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Horse Breeds and Breeding
in the Greco-Persian World
Horse Breeds and Breeding
in the Greco-Persian World:
1st and 2nd Millennium BC

By

Thomas Donaghy
 Horse Breeds and Breeding in the Greco-Persian World:
1st and 2nd Millennium BC,
by Thomas Donaghy

This book first published 2014

Cambridge Scholars Publishing

12 Back Chapman Street, Newcastle upon Tyne, NE6 2XX, UK

British Library Cataloguing in Publication Data

A catalogue record for this book is available from the British Library

Copyright © 2014 by Thomas Donaghy

All rights for this book reserved. No part of this book may be reproduced, stored in a retrieval system,
or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording or
otherwise, without the prior permission of the copyright owner.

ISBN (10): 1-4438-5363-1, ISBN (13): 978-1-4438-5363-7

 TABLE OF CONTENTS

List of Illustrations ................................................................................... viii

Maps ............................................................................................................ x

Acknowledgements ................................................................................... xii

Introduction ................................................................................................. 1

Part I: Evolution, Domestication, and Spread

Chapter One ................................................................................................. 8

Evolution of the Horse

Chapter Two .............................................................................................. 12

Domestication

Chapter Three ............................................................................................ 18

Early Horse-Riding

Chapter Four .............................................................................................. 26

Development and Spread of Chariotry

Chapter Five .............................................................................................. 36

Chariot Warfare

Chapter Six ................................................................................................ 40

Greek Chariot Warfare

Chapter Seven............................................................................................ 52
Horse-Riding during the Chariot Age

Chapter Eight ............................................................................................. 56

Development of Cavalry Forces

Chapter Nine.............................................................................................. 59
War Chariots during the Cavalry Age
vi Table of Contents

Part II: Breeds—The Greek World

Chapter Ten ............................................................................................... 72

The Troad

Chapter Eleven .......................................................................................... 79

Mainland Greece

Chapter Twelve ....................................................................................... 100

Thessaly

Chapter Thirteen ...................................................................................... 107

Thrace and Macedon

Chapter Fourteen ..................................................................................... 116

South Italy and Sicily (Magna Graecia)

Chapter Fifteen ........................................................................................ 131

North Africa

Part III: Breeds—The Eurasian Steppe and the Persian World

Chapter Sixteen ....................................................................................... 136

Scythia and the Eurasian Steppe

Chapter Seventeen ................................................................................... 142

The Heavenly Horses of Ferghana

Chapter Eighteen ..................................................................................... 147

The Lands of the Persian Empire

Chapter Nineteen ..................................................................................... 184

The Nisaean Horse

Chapter Twenty ....................................................................................... 191

The Caspian Miniature Horse

Chapter Twenty-One ............................................................................... 197

The Horse and Arabia
 Horse Breeds and Breeding in the Greco-Persian World vii

Conclusion ............................................................................................... 206

Abbreviations .......................................................................................... 209

Bibliography ............................................................................................ 212

Index ........................................................................................................ 225

 LIST OF ILLUSTRATIONS

Illustrations by Peter McCanney

Fig 10.1 - Hellenistic era statue of a bridled horse discovered at Pergamon.

Pergamon Museum. Berlin.

Fig 11.1 - Greek horses from the Parthenon frieze (ca. 438 - 432 BC).
Equestrian group from the west frieze. British Museum, London, West
Frieze II, 2 - 3.

Fig 12.1 - Coin of Larissa, Thessaly. Reverse of drachma (ca. 400 - 360
BC). Bridled stallion prancing right. Above and below ȁǹȇǿȈǹǿǹ
(Larissa). British Museum, London.

Fig 12.2 - Detail of Alexander and Bucephalas on the ‘Alexander

Sarcophagus’ from the Royal Cemetary of Sidon (ca. 310 BC).
Archaeological Museum, Department of Classical Antiquities, No. 68
(370), Istanbul.

Fig 13.1 - Thracian Horse Rython. Silver rython (fifth – fourth century
BC) from Basova Hill, Duvanli. Archological Museum, Plovdiv,
Bulkgaria.

Fig 13.2 - Detail of horse in Skudrian (Thracian?) procession. Eastern

Stairway, Apadana Palace, Persepolis, Iran. Fifth century BC.

Fig 13.3 - Coin of Philip II of Macedon (ca. 359 - 336 BC). Reverse of
tetradrachm. Young Rider on a large muscular horse. To either side,
ĭǿȁǿȆȆȅȊ = of Philip.

Fig 14.1 - Coins of Tarentum depicting ‘Tarentine’ cavalrymen.

Left: obverse of stater (ca. 333 – 331/0 BC).
Right: obverse of didrachm (ca. 290 – 281 BC).

Fig 14.2 - Bronze horse and rider from Armento, Basilicata, southern Italy
(ca. 560 – 550 BC). British Museum, London.
 Horse Breeds and Breeding in the Greco-Persian World ix

Fig 14.3 - Coin of Syracuse depicting a four horse chariot. Obverse of

tetradrachm (ca. 510 – 490 BC).

Fig 15.1 - Coin of Carthage. Reverse of a gold one-and-a-half stater (ca.

270 - 260 BC).

Fig 16.1 - Scythian horse from the Chertomlyk Vase discovered in the
Chertomlyk kurgan (Dnieper basin, Ukraine). Fourth century BC. The
State Hermitage Museum, St. Petersburg.

Fig 18.1 - Assyrian horse and groom. Detail of an Assyrian relief from
Nimrud (ca. 865 BC). British Museum, London, No. ANE124548.

Fig 18.2 - Detail of horse in Armenian procession. Eastern stairway,

Apadana Palace, Persepolis, Iran. Fifth century BC.

Fig 18.3 - Detail of horse in Cappadocian procession. Eastern stairway,

Apadana Palace, Persepolis, Iran. Fifth century BC.

Fig 18.4 - Nisaean horse from Apadana. Detail of a procession depicting

one of the king’s riding horses. Eastern stairway, Apadana Palace,
Persepolis, Iran. Fifth century BC.

Fig 18.5 - Rommelaere’s Longiligne type horse from the tomb of

Rekhmire at Thebes. Eighteenth Dynasty, reigns of Thoutmosis III (1490 -
1436 BC) and Amenhotep II (1438 - 1412 BC).

Fig 18.6 - Rommelaere’s Breviligne type horse from the tomb of Menna at
Thebes. Eighteenth Dynasty, reign of Thoutmosis IV (1412 - 1403 BC).

Fig 18.7 - Detail of horse in the Sogdianaean procession. Eastern

Stairway, Apadana Palace, Persepolis, Iran. Fifth century BC.

Fig 20.1 - Syrian (Caspian?) horse from Apadana. Detail of horses from
Syrian procession. Eastern stairway, Apadana Palace, Persepolis, Iran.
Fifth century BC.
Horse Breeds and Breeding in
n the Greco-Perrsian World xi
 ACKNOWLEDGEMENTS

This book developed out of my PhD thesis, “Horse Breeds and

Breeding in the Greco-Persian World: 2nd and 1st Millennia BC”, completed
at University College Dublin’s School of Classics in 2009. This book
would not have been possible without the assistance of the Irish Research
Council for the Humanities and Social Sciences (IRCHSS) whom I would
like to thank for funding of the final two years of my PhD.
I would especially like to thank my supervisor Dr. Philip de Souza for
his careful mentoring and insightful assistance throughout my years
researching this topic. I also wish to thank all the other members of staff
and postgraduate students of the School of Classics during my time there
for their kindness and constant willingness to help.
Many thanks to Dr. Kelly Fitzgerald for her continued encouragement
and to Peter McCanney for the excellent illustrations provided.
Last, and certainly not least, the greatest thanks must to go to my
family who always offer unconditional support for whatever I am doing.
 INTRODUCTION

After creating the heavens and the earth, the birds of the air and the fishes
of the sea, God found it good to bestow on man a supreme mark of his
favour: he created the horse. In the magnificent sequence of creation the
last phase, that of perfection, was reserved for this beautiful creature. The
horse was swifter than anything on the face of the earth; he could outrun
the deer, leap higher than the goat, endure longer than the wolf. Man,
encompassed by the elements which conspired to destroy him, by beasts
faster and stronger than himself, would have been a slave had not the
horse made him king. In the Garden of Eden the horse was of no service to
man, but the fall of man revealed to the horse his noble mission.1

The association between human and horse is one which has existed for
tens of thousands of years. From as early as thirty-five to forty thousand
years ago humans were hunting horses in large numbers for both food and
hides and, not long after this, horses began to appear quite frequently in
cave paintings or as figurines carved from bone and mammoth tusk. As the
post-glacial climate warmed and forests overtook steppe-lands wild horses
were gradually pushed eastwards and their habitat restricted to the
Eurasian steppe. Budiansky suggests that, had it not been for their
domestication, the remaining herds would have gradually disappeared and
the horse would be extinct today.2 That this may indeed have come to pass
is suggested by the fact that the last surviving herd of true wild horses, the
Przewalskii horse of Mongolia, disappeared late in the last century. The
last wild specimen of this breed was spotted in 1969.3
So it seems only fair that the horse, when it was saved from probable
extinction through domestication, repaid the favour by contributing so
much to the history of humankind. The domestication of the horse and the
subsequent development of chariotry and cavalry greatly changed human
lifestyle. Not only were horses available for food and hides they could also
be used for transportation and warfare.

