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John H. Langdon
Human
Evolution
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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some extent free of malarial poison, and certain parts have a climate
which some Europeans, even Englishmen enjoy.
CHAPTER XLVII
BRAZIL: THE CAPITAL, INDIVIDUAL STATES, CITIES
The Capital
Rio de Janeiro, the capital of Brazil, with a population of

1,200,000, the second city in South America and the fifth in the
Western Hemisphere, is generally conceded to have the finest
harbor in the world. It is also the most beautiful city in respect to
natural advantages, while the improvements which have recently
been carried out have made the artificial structure of the city in
harmony with its picturesque surroundings. In the early part of this
century the Government awoke to the disadvantages of the narrow
streets, the poor facilities for commerce, and the notoriously bad
conditions of health. Immense improvements were inaugurated and
speedily executed. Fine drainage canals were installed, mosquitoes
practically exterminated, wide avenues were cut through dense
quarters, and port works constructed, so that in health, beauty, and
commercial convenience, nothing remains to be desired. A new
Central Avenue, now renamed Rio Branco, is counted among the
finest in the world; other splendid avenues are lined with fine
commercial structures or beautiful private residences, perhaps with
royal palms; along the shores are magnificent boulevards unrivaled
in beauty, often thronged with luxurious automobiles, all amid
surroundings of ideal loveliness of sea-shore and of mountain. Hotel
accommodations, hitherto only fair, and inadequate, are improving
with the erection of needed new structures, as increasing travel for
business and pleasure demanded. Living expenses are called high
in all parts of Brazil, but on the whole Rio has seemed no dearer
than New York. The city may boast of a splendid opera house, said
to have cost $5,000,000, of the largest library in South America, and
the finest art gallery, a wonderful botanical garden, museums,
delightful sea and mountain resorts, etc., etc.
EASTERN BRAZIL
States and Cities
As the country of Brazil is so large and of such diversity in its

