and popularized by Robert Ardrey postulated that we are
by nature violent and bloodthirsty. We developed those instincts to survive on
the savanna and to compete with neighboring tribes, and they continue to
surface from within us when we find ourselves under threat. This thesis perco-
lated into popular literature through such novels as William Golding’s Lord of
the Flies (1954). The movie sequence, more in line with Darwin’s optimism,
ends with the image of the crude bone weapon transformed into a spaceship—
a symbol of peaceful progress. When Dart and Ardrey looked in the mirror,
they saw the perpetrators of two world wars, the Holocaust, and imminent
nuclear annihilation. Too often we have found that anthropology has not dis-
covered a deep understanding but reflected a single dimension that we wanted
or expected to see.
Nineteenth-century anthropologists, who had a very limited knowledge of
the apes, sought evidence of an evolutionary scale among living human popula-
tions. Their concepts of what represented a more highly evolved state reflected
what they valued—the technology and institutions of Western civilization.
Accordingly, non-European peoples were considered less evolved, biologically
as well as culturally. These values and prejudices existed well before Darwin, of
course, and scientific hypotheses of evolution served to justify the institutional-
ized discrimination and exploitation that was already practiced. Today, anthro-
pologists distance themselves from such ideas.
Scientific racism is an egregious example of how subjectivity creeps into sci-
ence, but bias can be more subtle. Consider, for example, the problem of defin-
ing genus Homo and distinguishing early fossil humans from the bones of
ancestors and cousins. Should we define humans anatomically, perhaps by fea-
tures of the dentition or brain size? These traits might be easier to recognize
among fragmented bones, but they do not capture the essence of being human,
at least not for non-anthropologists. Perhaps behavior is a more important
indicator, one that we can infer from tools and other archaeological remains.
Often, however, fossils are found without tools and vice versa. Anthropologists
have sometimes defined humans by drawing arbitrary boundaries that signify
an important break with the past. For Louis Leakey, a brain size of 600 cc or
higher defined Homo. Such decisions are consciously arbitrary.
Another source of subjectivity is what is known as “discoverer’s bias,” which
can be summed up in the sentiment, “My fossil is very important for under-
standing human evolution” or even “My fossil is more important than your
6 J. H. LANGDON
fossil.” Paleontologists need to justify their efforts and find recognition among
their peers, and some egos desire more recognition than others. It is not sur-
prising that a fossil that was highly touted as an ancestor at a critical transition
has since been relegated to dead-end lineage. These exaggerated claims may
appear to be benign; but they are intended to influence media attention around
a new discovery and they may divert funding from other projects.
Box 1.1 Some Essential Vocabulary

Human—A member of the genus Homo, which includes many species,
including our own, Homo sapiens.
Hominin (Subfamily Homininae)—A member of the group that
includes humans and our extinct relatives. A precise definition eludes us,
but bipedal posture and dental features provide a working approach.
(Before a change in classification in the 1980s, these were called
hominids).
Hominoid (Superfamily Hominoidea)—A member of the division of
primates that includes humans and apes.
The following terms should be considered informal common names for
grades of hominins. While they persist in standard use, they are now
divorced from formal taxonomic meaning and are retained because a
more precise sorting of species remains vague and controversial.
Australopiths—Early African hominins of the genera Australopithecus,
Kenyanthropus, and Paranthropus.
Habilines—Small-brained primitive humans belonging to Homo habi-
lis and a few more species that overlap in time, space, and morphology.
Erectines—Members of a more derived grade of genus Homo from the
Middle Pleistocene with medium-sized brains and more complex behav-
iors than habilines. Different authors recognize from one to four or more
species of erectines.
The following abbreviations are used for dates throughout the book:
Ma = millions of years ago.
Ka = thousands of years ago.
BP = years before the present.
CE = Common Era, replacing A.D. in historic dates.
Constructing Human Identity

Let us place our origin story in the broadest context. What is special about
humans? Of these traits, which are more important? What defines us and which
need to be explained? Let us begin by considering some of the more commonly
cited traits.
Brain Size Humans have large brains, our favorite organ in such discussions.
They are larger than those of any other living primate by far, but there are other
6900
0.027
6000
Brain volume (cc)
Brain:body ratio
0.02
4000 4300
0.012
0.01 0.010
2000
1346 0.006
797
349
0 0.00 0.001
chim h e d blu
pan uman lephan olphin e wha chim h e d blu
pan uman lephan olphin e wha
zee t le zee t le
Species Species
Fig. 1.2 Comparative brain sizes among larger-brained mammals. Comparisons of

