UNIT 11

EVOLUTION OF HORSE

Structure____________________________________________________
11.1 Introduction 11.3 Role of Climate in the Evolution of Horse
Expected Learning Outcomes 11.4 Activity
11.2 Evolution of Horse 11.5 Summary
Systematic Palaeontology 11.6 Terminal Questions
Place and Time of Origin 11.7 References
Major Evolutionary Transitions 11.8 Further/Suggested Readings
Phylogeny of Horse 11.9 Answers

11.1 INTRODUCTION
Vertebrates are a diverse group of organisms ranging from lampreys to human beings. The group
includes animals with backbone, such as fishes, amphibians, reptiles, birds and mammals. The
dinosaurs that have caught public attention by making their appearance in several films and books
too are vertebrates. Vertebrates have a long geological history on the planet Earth beginning more
than 500 million years (Myr) ago, starting from the Cambrian to the present.
They first appeared in the fossil record during the Cambrian period of the Palaeozoic era. Fishes,
amphibians and reptiles were the most dominant groups of vertebrates in the Palaeozoic and
Mesozoic eras. In the Mesozoic era, dinosaurs- a group of reptiles, and mammals made their
appearance. Dinosaurs had a wide geographic distribution, being reported from all continents and
were the largest animals to roam the Earth during the Mesozoic era. The mammals began to
diversify only after the demise of dinosaurs at the close of the Mesozoic era at around 66 Myr ago.
In the Cenozoic era, mammals rapidly occupied every niche and corner of the globe and therefore,
the Cenozoic era is also known as the “Age of Mammals”.
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Mammals of the Cenozoic era belong to three groups: placentals (that give birth
to young ones), marsupials (in which an offspring after birth continues to
develop within the pouch of the mother) and monotremes (egg laying
mammals). Among these three groups, the placentals like the horses,
elephants and humans have a well-documented fossil record.
Now we will deal with one of the most important groups of placental mammals
namely, horse. In this unit, we will discuss the systematics, evolution and
phylogeny of horse. In addition, we will also learn how climate has played a role
in the evolution of horse.

Expected Learning Outcomes_______________________

After reading this unit, you should be able to:
 outline the systematic positions of horse;
 describe the probable time and place of its origin;
 discuss the major evolutionary trends in horse;
 enumerate their phylogeny; and
 discuss the role played by the climate in the evolution of horse