various sections and even in single States, with many independent
systems of railways connected with its numerous harbors, it may be
desirable to speak of its ports and to some extent of its railways in
connection with a review of the different States. In this we may begin
with the Coastal States, coming up from the south, and conclude
with those of the interior.
Rio Grande do Sul, eleventh in area among the States of Brazil,
and equal in size to Indiana and Illinois together, borders on Uruguay
at the south. The Uruguay River separates it from Argentina on the
west and most of the way from the State of Santa Catharina on the
north. Its population is about the same as that of the State of South
Carolina, which is one-third of its size. It is called a pastoral State as
its chief source of wealth at present is cattle; 45 of the 70 million
acres are cattle ranges, 6¹⁄₄ million, farms, and 13³⁄₄ forests. The
nearly 9,000,000 head of cattle are called the best in Brazil. Blooded
stock is being introduced and packing houses are taking the place of
saladeros for charque or dried meat, which formerly were the
destination of most of the cattle. High class poultry is extensively and
profitably raised, $4,500,000 being invested in the industry.
With a temperate climate and well watered, the country is by no
means confined to stock raising, for its agricultural products are
more diverse than those of any other state. Not only cereals like
wheat and oats, but cotton, rice, sugar cane, tobacco, manioc,
alfalfa, etc., even tea are successfully raised, as well as temperate
zone fruits and vegetables. It has important coal mines, with some of
copper, gold, and wolfram also being operated, and a wide
distribution of these and other minerals. With 13,000 industrial
establishments its important factories include nine textile mills.
The capital, Porto Alegre, is the chief manufacturing centre, and
the State is third in such industries. The capital is the largest city in
southern Brazil, with a cosmopolitan population one fourth of
German descent. An important commercial and industrial centre, it
has fine public buildings, colleges, theatres, clubs, good shops,
parks, and hotels. The population is nearly all white with a large
percentage of Italians as well as of Germans.
Rio Grande is the chief port, 1000 miles south of Rio de Janeiro,
and 180 by sea north of Montevideo. The city lies just inside the
entrance of the Lagôa dos Patos at its south end. Unfortunately, the
entrance has been obstructed by a sand bar with a varying depth of
water above. Vessels drawing more than 11 feet could not pass;
uncertainty, often a few days’ waiting was the fate of others. An
attempt to dredge a passage was unsuccessful, but with building of
jetties improvement is manifest. Port works in the city, including a
mile of quays, have been established, and a rapid development of
the region is expected to follow. The city, population 40,000, has an
active commerce by rail and water. By rail it is connected at Cacequy
with the through line from São Paulo to Montevideo, and beyond this
line at Uruguayana (population 20,000), on the Uruguay River, with
the Argentine railways on the other side. There is regular steam
navigation to the port of Pelotas, three hours, population 32,000,
noted for its dried meat products, and to Porto Alegre at the north
end of the lagoon, 150 miles and 12 hours distant.
Santa Catharina, north of Rio Grande, less than one third of its
size, has rather similar characteristics. Bordering on the Atlantic, it
has Argentina on the west; on the north is the State of Paraná, from
which it is separated for half the distance by the Iguassú River. The
State is notable for wheat growing and other grains, for cattle and
dairy products, for its exports of fruits, for its coal mines, and for the
timber from its forests, especially the Paraná pines. It ranks second
in Brazil as a producer of yerba mate, in Portuguese herva matte,
14,000,000 pounds being its annual output. Its manufacturing
establishments (174) are of small size.
The capital and chief port is Florianopolis on an island of that
name. Improvements in the way of good port works are in hand, and
though without rail connection a tonnage of about 300,000 has
entered and cleared in a year. The port of São Francisco, a smaller
town in the State, is called the best port south of Santos. This
advantage will give the city future importance. Already it has rail
connection by way of the town Rio Negro with Curityba and
Paranaguá, and so with Ponta Grossa on the São Paulo-Uruguay
Railway which of course crosses the State, a distance of 225 miles.
The city is expected to be the terminus of a road which will pass
through União da Victoria to the Iguassú Falls and across Paraguay
to Asunción.
Paraná, about the size of Rio Grande, is between Santa Catharina
and São Paulo, having Argentina also on its southwest corner
beyond the Iguassú River and its great Falls. Along the western
border flows the larger river, the Alto Paraná (with the tremendous
La Guayra Falls) separating the State from Paraguay, and farther
north from Matto Grosso. Besides these boundary rivers the
Paranapanema, affluent of the Paraná, flows between this State and
São Paulo, while the Rio Negro and Iguassú are between it and
Santa Catharina. Many more tributaries of the Paraná are entirely
within the State. Matte is the chief industry at present, 100,000,000
pounds being annually exported; but with the best wheat land of
Brazil, its forests of Paraná pine, and other varieties of timber it may
have another leader before long. Paranaguá, the State’s chief port,
with a deep anchorage and improvements planned, is now visited by