absolute brain size (left) alone are misleading, because larger mammals can be expected
to have larger brains. A simple ratio of brain to body mass (right) does a better job of
capturing relative brain development among species, but there are many other factors
that will influence brain size. These will be explored at greater length in Chaps. 5 and
16. Data primarily from (Boddy et al., 2022). Source: author
mammals with brains even larger in absolute size, including elephants and most
whales (Fig. 1.2). Human brains are relatively larger than those of any other
living animal when scaled for body size, but they have the same internal anat-
omy as other mammals. No one would argue that the brains of whales or chim-
panzees are just like ours, but it is difficult to explain the human species from
the physical brain alone.
Bipedalism The fact that humans habitually stand and walk on two legs has
ramifications throughout the body, especially in the musculoskeletal system.
One of the more important consequences is the freeing of the upper limbs for
cultural activities. Darwin proposed that a positive feedback loop linked bipedal
posture, freeing of the hands, development of technology, and brain expan-
sion. We now know that our upright posture evolved before these other traits
and must have been selected for other reasons. Humans are one species among
many groups of animals that became fully or occasionally bipedal in different
contexts and in different ways. These include dinosaurs and birds, some lizards,
kangaroos, and kangaroo rats.
Loss of Body Hair Humans are “the naked ape,” such that hairlessness is one of
our most visually conspicuous traits. Of course, we have not completely lost
our body hairs—they are simply shorter, finer, and fewer in number. They con-
tinue to play an important role in sensation. Chimpanzees and orangutans have
relatively sparse hair by comparison with other primates, but not to the extent
seen in humans. Severe reduction or loss of body hair is observed in several
other mammalian groups, including all cetaceans (whales and relatives), hip-
popotamus, elephant, rhinoceros, many suids (pigs), and the naked mole rat.
8 J. H. LANGDON
Hands Human technology and material culture would not have been possible
without dexterous hands and opposable thumbs. However, opposable
thumbs—and first toes—are primitive primate traits shared among nearly all
monkeys and apes. Human hands are especially capable because of the propor-
tions of our digits and our neuromuscular control.
Meat-eating Meat-eating was considered to be a point of contrast with the
great apes until field studies revealed that chimpanzees regularly cooperate to
kill and eat a range of mammals, especially monkeys. The human proclivity for
hunting and carnivory and also for fishing has ramifications for our material
culture and provides windfalls of calories, protein, and other nutrients.
However, carnivorous behavior is not unique to humans and is far less universal
among us than the current Western culture would indicate.
Intellect As much trouble as we have defining intelligence, it is clear that
humans have mental abilities far beyond those of any other land animal and
probably sea animal as well. But whatever behavior we choose to measure intel-
ligence—tool use, culture, problem-solving, abstraction, social interaction, use
of language—at least the rudiments are observable in other species, and espe-
cially among the great apes.
Morality Darwin singled out a moral sense as a defining feature of humanity.
We may understand it as a “knowledge of good and evil,” or a recognition that
we should adhere to an abstract set of learned rules. Are elements of morality
found in other species? Individuals of many other species have demonstrated a
theory of mind—a recognition that other individuals have their own perspec-
tive in the world and are not merely projections of one’s own mind. With that
comes empathy, the ability to recognize and identify with the emotional state
of another individual. Add to these the ability to learn and anticipate the likely
consequence of one’s behavior, and the potential for moral behavior is present.
We can observe these behaviors, but we are unable to assess the thought pro-
cesses and motivations that link them.
Examination of the traits described above tells us that humans are not quali-
tatively different from other animals, but have merely developed certain ana-
tomical traits, behaviors, and abilities in different ways. The more comprehensive
list of traits in Box 1.2 identifies numerous ways in which humans diverge from
the typical mammalian condition, but very few of these traits are absent among
our close relatives. Their expression is a matter of degree. Our truly unique
attributes, e.g., mastoid process, chin, and arch of the foot, make a disappoint-
ing definition of human. The real lesson from these lists is not how humans
stand out, but how little we stand out. In all aspects—genes, anatomy, and
behavior—we are impressed by the continuity between ourselves and other
animals. The study of the fossil record blurs that line further. Nonetheless,
these are the features whose emergence we seek to track and explain.
Box 1.2 How Are Humans Different?

Most traits that appear to make humans different from other mammals
are unusual rather than unique. Here is a selected list of distinctive ana-
tomical characters possessed by humans. Unique traits are indicated with
an asterisk (*).
• Relatively enlarged brain.
• Relatively enlarged prefrontal cortex.
• Mastoid process*.
• Converging orbits—shared with all primates.
• Binocular vision—shared with all primates.
• Color vision—shared with most primates.
• Reduced olfactory sense and apparatus.
• Relatively flattened face and short jaws.
• Crowded squarish molars.
• Reduced canines.
• Chin*.
• Enlarged pharynx.
• Relatively complex articulation of sound.
• Forward position of foramen magnum.
• Relatively enlarged semicircular canals.
• Broad shallow thorax—shared with apes.
• Laterally facing shoulder joint—shared with apes.
• Greater humeral torsion.
• Relatively longer, dexterous thumb.
• Well-developed flexor pollicis longus muscle.
• Thumb opposition—shared with most primates.
• Fingernails and prints—shared with all primates.
• Structural lumbar lordosis*.
• Relatively inflated weight-bearing bones for shock absorption.
• Relatively short, broad ilium.
• Relatively curved iliac blade.
• Iliac pillar*.
• Relatively short space between acetabulum and sacroiliac joint.
• Relatively large gluteus maximus.
• Relatively short ischium.
• Relative enlargement of birth canal.
• Greater carrying angle of femoral shaft.
• Enlarged femoral head.
• Higher angle of femoral neck.
• Enlarged femoral condyles.
• Enlarged calcaneal tuberosity.
(continued)
10 J. H. LANGDON
Box 1.2 (continued)
• Lateral tubercle on the calcaneal tuberosity*.

• Reduced peroneal tubercle.
• Relatively enlarged hallux (first toe).
• Adducted hallux.
• Hyperextension of toes at the metatarsophalangeal joints*.
• Longitudinal arch of the foot*.
• Relative hair reduction.
• Relatively greater expansion of subcutaneous fat.
• Wider distribution of eccrine glands.
• Local thermal sensitivity of eccrine sweat glands*.
• Increased skin vascularity.
• Male facial hair.
• Prominent breasts.
• Relatively reduced gut size.
Behavioral traits in the following list also distinguish humans more by

degree of development and expression than by innovation. Unique traits
are indicated with an asterisk (*).
• Pair-bonding of sexual partners beyond a mating cycle.

• Sexual intercourse beyond that for conception.
• Sexual division of the economy*.
• Cooperative breeding.
• Resource sharing beyond kin.
• Intelligence, by any definition.
• Moral sense.
• Language.
• Learning language by immersion*.
• Culture.
• Tools and material culture.
• Art.
• Complex society involving individuals with defined roles.
• Omnivorous diet.
• Food production.
• Intentional alteration of the environment.
Genomes Humans are also distinct genetically. Comparisons of genomes are

not straightforward and estimates of similarity between the human and chim-
panzee genomes vary because different methods have been used. Chromosomes
contain long strands of DNA and we are familiar with the idea of mutations
changing out one nucleotide for another. However, many other kinds of
genetic change are possible, with varying effects on the actions of the genes.
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words and sentences which are its material embody an internally
coherent representation of human character and purpose. Apart from
this inner unity of meaning, mere uniformity of grammatical and
metrical construction would not of themselves constitute a work of
art. It will be one object of our later discussions to show that what is
thus obviously true of an æsthetic whole is universally true of every
genuine system or totality.
§ 7. The data or material of reality, then, are facts of experience,