11.2 EVOLUTION OF HORSE

Among all the vertebrates there is no other group whose fossil record is better
known than that of the horses, the odd-toed ungulate mammals. The word
ungulate refers to the mammals that bear hooves on their fingers and toes
rather than claws. Horses are odd-toed ungulates like the rhinoceros and tapirs
that have either 1, 3, or 5 counts of toes on their hand or foot and thus, belong
to the Order Perissodactyla. In contrast, cows and camels are even-toed
ungulates (2 or 4 toes) that are included in the order Artiodactyla. Equidae is
the family of horses. It comprises a single extant genus Equus, in which horses
are placed along with the donkeys and zebras
11.2.1 Systematic Palaeontology
Kingdom Animalia
Phylum Chordata
Subphylum Vertebrata
Class Mammalia
Order Perissodactyla
Family Equidae
Genus Equus
11.2.2 Place and Time of Origin
The story of evolution of horse started at the beginning of the Eocene epoch of
the Cenozoic era and North America is considered as the original home of the
horses. As stated earlier horses, rhinoceros and tapirs belong to the
mammalian group perissodactyls. It is believed that perissodactyls originated in
a forest environment by Early Eocene times. Whereas the tapirs and, to some
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extent, rhinoceroses, retained their forest specialisations, modern horses
adapted to life on grasslands. Because of their habit to live in large herds,
horses have been buried and fossilised in large numbers since the earliest
stage of their evolution. In North America, an almost complete sedimentary
sequence from the Eocene to the Recent is present that has yielded well-
preserved fossil horses since the early 19th century, because of which the
evolutionary history of horses is well-documented among all mammals.
Horse remains have also been found from different stratigraphic horizons in
Europe and Asia as we ascend the geological column, but the fossil record is
not as complete as in North America. The recent findings of the earliest horse
Hyracotherium from Europe and a speculated ancestor of horse Radinskya
from China questions the birth place of horses, although many palaeontologists
believe North America as the birth place of horses because the entire
evolutionary sequence of the horses is known from here. In addition, recent
studies have also shown that Hyracotherium was not a true horse.
Though the question of biogeographic origin of the equidae is currently
unresolved, it is well established that during the Early Eocene, the Equidae
could move between North America and Eurasia through a land bridge in the
North Atlantic ocean called as the DeGeer dispersal route (Fig. 11.1), which
by early Middle Eocene was severed thus restricting the movement of animals
either way. Thereafter, the Equidae became extinct in Europe, but continued to
evolve in isolation in North America until the Oligocene. The land connections
between North America and Europe were re-established in the Miocene, about
24 Myr ago and the equids again dispersed to Europe and from there to the Old
World. Though the DeGeer dispersal corridor was no longer available, these
migrations could take place during the times of lowered sea levels that exposed
the shallow Barrents shelf between Siberia and the Aleutian Islands, forming
the Bering land bridge (Fig. 11.1). Throughout the remainder of the Cenozoic
era whenever the sea level dropped the Bering land bridge was exposed and
the equids migrated from North America to Europe and the old world, such as in
the Middle Miocene, Pliocene and Pleistocene times. The horses also
dispersed to South America during the great faunal interchange that occurred
about 4 to 3 Myr ago when the land connection was established between the
Americas by way of the Isthmus of Panama (Fig. 11.1). By the end of the
Pleistocene, the horses became extinct in North America, but survived in other
parts of the world to continue till today. Horses were re-introduced in the New
World by early human migrants.
Do you know?
The “Old World” consists of Africa, Europe and Asia. The term originated in the
early 16th century after the discovery of North and South America called as the
“New World” by the Europeans who till then thought of the world as consisting
of Africa, Asia and Europe only.
The evolutionary history of the horse was one of the first to be developed and
cited as evidence for the evolutionary process soon after the theory of evolution
was proposed by Darwin in his book ‘Origin of Species’ in 1859. The story of
the horse soon became a classic example of evolution, appearing in every text
book on the subject.
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Fig. 11.1: Map showing the dispersal routes for the migration of horses during
different points of time in the Cenozoic era. (Source: simplified and
redrawn after MacFadden, 1992)

11.2.3 Major Evolutionary Transitions

The horse originated in the Eocene and through time it underwent a number of
changes from a dog-like ancestor to what it is today. The main changes which
are exhibited in the phylogeny of horse are listed below (after Colbert, 1980):
1. Increase in body size
2. Lengthening of legs and feet
3. Reduction of lateral toes with emphasis on the middle toe
4. Straightening and stiffening of the back
5. Widening of the incisor teeth
6. Molarisation of the premolars (i.e., in order to grind food premolars became
like molars)
7. Increase in height of the crown of the cheek teeth
8. Development of crests on molars
9. Deepening of the front portion of the skull and of the lower jaw to
accommodate the high crowned teeth
10. Lengthening of the face in front of eyes also to accommodate the high
crowned teeth
11. Increase in brain size
11.2.4 Phylogeny of Horse
The phylogeny of horses has been and still is one of the acceptable and most
frequently quoted examples of evolution. You can see summary of the
phylogenetic history of horse in Figs.11.2 and 11.3, which show the major
changes that took place in the skeleton of horses during the course of their
evolution.
Let us discuss the different evolutionary stages of horse during the
Cenozoic era.
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(i) Horses in Eocene
Eohippus: Eohippus, commonly known as the dawn horse, was a small animal
about 10-20 inches (25-50 cm) high at the shoulder, had an arched back, short
neck, short snout, short legs and a long tail (Fig. 11.3). Its legs were flexible
and rotatable with all major bones present and un-fused. It had 4 toes on each
of the front foot and 3 toes on hind feet. Vestiges of other toes were however
still present. Eohippus walked on dog-like padded feet and had only small
hooves on each toe. It was a forest dweller that looked more like a dog than a
horse. It browsed on soft foliage. Its teeth were low-crowned. Each side of the
jaw had 3 incisors, 1 canine, 4 premolars and 3 molars (i.e., its dental formula
was 3143/3143). The cusps of the molars were slightly connected by low
crests. It became extinct in the Early Eocene, around 49 Myr ago, but prior to its
extinction, it evolved into Orohippus (Fig. 11.2).