650 vessels a year and has a total annual trade of $6,000,000. A
smaller port, Antonina, is on the north side of the same bay.
The capital, Curityba, has rail connection with the two ports and
with Ponta Grossa on the line to Uruguay traversing these three
States. In the delightful four hours’ journey of 70 miles from
Paranaguá to the capital, the climb of the Serra do Mar up a steep
tropical valley is made without cogs or cables by means of high
trestles, bridges, and 17 tunnels; the ride rivaling if not surpassing in
beauty the more famous one from Santos to São Paulo.
São Paulo, a great and justly famous State, about the size of
Arizona, more than twice that of New York, has Minas Geraes on the
north, also east with a small extent of the State of Rio de Janeiro; the
ocean is southeast, Paraná south, and Matto Grosso west. This
enterprising State is the leader in agriculture, producing 60 per cent
of the world’s coffee supply, besides cotton, sugar, tobacco, cereals,
etc., in large quantities; it is second to Rio in manufacturing
industries, contains large mineral riches, is advanced in stock
raising, leads in educational advantages, and has the best railway
service of 4300 miles. The wealth of this State is estimated as at
$1,100,000,000 in agriculture, $500,000,000 in manufactures,
$170,000,000 in railways, and $2,230,000,000 miscellaneous. While
most of it is in the torrid zone, the altitude of the State averaging over
2000 feet gives it (except for the narrow coastal strip) a fine healthful
climate, a blessing enjoyed too by the States previously mentioned.
In the Falls of its rivers the State possesses 3,000,000 horse power
of which only 250,000 is employed.
The port of Santos, population 80,000, is regularly visited by the
large Transatlantic and North American steamers. It is called one of
the best and most important ports of the world, receiving 1600 ships
annually besides coasting vessels. The largest ocean steamers, 20-
40 foot draft, come up to the quay which extends for three miles
along the harbor front; fine granite walls rise five feet above high
water mark on a base 10-20 feet thick. Modern machinery is
provided, making it the best equipped port in Brazil, and nothing is
allowed to interfere with the efficiency of the service. It is a very busy
city though warm. The heat does not prevent people from rushing
about. A Brazilian writer has said, “People do not run, they fly.” The
reason for this unusual haste, by no means manifest in Rio, is that
many prominent business men come daily, others occasionally, from
São Paulo by morning train, returning at four p.m. A splendid railway
which will be referred to later, leads to that important city, a two
hours’ ride.
The capital city, São Paulo, 310 miles from Rio and 50 from
Santos, while lacking the charm of Rio’s scenic beauty, is preferred
by many as a place of residence on account of its cooler climate, the
greater business activity apparent, and the cosmopolitan society,
more than one third of the population being of foreign birth and
another one third direct descendants of Europeans. The city has
many splendid buildings, both public and private, including an opera
house superior to any in the United States, a large number of
excellent schools of various kinds, and all the attractions and
facilities, except good and adequate hotels, of a city of half a million
inhabitants.
The Federal District of Brazil, the capital, Rio de Janeiro, has
been sufficiently described except as a port city. From a commercial
point of view the harbor and port works are the chief interest. The
Guanabara Bay is a wonderful harbor, not merely on account of its
size, depth of water, absolutely safe anchorage, and the beauty of its
surroundings, but it is extraordinary from the fact that it is hardly a
mile from the ocean to the landing docks. Here a granite quay
extends for 2¹⁄₄ miles along the water front with a depth of water
alongside of 31 feet. The deep channel entrance is narrowed to a
mile in width by long narrow peninsulas extending on either side and
further by a small island blocking the waves. The docks have the
additional protection of a projecting point of the city, on the other side
of which, on rare occasions, the waves do break over upon the
splendid boulevard. The larger inner harbor is hardly noticeable,
being cut off by peninsulas and islands, of which last the bay
contains nearly a hundred. The entire bay is 18 miles long, and the
inner section is 15 miles wide. At the docks the most modern
machinery is provided for hoisting, loading and unloading ships, with
electric power for the work and for lighting. A width of 80 feet was left
for railway tracks; back of these are storage ware-houses,
administration offices, and customs, for which a space of 110 feet
was allowed. Behind these buildings is an avenue 125 feet wide,
with double tracks on which run electric cars. Two million dollars are
now (1921) to be expended for additional port works, an extension of
the granite quay or dock wall about 2000 feet and two breakwaters
of 800,000 cubic feet.
Rio de Janeiro. This State, though containing or surrounding the
capital, is distinct from it, with a capital of its own on the opposite
shore of the bay. As the direction of the coast line changes here, the
State has the ocean east and south; São Paulo is west, Minas
Geraes north, and Espirito Santo at the northeast. The State is
comparatively small, being only about one sixth the size of its
western neighbor, and less than one twelfth of its northern; it is even
a trifle smaller than Espirito Santo, its next coastal neighbor.
Nictheroy, the capital, is a comparatively small, quiet town of
65,000 population, which has some important manufactories. The