and nothing but facts of experience.[19] And experience, we have
said, means for our purposes immediate feeling or apprehension.
What immediacy means, as we have already seen, we cannot
further explain in psychological terms, except by saying that it is
what distinguishes an actual mental state from the mere thought of
that state. The reason why, in Psychology, we have to be content
with such an account is manifest. To characterise immediate feeling
further, we should have to identify the qualities by which it is
universally marked off from what is not immediate. We should, in
fact, have to describe it in general terms, and before we can do this
we must cease to feel or apprehend directly, and go on to reflect
upon and analyse the contents of our apprehension. What our
psychological description depicts is never the experience as it
actually was while we were having it, but the experience as it
appears from the point of view of subsequent reflection, interpreted
in the light of all sorts of conscious or unconscious hypotheses about
its conditions and its constituents. Thus our psychological
descriptions depend for their very possibility upon the recognition of
distinctions which are not present, as such, in the experience itself
as directly presented to us but created by later reflection about it
From the point of view of Metaphysics, however, it is possible to
specify one universal characteristic of immediate feeling, which is of
the utmost importance for our theories of reality and of knowledge.
When we reflect upon any psychical fact whatever, we may
distinguish within it two very different aspects. There is, in the first
place, the fact that it does happen, that it is a genuine psychical
occurrence,—the existence or that, as we may call it, of the piece of
psychical fact in question; and there is also the peculiar character or
quality which gives this mental occurrence its unique nature as
distinguished from any other which might conceivably have been
presented in its stead,—the content or what of the psychical fact.
Thus a simple colour-sensation, say that of green, has its that,—it is
actually present, and is thus distinguished from a merely
remembered or anticipated sensation; it has also its what,—the
peculiar quality by which it is distinguished, for example, from a
sensation of blue. So again with an imagined sensation; it is actually
imagined, the imagining of it is an actual occurrence with its
particular place in the course of the occurrences which together
make up my mental life; and again, it is the imagination of some
content with qualities of its own by which it is distinguished from any
other content.
The most striking illustration of the presence of these
distinguishable aspects in all psychical occurrences is, of course,
afforded by the case of error or illusion, the essence of which is the
false apprehension of the what. Thus, when an ignorant villager sees
a ghost, or a hypochondriac is tormented by “imaginary” symptoms
of disease, the ghost or the malady is not simply non-existent;
something is actually seen or felt, but the error consists in a mistake
as to the nature of what is seen or felt. Now, the peculiarity by which
direct and immediate apprehension is distinguished, for the
metaphysician, from subsequent reflection about the contents of
apprehension, is that in immediate apprehension itself we are not
conscious of the distinction between these two aspects of psychical
fact. The immediately experienced is always a this-what or process-
content[20] in which the distinction of the this from the what does not
enter into consciousness. In any act of reflection, on the other hand,
the what is explicitly distinguished from the that, and then ascribed to
it as something which can be truly said about it. The judgment or
proposition, which is the characteristic form in which the result of
reflection finds its expression, consists, in its most rudimentary
shape, of the embodiment of this distinction in the separation of
predicate from subject, and the subsequent affirmation of the first
about the second. The work of thought or knowledge in making our
world more intelligible to us essentially consists in the progressive
analysis of a content or what, considered in abstraction from the this
to which it belongs. The this may, as in the singular judgment or the
particular judgment of perception, actually appear in our propositions
as the subject to which the what is explicitly ascribed; or again, as in
the true universals of science, both the predicate and the ostensible
subject of the proposition may belong to the content analysed, and
the this, or directly apprehended reality of which the content forms
an attribute, may not appear in the proposition at all. This is why the
true universal judgment has long been seen by logicians to be
essentially hypothetical, and why, again, thought or knowledge
always appears to the common-sense man to be dealing with
realities which have previously been given independently of the
“work of the mind.” He is only wrong in this view because he forgets
that what is given in this way is merely the that or existence of the
world of real being, not its what or content in its true character as
ultimately ascertained by scientific thought.[21]
§ 8. The fundamental characteristic of experience, then, for the
metaphysician, is its immediacy: the fact that in experience as such
the existence and the content of what is apprehended are not
mentally separated. This immediacy may be due, as in the case of
mere uninterpreted sensation, to the absence of reflective analysis of
the given into its constituent aspects or elements. But it may also be
due, as we shall have opportunities to see more fully later on, to the
fusion at a higher level into a single directly apprehended whole of
results originally won by the process of abstraction and reflection.
There is an immediacy of experience which is below mediate
reflective knowledge but there is also a higher immediacy which is
above it. To explain and justify this statement will be the work of
subsequent chapters; for the present we may be content to illustrate
it by a simple example. A work of art with an intricate internal
structure, such, for instance, as a musical composition or a chess
problem, as directly presented to the artistically uncultivated man, is
little more than a mere succession of immediately given data in
which the aspects of existence and content are as yet hardly
separated; it has no significance or meaning, but merely is. As
education in the perception of artistic form proceeds, the separation
becomes at first more and more prominent. Each subordinate part of
the structure now acquires a meaning or significance in virtue of its
place in the whole, and this meaning is at first something over and
above the directly presented character of the part, something which
has to be grasped by reflective analysis and comparison of part with
part. The individual part has now, through analysis of its content,
come to mean or stand for something outside itself, namely, its
relation to all the other parts. But with the completion of our æsthetic
education the immediacy thus destroyed is once more restored. To
the fully trained perception the meaning of the composition or the
problem, its structure as an artistic whole, is no longer something
which has to be pieced together and inferred by reflective
comparison: it is now directly apprehended as a structural unity. The
composition has a meaning, and thus the results of the intermediate
stage of reflection and comparison are not lost, but taken up into the
completed experience. But the meaning is no longer external to the
existence of the composition; it is what it means, and it means what
it is.[22] We may subsequently see that what is thus strikingly
illustrated by the case of artistic perception holds good, to a greater
or less degree, of all advance in the understanding of reality. It is
perhaps the fundamental philosophical defect of what is popularly
called Mysticism that it ignores this difference between a higher and
a lower immediacy, and thus attempts to restore the direct contact
with felt reality which scientific reflection inevitably loosens by simply
undoing the work of analytic thought and reverting to the standpoint
of mere uninterpreted feeling.[23]
§ 9. We may perhaps specify one further characteristic which
seems, at least, to belong to every datum of immediate experience.
Every experience seems to be implicitly complex, that is, its aspect
of content appears never to be absolutely simple, but always to
contain a plurality of aspects, which, as directly felt, are not distinct,
but are at the same time distinguishable as soon as we begin by
reflection to describe and analyse it. From the nature of the case this
complexity cannot be directly ascertained by inspection, for the
inspection itself presupposes that we are dealing with the experience
not as immediately felt, but as already sufficiently analysed and
reflected upon to be described in general terms. Indirectly, however,
our result seems to be established by the consideration that, as soon
as we reflect upon the given at all, we find these distinguishable
aspects within its content, and that, unless they were there implicitly
from the first, it is hard to see how the mere process of reflection
could have given birth to them. Thus, for instance, in even the most
rudimentary experience there would appear to be something
answering to the distinction between the presentational quality of a