Fig. 11.2: Phylogeny of horse (Source: simplified and redrawn after MacFadden,
1992), Pal stands for Palaeocene, Plio for Pliocene and Qt for
Quaternary.
Do you know?
A dental formula is a summary of a mammal’s teeth. The number of teeth of
each type is written for one side of the mouth with the upper and lower teeth
shown on separate rows.
Orohippus: Orohippus means “mountain horse”, although it never lived in the
mountainous region. It looked largely like Eohippus. The major changes that
took place in Orohippus were the loss of the toe vestiges and change in dental
morphology. It was about 10-20 inches (25-50 cm) high at the shoulder with a

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dog-like look and padded feet. It had a small brain, arched back, and short legs,
neck and snout (Fig. 11.3). Its diet comprised tougher plant material than that of
Eohippus and this brought in molarisation of the 4th premolar and more
pronounced development of the crests on the teeth. Orohippus survived until 46
Myr ago.
Epihippus: It arose from Orohippus in the Middle Eocene times (Fig. 11.2)
about 47 Myr ago. Like Orohippus and Eohippus, Epihippus was a small, dog-
like, pad-footed and small-brained horse. It had 4 toes in front and 3 behind.
Meanwhile, the evolution of teeth continued. In Epihippus, the last two
premolars became molarised, making five cheek teeth grinding type. The teeth
however, remained low-crowned, but with well-developed crests. It has been
seen that Epihippus was slightly larger than Orohippus and became extinct by
the end of Eocene.
Mesohippus: By the Late Eocene time, approximately 40 Myr ago the horse
genus Mesohippus descended from Epihippus (Fig. 11.2). Mesohippus also
known as an intermediate horse was slightly larger than Epihippus. Standing 24
inches (71 cm) high at the shoulder, it was of the size of a sheep. Its back was
less arched, and the legs, neck, snout and face a little longer (Fig. 11.3).
However, Mesohippus retained three toes on its hind feet whereas on its front
feet the 4th toe was reduced. Mesohippus like its ancestors was still pad-footed.
The third last premolar however continued the evolutionary trend of
molarisation, thereby resulting in six grinding cheek teeth in Mesohippus as well
as all the later horses. Mesohippus continued to survive until 29 Myr in the Late
Oligocene.
(ii) Horses in Oligocene
Miohippus: It arose from Mesohippus in the Late Oligocene time (Fig. 11.2)
about 37 Myr ago. It was a larger horse with a slightly longer skull than
Mesohippus. A characteristic feature of Miohippus was the development of a
variable extra crest on its upper cheek teeth, a trait that was retained by all the
later horses. The ankle joint of Miohippus also changed a little. Miohippus
continued for a while as it was, and then by the Early Miocene, it had split into
two branches. One of the branches continued as three-toed browsing horses
such as Kalobatippus, Anchitherium, Megahippus and the Asian Sinohippus.
The other branch that arose from Miohippus was that of Parahippus.
(iii) Horses in Miocene
Parahippus: As stated above, Parahippus evolved from Miohippus (Fig. 11.2).
It was both a browser and a grazer (a mixed feeder). Appearing in Early
Miocene around 25 Myr ago, Parahippus was a little larger than Miohippus, with
about the same brain size and body form (Fig. 11.3). However, as this horse
started eating grass its teeth became slightly high-crowned (hypsodont).
Parahippus adapted to a running mode, which in turn developed springy foot
ligaments, although the foot still remained three-toed. Parahippus went extinct
by 15 Myr ago.
Do you know?
Hypsodont dentition comprises high-crowned and short-rooted teeth. This
dentition is characteristic of those animals that feed on gritty and fibrous food
material such as grasses.
