Evolution

Human Origin & Evolution

Dr Tanushree Saxena
Associate Professor
Department of Zoology
Swami Shraddhanand College
Alipur Village,Delhi-110036

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 “What a piece of work is a man! How noble in reason! How infinite in faculties! Inform and
in moving, how express and admirable! In action how like an angel! In apprehension how like
a God! The beauty of the world! The paragon of animals!”

- William Shakespeare

Hamlet, II, ii (1564 – 1616)

Indeed, man occupies the loftiest pinnacle of the evolutionary fabric and is considered to be a
paragon of animals but the story of our origin is still clouded with many unanswered queries
(shrouded) .Where did man come from? This question has preoccupied human thought for tens
of thousands of years but the picture is still hazy because of the maddening scarcity of human
fossils in sedimentary rocks and excavation work at sites is time consuming and demanding.
Nevertheless, the evidences, new informations about missing links are growing faster than it
can be analyzed, but there are still surprises in store and unsolved problems. All this makes the
understanding of the story of human evolution still more fascinating and exciting.

Primate Evolution and Classification (The classification of Homo)

In 1859, when the “Origin of Species” was published, it was already obvious that if humans
shared a common ancestry with other species, their closest relatives must be great apes of the
order Primates of class Mammalia.

Characteristics of Primates

• Independent mobility of the digits

• An opposable first digit in both hands and feet (thumb, big toe)
• Replacement of claws by nails to support the digital pads on the last phalanx of each
finger and toe.
• Teeth and digestive tract adapted to an omnivorous diet – i.e. simple cusp pattern in
molar teeth 32- 36 teeth.
• A semierect posture that enables hand manipulation and provides a favourable
position preparatory to leaping.
• Centre of gravity positioned close to the hind legs.
• Well developed hand - eye motor coordination.
• Optical adaptations that include overlap of the visual fields to gain precise three –
dimensional information on the location of food objects and tree branches.

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 • An eye completely (anthropoids) or fractionally (prosimians) encased by bone (bony
orbits).
• Shortening of the face accompanied by reduction of the snout.
• Diminution of the olfactory apparatus in diurnal forms.
• Compared with practically all other mammals, a very large and complex brain in
relation to body size.
• Lengthened period of maturation, low reproductive rate.

Two alternative classifications of living primates

I. Traditional classification (Fleagle, 1988)

Order Primates
Sub-order Prosimii (nocturnal, arboreal, low organization claw on the second digit, long
muzzles and furry faces).
Infra-order – Lemuriformes [lemurs or “half-monkeys” (most ancient primates) lorises.
- Tarsiformes (Tarsiers).
Sub-order Anthropoidea
- 32-36 teeth, closed orbit, 2 pectoral mammae, prehensile feet, convoluted cerebral
hemispheres covering cerebellum.
Infra-order: Platyrrhini (New-World monkeys) e.g. Marmosets etc.)
- wide nasal cartilage between the mostrils, prehensile tails, restricted to Central & South
America probably descended from Eocene lemurs.
Infra-order: Catarrhini (Old-World monkeys)- Apes & Hominids
- narrow nasal septum, non-pre-hensile tails often short, natives of Africa & Eurasia.
Super-Family: Cercopithecoidea (Old World monkeys, Macaca, baboons)
“ “ : Hominoidea (Apes & Hominids)
Family : Hylobatidae (Lesser apes e.g. Gibbon (Hylobatus)
“ : Pongidae (Great apes e.g. Chimpanzee Pan Gorilla, Orangutan (Pongo)
“ : Hominidae (Only 1, Species extinct)
Homo sapiens
4 recognized races – Australian, Negroid, Mongoloid & Caucasians
(India – Caucasians in North & Australoid in South)

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II Recently Proposed Classification

Order: Primates

Sub order: Prosimii

Super family: Lemuroidea: Lemurs

“ : Lorisoidea: Lorises, galagos (bush babies)

“ : Tarsioidea: Tarsiers.

Sub order: Anthropoidea

Infra order: Platyrrhini (New World)

Super family: Ceboidea

Family: Callitrichidae: marmosets tamarins

“ : Cebidae: Capuchins, howler monekys, spider

monkeys

Infra order: Catarrhini (Old World)

Super family: Cercopithecoidea

Family: Cercopithecidae: macaques, baboons, vervet monkeys

Family: Colobidae: langurs, colobines

Super family: Hominoidea

Family: Hylobatidae: gibbons, siamangs

“ : Pongidae*: Orangutans, gorillas, chimpanzees

“ : Hominidae: Humans
*In some molecular classifications based on immunological studies and DNA sequencing
(Miyamoto et al., 1987) researchers put all the great apes together with humans in a single

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family, Hominidae, which they then divide into 2 subfamilies

Ponginae (Orangutans) and Homininae (Humans, Chiampanzees & Gorillas)

(Tattersall et al., & Ruvolo, 1997, Goodman et al., Shoshani et. al., 1996).