State has other smaller towns, as Petropolis, the so-called summer
capital, population 30,000, at an elevation of 3000 feet. The
Parahyba Rio do Sul, which flows northeast, back of the mountain
range, is navigable to the city of Campos, population 30,000, 60
miles from its mouth. The agricultural products are important, the
State ranking third in coffee and second in sugar. Large quantities of
corn and rice are raised, the coast lands with lakes and lagoons
being well suited to the latter. Valuable forests and minerals exist,
and in manufacturing industries of great variety the State stands first.
Besides the harbor of Rio there are a number of good though small
ports.
Espirito Santo. Little Espirito Santo, northeast of Rio State, has
Minas at the west and Bahia north. The principal products of the
State are coffee, rice, and other cereals, sugar, cotton, and
mandioca; while the export of fine timber, rosewood, satin and brazil
wood, is increasing. Gold and precious stones are its chief minerals.
Its factories are few, but the town of Itaperim on a navigable river has
cheap electric power, which makes it a fine centre for industry in the
future.
The capital and chief city, Victoria, population 20,000, is the first
port of importance north of Rio. On a fine bay 2¹⁄₂ miles wide with a
narrow entrance of less than ¹⁄₂ a mile, it is the outlet of the eastern
part of the State of Minas, which contains the richest mineral
deposits so far exploited in Brazil. A railway connects the port with
interior cities. Works of importance have been planned, a quay ⁵⁄₈ of
a mile long with 28 feet of water and with suitable equipment; the
widening and deepening of the channel entrance, and the building of
a steel bridge to the main land from the island on which the city
stands. Over 500 steamers and 200 sailing vessels clear yearly.
Bahia, an important and well known State, the largest yet
considered, exceeding California in area, touches three smaller
States on the north, Piauhy, Pernambuco, and Sergipe; it has
Espirito Santo and Minas on the south and Goyaz west. All kinds of
tropical and subtropical products are found here, cacao, sugar, and
coffee in large quantities, rubber of the maniçoba and mangabeira
varieties, cotton, vanilla, the finest kind of oranges and pine apples,
and other fruits. Bahia has the principal whale fisheries of the
country and the best grounds for table fish. Its mineral wealth
includes even the sand, monazite, the most important supply in the
world.
The capital and chief port, 720 miles northeast of Rio, about 60
hours sail, is generally called by the name of the State, though its
proper title is São Salvador. It has an excellent and beautiful harbor,
though over-shadowed by the more remarkable picturesqueness of
Rio. The bay, Bahia de Todos os Santos, about 25 miles wide (three
at the entrance), and 20 miles deep, provides good anchorage for
large steamers, 40 feet close in shore. It is a port of call for Trans-
Atlantic liners and for steamers from North America, the only one
north of Rio for some of the Lines. A company has undertaken port
works which will greatly advance the prosperity of City and State, the
works to include three breakwaters and two quay walls, the levelling
of a large wharf space, erection of store-houses, laying railway lines,
installing electric cranes, and constructing a floating dock and a
lighthouse. Over 1000 large steamers with 2,300,000 tonnage enter
the port annually. The imports are $13,000,000, the exports
$20,000,000. The city is connected by rail with various cities in the
interior of the State, and with the São Francisco River at a point
above the Falls, from which there is navigation upstream a distance
of 990 miles. The river has a length in the State of 850 miles. Bahia
is a great cocoa port, shipping about one-fifth of the world’s supply;
the State produces about as much tobacco as Cuba.
Sergipe, the smallest of all the States, is larger than nine of ours,
a trifle bigger than Maryland and Delaware together. The São
Francisco River separates it from Alagôas on the north, it has Bahia
on the west and south. Its chief products are cotton, sugar, and rice;
the cattle industry is important; the State has profitable
manufacturing industries; cotton mills, sugar, shoes, soap, and other
factories, and unworked mineral deposits.
The capital, Aracajú, population 40,000, is a small port with 95,000
tons of shipping yearly; but it suffers the disadvantage of receiving
ships of no more than 8 feet draft, and needs the improvements now
planned.
Alagôas, of triangular shape, has Sergipe on one side and
Pernambuco on the other, the ocean on the third. Its main products
are sugar and cotton; the cattle industry is prosperous; it has copper,
lead and iron deposits, not operated, and very important milling
industries, particularly of cotton. The São Francisco River is regularly
navigated 175 miles to the Paulo Affonso Falls, around which there
is a railway 52 miles long to navigation above.
The capital, Maceió, is a modern city; its suburb, the port, Jaraguá,
with a tonnage of 600,000. The State is the most thickly populated
next to Rio de Janeiro.
Pernambuco, the twelfth State in size, with area equal to that of
New York, extends well into the interior, though with a coast line of
112 miles only. Ceará and Parahyba are on the north, Alagôas and
Bahia on the south, and Piauhy on the west. The State leads in the
production of cotton and sugar, sometimes exporting 150,000 tons of
the latter, and raw cotton worth $5,000,000. Other agricultural
products are secondary, but cattle and dairying are important, still
more the milling industry. Minerals, coal, iron, saltpetre, kaolin, and