sensation and its accompanying tone of pleasure or pain. It is
difficult, again, not to think that in any sentient experience there must
be some difference between elements which correspond to more or
less stable conditions of the sentient organism itself (“organic
sensation”) and those which correspond to relatively novel and
infrequent features of the environment. Some philosophers would
indeed be prepared to go further, and to maintain that a more or less
explicit consciousness of distinction between self and not-self, or
again between subject and object, is logically involved in the very
possibility of an experience. The question, as a psychological one,
need not be raised here; it must, however, be carefully remarked that
whatever view we may adopt as to the number and character of the
aspects which analysis reveals within the contents of the simplest
experience, those aspects, as directly apprehended, originally
constitute an unanalysed whole. Our various subsequent analyses
all presuppose theories as to the ultimate what of experience which it
is the business of Metaphysics to test.
§ 10. Our foregoing discussion of the metaphysical criterion will
suggest a fairly definite ideal of what a completely adequate
apprehension of the whole of reality would be. A completely
adequate apprehension of reality would be one which contained all
reality and nothing but reality, and thus involved no element
whatever of deceptive appearance. As such it would, in the first
place, be all-embracing; it would include in itself every datum of
direct experience, and, since nothing but data of experience, or, as
we have also called them, matters of psychical fact, are the materials
of reality, it would contain nothing else. In the second place, it would
contain all its data without contradiction or discrepancy as part of a
single system with a harmonious internal structure of its own. For
wherever there is discrepancy, as we have already seen, there is
imperfect and therefore partially false appearance. And, in the third
place, such an all-embracing harmonious apprehension of the whole
data of experience would clearly transcend that separation of
existence from content which is temporarily effected by our own
efforts to restate our experience in a consistent form. It would,
because complete in itself, involve at a higher level that immediacy
which, at a lower level, we know as characteristic of feeling. It would
thus experience the whole of real existence directly as a system with
internal consistency and structure, but without any reference to
anything beyond itself. As we said of the artistic whole, so we may
say of the whole of existence as it might be apprehended by a
completed insight, it would be what it meant, and mean what it was.
To such an ideally complete experience of reality as a single system,
by way of marking its exclusively experiential nature, we may give
the name, introduced into Philosophy by Avenarius, of a “pure”
experience, that is, an experience which is in all its parts experience
and nothing else. Of course, in adopting the name, we are not
necessarily identifying ourselves with the further views of Avenarius
as to what in particular the structure of such an experience would be.
Our own human experience clearly falls far short of such an ideal,
and that for two reasons. To begin with, our experience is incomplete
in respect of its data: there is much in reality which never directly
enters into the structure of our experience at all. Of much of what
falls within the scope of our knowledge we can only say, in a general
way, how it would appear to ourselves supposing certain conditions
of its perceptibility to be realised, and even these conditions are
usually only most imperfectly known. What the actual matters of
psychical fact corresponding to these conditions and to the
appearance which they would determine for us are, we are totally
unable to say. Again, there may well be much in the real world which
never, even in this indirect way, enters into the structure of human
knowledge at all. Hence our human experience and the intellectual
constructions by which we seek to interpret it have always the
character of being piecemeal and fragmentary. Perfect apprehension
of systematic reality as a whole would be able to deduce from any
one fact in the universe the nature of every other fact. Or rather, as
the whole would be presented at once in its entirety, there would be
no need for the deduction; every fact would be directly seen as
linked with every other by the directly intuited nature of the system to
which all facts belong. But in our imperfect human apprehension of
the world our facts appear to be largely given us in isolation and
independence of one another as bare “casual conjunctions” or
“collocations,” and the hypotheses by which we seek to weld them
into a system, however largely determined by the character of our
data, never quite get rid of an element of arbitrary “free” construction.
They are never fully necessitated as to their entirety by the nature of
the facts they serve to connect. Hence we can never be certain that
our hypothetical constructions themselves are true in the sense of
consisting of statements of what for a completed experience would
be matters of fact. Our ideal is to connect our presented facts by
constructions in which each link is itself matter of fact, or experience,
in the sense that it would under known conditions form the content of
a direct apprehension. But it is an ideal which, owing to the
fragmentary character of our own experience, we are never able
adequately to realise. In all our sciences we are constantly
compelled to use hypothetical constructions, which often are, and for
all we know always may be, merely “symbolic,” in the sense that,
though useful in the co-ordination of experienced data, they could
never themselves become objects of direct experience, because
they conflict either with the general nature of experience as such, or
with the special nature of the particular experiences in which they
would have to be presented. Our scientific hypotheses thus present
a close analogy with the uninterpretable stages in the application of
an algebraical calculus to a numerical or geometrical subject-matter.
Their usefulness in enabling us to co-ordinate and predict facts of
direct experience need no more guarantee their own reality, than the
usefulness of such a calculus guarantees our ability to find an
intelligible interpretation for all the symbolic operations it involves.[24]
In a pure or completed experience, at once all-comprehending and
systematic, where existence and content, fact and construction, were
no longer separated, there could of course be no place for such
ultimately uninterpretable symbolism.
Our fundamental metaphysical problem, then, is that of
discovering, if we can, the general or formal characteristics of such a
complete or “pure” experience, i.e. those characteristics which
belong to it simply in virtue of its all-containing and completely
systematic nature. Further, it would be the work of a completed
Metaphysic to ascertain which among the universal characteristics of
our own human experience of the world are such as must belong to
any coherent experience in virtue of its nature, and are thus
identifiable with the formal characteristics of a “pure” experience.
Also, our science would have to decide what features of human
experience, among those which do not possess this character,
approximate most nearly to it, and would thus require least
modification in order to enable them to take their place in an
absolutely complete and harmonious experience of reality. If we
could completely carry out our programme, we should, in the first
place, have a general conception of what in outline the constitution
of experienced reality as a systematic whole is; and, in the second,
we should be able to arrange the various concepts and categories by
which we seek, alike in everyday thinking and in the various
sciences, to interpret the world of our experience, in an ascending
order of degrees of truth and reality, according to the extent to which
they would require to be modified before they could become
adequate to express the nature of a systematic experienced reality.
The knowledge conveyed by such a science would, of course, not be
itself the pure or all-embracing experience of Reality, but merely
mediate knowledge about the general nature of such an experience,
and would therefore, so far, be like all mere knowledge about an
object, abstract and imperfect. It would still refer to something
beyond itself, and thus have a meaning other than its own existence.
But, unlike all other knowledge, our metaphysical knowledge of the
formal character of an all-inclusive experienced whole would be final,
in the sense that no addition of fresh knowledge could modify it in
principle. Fresh knowledge, which in all other cases involves at least
the possibility of a transformation of existing theories, would here do
no more than fill in and make more concrete our conception of the
system of Reality, without affecting our insight into its general
structure.
We may perhaps illustrate this conception of a knowledge which,
though imperfect, is yet final, by an instance borrowed from
elementary Mathematics. We know absolutely and precisely, e. g.,
what the symbol π stands for. π is completely determined for us by
the definition that it is the ratio of the circumference of a circle to its