1
Eighteenth century French explanation of man’s partnership with the horse.
[Chenevix-Trench, (1970), 7].
2
Budiansky, (1997), 35.
3
Przewalski horses bred from captive specimens have since been successfully
reintroduced into the wild.
2 Introduction

It was for the peoples of the Eurasian Steppe that horses seem to have
facilitated the most radical change. Possession of horses permitted these
peoples to abandon their sedentary ways on the fringes of the great plains
and to embrace a fully nomadic lifestyle. Throughout history such
nomadic cultures regularly came into conflict with the settled cultures of
non-steppe lands and the former played an important role in the
development and evolution of the latter. From the steppes of the Far East
the nomadic Hsiung-Nu proved a constant menace to the Chinese
kingdoms throughout the second half of the first millennium BC while in
Western Europe the nomadic Huns played a large role in the troubles faced
by the Roman Empire during the fourth and fifth centuries AD. The largest
land empire which the world has ever known was created in the late
twelfth and early thirteenth century AD by the nomadic Mongols. When
horses were introduced into the Americas by European explorers and
conquistadors during the fifteenth and sixteenth centuries AD they swiftly
affected cultural lifestyles and within two centuries the Plains Indians of
North America had embraced a fully nomadic lifestyle similar to that of
their Eurasian counterparts.
Horses also greatly influenced the history and development of
sedentary civilisations. Indo-European peoples (possibly originating from
the border-lands of the steppes), with their newly developed chariot
technology, spread their culture through a wide swathe of the ancient
world through a series of invasions and/or migrations. By the middle of
the second millennium BC chariot-using Indo-European cultures had
established themselves as Hittites in Asia Minor, Mitannians in Armenia,
Kassites in Babylonia, and Aryans in northwest India to name but a few.
Chariot forces were to play a vital role in the military might of many
kingdoms in the eastern Mediterranean and Near East until they were
supplanted by cavalry in the first millennium BC. Chariotry was to play an
even greater role in China and India where it continued in military use
down to the latter centuries of the first millennium BC.
The development of cavalry forces continued the horse’s contribution
to history and the rise of great empires, such as those of the Assyrians,
Medes, and Persians, owed much to their ability to breed and raise large
forces of cavalry. When Alexander the Great invaded the Persian Empire it
was cavalry which played a vital role in his success. The great importance
of cavalry continued throughout medieval times and well into the modern
era. As late as the Second World War horses played a vital military role.
Although by WWII they had mostly been replaced by mechanised forces
many hundreds of thousands of horses were still used for transporting
troops to the front and for the transport of provisions and heavy artillery.
 Horse Breeds and Breeding in the Greco-Persian World 3

Although cavalry forces do not generally form part of modern armies

(with the exception of some ceremonial units) they are still occasionally
used. Most notable is their use by the Janjaweed militiamen in Darfur.
Probably less well known is the fact that cavalry forces were recently in
use in America’s war against the Taliban in Afghanistan. When a Special
Forces Operational Detachment was air-dropped into Afghanistan in the
fall of 2001 they were surprised to discover that the Northern Alliance
force of their ally General Dostum consisted of a mounted unit of two
thousand horsemen armed with modern small arms. Since the Taliban
possessed relatively modern tanks and artillery the Americans used GPS,
laser range finders, and digital satellite radios to destroy the Taliban’s
bunkers and military hardware after which the Afghan horsemen would
charge the enemy positions.4
Due to the vital role which the horse has played in human history there
have been many publications over the years detailing the history of the
association between human and horse. The majority of such works,
however, concentrate mostly on the horse’s military contribution without
making any real attempt to examine the origin of the horse itself. That is to
say that, although much has been written on the construction and
manufacture of chariots, the origin and development of cavalry equipment,
and the tactics and manoeuvres used by chariots and cavalry in battle,
there has been much less written concerning the origin and development of
the various breeds of horse and how they spread across the ancient world.
In order to address this imbalance this book takes as its topic an
examination of the origins and development of the variously attested horse
breeds of the Greco-Persian World of the second and first millennia BC.
This particular time period has been selected for examination because it
was during these two millennia that the vital role which the horse was to
play in human history became fully apparent. The second millennium BC
saw the development of the chariot and the subsequent creation of vast
chariot forces which were to form an important part of the armed forces of
numerous lands from Mycenaean Greece in the west as far as India and
China in the east. The first millennium BC saw the gradual replacement of
these chariots with cavalry forces which continued to play a vital role in
military warfare right up until the beginnings of the twentieth century AD.
This book will focus its examination upon the region of the Greco-
Persian world because of the great interaction which these lands
experienced during this period. For the purposes of this book the Greek
World is taken to encompass those main regions of the ancient
Mediterranean that were either settled by Greeks or had close contact with
4
DiMarco, (2008), 352 - 354.
4 Introduction

(or influence from) Greek lands. The Persian World encompasses all those
lands which were later to fall under the sway of the Persian Empire when
it was at the height of its power (ca. 480 BC).
The interaction between these two regions during the second and first
millennia BC was constant and extensive. There was continual trade
between the many nations of the eastern Mediterranean and the Near East
throughout this period as well as cultural and military interaction. The
Mycenaean Greeks had strong connections with both Egypt and western
Anatolia and often were engaged in military actions against Hittite
dependencies. There was close contact between the major powers of the
Near East at this time - Hatti, Mitanni, Assyria, Babylon, and Egypt. From
the mid-fifth century BC the Persian Empire was regularly engaged in
military conflict with the Greek World while the late-fourth century BC
witnessed the conquest of the Persian Empire by the Macedonian and
Greek armies of Alexander the Great.
Existing publications on the topic of ancient horses can be broadly
divided into two main categories. The first category consists of works
whose focus is to provide a general account of the history of the horse over
a very large period of time - such as from its evolution, domestication, or
from a particular time (e.g. Classical Greece or the Roman Republic) up
until the present day. Due to the lengthy time span covered any examination
of horses during the second and first millennia BC tends to form but a small
portion of the whole with the greater part focusing on the period from late
medieval times to the present. Examples of such publications include C.
Chenevix-Trench, (1970), A History of Horsemanship, New York; A. Dent,
(1974), The Horse Through Fifty Centuries of Civilisation, London; D. M.
Goodall, (1977), A History of Horse Breeding, London; H. B. Barclay,
(1980), The Role of the Horse in Man’s Culture, London; and J. Clutton-
Brock, (1992), Horse Power: A History of the Horse and Donkey in
Human Societies, London.
The second category of publications focus more closely on the horses
of the ancient world and, therefore, tend to contain more detailed
information regarding the period in question. While some of these works
may indeed contain information and discussions in regard to ancient
breeds, in the majority of cases the focus is more on an analysis of
harnessing and tack (saddles / saddle-cloths, bits, bridles, harnessing
equipment for chariot horses) and the role horses played in military
matters (tactics, manoeuvres, cavalry armour and weaponry). Such works
can also be divided into two main categories - those concerned with the
ancient world in general and those in which specific regions or nations are
examined. Examples of the former include A. Hyland, (2003), The Horse
 Horse Breeds and Breeding in the Greco-Persian World 5

in the Ancient World, Surry; A. Cotterell, (2005), Chariot: The Astounding

Rise and Fall of the World’s First War Machine, London; and P. Sidnell,
(2006), Warhorse: Cavalry in Ancient Warfare, London. Examples of the
latter include A. Hyland, (1990), Equus: The Horse in the Roman World,
London; J. H. Crouwel, (1992), Chariots and other wheeled vehicles in
Iron Age Greece, Amsterdam; R. E. Gaebel, (2002), Cavalry Operations
in the Ancient Greek World, Norman, Oklahoma.
Although the categories of publications mentioned above constitute the
larger portion of works dealing with the history of the horse, there are a
small number of publications covering the period dealt with by this book
which do attempt to examine the origin and development of particular
breeds. Works which include an analysis of ancient breeds include C.
Rommelaere, (1991), Les Chevaux du Nouvel Empire Egyptien: Origines,
Races, Harnachement, Bruxelles and P. G. Gonzaga, (2004), A History of
the Horse Volume I: The Iberian Horse from Ice Age to Antiquity, London.
The former contains an excellent analysis of Egyptian horse paintings
from the New Kingdom Period which indicate that two separate breeds of
horse may have entered Egypt during this period. The author attempts to
identify and trace the origins of both these breeds. The latter consists of a
detailed investigation into the possible origins and development of Iberian
horses through an examination of paintings, sculptures, archaeological
remains, and the movement of horses throughout the ancient world
through trade, migration, and warfare.
Such being the current state of this field of research, this book will
examine humankind’s influence on the origin and development of the
various breeds of ancient horse which are known from the Greco-Persian
World of the second and first millennia BC. Such an examination will help
provide a better understanding of the history of humankind’s association
with the horse. Our understanding of the huge role which the horse played
in human history can only be improved by our gaining an understanding of
the equally huge role which humans played when they took horses from
the wild and, through many hundreds of years of daily interaction, cross-
breeding, and training, facilitated the development and spread of many
different breeds throughout the ancient world. The examination of breeds
will attempt, through an analysis of surviving archaeological,
iconographical, and literary evidence, to determine what those described
physical characteristics and given attributes of the various breeds may
reveal about their possible geographical and racial origins.
This book is divided into three main parts. Part One provides an
historical overview of the origin and spread of the horse including an
overview of its evolution, domestication, spread, and the origins and
6 Introduction

development of horse riding and chariotry.