manganese exist in paying quantities. There are two good ports
besides the capital; railways connect Recife with Maceió and with
cities inland.
Recife (often called Pernambuco), capital of the State, is a port
protected by a coral reef parallel to the shore, where fine works are
being constructed: these include a breakwater three-fourths of a mile
long, a stone jetty one-half a mile; quays, one with 33 feet of water,
one with 28-30, together three-fourths of a mile long, also other
equipment. One thousand steamers are its annual quota, with
tonnage of 1,750,000; the city has obviously great commercial
importance, exporting especially cotton and sugar, also rum, hides,
and cereals. Its imports surpass those of any other Brazilian city
except Rio. The cost of living is high.
Parahyba has on the north Rio Grande do Norte, Pernambuco
south, Ceará west, and 72 miles of coast east. Cotton is the chief
product, and mandioca is important. Vast coconut groves of trees,
growing wild, should be taken advantage of. The pastoral industry is
important, with goats a specialty as with its neighbors.
A little below the capital, Parahyba, at the mouth of the Rio
Parahyba do Norte, is the seaport Cabedello, where port works are
planned.
Rio Grande do Norte is the first of several States which have the
Atlantic Ocean on the north. It is on the east also, forming a coast
line of 290 miles. Parahyba is south and Ceará west. Here, too,
cotton and sugar are the chief products, though 180 tons of rubber
annually, from the maniçoba trees, are of value; the cattle and goats
are important; carnaüba wax and vegetable oils come from the
forests; the State, from natural salines, supplies much of the salt
used in Brazil. Dried and salt fish are supplied to the rest of the
northern States, and much cotton cloth and thread are exported.
Natal, the capital, is near the mouth of the Rio Grande. A great
reservoir with a dam 160 feet high and a capacity of 2,200,000,000
cubic metres of water is to be constructed, the irrigation of 250,000
acres thereby transforming a large district; others here and in Ceará.
Ceará, smaller than Pernambuco, is nearly twice the size of Rio
Grande do Norte, which with Parahyba is on the east; Pernambuco
is south and Piauhy west. Cotton is the chief product though coffee,
sugar, cacao and cereals are also raised. Cultivated rubber is
exported as from Bahia; the cattle industry is important, though
affected by occasional severe droughts. However there are
approximately 2,000,000 head of cattle. Minerals and precious
stones are found in variety.
The capital and port, Fortaleza, is one of the worst on the coast.
Three powerful dredges are maintained by the Government for the
continual dredging of the channel into which sand from the dunes is
ever drifting. Two smaller ports require similar dredging.
Piauhy, eighth of the States in size, with a deep interior has the
smallest coast line, only 43 miles. It has Ceará and Pernambuco
east, Bahia south, and Maranhão west, from which it is separated by
the Parnahyba River. The State has the expected agricultural
products, vast herds of cattle and large flocks of goats; important
forest wealth of rubber, timber, wax, and medicinal plants, and a
variety of minerals. It has one seaport, Amarraçao, but a good deal
goes out from a port of the next State, Tutoya, on the other side of
the river Parnahyba. Therezina, the capital, is of some importance.
Maranhão, a little larger, has Piauhy east and south, Goyaz south
and west, almost touching Bahia between these two; it has Pará also
on the west. Its coast line is 100 miles more than Piauhy’s. Cotton is
the chief agricultural product, but others exist; the cattle industry is
important, much live stock going to Amazonas; there is good forest
wealth, various minerals, and important cotton factories.
São Luis do Maranhão, the capital, population 50,000, is called the
chief port though troubled by sand, while Tutoya is excellent.
Pará, a name at last familiar to all, is third of the States in area,
equal to Texas and California, with West Virginia thrown in. It has
Guiana on the north, with the Atlantic northeast; east are Maranhão
and Goyaz. Matto Grosso is south, and Amazonas west. Its chief
products are of the forest, particularly rubber, also Brazil nuts,
medicinal plants, oils and timber. Little is done in agriculture though
many plants grow freely such as cotton, rice, tobacco, and especially
cacao, of which 3500 tons a year have been exported. There are
large herds of cattle, perhaps 2,000,000 head, and various minerals
are found. One railway, with branches, leads from Pará to Bragança,
near the ocean, and to other towns; and one is open for a short
distance in the valley of the Tocantins, the beginning of an ambitious
project.
The capital, Belém, usually called by foreigners Pará is a fine city
of 200,000 and the only considerable seaport, if so it may be termed,
as it is on a bay of the Pará River 80 miles from the ocean. Mean
temperature 78°. Harbor works, begun in 1906, and costing over
$60,000,000, have been of immense value. They include a fine quay
a mile and a half long with a depth of water part of the way of 30
feet, docks and storehouses, two floating docks, a Custom House,
oil storage tanks, etc. One thousand steamers formerly entered
yearly with tonnage of 1,500,000; imports at one time were valued at
$15,000,000 and exports at $30,000,000. A channel from the outer
river, 30 feet in depth, is marked by 26 buoys lighted by acetylene
gas. The city is notable with attractive plazas, a unique forest park, a
museum, a white marble theatre, and a good hotel. The various
cities or villages are coast or river ports, some on the Amazon,
others on the Tapajós, Xingú, Tocantins, Araguary, or smaller