diameter. And again, we can define unequivocally both the terms,
circumference and diameter of the circle, which we have employed
in our definition of π. Thus our knowledge of the meaning of the
symbol is clearly final; no fresh accretion to our knowledge will make
any modification in it. At the same time, our knowledge of π, though
final, is imperfect. For the quantity π is incommensurable, and thus
we can never precisely evaluate it. All we can do is to assign its
value correctly within any desired degree of approximation. Again,
while no approximation gives an absolutely correct value for the
quantity, one approximation is, of course, closer than another.
Because no approximation is more than approximately the truth, it by
no means follows that all are equally wide of the mark. Similarly, it
may well be that, though we can say with finality what the general
nature of experience and experienced Reality as a systematic whole
is, yet, when we come to ask after the character of the system in
detail, we have to depend on sciences which are merely
approximate in their results; it will not follow, as is sometimes
assumed, that the categories of one science do not present us with a
nearer approximation to the absolute truth than those of another.[25]
§ 11. We may end this chapter with some general reflections on
the method required for such a science of Metaphysics as we have
described in the preceding paragraphs. The true character of any
scientific method can, of course, only be discovered by the actual
use of it; a preliminary disquisition on the nature of a method not
previously exhibited in actual use is apt to be at best sterile, and at
worst a positive source of prejudices which may subsequently
seriously hamper the process of investigation. Still, there are certain
general characteristics of the method imposed on us by our
conception of the problems to be solved which may conveniently be
pointed out at this stage of our inquiry. Our method will, in the first
place, clearly be analytical and critical in its character. We analyse
experience with a view to discovering its implications, and we
analyse our various scientific and unscientific theories of the
contents of the world-system for the same purpose. Also, once
having determined what are the formal characteristics of an all-
embracing, systematic whole of experienced fact, we criticise our
various concepts and theories by reference to these characteristics
as an ultimate standard of reality and truth. Negatively, we may add
that our method is non-empirical, and also non-inductive, in the
same sense in which pure Mathematics, for instance, may be called
non-inductive. It is non-empirical inasmuch as we are called upon to
analyse all our data and criticise all our pre-conceived theories. We
are not allowed to accept any fact without analysis, or any concept
without criticism, as an unchallenged datum upon which we may
build without preliminary justification. Hence our method is non-
empirical. Also, as our analysis is concerned entirely with the internal
character and self-consistency of the data analysed, it is, like the
reasonings of pure Mathematics, independent of external
confirmation outside the analysed data themselves, and is therefore
non-inductive.[26] In precisely the same sense our method and its
results may be called, if we please, a priori; that is to say, we
proceed entirely by internal analysis of certain data, and are, alike in
procedure and result, independent of experience outside the
experience we are concerned with analysing. We can, of course, add
that our method is constructive, that is, if successfully carried out it
would culminate in an intellectual attitude towards the world which,
as an intellectual attitude, we did not possess before entering on our
study of Metaphysics; but as construction, in this sense, is
characteristic of all scientific method, it does not seem necessary to
specify it as a peculiarity of metaphysical procedure in particular.
Historically, our conception of metaphysical method as
fundamentally analytical and directed to the detection and removal of
internal contradictions in the categories of ordinary thought, is
perhaps nearer to the view of Herbart than to that of any other great
philosopher of the past. In our insistence upon the non-empirical
and, in a sense, a priori character of Metaphysics, we are again, of
course, largely in agreement with the position of Kant. There is,
however, a most important difference between our own and the
Kantian conception of the a priori upon which it is essential to insist.
A-priority, as we have used the term, stands merely for a peculiarity
of the method of Metaphysics; by an a priori method we understood
one which is confined to the internal analysis of a datum and
independent of external reference to outside facts. With Kant the a
priori is a name for certain forms of perception and thought which,
because revealed by analysis as present in every experience, are
supposed to be given independently of all experience whatsoever,
and so come to be identified by him as “the work of the mind,” in
opposition to the empirical factor in experience, which is held to be
the product of an external system of “things-in-themselves.” Hence
Kant’s whole discussion of the a priori is vitiated by a constant
confusion between what is metaphysically necessary (i.e. implied in
the existence of knowledge) and what is psychologically primitive.
This confusion, perplexing enough in Kant, reaches a climax in the
works of writers like Mr. Spencer, who appear to think that the whole
question of the presence of a non-empirical factor in knowledge can
be decided by an appeal to genetic Psychology. It is clear that, from
our point of view, the identification of the a priori with the “work of the
mind” would involve a metaphysical theory as to the constitution of
experience which we are not entitled to adopt without proof.[27]
A word ought perhaps to be said about our attitude towards the
“dialectical” method as employed by Hegel and his followers. It was
Hegel’s conviction that the whole series of concepts or categories by
which the mind attempts to grasp the nature of experienced Reality
as a whole, from the most rudimentary to the most adequate, can be
exhibited in a fixed order which arises from the very nature of
thought itself. We begin, he held, by the affirmation of some rude
and one-sided conception of the character of what is; the very
imperfection of our concept then forces us on to affirm its opposite
as equally true. But the opposite, in its turn, is no less one-sided and
inadequate to express the full character of concrete reality. Hence
we are driven to negate our first negation by affirming a concept
which includes both the original affirmation and its opposite as
subordinate aspects. The same process repeats itself again at a
higher stage with our new category, and thus we gradually pass by a
series of successive triads of categories, each consisting of the three
stages of affirmation, negation, and negation of the negation, from
the beginning of an intellectual interpretation of the world of
experience, the thought of it as mere a “Being,” not further defined,
to the apprehension of it as the “Absolute Idea,” or concrete system
of spiritual experience. It was the task of abstract Metaphysics
(called by Hegel, Logic) to exhibit the successive stages of this
process as a systematic orderly advance, in which the nature of
each stage is determined by its place in the whole. As Hegel also
held that this “dialectic” process is somehow not confined to the
“subjective” or private intelligence of the student of Philosophy, but
also realised in the structure of the “objective” universe, it followed
that its successive stages could be detected in physical nature and
in History in the same order in which they occur in “Logic,” and many
of Hegel’s best-known works are devoted to exhibiting the facts of
Physics, Ethics, Religion, and History in the light of this doctrine. The
subsequent advances of the various sciences have so completely
proved the arbitrariness and untrustworthiness of the results
obtained by these “deductions” that some of the best exponents of
the Hegelian type of Philosophy are now agreed to abandon the
claim of the Dialectic to be more than a systematisation of the stages
through which the individual mind must pass in its advance towards
a finally satisfactory conception of Reality. But even within these
limits its pretensions are probably exaggerated. No satisfactory proof
can be produced that, even in abstract Metaphysics, the succession
of categories must be precisely that adopted by Hegel. There are
some categories of the first importance, e.g., that of order in
Mathematics, which hardly get any recognition at all in his system,
and others, such as those of “Mechanism” and “Chemism,” which
play a prominent part, are obviously largely dependent for their
position upon the actual development of the various sciences in
Hegel’s own time. Hence the method seems unsuitable for the
original attainment of philosophical truth. At best it might serve, as
Lotze has remarked, as a convenient method for the arrangement of
truth already obtained by other means, and even for this purpose it
seems clear that the succession of categories actually adopted by
Hegel would require constant modification to adapt the general
scheme to later developments of the various special sciences.
Consult further:—F. H. Bradley, Appearance and Reality, chaps.