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Merychippus: Parahippus by about 18.5 Myr ago evolved into a spring-footed,
hypsodont horse, named Merychippus (Fig. 11.2). Merychippus was about 40
inches (1 m) high at the shoulder. It was still three-toed, but a fully spring footed
horse that stood permanently on the tiptoe (Fig. 11.3). The side toes though
present were of variable size whereas the central toe was a little larger with a
well developed large hoof. Also the leg bones were fused and reduced in size
to eliminate leg rotation. These transitions occurred as Merychippus adapted to
rapid running over hard ground. Another major transformation that took place in
Merychippus was the increase in the height of its teeth as it fed dominantly on
grass. It was the first member of the line of descent of the grazing horses that
marked the transition of horses from primitive browsers to the modern grazers.
It is, therefore, also known as a ruminant horse. However, to accommodate the
large teeth, the jaw became deeper, and the muzzle (the projecting part of the
face, including the nose and mouth) became elongated. This gave Merychippus
a new look and so it was aptly named as Merychippus, “the horse with a new
look”.
Merychippus underwent rapid speciation called “merychippine radiation”
giving rise to new horse species that fall in three major groups. These groups
are hipparions, protohippines and true equines.
Hipparions were small to large pony like, three-toed grazing horses that
emerged around 16 Myr ago. They include forms like Comohipparion,
Pseudhipparion, Neohipparion, Hipparion and Nannippus. These horses spread
from North America to the Old World, reaching the Indian subcontinent about
14-13 Myr ago.
Protohippines evolved by 16 Myr ago. Protohippus and Calippus were the
older, smaller forms that retained three toes in their feet. By the Middle Miocene
the protohippines line of descent was represented by Pliohippus. Initially, a
three-toed horse, Pliohippus, through three successive stages in the Early
Pliocene, lost side toes, ultimately culminating into a single-toed Pliohippus
(Fig. 11.3). Pliohippus gave rise to the “hippidions” which were stocky, short-
legged, single-toed horses with odd boxy skulls. These horses travelled from
North America to South America and thrived there at least up to 2-1 Myr ago.
True equines include the present day horse and its ancestors. They comprise
both three-toed as well as single-toed horses. The three-toed horses were
Merychippus in which the side toes were smaller than the central toe, whereas,
the single-toed horses are Dinohippus and Equus. By 12 Myr ago, Dinohippus
descended from Merychippus (Fig. 11.2). Dinohippus the one-toed horse had
slightly straighter teeth than those of Merychippus. It looked very similar in foot,
tooth, and skull morphology to the modern horse (Fig. 11.3).
(iv) Horses in Pliocene
Descendents of “hipparions” and “protohippines” stock continued into the
Pliocene and spread to the old world and South America. Dinohippus the true
equine was the most common horse in North America in the late Pliocene. It
gave rise to the modern day horse, Equus.

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(v) Modern Horse
Equus: It is the genus of all the modern equines that first arose around 5 Myr
ago from Dinohippus. The first Equus was 52 inches (1.3 m) tall (the size of a
pony), with a rigid spine, long neck and legs, fused leg bones with no rotation,
long nose, flexible muzzle and a deep jaw. The brain was a bit larger than that
of Dinohippus. Equus soon diversified into several species that co-existed with
other horses. Equus like Dinohippus is single-toed (Fig. 11.3). However, its
teeth are straight and higher-crowned than of Dinohippus (Colbert and others,
2001).