streams.
Amazonas, first of the inland States, and the largest of all, has
Colombia and Venezuela north, Pará east, Matto Grosso, Bolivia,
and Acre south, and Peru and Colombia west. Its area is equal to
that of our three Pacific Coast States with Idaho, Montana, Nevada,
and Colorado. In this great space the population, estimated at
435,000, is about one person to 1¹⁄₂ square miles. Manaos, the
capital, and the various other centres of population by the river side
mean rubber. There are no roads except of water, no paths save
those made by rubber gatherers, with a few by Indians, the number
of whom is a mere guess. All tropical products thrive but their
production is negligible save that of rubber. Various minerals are
unexploited. Manaos, 924 miles from Pará, 1030 from the ocean,
and 2000 from Rio, is the first real city in the wilderness, though
Obidos, still in the State of Pará, is a port where 300 vessels call in a
year, and a number of smaller places claim that title.
Manaos, a city of 60,000 population, is located just off the Amazon
seven miles up the Rio Negro, on a large quiet bay. Port works have
been constructed, a fine stone wall over 1500 feet long, with floating
docks to fit the 50-foot rise and fall of the river, and 16 electrically
equipped warehouses conveniently arranged. Considering its
location, it is a wonderful city, well lighted, with a splendid opera
house, expensive of course, a fine cathedral, schools, a public
library, museum, and good sanitation. One thousand four hundred
miles farther is the Peruvian city of Iquitos, near the limit of
navigation for ocean vessels on the main stream. Some distance
below Manaos the Amazon is entered by the Madeira River which
gives access to the State of Matto Grosso and to Bolivia.
Matto Grosso is second in size, equal to the States mentioned
above without Nevada and Colorado. Estimated population 275,000.
This State has Amazonas and Pará on the north, Goyaz, Minas
Geraes, São Paulo, and Paraná on the east, Paraguay and Bolivia
are on the south, and Paraguay and Bolivia west. The State is much
more open than Amazonas; its chief industry is cattle. Forest
products however abound, with all kinds of rubber and magnificent
timber. There are large agricultural possibilities and considerable
matte is exported. Diamonds and auriferous sands are exploited
though their origin is uncertain. The river systems north and south
meet in the highlands, and connection might be made by a short
canal, opening a way from Pará or Manaos to Buenos Aires.
Cuyabá, the capital, on a river of the same name, an affluent of
the Paraguay, is quite a city, population 32,000, although 1045 miles
above Asunción, while Corumbá, lower down, on the Paraguay, is of
nearly equal size. The river at Corumbá is 1000 feet wide, and 6 feet
deep at the docks at low water. Corumbá is regularly visited by
steamers, though 1800 miles by river from Montevideo. Its export
and import trade amounts to at least $4,000,000 annually.
Goyaz, the fourth State in size, following Pará, has Maranhão on
the north, Maranhão, Bahia, and Minas Geraes are on the east,
Minas and Matto Grosso are south, and Matto Grosso and Pará
west. Three hundred thousand is the estimated population with a
guess at the number of Indians of many tribes. The principal industry
is stock raising and many cattle are exported to neighboring States.
The forests have the maniçoba and the mangabeira rubber, also a
vegetable silk, paina. There are great mineral riches, placer gold in
many streams, and veins in the hills; many diamonds and rock
crystal are also produced. Other metals as iron and copper exist.
Navigable rivers are the means of communication as in the
neighboring States, but a railroad is coming and more rapid
development will follow.
The capital, Goyaz, is not much of a place, though of late evincing
progress. Several other cities have from 5000 to 10,000 inhabitants.
Minas Geraes, the fourth inland State and the fifth in size, has
Bahia on the north, Bahia and Espirito Santo east, Rio de Janeiro
southeast, São Paulo southwest, and Goyaz west, a small southwest
corner about reaching Matto Grosso below. Although without a
seaport and with no large city, Minas is the most populous of the
States, with an active industrious population. The State leads in
mineral riches of almost every kind; it is one of the foremost in
agriculture of all varieties, being second to São Paulo in coffee; it
has vast pasture lands, exporting 300,000 head of cattle a year, a
sugar refinery, flour mills, and a great dairy industry. It has increasing
railway service as well as river transportation. It is believed to have a
mineral future rivaling that of the best region in the world. Its
factories are important and there is a great store of water power.
Bello Horizonte, the present capital, was made to order in 1897,
and is well laid out with broad streets, water supply, sewerage,
everything of the best type: a Government Palace, the finest State
building of Brazil, and a fine Agricultural School. Ouro Preto, the old
capital, has a free Mining School, said to be one of the best in the
world.
The Acre Territory, triangular in shape, has Amazonas north,
Bolivia south, and Peru west. It is naturally similar to the neighboring
sections. There are three Districts: Juruá, of which the capital is
Cruzeiro do Sul, population 2000, 1351 miles from Manaos; Purús,
capital, Senna Madureira, population 4000, 1320 miles from
Manaos; and Acre, capital Rio Branco, population 2000, 1351 miles
from Manaos.
CHAPTER XLVIII
BRAZIL: TRANSPORTATION—OCEAN, RIVER, AND RAILWAY
Ocean and River Traffic
Brazil has a considerable navy, several warships; and a merchant