13, 14; B. Bosanquet, Essentials of Logic, Lect. 2; Shadworth
Hodgson, Metaphysic of Experience, bk. i. chap. 1; J. S. Mackenzie,
Outlines of Metaphysics, bk. i. chaps. 2 and 4. And for criticism of the
Hegelian dialectic: J. E. M‘Taggart, Studies in Hegelian Dialectic,
chaps. 1-3; J. B. Baillie, The Origin and Significance of Hegel’s
Logic, chaps. 8-12, especially chap. 12; and also Adamson,
Development of Modern Philosophy, bk. i. pp. 271 ff.
10. See Bosanquet, Essentials of Logic, Lect. 8. As an illustration

we may take an extreme case: “The Jabberwock was not killed
yesterday.” What is the ground of this denial? At first sight it appears
to be merely negative, “there are no such things as Jabberwocks to
kill.” But before I can say “there are no such things as Jabberwocks”
with confidence, I must have enough positive information about the
structure and habits of animals to be aware that the qualities
ascribed to the Jabberwock conflict with the laws of animal life. Or, if
I deny the existence of Jabberwocks simply on the ground that I
have never come across a specimen, this involves a positive
judgment as to the relation between the animal world and the part of
it I have examined, such as, “if there were Jabberwocks, I should
have come across one”; or, “my acquaintance with the varieties of
animals is sufficiently exhaustive to afford ground for a valid
generalisation.” The fact that symbolic Logic finds it convenient to
treat the universal affirmative as a double negative must not mislead
us as to its actual priority in thought.
11. To meet the kind of criticism which finds it humorous to jest at
the expense of those who “take consolation from spelling Reality
with a big R,” may I once for all say that when I spell Reality thus it is
simply as a convenient way of distinguishing the ultimately from the
merely relatively real?
12. We shall meet this same difficulty again later on as the
principle of the famous Kantian objection to the “ontological proof” of
God’s existence. Infra, Bk. IV. chap. 5. § 8.
13. Take a concrete example. A theory as to the early religious
history of the Hebrews, let us say, is put forward upon grounds
derived from Semitic philology. Though unacquainted with Semitic
philology in particular, I may be able to form some sort of estimate of
the cogency of the professed reasoning if I already have an
adequate acquaintance with the use and value of philological
evidence in parallel cases, say, in the study of Greek antiquities. But
if I have no positive acquaintance at all with the use of philology in
antiquarian research, it would be the merest impertinence for me to
offer any opinion whatever.
14. What follows must be regarded as a mere outline which awaits
subsequent filling up by the more concrete results of Bk II. chap. 1.
15. For the modern logical doctrine of possibility consult Bradley,
Principles of Logic, 192-201; Bosanquet, Logic, i. chap. 9.
16. This is—apart from non-essential theological accretions—the
principle of Berkeley’s argument for the existence of God (Principles
of Human Knowledge, §§ 146, 147).
17. I should explain that I use “feeling” and “apprehension”
indifferently for immediate and non-reflective awareness of any
psychical content. The exclusive restriction of the term to awareness
of pleasure and pain seems to me to rest on a serious mistake in
psychology, and I therefore avoid it.
18. In fact, we shall see in Bk. II. chap. 1 that in virtue of its unity
with immediate feeling, all experience is essentially connected with
purpose.
19. I take “fact” as equivalent to “what is directly apprehended in a
single moment of consciousness.” In a previous work (The Problem
of Conduct, chap. 1) I used the word in a different sense for “the
contents of a true description of experience.” This employment of the
word, however, seems at variance with established philosophical
usage, and I therefore abandon it as likely to lead to
misapprehension.
20. Of course, the apprehended “content” may itself be a
“process,” as is the case in all instances of the apprehension of
change; but the apprehended process is always distinguishable from
the process of apprehension.
21. We shall see in Bk. II. chap. 1 that the “that” of an experience
implies relation to a unique individual interest or purpose.
22. Of course this is only partly true. As we shall see in the sequel,
to “be what it means and mean what it is” is an ideal never fully
realised in the structure of any finite piece of reality, precisely
because the finite, as its name implies, is never a completely
systematic whole.
23. On the psychological processes by which meaning is acquired,
see Stout, Manual of Psychology3, bk. i. chap. 3; and on the
apprehension of form, the same author’s Analytic Psychology, bk. i.
chap. 3. Much interesting discussion of the difference between
“external” and “internal” meaning will be found in Royce, The World
and the Individual, First Series.
24. For some good observations on the fallacy of assuming that
mathematical symbolism must always be interpretable, see B.
Russell, Foundations of Geometry, p. 45-46; or Whitehead,
Universal Algebra, vol. i p. 10 ff. For a further elaboration of the
argument of the foregoing section I may refer to my Problem of
Conduct, pp. 14-21. I need hardly warn the reader against confusing
a “symbolic” concept in my sense of the word, i.e. one which cannot
be fully interpreted in terms of direct experience, with a “symbolic”
idea in Mr. Spencer’s sense, i.e. one which is not, psychologically, a
copy of the presentations for which it stands. Our use of the word is,
of course, purely logical, and has nothing to do with the
psychological character of mental images, but only with their
meaning.
25. Compare my Problem of Conduct, pp. 22-39.
26. The fundamental peculiarity of “inductive” procedure, in fact, is
that, while its object is the internal analysis of its data, which, if
completed, would permit of a universal conclusion being drawn from
the single case, it is never able to effect the analysis, and is driven to
reinforce it by external comparison with “similar” cases.
27. On the confusion between the metaphysical and psychological
standpoint in Kant’s own treatment of the a priori, see B. Russell,
Foundations of Geometry, pp. 1-4, and Adamson, Development of
Modern Philosophy, bk. i. pp. 244-247.
CHAPTER III
THE SUB-DIVISIONS OF METAPHYSICS