Part Two entails a detailed overview of the main regions of the Greek
World which were famed for producing high-quality horse breeds during
the second and first millennia BC. For the purposes of this thesis the term
‘Greek World’ encompasses the main regions of the ancient
Mediterranean world that were either settled by Greeks or else had close
contacts with (or influence from) Greek lands. Drawing upon a wide range
of source material (including archaeology, iconography, literary sources,
geographical and agricultural surveys) this section aims to gather together
information which explains why these particular regions were capable of
breeding high-quality horses, the number of horses that each region was
capable of breeding, and what the physical appearance of these breeds (as
well as the particular attributes associated with them) can tell us about
their possible origins.
Part Three takes the same approach as Part Two but applies it to the
horse-breeding lands of the Eurasian steppe and the Persian World. The
analysis of the Eurasian Steppe covers those breeds used by nomadic
horsemen from the plains of Eastern Europe right the way across to the
plains of Mongolia. This was a region which, despite its enormous extent,
possessed a relatively uniform culture as well as relatively uniform breeds
of horse. For the purposes of this thesis the term ‘Persian World’ is used to
encompass all those lands which were later to fall under the sway of the
Persian Empire when it was at the height of its power (ca. 480 BC). The
majority of the narrative, however, is based around ancient texts detailing
the Assyrian rise to power in the early centuries of the first millennium BC
(supplemented by a similar range of source material as used in Part Two).
The section has been structured this way because it was the Assyrians who
were the first to join under one rule the all the major horse-breeding lands
of the Near East which lay between the eastern Mediterranean and the
Zagros Mountains. It was the control of these core breeding-grounds upon
which the basis of the Assyrian Empire’s military might depended as well
as that of the subsequent Median and Persian Empires.
 PART I:

EVOLUTION, DOMESTICATION, AND SPREAD

 CHAPTER ONE

EVOLUTION OF THE HORSE1

The horse (Equus Caballus) is just one member of the larger family of
equus that also includes the ass (Equus Asinus), the onager (Equus
Hemionus), and the three species of zebra (Equus Zebra, Equus Burchelli,
and Equus Grevyi). For the origins of the Equus genera one must go back
to the Eocene Period (ca. 54 – 38 million years ago) in America.2 It was
here that the first equids originated and, over the next fifty-four million
years, evolved and spread across the world.
The first equid to be found in the fossil record is aptly named Eohippus
(Greek for ‘Dawn Horse’).3 Eohippus was a small, forest-dwelling animal
which stood between 10 and 20 inches (25 and 50 cm) at the shoulder and
weighed approximately 5.4 kg (12 lbs). It had a flexible, arched back (in
contrast to the rigid spine of today’s horse), a short neck, snout, and legs,
and a long tail.
Unlike modern horses Eohippus was toed rather than hoofed. It had
four toes on its front feet and three on its hind (with visible traces of two
more). Each toe terminated in a small hoof-like nail. Its under-foot was
protected by a dog-like pad. In the forest environment of Eocene North

1
This overview of horse evolution draws much from the following works: Hulbert,
(1996), 11 - 34; MacFadden, (1988), 131 - 158; Edwards, (1987), 17 - 25; Clabby,
(1976), 6 - 12.
2
The following geological periods cover the evolution of the horse. Eocene (ca. 54
- 38 million years ago), Oligocene (ca. 38 - 26 million years ago), Miocene (ca. 26
- 7 million years ago), Pliocene (ca. 7 - 3 million years ago), Pleistocene (ca. 3
million years ago - recent). [Clabby, (1976), 1].
3
Eohippus is also commonly referred to as Hyracotherium due to a mistake made
in the early identification of its fossil finds. When partial remains of a skull were
discovered in Kent, England in the early 1800’s they were identified as belonging
to a cony or hyrax and therefore the find was given the scientific name
Hyracotherium. In 1867 a more complete skeleton was discovered in Eocene rock
structures in the southern part of the United States and named Eohippus. It was not
until 1932, however, that the connection between the two finds was made.
[Edwards, (1987), 17 - 18].
 Evolution of the Horse 9

America Eohippus would have browsed on fruit and soft foliage and so its
teeth were not adapted for rough grazing. They were low-crowned and
consisted of three sets of incisors, one set of canines, four sets of
premolars and three sets of ‘grinding’ molars. The modern horse, due to its
grazing orientation, has six sets of ‘grinding’ molars.
Having evolved in North America Eohippus soon spread across the
existing land-bridges into Eurasia. At the end of the early Eocene
continental drift led to the connection between North America and Europe
being severed while that between North America and Asia was
periodically cut off by shallow seas. Although Eohippus diversified into a
number of different species in Europe only a few of them survived into the
middle Oligocene (ca. 30 million years ago) when they finally died out.
Those species in Asia died out shortly before their counterparts in Europe.
In North America, however, the evolution of the horse continued apace.
Within the Eocene the immediate descendants of Eohippus were
Orohippus and Epihippus. Both equids retained the same foot structure as
their predecessor and were of similar size. Their teeth, however, had
developed further. Orohippus now had four sets of ‘grinding’ cheek teeth
while Epihippus had five.
The beginning of the Oligocene (38 – 26 million years ago) saw the
appearance of the larger Mesohippus and Miohippus some specimens of
which were up to 30 inches (75 cm) tall and weighed about 55 kg (121
lbs). Both species were three-toed on all limbs although most of their
body-weight rested on the middle toe. Their feet still retained the same
‘dog-like’ pad behind the toes. Both equids also sported larger incisors and
had the recognisably ‘horse-like’ six sets of ‘grinding’ molars.
Although different species of Mesohippus and Miohippus coexisted at
various times during the mid to late Oligocene it seems to have been from
Miohippus that the later Miocene equids (ca. 26 - 7 million years ago)
descended. There appears to have been two main lines of descent.

1. Three-toed browsers called Anchitheres. These retained the same

dental and foot structure of Miohippus but grew to much larger sizes. Their
body weights are estimated to have ranged between 200 to 400 kg (441 to
882 lbs). This branch was very successful spreading to all corners of North
America and across the, by then re-established, land-bridges into Eurasia.
They became extinct in both places, however, by the late Miocene / early
Pliocene (ca. 9 - 7 million years ago).

2. A line of equids, which began to specialise in eating grasses. These

equids developed teeth more suitable for chewing grass (small crests on
their teeth enlarged and joined together into a series of ridges for grinding).
They also began to develop into specialised runners. The fleshy pads
10 Chapter One

behind the toes were lost and during normal movement on hard ground
their entire body-weight was supported by the middle toe. The side toes
were only used on soft ground and at maximum speeds.

Parahippus arose from this second branch early in the Miocene and
seems to have been a transitional form between browsers and grazers. It
was larger than Miohippus and still three-toed but was beginning to
develop springy ligaments under the foot. By about seventeen million
years ago, however, Parahippus had greatly declined in numbers and
another equid Merychippus began to take centre-stage. Merychippus was
still three-toed but was fully spring-footed and stood on its tiptoes with
most of its weight borne on the middle toe. The side toes now began to be
of varying sizes (some were fully formed while others barely touched the
ground) while the nail of the central toe began to develop into a hoof.
By the end of the Miocene there were at least six distinct groups of
equids existing in the Americas. The two main groups were the three-toed
grazers called Hipparions and a line of equids in which the side toes
gradually began to reduce in size and disappear. Hipparions dominated the
fauna of North America for millions of years and also spread throughout
Eurasia and into Africa. They seem to have been the first equids to
penetrate the latter continent. They survived in Eurasia and Africa into the
Pleistocene (3 million years ago - present) when they died out.
Back in North America close ancestors of today’s horse were
beginning to emerge (ca. 17 million years ago) from the line of equids in
which the side toes began to disappear. While the earliest of these equines
were all three-toed the side toes were eventually lost in some branches by
the late Miocene. This condition of having one digit per foot
(monodactyly) seems to have evolved only twice in equids - in Pliohippus
and in the branch of equids that included Dinohippus and Equus. Due to
dental similarities it was originally thought that Equus may have
descended from Pliohippus, however, more recent analysis of the
development of facial fossae has shown that Dinohippus is a more likely
candidate. Dinohippus was the most common equid in North America in
the late Pliocene.
Equus finally developed in North America about four million years
ago. It stood approximately 13.2 hands (54 inches) at the shoulder and
possessed a recognisably ‘horse-like’ body with a rigid spine, long neck,
legs, and nose. The earliest known specimens had zebra-like bodies with
short, narrow, donkey-like skulls. Equus was soon to evolve into many
new and varied species.
During the first major glaciations of the late Pliocene (ca. 3 million
years ago) certain equus species spread across the world and, up until a
 Evolution of the Horse 11

million years ago, were to be found in Africa, Asia, Europe, and North and
South America. The Pleistocene (3 million years ago - present) saw a
number of extinctions that killed off most of the large mammals in North
and South America including all equids.4 It is not known why these
extinctions took place. They may have been due to climate changes in the
post-glacial period, over-hunting by humans (who had just then arrived in
the Americas), sickness, or a combination of all three. Budiansky believes
that equus would also have eventually died out in Eurasia had it not been
for its domestication by humankind.