Fig. 11.3: Morphological changes that took place in horses during the course of
evolution. (Source: modified after Benton, 2005)

During the first major glaciation of the Late Pliocene, about 2.5 Myr ago, certain
Equus species crossed to the Old World. Some entered Africa and diversified
into the modern zebras. Others spread across Asia, the Middle East and Africa
where they adapted to desert life such as Kiangs, Onagers and wild asses
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including donkeys. Still others spread across Asia, Africa, South America, and
Europe as the true horse, Equus caballus.
It is worth mentioning here that Equus arose in North America and at the end of
Pleistocene epoch (11000 years ago), it became extinct in North America
probably as a result of climate change and overhunting by humans. Later in the
16th century the horse was reintroduced in North America by humans.
Do you know?
The Miocene to Pliocene fossil record of the Siwalik Hills of the Indian
Subcontinent yields abundant material of hipparionine equids. The oldest fossil
comes from the Chinji Formation and its equivalent beds dated around 14-13
Myr old. The fossil horse recovered is Hipparion which was an immigrant from
North America. Though, Hipparion was not the dominant component of
mammalian taxa initially, its number and diversity increased by the upper part of
the Middle Siwaliks. However, by the Upper Siwalik times it became extinct and
was replaced by Equus. Species of fossil Equus are also known from the
Karewa deposits of the Kashmir valley and from Pleistocene alluvial deposits of
the Narmada valley.

11.3 ROLE OF CLIMATE IN THE EVOLUTION OF

HORSE
You now know that the beginning of the evolution of horse took place in the
Cenozoic era, close to the Palaeocene-Eocene boundary. You may find it
interesting to note that the Palaeocene–Eocene boundary witnessed an abrupt
global warming, which is commonly referred to as the Palaeocene–Eocene
Thermal Maximum (PETM). At this boundary many modern mammalian taxa
appeared and so did the first horse Eohippus. The major changes that took
place among horses during the course of evolution were the reduction in the
number of toes and deepening of the cheek teeth (i.e., premolars and molars).
These changes are linked to changes in environmental conditions which
brought retreat in forest cover and spread of grasslands. The Eocene warming
trend continued at least up to its earliest part which brought in only minor
evolutionary changes in Eohippus and its near descendants. The body and feet
stayed mostly the same, with slight changes in the toes. The major change was
in the teeth of Eocene equids as they started to eat more plant browse and less
fruit. They also developed more grinding teeth to eat the slightly tougher food.
With the onset of the Late Eocene, the cooling of the Earth began and small
ephemeral ice sheets developed and the climate became drier. This affected
the vegetation patterns with vast forests shrinking and grass evolving during the
Late Oligocene–Early Miocene times. So as we move toward the Oligocene,
horses become as mixed feeders and by Miocene horses changed their dietary
mode from browsing to grazing. Since grass is difficult to chew and wears down
teeth rapidly due to its high silica content, horse teeth increased the height of
the tooth crowns so that the teeth remain out of the gum as their tops were
continuously worn down. There was an increase in body size and length of the
face as well.
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The grazing horses also adapted to rapid running on open grasslands. They
started to become leggier for better speed out in the open eventually becoming
specialised runners. The bones of the legs began to fuse together, and the leg
bones became specialised for efficient forward-and-back strides, with flexible
leg rotation being eliminated. Most significantly, the horses began to stand
permanently on tiptoe i.e., they changed from three-toed to single-toed horse,
another adaptation for speed, instead of walking on dog-like pads.
In Pleistocene times, the climate again changed as Earth went into a glacial
mode. The Pleistocene climate change accompanied by the overhunting by
man led to the extinction of numerous mammals in North America including the
modern horse, Equus. The warming and cooling episodes of the Cenozoic had
also brought in global sea level changes that formed land bridges during the fall
of sea level which facilitated the dispersal of horse from North America to the
rest of the world. The horses, though became extinct in North America where
its entire lineage proliferated, survived in other parts of the world.
The fossil history of equids reveals that the progression between different equid
species was not as smooth and consistent as was once believed. Though some
transitions like molarisation of premolars were gradual, others like reduction in
toes and increase in body size were abrupt. Since these changes were the
adaptations required by the horse to survive the changing climate, it can be
said that the evolution of horse was driven by climate change.