§ 1. The traditional sub-division of Metaphysics into Ontology, Cosmology,
Rational Psychology, common to all the great modern constructive systems. §
2. Precise sense in which we adopt these divisions for the purposes of our
own treatment of the subject. § 3. Relation of Cosmology and Rational
Psychology to the empirical sciences.
§ 1. English philosophers, who have usually been imbued with a

wholesome distrust of deliberate system-making, have commonly
paid comparatively little attention to the question of the number and
character of the sub-divisions of metaphysical philosophy. They have
been content to raise the questions which interested them in the
order of their occurrence to their own minds, and have gladly left it to
the systematic historians of Philosophy, who have rarely been
Englishmen, to discuss the proper arrangement of the parts of the
subject. Continental thinkers, who are naturally more prone to
conscious systematisation, have bestowed more thought on the
problem of method and order, with the result that each great
independent philosopher has tended to make his own special
arrangement of the parts of his subject. The different arrangements,
however, seem all to agree in conforming to a general type, which
was most clearly exhibited by the otherwise rather arid Wolffian
dogmatism of the eighteenth century. All the constructive systems
(those, e.g., of Hegel, Herbart, Lotze,) feel the necessity of giving the
first place to a general discussion of the most universal
characteristics which we find ourselves constrained to ascribe in
thought to any reality which is to be an intelligible and coherent
system and not a mere chaos. This division of the subject is
commonly known by the title it bears alike in the Wolffian Metaphysic
and the systems of Herbart and Lotze, as Ontology,[28] or the general
doctrine of Being; with Hegel it constitutes, as a whole, the contents
of the science of Logic, as distinguished from the other two great
departments of speculative thought, the Philosophies of Nature and
Mind; and its most formal and general parts, again, compose, within
the Hegelian Logic itself, the special first section entitled “Doctrine of
Being.”
Further, every system of metaphysical philosophy is bound to deal
with more special problems, which readily fall into two principal
classes. It has to consider the meaning and validity of the most
universal conceptions of which we seek to understand the nature of
the individual objects which make up the experienced physical world,
“extension,” “succession,” “space,” “time,” “number,” “magnitude,”
“motion,” “change,” “quality,” and the more complex categories of
“matter,” “force,” “causality,” “interaction,” “thinghood,” and so forth.
Again, Metaphysics has to deal with the meaning and validity of the
universal predicates by which we seek to interpret the nature of the
experiencing mind itself, and its relation both to other minds and to
the objects of the physical world, “the soul,” “the self,” “the subject,”
“self-consciousness,” “ethical purpose,” and so forth. Hence it has
been customary to recognise a second and third part of
Metaphysics, dealing respectively with the most general
characteristics of external Nature and of conscious Mind. These
sections of the subject are commonly known as Cosmology and
Rational Psychology. In Hegel’s system they appear in a double
form: in their most abstract generality they constitute the “Doctrine of
Essence,” and the “Doctrine of the Notion” in the Hegelian Logic; in
their more concrete detail they form the second and third parts of his
complete system or “Encyclopædia” of the philosophical sciences,
the previously mentioned Philosophies of Nature and Mind.
In the pre-Kantian eighteenth century it was not unusual to add yet
a fourth division to Metaphysics, Rational Theology, the doctrine of
the existence and attributes of God, so far as they can be deduced
from general philosophical principles apart from the appeal to
specific revelation. Kant’s onslaught on the whole Wolffian scheme in
the “Dialectic of Pure Reason,” while profoundly modifying for the
future the view taken by metaphysicians of Cosmology and Rational
Psychology, proved annihilating so far as eighteenth-century Deism
and its philosophical offspring, Rational Theology, were concerned,
and that sub-division may fairly be said to have disappeared from
subsequent philosophical systems.[29]
§ 2. There are good and obvious reasons why we should adhere,
in the form of our inquiry, to the main outlines of this traditional
scheme. It is true that it is largely a question of simple convenience
what order we adopt in a systematic metaphysical investigation. A
genuinely philosophical survey of the general character of
knowledge and experience would exhibit so complete a systematic
unity, that you might start from any point in it and reach the same
results, much as you may go round a circle equally well from any
point of the periphery. But for the beginner, at any rate, it is
advantageous to start with the general question what we mean by
Being or Reality, and what character is to be ascribed to the whole of
Being as such, before attacking the problem of the particular kind of
Being which belongs to the various “realities” of common life and the
special sciences. Thus we have to discuss in the first part of our
programme such questions as the relation of Being in general to
experience, the sense in which Being may be said to be inseparable
from, and yet again to transcend, experience; the problem of the
existence of different kinds or degrees of Being; the question
whether Being is ultimately one or many; the relation between Real
Being and its appearances. All these problems correspond with
reasonable closeness to the contents of what was traditionally
known as Ontology.
It is only when we have reached some definite conclusion on
these most fundamental questions that we shall be in a position to
deal with the more special problems suggested by the various
departments of science and common life; hence we shall do well to
acquiesce in the arrangement by which Ontology was made to
precede the other divisions of the subject. Again, in dealing with the
more complex special problems of Metaphysics, it is natural to
recognise a distinction corresponding to the separation of
Cosmology from Rational Psychology. Common language shows
that for most of the purposes of human thought and action the
contents of the world of experience tend to fall into the two groups of
mere things and things which are sentient and purposive—Physical
Nature on the one hand, and Minds or Spirits on the other. We must,
of course, be careful not to confuse this division of the objects of
experience with the distinction between an experienced object as
such and the subject of experience. We are to start, in our critical
investigation, not with the artificial point of view of Psychology, which
sets the “subject” of presentations over-against the presentations
considered as conveying information about “objects of knowledge,”
but with the standpoint of practical life, in which the individual agent
is opposed to an environment itself consisting largely of similar
individual agents. It is not “Nature” on the one side and a “perceiving
mind” on the other, but an environment composed partly of physical
things, partly of other human and animal minds, that furnishes the
antithesis on which the distinction of Cosmology from Rational
Psychology is founded. There is no confusion against which we shall
need to be more on our guard than this fallacious identification of
Mind or Spirit with the abstract subject of psychological states, and
of the “environment” of the individual with Physical Nature. Of
course, it is true that we necessarily interpret the inner life of other
minds in terms of our own incommunicably individual experience, but
it is equally true that our own direct experience of ourselves is
throughout determined by interaction with other agents of the same
type as ourselves. It is a pure delusion to suppose that we begin by
finding ourselves in a world of mere physical things to some of which
we afterwards come by an after-thought, based on “analogy,” to
ascribe “consciousness” akin to our own. Hence, to avoid possible
misunderstandings, it would be better to drop the traditional
appellations “Cosmology” and “Rational Psychology,” and to call the
divisions of applied Metaphysics, as Hegel does, the Philosophy of
Nature and of Spirit or Mind respectively.[30]
In recognising this sub-division of applied Metaphysics into two
sections, dealing respectively with Physical Nature and with Mind or
Spirit, we do not mean to suggest that there is an absolute disparity
between these two classes of things. It is, of course, a matter for
philosophical criticism itself to decide whether this difference may not
in the end turn out to be merely apparent. This will clearly be the
case if either minds can be shown, as the materialist holds, to be
simply a peculiar class of highly complex physical things, or physical
things to be, as the idealist contends, really minds of an unfamiliar
and non-human type. It is sufficient for us that the difference,
whether ultimate or not, is marked enough to give rise to distinct
classes of problems, which have to be treated separately and on
their own merits. We may feel convinced on general philosophical
grounds that minds and physical things are ultimately existences of
the same general type, whether we conceive that type after the
fashion of the materialist or of the idealist, but this conviction does
not in the least affect the fact that the special metaphysical problems
suggested by our experience of physical things are largely different
from those which are forced on us by our interest in the minds of our
fellows. In the one connection we have, for instance, to discuss the
questions connected with such categories as those of uniform spatial
extension, uniform obedience to general law, the constitution of a
whole which is an aggregate of parts; in the other, those connected
with the meaning and value of ethical, artistic, and religious
aspiration, the concept of moral freedom, the nature of personal
identity. Even the categories which seem at first sight most readily
applicable both to physical things and to minds, such as those of
quality and number, lead to special difficulties in the two contrasted
cases. This consideration seems to justify us in separating the
metaphysics of Mind from the metaphysics of Nature, and the
superior difficulty of many of the problems which belong to the
former is a further reason for following the traditional order of the two
sub-divisions, and placing Rational Psychology after Cosmology. In
so far as the problems of Rational Theology can be separated from
those of general Ontology, the proper place for them seems to be
that section of Rational Psychology which deals with the meaning
and worth of our religious experiences.
§ 3. It remains, in concluding the present chapter, to utter a word
of caution as to the relation between the two divisions of applied
Metaphysics and the body of the empirical sciences. It is perhaps
hardly necessary to warn the student that Rational Cosmology and
Psychology would become worse than useless if conceived of as
furnishing in any sense a substitute for the experimental study of the
physical, psychological, and social sciences. They are essentially
departments of Metaphysics, and for that very reason are incapable
of adding a single fact to the sum of our knowledge of ascertained
fact. No doubt the discredit into which Metaphysics—except in the
form of tacit and unconscious assumption—has fallen among