As the climate warmed and forests overtook the open grasslands, the herds
fled ever eastward, vanishing from the British Isles and from France and
Spain … until all that remained were remnant herds in the still-open
grassland steppes of Ukraine and Central Asia … But for domestication,
the Eurasian wild horse would likely have shared the fate of its New World
cousins. This rescue of the horse follows a pattern that has been repeated
time and again in domestic species. The aurochs, the wild progenitor of the
domestic cow, is gone: wild sheep and goats teeter on the brink of
extinction throughout the world: the last remnant of the Asian wild horse,
Przewalskii’s horse, survives only in captive and artificially managed
populations rescued from oblivion at the end of the last century.5

By the time the ice caps had retreated some ten thousand years ago
only eight species of equus had survived:

1. Horses (Equus Caballus) in Europe and Asia.

2. The onager (Equus Hemionius) in the Near East.
3. The Tibetan wild ass (Equus Kiang) – now extinct.
4. The African wild ass (Equus Asinus) in North Africa.
5. The Mountain zebra (Equus Zebra) in South Africa.
6. The Plains zebra / Burchell’s zebra (Equus Burchelli) in South Africa.
7. Grevy’s zebra (Equus Grevyi) in Kenya / Ethiopia.
8. The quagga (Equus Quagga) in South Africa – now extinct.

4
Horses were to become unknown in the Americas until their re-introduction by
Europeans in the late fifteenth century A.D.
5
Budiansky, (1997), 35.
 CHAPTER TWO

DOMESTICATION

Of the eight species of equus in existence after the last ice age it was
Equus Caballus that was to have the greatest impact on human history. In
the post-glacial period it appears that a number of distinct breeds of Equus
Caballus roamed a wide range of territory across northern Europe and
Asia.

Because of the wide variation in size and conformation of modern

domestic horses, with small stocky ponies in northern Europe and slender-
limbed Arabian horses in the south, it has often been suggested that at the
end of the Pleistocene there were several different species of wild equids
that were domesticated.1

It is generally believed that Equus Caballus had developed into three

distinct breeds by the post-glacial period.

1. The Takhi or Asiatic Wild Horse (Equus Caballus Przewalskii).2 The

Przewalskii was the last truly wild horse in existence. Its range was
generally located east of the 40th degree of longitude across the Russian
and Mongolian steppes.3 It was last seen free in 1969 and died out in the
wild sometime after this. It continued to be bred in captivity, however, and
in the early 1990s was successfully reintroduced into the national parks of
Mongolia. The Przewalskii is a pony-sized horse of about 13 hands with a
large head, short and fleshy neck, and narrow legs. Its general body colour
is a light bay with the lower part a smooth white. It has dark stripes on its
chest, its mane and tail are black or dark brown (the mane is stiff and
upright without a forelock), and the lower parts of its legs have dark
stripes. There is also a dark dorsal stripe visible running from mane to tail.

1
Here the author uses ‘species’ to denote various ‘breeds’ of horse rather than an
actual separate species like the ass or onager. [Clutton-Brock, (1981), 82].
2
Named after the famous Russian geographer and explorer Nikolaj Michaljovitch
Przewalskii who rediscovered this wild horse while travelling in Mongolia in 1879.
3
Hilzheimer, (1935), 136.
 Domestication 13

2. The Tarpan (Equus Caballus Gmelini). The Tarpan ranged across

northern Europe and the western steppe as far east as the 40th degree of
longitude. It is not known for certain how far to the west its range
extended.4 It existed in a wild state until the mid 1800s when it became
extinct due to over-hunting. As their natural forest and steppe habitat was
destroyed by the encroaching human population Tarpans came into
increasing conflict with farmers and were often shot in order to prevent
them eating crops and enticing away domesticated mares. The last true
Tarpan died out in a Russian game preserve at Askania Nova in 1876. It
has since been genetically recreated from domesticated descendants and
this recreated breed is sometimes referred to as the Polish Primitive Horse.
The Tarpan is mousy dun / light grey in colour with face and legs being
darker than the body. Its mane and tail are flaxen but dark in the centre
where the dorsal stripe passes through.5 It stands at about 13 – 13.2 hands
and, like the Przewalskii, has a large head and thick neck.

3. A heavy ‘forest’ horse from northwest / west Europe (Equus

Caballus Silvaticus) that perhaps was the ancestor of today’s ‘heavy’
breeds of horse.

Chenevix-Trench mentions a possible fourth breed, that of a white

horse, inhabiting the northern tundra of Eurasia. It was contemporary with
the mammoth (carcases have been found together in the same ice) and
native hunters spoke of sightings as late as 1926.6 Possibly these were
similar to the wild white horses Herodotus described as grazing in the
region of the Hypanis river.7 Chenevix-Trench believes, however, that this
breed of white horse, along with that of the European ‘forest’ horse, may
have been a variety of the more familiar Przewalskii or Tarpan rather than
separate breeds.8 Bokonyi suggests that Herodotus’ wild white horses were
Tarpans in their winter coats.

In the autumn, [the Tarpan] grew quantities of white downy hair and in
winter its body was nearly grey, although the head, legs, mane and tail
remained dark. In this way the animal’s colour shaded into the snow-
covered landscape. Owing to this quality, the tarpan may be identified as
the wild white horse portrayed by Herodotus (IV: 52) as living beside the

4
ibid 135 - 136.
5
Originally the wild Tarpan would have had a stiff upright mane like the
Przewalskii. This trait, however, does not often pass on to its genetically recreated
descendants.
6
Chenevix-Trench, (1970), 8.
7
Hdt. 4.52.
8
Chenevix-Trench, (1970), 8.
 Another Random Document on
Scribd Without Any Related Topics
 Subclass I. Ornithodelphia or Prototheria.
This subclass contains only a single order, the Monotremata, and
the following characteristics are equally applicable to the subclass
and to the order. The vertebral centra have no epiphyses, and the
odontoid process remains for a long time free from the centrum of
the second vertebra. With the exception of the atlas of Echidna the
cervical vertebrae are without zygapophyses. The cranial walls are
smooth and rounded, and the sutures between the several bones
early become completely obliterated as in birds. The mandible is a
very slight structure, with no ascending ramus, and with the
coronoid process (see p. 398) and angle rudimentary. The auditory
ossicles show a low state of development. The tubercula of the ribs
articulate with the sides of the centra of the thoracic vertebrae, not
with the transverse processes. Some of the cervical ribs remain for a
long time separate from the vertebrae. Well ossified sternal ribs
occur. No true teeth are present in the adult. The young
Ornithorhynchus has functional molar teeth, but in the adult their
place is taken by horny plates. In the Echidnidae neither teeth nor
horny plates occur.
The coracoid (fig. 66, 3) is complete and well developed, and
articulates with the sternum. A precoracoid (epicoracoid) occurs in
front of the coracoid, and there is a large interclavicle (fig. 66, 6).
The ridge on the scapula, corresponding to the spine of other
mammals, is situated on the anterior border instead of in the middle
of the outer surface. Epipubic bones are present. In the Echidnidae,
but not in Ornithorhynchus[112], the central portion of the
acetabulum is unossified as in birds. The humerus has a prominent
deltoid crest; its ends are much expanded, and the distal end is
pierced by an ent-epicondylar foramen. The fibula has a broad
proximal process resembling an olecranon. The limbs and their
girdles bear a striking resemblance to those of some Theromorphous
reptiles.
 Fig. 66. Ventral view of the shoulder-girdle and sternum of a Duckbill
(Ornithorhynchus paradoxus) × ¾ (after Parker).
1 and 2. scapula. 8. presternum.
 3. coracoid. 9. third segment of
4. precoracoid (epicoracoid). mesosternum.
5. glenoid cavity. 10. sternal rib.
6. interclavicle. 11. intermediate rib.
7. clavicle. 12. vertebral rib.

The order Monotremata includes only two living families, the

Echidnidae and Ornithorhynchidae.