SAQ 1
a) Fill in the blanks.
1. ______________________ is the family of horses.
2. ______________________was the first horse genus.
3. ______________________ is the possible place of origin of horse.
b) Match the following:
(i) Dog-like padded feet (a) Merychippus
(ii) Mountain horse (b) Parahippus
(ii) A mixed feeder (c) Orohippus
(iv) The horse with a new looks (d) Eohippus
c) What is Equus?

11.4 ACTIVITY
You will find the generalised evolutionary sequence of horse from
Hyracotherium to Equus through the Cenozoic era in the Fig. 11.6. In this
figure, some evolutionary stages of horse are missing. Here you have to do
following three things:
Here you have to do three things:
i) identify the missing stages,
ii) write them at an appropriate level in the sequence and
iii) mark the boundary between browsing and grazing horses.
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Fig. 11.6: Evolutionary sequence of horse. Pal stands for Palaeocene, Plio for
Pliocene and Qt for Quaternary.

11.5 SUMMARY
Let us quickly summarise what you have learnt in this unit.
 Horses are odd toed ungulates like the rhinoceros and tapirs that have 1, 3
or 5 numbers of toes on their hand and feet and thus, belong to the
order Perissodactyla.
 Equidae is the family of horses. It comprises a single extant genus Equus.
 Ancestor of the horses first appeared in North America during the Eocene
epoch.
 The evolutionary sequence of Equidae from the oldest to the modern horse
comprises Eohippus – Orohippus – Epihippus – Mesohippus – Miohippus –
Parahippus –Merychippus – Dinohippus – Equus.
 The evolution of horse involves increase in body size, lengthening of legs
and feet, reduction of toes, modification of dentition, deepening of the front
portion of the skull and of the lower jaw, lengthening of the face and increase
in brain size.
 The major evolutionary changes among the horse were brought by change in
environmental conditions such as retreat in forest cover and spread of
grasslands.
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11.6 TERMINAL QUESTIONS

1. Write in brief on the major trends of evolution of horse.
2. Describe the phylogeny of horse.
3. Discuss the role of climate in the evolution of horse.

11.7 REFERENCES
 Benton, M.J. (2005) Vertebrate Palaeontology, 3rd edition, Blackwell Science
Ltd., USA.
 Colbert, E.H. (1980) Evolution of the Vertebrates – A history of the
backboned animals through time. John Wiley & Sons, USA, Pp. 379-384 and
425-437.
 Colbert, E.H. and others (2012) Colbert’s Evolution of the Vertebrates – A
history of the backboned animals through time, Wiley India (Pvt.) Ltd., New
Delhi (reprinted).
 MacFadden, B.J. (1992) Fossil Horses: Systematics, Paleobiology, and
Evolution of the Family Equidae, Cambridge University Press, Cambridge.

11.8 FURTHER/SUGGESTED READINGS

 Colbert, E.H. and others (2012) Colbert’s Evolution of the Vertebrates – A
history of the backboned animals through time, Wiley India (Pvt.) Ltd., New
Delhi (reprinted).
 Jain, P.C. and Anantharaman, M.S. (1996) Palaeontology – Evolution and
Animal Distribution, Fourth Edition, Vishal Publications, Jalandhar.

11.9 ANSWERS
Self Assessment Questions
1. (a) Equidae.
(b) Eohippus.
(c) North America.
2. (i) - (d), (ii) - (c), (iii) - (b), (iv) - (a)
3. Equus is a genus of all modern equines. It first arose around 5 Myr
ago from Dinohippus. The first Equus was 52 inches (1.3 m) tall (a
size of pony), with a rigid spine, long neck and legs, fused leg bones
with no rotation, long nose, flexible muzzle and a deep jaw. The
brain was a bit larger than that of Dinohippus. Equus soon diversified
into several species that co-existed with other horses. Equus like
Dinohippus is single-toed. However, its teeth are straight and higher-
crowned than Dinohippus.
Terminal Questions
1. Refer to subsection 11.2.3.
2. Refer to subsection 11.2.4.
3. Refer to section 11.3.
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