marine of 450,000 gross tonnage. The leading national line is the
Brazilian Lloyd which with 62 ships has a service to the United
States as well as a coastwise, the latter shared by the Navegaçao
Costeira and other companies. Rio de Janeiro has three dry docks,
one with a capacity for the largest battleships, and a yard where
ships are constructed. The country has 30,000 miles of navigable
rivers, with boats for these having a tonnage of 75,000.
Coastwise and river steamers have service in the Amazon Basin,
the most important as follows: Pará-Obidos-Manaos, 975 miles;
Pará-Santarem-Itaituba (Rio Tapajós) 729 miles; Pará-Maués, Pará
Tocantins, 1544 miles; Tabatinga (Frontier) Remate dos Males, 1743
miles; Pará-Santo Antonio (Madeira) 1617 miles; Pará-Rio, Peru-
Senna-Madureira, 1934 miles; Pará-Chaves (Marajos Island),
Oyapock River; Manaos and Rio Negro to Santa Isabel, 423 miles;
Manaos to the Yapurá River—to the Juruá River, Cruzeiro do Sul,
1090 miles.
Other companies have service, Maranhão north to Pará, 599
miles, south to Pernambuco 884 miles; Recife north to Maranhão
803 miles, south to Bahia 385 miles, east to Fernando do Noronha
Island, 239 miles. On the São Francisco River, Januaria to Pirapora.
Other Lines serve from Rio de Janeiro to ports south to Laguna and
Ribeira de Iguapé, São Paulo; also on the Paraná and the Rio
Grande, and on the Lakes Patos and Mirím. Service from Rio de
Janeiro to Corumbá, 765 miles above Asunción, and 280 miles
farther to Cuyabá on the Cuyabá River, in time of high water is
continued on the Paraguay 250 miles beyond to São Luiz de
Caceres.
Railway Transportation
While the great rivers of Brazil with their 27,000 miles of navigable
waters have been a large factor in the development of the country, of
equal importance for the future is transportation by rail. The total
present mileage, second to that of Argentina, exceeds 16,500, these
having developed from 9¹⁄₂ miles in 1854 and about 10,000, 50 years
later. About 90 per cent are of metre gauge. For the economic unity
of the great Republic, the Government policy is favorable to a rapid
extension of the present railway systems, a matter as important as
was the creation of our railways to the Pacific 50 years ago. The
development in Brazil has been retarded by the difficult topography
of the country, in striking contrast to the Argentine plains. Along the
greater and better part of the coast is the high steep wall of the
plateau region, which must be climbed to enter the interior. Once at
the top the way in some sections is easier, but in others there are
additional mountain ranges. The wall, obviously low in comparison
with the mighty rampart of the Andes, presents difficulties, but none
to compare with those experienced in Peru.
The first railway, built and operated in 1854, was from Mauá on the
north shore of the Rio harbor to Fragosa. Soon after carried to
Petropolis, it is now a part of the Leopoldina Railway system. Only
two South American railways antedate this, the Demarara of British
Guiana and the Caldera-Copiapó in Chile.
The Central of Brazil Railway was the first of much importance,
its first section, 32 miles, opened in 1858. For this road leading out of
Rio expenses were heavy. Many were the difficulties of construction,
tunnels, bridges, etc., the boring of one tunnel, only 1¹⁄₂ miles long,
requiring seven years. The system, now in Government ownership,
has a network of lines extending from Rio to the city of São Paulo, to
Bello Horizonte in Minas, and to other points. It has express trains
with sleeping cars (state rooms), and a heavy suburban traffic. It is
intended to carry this system northward to the city of Pará, with
branches to new territory. Twenty-five million passengers are carried
annually with much freight of coffee, lumber, iron, manganese, etc.
The railway is to be electrified near the capital with a Government
appropriation of $32,760,000.
The São Paulo Railway. The most successful, financially and
otherwise, of the railways of Brazil is no doubt the São Paulo,
operating a double track, broad gauge line (5 feet 6 inches) from
Santos to the town and junction, Jundiahy (population 20,000), 86
miles; it passes through the city, São Paulo, and has a branch to
Bragança, 65 miles. This road has a granted monopoly of trade
between the two cities, Santos and São Paulo, 50 miles apart. This,
in spite of the great expense of the railway construction, has insured
them large profits, dividends in 1912-13 being 14 per cent. The train
mile earnings are the largest in South America and normally are
greater than any reported in the United States. With the construction
of more and more lines in this State and beyond, a steady increase
in the traffic seems assured, as nearly all must pass in and out by
way of Santos. The road now carries annually more than half of the
world’s coffee supply.
ENVIRONS OF SÃO PAULO AND RIO DE JANEIRO