Mesozoic Mammalia[113].
It will be well here to briefly refer to certain mammals of small
size, the remains of which have been found in deposits of Mesozoic
age. In the great majority of cases they are known only by the lower
jaw, or sometimes only by isolated teeth. A large number of them
are commonly grouped together as the Multituberculata, and are
sometimes, partly owing to the resemblance of their teeth to those
of Ornithorhynchus, placed with the Prototheria, sometimes between
the Prototheria and the Metatheria. They are characterised by having
a single pair of large incisors in the lower jaw, and one large with
one or two smaller incisors in each premaxillae. The lower canines
are very small or altogether wanting. The incisors are separated by a
diastema from the grinding teeth, which are sometimes (Tritylodon)
characterised by the possession of longitudinal rows of little
tubercles separated by grooves, sometimes by having the premolars
provided with high cutting edges, whose surfaces are obliquely
grooved. Some of the Mesozoic mammals found associated with the
Multituberculata, have however a dentition of an altogether different
type, with at least three lower incisors, well developed canines and
premolars, and numerous molars with peculiar three-cusped or
tritubercular grinding surfaces. These mammals, one of the best
known of which is Phascolotherium, are commonly separated from
the Multituberculata, and are divided by Osborn into two groups,
one allied to the Marsupials, and one to the Insectivores. The group
showing Marsupial affinities is further subdivided into carnivorous,
omnivorous, and herbivorous subgroups. The members of both
groups commonly have four premolars, and six to eight molars in
each mandibular ramus.
Subclass II. Didelphia or Metatheria.
This subclass, like the previous one, contains only a single order,
viz. the Marsupialia[114]; but the forms referable to it are far more
numerous than in the case of the Monotremata.
The integument is always furry, and the teeth are always
differentiated into incisors, canines, premolars and molars. Except in
Phascolomys, the number of incisors in the upper and lower jaws is
never equal, and the number in the upper usually exceeds that in
the lower jaw. There is no such regular succession and displacement
of teeth as in most mammals. Sometimes the anterior teeth are
diphyodont, and as a general rule the tooth commonly regarded as
the last premolar is preceded by a milk tooth. The majority of the
permanent teeth of most Marsupials are regarded as belonging to
the milk series for two reasons, (1) they are developed from the
more superficial tissues of the jaws, (2) a second set, the permanent
teeth, begin to develop as outgrowths from them, but afterwards
become aborted[115].
The odontoid process at an early stage becomes fused with the
centrum of the second cervical vertebra, and the number of thoraco-
lumbar vertebrae is always nineteen. The skull has several
characteristic features. The tympanic bone remains permanently
distinct, and the anterior boundary of the tympanic cavity is formed
by the alisphenoid. The carotid canal perforates the basisphenoid,
and the lachrymal canal opens either outside the orbit or at its
margin. There are generally large vacuities in the palate. The angle
of the mandible is (except in Tarsipes) more or less inflected; and as
a rule the jugal furnishes part of the articular surface for the
mandible. There is no precoracoid (epicoracoid) or interclavicle, and
the coracoid is reduced to form a mere process of the scapula, not
coming near the sternum.
 Epipubic, or so-called marsupial bones[116], nearly always occur,
and a fourth pelvic element, the acetabular bone, is frequently
developed. The fibula is always complete at its distal end, sometimes
it is fused with the tibia, but often it is not only free but is capable of
a rotatory movement on the tibia. This is the case in the families
Phascolomyidae, Didelphyidae, and Phalangeridae.
The Marsupialia can be subdivided into two main groups,
according to the character of the teeth:—
1. Polyprotodontia.
In this group the incisors are small, subequal and numerous, not
less than 4/3. The canines are larger than the incisors, and the
molars have sharp cusps. The members of this group are all more or
less carnivorous or insectivorous. The group includes the families
Didelphyidae, Dasyuridae, Peramelidae, and Notoryctidae[117].
2. Diprotodontia.
In this group the incisors do not exceed 3/3, and are usually 3/1,
occasionally 1/1. The first upper and lower incisors are large and
cutting. The lower canines are always small or absent, and so in
most cases are the upper canines. The molars have bluntly
tuberculated, or transversely ridged crowns. The group includes the
families Phascolomyidae, Phalangeridae, Macropodidae, and
Epanorthidae.
Subclass III. Monodelphia or Eutheria.
This great group includes all the Mammalia except the orders
Monotremata and Marsupialia. Coming to their general
characteristics—as in the Didelphia the odontoid process and cervical
ribs early become fused with the centra which bear them, while the
coracoid is reduced so as to form a mere process on the scapula,
and there is no precoracoid (epicoracoid), such as is found in
Ornithodelphia. Clavicles may be present or absent; when fully
developed they articulate with the sternum, usually directly, but
occasionally, as in some Rodents and Insectivores, through the
remains of the sternal end of the precoracoid. There is never any
interclavicle in the adult, though sometimes traces of it occur during
development. In the pelvis the acetabula are imperforate; and well-
developed epipubic bones are never found in the adult, though
traces of them occur in some Carnivores and foetal Ungulates.

Order 1. Edentata[118].
Teeth are not, as the name of the order seems to imply, always
wanting; and sometimes they are very numerous. They are,
however, always imperfect, and, with very few exceptions, are
homodont and monophyodont. They have persistent pulps, and so
grow indefinitely and are never rooted. In all living forms they are
without enamel, consisting merely of dentine and cement, and are
never found in the front part of the mouth in the situation occupied
by the incisors of other mammals. These characters derived from the
teeth are the only ones common to the various members of the
order, which includes the living sloths, ant-eaters, armadillos,
pangolins and aard varks, together with various extinct forms, chiefly
found in beds of late tertiary age in both North and South America,
the best known being the Megatheridae and Glyptodonts.

Order 2. Sirenia[119].
The skeleton of these animals has a general fish-like form, in
correlation with their purely aquatic habits. The fore limbs have the
form of paddles, but the number of phalanges is not increased
beyond the normal. There are no external traces of hind limbs.
The whole skeleton and especially the skull and ribs is remarkably
massive and heavy. The dentition varies; in the two living genera
Manatus and Halicore, incisor and molar teeth are present, in one
extinct genus, Rhytina, teeth are entirely absent, while in another,
Halitherium, the dentition is more decidedly heterodont than in living
forms. In the two living genera the dentition is monophyodont, but
in Halitherium the anterior grinding teeth are preceded by milk
teeth. The tongue and anterior part of the palate and lower jaw are
covered with roughened horny plates. The skull is noticeable for the
size and backward position of the anterior nares, also for the
absence or small size of the nasal bones. There is no union of
certain of the vertebrae to form a sacrum, and in living forms the
centra are not terminated by well-formed epiphyses[120].
The cervical vertebrae are much compressed, but they are never
ankylosed together. In Manatus there are only six cervical vertebrae.
The caudal vertebrae have well-developed chevron bones. The
humerus is distinctly articulated to the radius and ulna, and these
two bones are about equally developed, and are often fused
together. There are no clavicles, and the pelvis is vestigial, consisting
of a pair of somewhat cylindrical bones suspended at some distance
from the vertebral column. In living forms there is no trace of a
posterior limb, but in Halitherium there is a vestigial femur
connected with each half of the pelvis.

Order 3. Cetacea[121].
In these mammals the general form is more fish-like than is the
case even in the Sirenia. The skin is generally almost completely
naked, but hairs are sometimes present in the neighbourhood of the
mouth, especially in the foetus. In some Odontoceti vestiges of
dermal ossicles have been described, and in Zeuglodon the back was
probably protected by dermal plates. The anterior limbs have the
form of flattened paddles, showing no trace of nails, the posterior
limb bones are quite vestigial or absent, and there is never any
external sign of the limb. Teeth are always present at some period of
the life history, but in the whalebone whales they are only present
during foetal life, their place in the adult animal being taken by
horny plates of baleen. In all living forms the teeth are simple and
uniform structures without enamel; they have single roots, and the
alveoli in which they are imbedded are often incompletely separated
from one another. As in some forms traces of a replacing dentition
have been described, it has been concluded that the functional teeth
of Cetacea belong to the milk dentition.
The texture of the bones is spongy. The cervical vertebrae are
very short, and though originally seven in number, are in many
forms completely fused, forming one solid mass (fig. 67). The
odontoid process of the axis is short and blunt, or may be
completely wanting. The lumbar and caudal vertebrae are large and
numerous, and as zygapophyses are absent, are very freely movable
on one another; zygapophyses are also absent from the posterior
thoracic vertebrae. The lumbar vertebrae are sometimes more
numerous than the thoracic. The epiphyses are very distinct, and do
not unite with the centra till the animal is quite adult. None of the
vertebrae are united to form a sacrum, but the caudal vertebrae
have large chevron bones.
 Fig. 67. Cervical vertebrae of a Ca'ing Whale (Globicephalus melas) × ¼. (Camb.
Mus.)
1. centrum of seventh cervical 4. foramen for exit of first spinal
vertebra. nerve.
2. neural arch of seventh 5. transverse process of axis.
cervical vertebra. 6. fused neural spines of atlas
3. transverse process of atlas. and axis.

