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Springer Texts in Social Sciences
John H. Langdon
Human
Evolution
Bones, Cultures, and Genes
Springer Texts in Social Sciences
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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
© The Editor(s) (if applicable) and The Author(s), under exclusive licence to Springer Nature
Switzerland AG 2022
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To all my ancestors, remembered, forgotten, or undiscovered.
Preface
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.
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
4 Miocene Hominoids 73
7 Understanding Australopiths165
xi
xii Contents
13 Leaving Africa359
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
3.6 Ma (cast). Photo by Katherine Langdon 136
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
Wikimedia Commons 150
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
Katherine Langdon 175
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
Wikimedia Commons 193
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. 8.16 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
CC BY-SA 2.0, via Wikimedia Commons 233
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. 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
Wikimedia Commons 289
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,
via Wikimedia Commons 292
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
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,
via Wikimedia Commons 401
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
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
Wikimedia Commons 435
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,
via Wikimedia Commons 436
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.
CC BY-SA 4.0, via Wikimedia Commons 445
Fig. 15.19 Cranium from Steinheim, Germany, an example of a gracile
specimen of H. heidelbergensis, about 150 Ka (model). Photos by
Katherine Langdon 446
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
Wikimedia Commons 466
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
Wikimedia Commons 477
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
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
Katherine Langdon 526
Fig. 17.21 LH17 Ngaloba cranium from Ndutu Beds near Laetoli, Tanzania
(cast). Photo by Katherine Langdon 526
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.
CC BY-SA 2.0, via Wikimedia Commons 527
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
Wikimedia Commons 546
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.
CC BY-SA 3.0, via Wikimedia Commons 549
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
BY-SA 4.0, via Wikimedia Commons 551
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
BY-SA 4.0, via Wikimedia Commons 560
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
BY-SA 4.0, via Wikimedia Commons 560
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.
CC BY-SA 4.0, via Wikimedia Commons 584
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,
via Wikimedia Commons 584
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
Wikimedia Commons 598
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
BY-SA 4.0, via Wikimedia Commons 599
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
Wikimedia Commons 601
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.
CC BY-SA 2.5, via Wikimedia Commons 603
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.
CC BY-SA 4.0, via Wikimedia Commons 610
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
Wikimedia Commons 629
List of Figures xxxiii
xxxv
xxxvi List of Tables
Introduction
CHAPTER 1
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.
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
1 THE REFLECTION IN THE MIRROR 7
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
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.
1 THE REFLECTION IN THE MIRROR 9