One of many engineering triumphs in South America, the railway
is ranked among the greater achievements of the world. An ascent of
2600 feet is made within seven miles. Beginning 15 feet above the
sea not far from Santos five inclined planes with 8 per cent grade,
each 1¹⁄₂ miles long, serve for the rapid climb. A stationary engine at
the top of each plane runs the cables; to grip these a small engine is
attached to each car. The tracks seem unique. The double track on
the inclines has but three rails for up and down cars, which therefore
cannot meet there, but may on the intermediate levels of which there
are four, each 600 feet long. On each side between the middle and
the outside rails are the pulleys which carry the cable, an endless
steel wire of enormous strength run by a 1000 horse power engine,
capable of carrying six freight or three passenger cars at the same
time. In this short section are 16 viaducts, 15 tunnels, and two miles
of retaining wall. One viaduct is 334 feet long, and nearly 150 feet
high in the centre. There are now two of these double roads, one just
above the other, as I have seen, and not on a different route as
stated elsewhere. The first soon proving insufficient to accommodate
the freight, the second was built with a few technical improvements.
It is a wonderful ride through tropical forests, with scenes of
picturesque beauty.
The Sorocabana Railway, 864 miles, which traverses a rich and
progressive part of Brazil opening up new territory, has important
connections with other lines. This Company has several lines
running west and northwest from São Paulo. One of the most
important is to Baurú, a city also on the Paulista Railway. From this
point the North Western Railway has continued the line to Itapura on
the Paraná River and across the State of Matto Grosso to Porto
Esperança on the Paraguay, about 40 miles by land from Corumbá
on the west side of the River, and 80 by water. Probably a train-car
ferry will later be provided at Esperança and the road continued on
the other side to Corumbá. By the present road connection is made
with Bolivia opposite, and by water with Paraguay farther down, the
capital Asunción being about 700 miles distant, Montevideo nearly
1800. Previously to the completion of this railway Corumbá and that
section of Brazil was reached from Rio only by a long, long sail,

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Human Evolution: Bones, Cultures, and

Genes John H. Langdon

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

• Pair-bonding of sexual partners beyond a mating cycle.

Genomes Humans are also distinct genetically. Comparisons of genomes are

Rio de Janeiro, the capital of Brazil, with a population of

States and Cities

As the country of Brazil is so large and of such diversity in its

Ocean and River Traffic

Brazil has a considerable navy, several warships; and a merchant

ENVIRONS OF SÃO PAULO AND RIO DE JANEIRO

You might also like

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