The skull is peculiarly modified; the bones forming the occipital

segment show a specially strong development, and the cranial cavity
is short, high, and almost spherical. The supra-occipital is very large
and rises up to meet the frontals, thus with the interparietal
completely separating the parietals from one another.
 The frontals are expanded, forming large bony plates, which roof
over the orbits. The zygomatic process of the squamosal is
extremely large and extends forwards to meet the supra-orbital
process of the frontal; the zygomatic process of the jugal is on the
contrary very slender. The face is drawn out into a long rostrum,
formed of the maxillae and premaxillae surrounding the vomer and
the mesethmoid cartilage. The maxillae are specially large, and
extend backwards so as to partially overlap the frontals. The nasals
are always small, and the anterior nares open upwards between the
cranium and rostrum. The periotics are loosely connected with the
other bones of the skull and the tympanics are commonly large and
dense. The mandible has hardly any coronoid process, and the
condyles are at its posterior end.
There are no clavicles, but the scapula and humerus are well
developed. The humerus moves freely in the glenoid cavity, but all
the other articulations of the anterior limb are imperfect; the various
bones have flattened ends, and are connected with one another by
fibrous tissue, which allows of hardly any movement. Frequently the
carpus is imperfectly ossified.
The number of digits in the manus is generally five, sometimes
four, and when there are four digits it is the third and not the first
that is suppressed. The number of phalanges in the second and third
digits almost always exceeds that which is normal in mammals, and
the phalanges are also remarkable for having epiphyses at both
ends. The pelvis is represented by two small bones which lie
suspended horizontally at some distance below the vertebral
column; in some cases vestiges of the skeleton of the hind limb are
attached to them.
The Cetacea are divided into three suborders.
Suborder (1). Archaeoceti.
The members of this group are extinct; they differ from all living
Cetacea in having the dentition heterodont and in the fact that the
back was probably protected by dermal plates. The skull is elongated
and depressed, and the brain cavity is very small. The temporal
fossae are large, and there is a strong sagittal crest. The nasals and
premaxillae are a good deal larger than they are in living Cetacea,
and the anterior nares are usually far forward. The cervical vertebrae
are not fused with one another, and the lumbar vertebrae are
unusually elongated.
The limbs are very imperfectly known, but while the humerus is
much longer than in modern Cetaceans, it is nevertheless flattened
distally, indicating that the limb was paddle-like, and that there was
scarcely any free movement between the fore-arm and upper arm.
The best known genus is Zeuglodon, which is found in beds of
Eocene age in various parts of Europe, and in Alabama.
Suborder (2). Mystacoceti or Balaenoidea.
These are the Whalebone Whales or True Whales.
Calcified teeth representing the milk dentition occur in the foetus,
but the teeth are never functional, and always disappear before the
close of foetal life. There is a definite though small olfactory fossa.
The palate is provided with plates of baleen or whalebone. The skull
is symmetrical, and is extremely large in proportion to the body. The
nasals are moderately well developed, and the maxillae do not
overlap the orbital processes of the frontals. The lachrymals are
small and distinct from the jugals. The tympanics are ankylosed to
the periotics, and the rami of the mandible do not meet in a true
symphysis. The ribs articulate only with the transverse processes,
and the capitula are absent or imperfectly developed. Only one pair
of ribs meets the sternum, which is composed of a single piece.
The group includes among others the Right whale (Balaena),
Humpbacked whale (Megaptera), and Rorqual (Balaenoptera).
Suborder (3). Odontoceti.
 Teeth always exist after birth and baleen is never present. The
teeth are generally numerous, but are sometimes few and
deciduous; the dentition is homodont (except in Squalodon). The
dorsal surface of the skull is somewhat asymmetrical, there is no
trace of an olfactory fossa, the nasals are quite rudimentary, and the
hind ends of the maxillae cover part of the frontals; in all these
respects the skull differs from that of the Mystacoceti. The lachrymal
may either be united to the jugal or may be large and distinct. The
tympanic is not ankylosed to the periotic. The rami of the mandible
are nearly straight and become united in a long symphysis. Some of
the ribs have well developed capitula articulating with the vertebral
centra. The sternum is almost always composed of several pieces as
in other mammals, and several pairs of ribs are connected with it.
There are always five digits to the manus, though the first and fifth
are usually very little developed.
The suborder includes the Sperm Whale (Physeter), Narwhal
(Monodon), Dolphin (Delphinus), Porpoise (Phocoena), and many
other living forms as well as the extinct Squalodon which differs
from the other members of the suborder in its heterodont dentition.
Order 4. Ungulata.
This order includes a great and somewhat heterogeneous group of
animals, a large proportion of which are extinct. They all (except
certain extinct forms) agree in having the ends of the digits either
encased in hoofs or provided with broad flat nails. The teeth are
markedly heterodont and diphyodont, and the molars have broad
crowns with tuberculated or ridged surfaces. Clavicles are never
present in the adult except in a few generalised extinct forms such
as Typotherium, and it is only recently that vestigial clavicles have
been discovered in the embryo[122]. The scaphoid and lunar are
always distinct.
The order Ungulata may be subdivided into two main groups,
Ungulata vera and Subungulata.
 Section I. Ungulata vera
[123].

The cervical vertebrae except the atlas are generally

opisthocoelous. The feet are never plantigrade[124]. In all the living
and the great majority of the extinct forms the digits do not exceed
four, the first being suppressed. In the carpus the os magnum
articulates freely with the scaphoid, and is separated from the
cuneiform by the lunar and unciform. In the tarsus the cuboid
articulates with the astragalus as well as with the calcaneum, and
the proximal surface of the astragalus is marked by a pulley-like
groove. All the bones of the carpus and tarsus strongly interlock.
These characters with regard to the carpus and tarsus do not hold in
Macrauchenia and its allies. The humerus never has an ent-
epicondylar foramen.
The group is divided into two very distinct suborders:—
Suborder (1). Artiodactyla.
The Artiodactyla have a number of well marked characters, one of
the most obvious being the fact that many of the most characteristic
forms have large paired outgrowths on the frontal bones. These may
be (1) solid deciduous bony antlers, or (2) more or less hollow bony
outgrowths which are sheathed with permanently growing horn.
The premolar and molar teeth are usually dissimilar, the premolars
being one-lobed and the molars two-lobed; the last lower molar of
both the milk and permanent dentitions is almost always three-
lobed.
The grinding surfaces of the molar teeth have a tendency to
assume one of two forms. In the Pigs and their allies the crowns are
bunodont[1], while in the more highly specialised Ruminants the
crowns are selenodont[125]. The nasals are not expanded posteriorly,
and there is no alisphenoid canal[126]. The thoraco-lumbar vertebrae
are always nineteen. The symphysis of the ischia and pubes is very
elongated, and the femur has no third trochanter. The limbs never
have more than four digits, and are symmetrical about a line drawn
between the third and fourth digits; the digits, on the other hand,
are never symmetrical in themselves. The astragalus has pulley-like
surfaces both proximally and distally, and articulates with the
navicular and cuboid by two nearly equal facets. The calcaneum
articulates with the lower end of the fibula when that bone is fully
developed.
In the Artiodactyla are included the following living groups:—
a. Suina. Pigs and Hippopotami.
b. Tylopoda. Camels and Llamas.
c. Tragulina. Chevrotains.
d. Ruminantia or Pecora. Deer, giraffes, oxen, sheep and
antelopes.

Suborder (2). Perissodactyla[127].

In this group there are never any bony outgrowths from the
frontals. The grinding teeth form a continuous series, the posterior
premolars resembling the molars in complexity, and the last lower
molar generally has no third lobe. The cervical vertebrae with the
exception of the atlas almost always have markedly opisthocoelous
centra, but in Macrauchenia they are flat. The nasals are expanded
posteriorly, and an alisphenoid canal is present. The thoraco-lumbar
vertebrae are never less than twenty-two in number and are usually
twenty-three. The femur has a third trochanter (except in
Chalicotherium). The third digit of the manus and pes is symmetrical
in itself, and larger than the others, and in some cases the other
digits are quite vestigial. The number of the digits of the pes is
always odd. The astragalus is abruptly truncated distally, and the
facet by which it articulates with the cuboid, is much smaller than
that by which it articulates with the navicular. The calcaneum does
not articulate with the fibula, except in Macrauchenia. The group
includes many extinct forms, and the living families of the Tapirs,
Horses and Asses, and Rhinoceroses.
Section II. Subungulata.
In this group is placed a heterogeneous collection of animals, the
great majority of which are extinct. There is really no characteristic
which is common to them all, and which serves to distinguish them
as a group from the Ungulata vera. But the most distinctive
character common to the greatest number of them is to be found in
the carpus, whose bones in most cases retain their primitive relation
to one another, the os magnum articulating with the lunar and
sometimes just meeting the cuneiform, but in living forms at any
rate not articulating with the scaphoid. The feet frequently have five
functional digits, and may be plantigrade. The proximal surface of
the astragalus is generally flattened instead of being pulley-like as in
Ungulata vera.
Suborder (1). Toxodontia.
This suborder includes some very aberrant extinct South American
ungulates, which have characters recalling the Proboscidea, both
groups of Ungulata vera, and the Rodentia. The limbs are
subplantigrade or digitigrade, and the digits are three, rarely five, in
number, the third being most developed. The carpus resembles that
of the Ungulata vera, in that the bones interlock and the magnum
articulates with the scaphoid. In the tarsus, however, the bones do
not interlock. The astragalus has a pulley-like proximal surface
(except in Astrapotherium, in which it is flat), and articulates only
with the navicular, not meeting the cuboid. The calcaneum has a
large facet for articulation with the fibula, as in Artiodactyla. There is
no alisphenoid canal, and the orbit is confluent with the temporal
fossa. Some of the forms (e.g. Nesodon) referred to this group have
the typical mammalian series of forty-four teeth, but in others the
canines are undeveloped. In Toxodon all the cheek teeth have
persistent pulps, while in Nesodon and Astrapotherium they are
rooted. A clavicle is sometimes present (Typotherium), and the
femur sometimes has a third trochanter (Typotherium and
Astrapotherium), sometimes is without one (Toxodon).
The remains of these curious Ungulates have been found in beds
of late Tertiary age in South America.