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Human Evolution: Bones, Cultures, and

Genes John H. Langdon

On Life: Cells, Genes, and the Evolution of Complexity

Landscapes of Human Evolution: Contributions in Honour

Disembodied Brains: Understanding our Intuitions on

Artificial Intelligence and Human Evolution:

Information Theory and Evolution, 3rd Edition John

Nuclear Decisions Lisa Langdon Koch

Hearts and Bones Niamh Mulvey

The Pleistocene Social Contract: Culture and

ISSN 2730-6135 ISSN 2730-6143 (electronic)

I find the study of paleoanthropology both fascinating and humbling. The

Indianapolis, IN, USA John H. Langdon

1 The Reflection in the Mirror 3

2 Background: Evolutionary Classification and Fossil Dating 31

3 A Primate Heritage 51

Part II The First Hominins 71

5 The Early Hominins: Australopiths103

6 The Ecological Context of Early Hominin Evolution145

8 The Evolution of Bipedality191

Part III The First Humans 249

9 The Earliest Humans in Africa251

10 The First Technology277

11 Diet of Early Homo299

12 Evolution of the Brain321

Part IV The Middle Pleistocene 389

14 The Erectines of Asia391

15 Erectines of the West419

16 Behavior in the Middle Pleistocene461

Part V The Emergence of Modern People 495

17 Late Pleistocene Homo and the Emergence

18 From the Middle Paleolithic to the Modern Mind539

19 Modern Humans Disperse From Africa581

20 Distributing Modern Peoples625

21 Life History and Reproduction651

22 Evolution Today and Tomorrow683

Appendix of Anatomical Terms697

Fig. 8.7 OH 8 partial foot of Paranthropus boisei from Olduvai Gorge,

Fig. 9.1 The geographic distribution of early fossils of genus Homo in

each has been averaged from many specimens to indicate to

File:Meganthropus_palaeojavanicus.jpg. CC BY-SA 4.0, via

Sediment_1997_%C2%A9_P._Pfarr_NLD.jpg. CC BY-SA 3.0,

Fig. 20.2 Reconstructed phylogeny of human lice (red) mapped onto

(green). The rest of the reproductive lifespan consists of cycles of

Table 1.1 Commonly recognized or recently proposed species of hominins

negative values represent the opposite, contributing to a lordosis.

Table 16.1 The spread of Acheulean technologies by early appearance 467

The Reflection in the Mirror

Key Ideas in this Chapter

1. Human evolution is told as a narrative, with characters, motivations, and a

© The Author(s), under exclusive license to Springer Nature 3

Box 1.1 Some Essential Vocabulary

Constructing Human Identity

Fig. 1.2 Comparative brain sizes among larger-brained mammals. Comparisons of

Box 1.2 How Are Humans Different?

Box 1.2 (continued)

• Lateral tubercle on the calcaneal tuberosity*.

Behavioral traits in the following list also distinguish humans more by

Indianapolis, IN, USA John H. Langdon

1 The Reflection in the Mirror 3

2 Background: Evolutionary Classification and Fossil Dating 31

3 A Primate Heritage 51

Part II The First Hominins 71

5 The Early Hominins: Australopiths103

6 The Ecological Context of Early Hominin Evolution145

8 The Evolution of Bipedality191

Part III The First Humans 249

9 The Earliest Humans in Africa251

10 The First Technology277

11 Diet of Early Homo299

12 Evolution of the Brain321

Part IV The Middle Pleistocene 389

14 The Erectines of Asia391

15 Erectines of the West419

16 Behavior in the Middle Pleistocene461

Part V The Emergence of Modern People 495

17 Late Pleistocene Homo and the Emergence

18 From the Middle Paleolithic to the Modern Mind539

19 Modern Humans Disperse From Africa581

20 Distributing Modern Peoples625

21 Life History and Reproduction651

22 Evolution Today and Tomorrow683

Appendix of Anatomical Terms697

Table 16.1 The spread of Acheulean technologies by early appearance 467