Suborder (2). Condylarthra[128].

This group includes some comparatively small extinct ungulates,
which are best known from the Lower Eocene of Wyoming, though
their remains have also been found in deposits of similar age in
France and Switzerland. Their characters are little specialised, and
they show relationship on the one hand to the Ungulata vera and on
the other to the Hyracoidea. They also have characters allying them
to the Carnivora. They generally have the typical mammalian series
of forty-four teeth, the molars being brachydont and generally
bunodont. The premolars are more simple than the molars. The
limbs are plantigrade, and have five digits with rather pointed ungual
phalanges. The os magnum, as in living Subungulates, articulates
with the lunar, not reaching the scaphoid. The astragalus has an
elongated neck, a pulley-like proximal and a convex distal articular
surface, and does not articulate with the cuboid. The humerus has
an ent-epicondylar foramen, and the femur has a third trochanter.
The best known genus is Phenacodus; it is perhaps the most
primitive ungulate whose skeleton is thoroughly well known, and is
of special interest from the fact that it is regarded as the lowest
stage in the evolutionary series of the horse. Its remains are found
in the Lower Eocene of Wyoming.

Suborder (3). Hyracoidea[129].

This group of animals is very isolated, having no very close allies,
either living or extinct. The digits are provided with flat nails, except
the second digit of the pes, which is clawed. Canine teeth are
absent, and the dental formula is usually given as i ½, c 0/0, pm 4/4,
m 3/3. The upper incisors are long and curved, and have persistent
pulps as in Rodents; their terminations are, however, pointed, not
chisel-shaped, as in Rodents. The lower incisors have pectinated
edges. The grinding teeth have a pattern much like that in
Rhinoceros. In the skull (fig. 83) the postorbital processes of the
frontal and jugal almost or quite meet. The jugal forms part of the
glenoid cavity for articulation with the mandible, and also extends
forwards so as to meet the lachrymal. There is an alisphenoid canal.
There are as many as twenty-one or twenty-two thoracic vertebrae,
and the number of thoraco-lumbar vertebrae reaches twenty-eight
or thirty. There are no clavicles, and the scapula has no acromion;
the coracoid process is, however, well developed. The ulna is
complete. In the manus the second, third and fourth digits are
approximately equal in size, the fifth is smaller, and the first is
vestigial. The femur has a slight ridge representing the third
trochanter. The fibula is complete, but is generally fused with the
tibia proximally. There is a complicated articulation between the tibia
and astragalus, which has a pulley-like proximal surface. In the pes
the three middle digits are well developed, but there is no trace of a
hallux, and the fifth digit is represented only by a vestigial
metatarsal.
The only representatives of the suborder are some small animals
belonging to the genus Procavia (Hyrax), which is found in Africa
and Syria; some of the species are by many authors placed in a
distinct genus Dendrohyrax.

Suborder (4). Amblypoda[130].

This suborder includes a number of primitive extinct Ungulates,
many of which are of great size. Their most distinguishing
characteristics are afforded by the extremities. In the carpus the
bones interlock a little more than is the case in most Subungulata,
and the corner of the os magnum reaches the scaphoid, while the
lunar articulates partially with both magnum and unciform, instead
of only with the magnum. In the tarsus the cuboid articulates with
both the calcaneum and the astragalus, which is remarkably flat.
The manus and pes are short, nearly or quite plantigrade, and have
the full number of digits. The cranial cavity is singularly small.
Canine teeth are present in both jaws, and the grinding teeth have
short crowns, marked by V-shaped ridges. The pelvis is large, the
ilia are placed vertically, and the ischia do not take part in the ventral
symphysis.
The best known animals belonging to this suborder are the
Uintatheriidae (Dinocerata)[131], found in the Upper Eocene of
Wyoming. They are as large as elephants, and are characterised by
the long narrow skull drawn out into three pairs of rounded
protuberances, by the strong occipital crest, and by the very large
upper canines.
Suborder (5). Proboscidea.
This suborder includes the largest of land mammals, the
Elephants, and certain of their extinct allies. The limbs are strong,
and are vertically placed; the proximal segment is the longest, and
the manus and pes are pentedactylate and subplantigrade. The
digits are all enclosed in a common integument, and each is
provided with a broad hoof. The vertebral centra are much flattened
and compressed, especially in the cervical region. The number of
thoracic vertebrae is very great, reaching twenty. The skull (figs. 96
and 97) is extremely large, this being due to the great development
of air cells, which takes place in nearly all the bones of the adult
skull. In the young skull there are hardly any air cells, and the
growth of the cranial cavity does not by any means keep pace with
the growth of the skull in general. The supra-occipital is very large,
and forms a considerable part of the roof of the skull. The nasals
and jugals are short, and the premaxillae very large. The rami of the
mandible meet in a long symphysis, and the ascending portion is
very high. Canine teeth are absent, and the incisors have the form of
ever-growing tusks composed mainly of dentine; in living forms they
are present in the upper jaw only. The grinding teeth are large, and
in living forms have a very complex structure and mode of
succession. In some of the extinct forms, such as Mastodon and
especially Dinotherium, the teeth are much more simple. In every
case the teeth have the same general structure, consisting of a
series of ridges of dentine, coated with enamel. In the more
specialised forms the valleys between the ridges are filled up with
cement. The acromion of the scapula has a recurved process, similar
to that often found in rodents. Clavicles are absent. The radius and
ulna are not ankylosed, but are incapable of any rotatory movement.
All the bones of the extremities are very short and thick; the
scaphoid articulates regularly with the trapezoid and the lunar with
the magnum. The ilia are vertically placed, and are very much
expanded; the ischia and pubes are small, and form a short
symphysis. The femur has no third trochanter, and the tibia and
fibula are distinct. The fibula articulates with the calcaneum, and the
astragalus is very flat.

Here brief reference may be made to the Tillodontia[132], a group

of extinct mammals found in the Eocene beds of both Europe and
North America. They seem to connect together the Ungulata,
Rodentia, and Carnivora.
The skull resembles that of bears, but the grinding teeth are of
Ungulate type, while the second incisors resemble those of rodents,
and have persistent pulps. The femur has a third trochanter, and the
feet resemble those of bears in being plantigrade and having pointed
ungual phalanges, differing, however, in having the scaphoid and
lunar distinct.
Order 5. Rodentia.
The Rodents form a very large and well-defined group of
mammals easily distinguishable by their peculiar dentition. Canines
are absent, and the incisors are very large and curved, growing from
persistent pulps. They are rectangular in section and are much more
thickly coated with enamel on their anterior face than elsewhere;
consequently, as they wear down they acquire and retain a chisel-
shaped (scalpriform) edge. There is never more than one pair of
incisors in the mandible, and except in the Hares and Rabbits, there
is similarly only a single pair in the upper jaw. These animals are,
too, the only rodents which have well developed deciduous incisors.
There is always a long diastema separating the incisors from the
grinding teeth. The grinding teeth, which are arranged in a
continuous series, vary in number from two to six in the upper jaw,
and from two to five in the lower jaw. The number of premolars is
always below the normal, often they are altogether wanting, but
generally they are 1/1. Sometimes the grinding teeth form roots,
sometimes they grow persistently.
The premaxillae are always large, and the orbits always
communicate freely with the temporal fossae. The condyle of the
mandible is elongated from before backwards, and owing to the
absence of a postglenoid process to the squamosal, a backward and
forward motion of the jaw can take place. The zygomatic arch is
complete, but the jugal is short and only forms the middle of it. The
palate is small, being sometimes, as in the hares, narrowed from
before backwards, sometimes as in the mole-rats (Bathyerginae)
narrowed transversely.
The thoraco-lumbar vertebrae are usually nineteen in number.
Clavicles are generally present, and the acromion of the scapula is
commonly very long. The feet are as a rule plantigrade, and
provided with five clawed digits.
There are two main groups of Rodentia; the Duplicidentata, or
Hares and Rabbits, which have two pairs of upper incisors, whose
enamel extends round to the posterior surface; and the
Simplicidentata, in which there is only a single pair of upper incisors,
whose enamel is confined to the anterior surface. This group
includes all the Rodents except the Hares and Rabbits.
Order 6. Carnivora.
 The living Carnivora form a natural and well-marked group, but as
is the case with so many other groups of animals, when their extinct
allies are included, it becomes impossible to readily define them.
The manus and pes never have less than four well-developed
digits, and these are nearly always provided with more or less
pointed nails, generally with definite claws. The hallux and pollex are
never opposable. The dentition is diphyodont and markedly
heterodont. The teeth are always rooted, except in the case of the
canines of the Walrus. The incisors are generally 3/3, and are
comparatively small, while the canines are large, pointed, and
slightly recurved. The cheek teeth are variable, and are generally
more or less compressed and pointed; sometimes their crowns are
flattened and tuberculated, but they are never divided into lobes by
deep infoldings of enamel. The squamosal is drawn out into a
postglenoid process, and the mandible has a large coronoid process.
The condyle of the mandible is transversely elongated, and the
glenoid fossa is very deep; in consequence of this arrangement the
mandible can perform an up and down movement only, any rotatory
or back and fore movement being impossible. The jugal is large, and
the zygomatic arch generally strong, while the orbit and temporal
fossa are in most cases completely confluent. The scapula has a
large spine. The clavicle is never complete and is often absent, this
forming an important distinction between the skeleton of a Carnivore
and of any Insectivore except Potamogale. The humerus often has
an ent-epicondylar foramen, and the radius and ulna, tibia and fibula
are always separate. The manus is often capable of the movements
of pronation and supination, and the scaphoid, lunar and centrale
are in living forms always united together.
The order Carnivora includes three suborders.

Suborder (1). Creodonta[133].

This suborder contains a number of extinct Carnivora, which
present very generalised characters.
 The cranial cavity is very small; and the fourth upper premolar and
first lower molar are not differentiated as carnassial teeth[134], as
they are in modern Carnivora. The Creodonta also differ from
modern Carnivora in the fact that the scaphoid and lunar are usually
separate, and that the femur has a third trochanter. The feet are
plantigrade.
They resemble the Condylarthra, another very generalised group,
in having an ent-epicondylar foramen.
They occurred throughout the Tertiary period in both Europe and
North America, and have also been found in India. One of the best
known genera is Hyaenodon.
Suborder (2). Carnivora vera or Fissipedia.
The skeleton is mainly adapted for a terrestrial mode of life, and
the hind limbs have the normal mammalian position. In almost every
case the number of incisors is 3/3. Each jaw always has one specially
modified carnassial or sectorial tooth which bites like a scissors blade
against a corresponding tooth in the other jaw. In front of it the
teeth are always more or less pointed, while behind it they are more
or less broadened and tuberculated. In the manus the first digit, and
in the pes the first and fifth digits are never longer than the rest,
and the digits of both limbs are almost invariably clawed. Some
forms are plantigrade, some digitigrade, some subplantigrade. The
group includes all the ordinary terrestrial Carnivora, and is divided
into three sections:—

Æluroidea[135], including the cats, civets, hyaenas, and allied

forms.
Cynoidea, including the dog tribe.
Arctoidea, including the bears, raccoons, weasels, and allied
forms.

Suborder (3). Pinnipedia[136].

 In this suborder the limbs are greatly modified and adapted for a
more or less purely aquatic life, the proximal and middle segments
of the limbs are shortened, while the distal segment, especially in
the leg, is much elongated and expanded. There are always five
well-developed digits to each limb, and in the pes the first and fifth
digits are generally larger than the others. The digits generally bear
straight nails instead of claws, but even nails are sometimes absent.
There is no carnassial tooth, and the teeth in other ways differ
considerably from those of Carnivora vera. The incisors are always
fewer than 3/3; while the cheek teeth generally consist of four
premolars and one molar, all of very uniform character, being
compressed with conical crowns, and never more than two roots.
The suborder includes three families—Otariidae (Eared Seals),
Trichechidae (Walrus), and Phocidae (Seals).

Order 7. Insectivora[137].
This order contains a large number of small generally terrestrial
mammals. The limbs are plantigrade or subplantigrade, and are
generally pentedactylate. All the digits are armed with claws, and
the pollex and hallux are not opposable. The teeth are diphyodont,
heterodont, and rooted. The cheek teeth have tuberculated crowns,
and there are never less than two pairs of incisors in the mandible;
often the incisors, canines, and premolars are not clearly
differentiated from one another, and special carnassial teeth are
never found. The cranial cavity is small, and the facial part of the
skull is generally much developed; often the zygomatic arch is
incomplete. Clavicles are well developed (except in Potamogale), and
the humerus generally has an ent-epicondylar foramen. The femur
frequently has a ridge representing the third trochanter. There are
two suborders:
Suborder (1). Dermoptera.
This suborder includes only a very aberrant arboreal genus
Galeopithecus, remarkable for its greatly elongated limb bones, and
peculiar dentition. The incisors of the lower jaw are deeply
pectinated or divided by several vertical fissures, the canines and
outer upper incisors have two roots. Ossified inter centra occur in
the thoraco-lumbar region of the vertebral column.
Suborder (2). Insectivora vera.

This suborder includes all the ordinary Insectivora, such as moles,

shrews and hedgehogs. The upper and lower incisors are conical,
not pectinated.

Order 8. Chiroptera[138].
This order is perhaps the best marked and most easily defined of
all the orders of mammals. The anterior limbs form true wings and
the whole skeleton is modified in relation to flight.
The anterior limbs are vastly larger than the posterior; for all the
bones except the carpals are much elongated, and this applies
specially to the phalanges of all the digits except the pollex.
The pollex is clawed and so is sometimes the second digit; the
other digits of the manus are without nails or claws. The teeth are
divisible into the four usual types and the series never exceeds i 2/3
c 1/1 pm 3/3 m 3/3 × 2, total 38. The milk teeth are quite unlike the
permanent teeth. The orbit is not divided by bone from the temporal
fossa. The vertebral column is short, and in old animals the trunk
vertebrae have a tendency to become partially fused together. The
cervical vertebrae are remarkably wide, and the development of
spinous processes is everywhere slight. The presternum has a
prominent keel for the attachment of the pectoral muscles. The
clavicles are very long and strong, and the scapula has a long spine
and coracoid process. The ulna is vestigial, consisting only of a
proximal end ankylosed to the radius. All the carpals of the proximal
row—the scaphoid, lunar and cuneiform—are united, forming a
single bone. The pelvis is very weak and narrow, and only in the
Rhinolophidae do the pubes meet in a symphysis. The anterior
caudal vertebrae are frequently united to the ischia. The fibula is
generally vestigial, and the knee joint is directed backwards instead
of forwards. The pes has five slender clawed digits, and the
calcaneum is often drawn out into a spur which helps to support the
membrane connecting the hind limbs with the tail.
There are two suborders of Chiroptera:
1. The Megachiroptera or Flying foxes, which almost always have
smooth crowns to the molar teeth, and the second digit of the
manus clawed.
2. The Microchiroptera including all the ordinary bats which have
cusped molar teeth, and the second digit of the manus clawless.
Order 9. Primates.
The dentition is diphyodont and heterodont, the incisors generally
number 2/2, and the molars, except in the Hapalidae (Marmosets),
are 3/3. The cheek teeth are adapted for grinding, and the molars
are more complex than the premolars. A process from the jugal
meets the postorbital process of the frontal completing the
postorbital bar.
The clavicle is well developed, and the radius and ulna are never
united. The scaphoid and lunar of the carpus, and commonly also
the centrale, remain distinct from one another. As a rule both manus
and pes have five digits, but the pollex may be vestigial. The pollex
is opposable to the other digits, and so is the hallux except in Man;
the digits are almost always provided with flat nails. The humerus
has no ent-epicondylar foramen and the femur has no third
trochanter.
The order Primates is divisible into two suborders:
Suborder (1). Lemuroidea.
 The skull has the orbit communicating freely with the temporal
fossa beneath the postorbital bar (except in Tarsius). The lachrymal
foramen is external to the margin of the orbit. Both pollex and hallux
are well developed. In the pes the second digit is terminated by a
long pointed claw, and so is also the third in Tarsius. The lumbar
region of the vertebral column is long, sometimes including as many
as nine vertebrae. Besides the Lemurs the group includes the
aberrant Tarsius and Chiromys.
Suborder (2). Anthropoidea.
The skull has the orbit almost completely shut off from the
temporal fossa, and the lachrymal foramen is situated within the
orbit. The pollex is sometimes vestigial or absent. The second digit
of the pes has a flattened nail except in the Hapalidae, in which all
the digits of the pes except the hallux are clawed.
The Anthropoidea are divided into five families:
1. Hapalidae or Marmosets.
2. Cebidae or American Monkeys.
3. Cercopithecidae or Old World Monkeys.
4. Simiidae or Anthropoid Apes.
5. Hominidae or Men.
 CHAPTER XXI.
THE SKELETON OF THE DOG[139] (Canis

familiaris).

I. EXOSKELETON.
The exoskeleton of the dog includes three sets of structures: 1.
hairs, 2. claws, 3. teeth. Hairs and claws are epidermal exoskeletal
structures, while teeth are partly of dermal, and partly of epidermal
origin.
1. Hairs are delicate epidermal structures which grow imbedded
in little pits or follicles in the dermis. Specially large hairs forming the
vibrissae or whiskers grow attached to the upper lip.
2. Claws are horny epidermal sheaths, one of which fits on to the
pointed distal phalanx of each digit. They are sharply curved
structures, and being in the dog non-retractile, their points are
commonly much blunted by friction with the ground. The claws of
the pollex, and of the hallux when it is present, however do not
meet the ground, and therefore remain comparatively sharp.

3. Teeth[140]. Although as regards their mode of origin, teeth are

purely exoskeletal or tegumentary structures, they become so
intimately connected with the skull that they appear to belong to the
endoskeleton.
Each tooth, as has been already described, consists of three
distinct tissues, dentine and cement of dermal origin, and enamel of
epidermal origin.
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