Human Phylogeny/ Phylogenetic Relationships

About 65-70 million years ago, in the Paleocene epoch, some early insectivorous mammals
became arboreal, (like modern tree shrews) presumably because of intense competition on the
ground and availability of rich, untapped source of food on the trees. This brought about
radical changes in them viz. replacement of claws by flat nails, development of limbs into
grasping organs, opposable, more flexible digits with tactile pads, shortening of snout,
rounding of skull, development of stereoscopic vision, growing importance of sight over
smell, anteriorly directed eyes and so on. Gradually, as the years passed by the climate became
colder in Oligocene era causing the forests to dwindle. This triggered a splitting or divergence
in the parental stock of Primates. Some retained their arboreal mode of living while others
such as man took to a more adventurous and progressive route – the terrestrial habitat. This
descent from trees in turn brought about several changes in them. Thus, we are a product of
the primate arboreal adventure Characteristics of Homo sapiens

Despite our place in the animal kingdom, humans are unique among animals and dominate the
entire planet, because of some very special and unique characteristics which are the following:

(1) Bipedal Locomotion – Balanced bipedalism is uniquely human. Bipedal locomotion

coupled with upright posture gave freedom to the forelimbs (arms) which were
subsequently used for more complicated manipulative and creative tasks – for e.g.
gripping a tool, holding, gesturing, playing a violin etc. or simply feeling (as it is also a
prime organ of touch). Thus, the hands were liberated from the locomotor function and
used for many different purposes.
(2) Upright posture – It involved many anatomical changes i.e.
a) balancing of skull on the upper part of vertebral column.
b) Therefore, the foramen magnum shifted forward.
c) Development of spinal curvatures – There are four such curvatures developed in the
spinal column, making it S – shaped. These curvatures acted as shock – absorbers.
d) Basin-shaped pelvis to support the viscera.
e) relatively short forelimbs (in contrast to apes where they reach below knees).
f) Plantigrade foot
g) loss of opposability of great toe (et. Anthropoids)
h) development of shock absorbing arch.
i) tendency towards monodactyly.

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 j) widened but shortened ilium bringing sacrum closer to acetabulum.
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(3) Brain: Human brain has a large cranial capacity ~1350cm on an average, with the
cortical matter, considered to be the seat of intelligence, reasoned behaviour, memory,
abstract thought and delicate as well as accurate muscular movements, is highly
“convoluted” complex and well developed. Further, there is increase in the size of
frontal lobes of cerebral hemispheres resulting in a relatively high forehead.

Warren McCulloch, an American student of brain has put it, “The brain is like a
computing machine, but there is no computing machine like the brain”.

(4) Vision: Only human beings and their nearest primate relatives have the special
combination of full stereoscopic vision and Colour vision which enables them to focus
in fine detail upon particular and distant objects.
(5) Teeth & Jaws of humans have undergone immense modifications to suit an omnivorous
diet. The differences between apes and man are underlined below:
Apes Man

Incisors Large Small

Canines Large projecting Small, not projecting

Premolars & Molars straight rows rounded rows

jaw shape u – shaped rounded parabola

Diastema (gap between yes no

incisors and canines)

Simian shelf (supporting Yes no

shelf behind incisors &
canines)

1st pre-molar cutting type not cutting type

face (facial angle) prognathus orthognathus(nearly vertical)

(protruding)

Chin no chin, since upper edge of Present, because lower

lower jaw extends further margin of jaw protrudes
than does the lower

tori present-these provided No tori

support for chewing

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 (6) Language: The great significance of the combination of human brain and body is
perhaps best shown by its most important innovation: the language/articulate speech.
Apes lack the kind of pharynx, (long), tongue (thick), soft palate to articulate. It was
the major biological characteristic to evolve.
(7) Cultural Adaptation – With the gift of speech, we acquired an immensely powerful
tool for cultural evolution. The ability to speak facilitated and speeded up, cultural
evolution – to communicate, to hunt, to modify environment to suit themselves to
make weapons etc. but most importantly to learn from the accumulated experience of
other people and groups.
This in turn, resulted in development of communal life, use of fire and gave
independence from climatic conditions and hence led to dispersal.
(8) Other Characteristics
a) loss of hair in some parts – need of clothing to overcome winter.
b) loss of pigment in some races
c) vestigial organs – viz; vermiform appendix reduced (caecum), muscle etc.

Therefore, the evolutionary trends that one observes in the phylogeny of human beings are:
- Increase in size and complexity of brain specially frontal lobes.
- Erect posture
- Shorter arms
- Perfection of thumb opposability
- Reduction in size of teeth and muzzle

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 - Diminution of brow ridges
- Loss of jaw power
- Development of chin prominence
- Increase of skull capacity
- Diminution in strength of zygomatic arch
- Development of articulate speech.

The Early Apes/Human-Ape Comparisons

Table 20-4, Pg. 469 – Strickberger

The Fossil Record

Although scanty, the story of human origin and evolution is remarkably complete.

According to the fossil record, many Mesozoic mammals were very much like extinct tree
shrews adapted to insectiveorous lifestyle inhabiting both the forest floor and tree shrubs.
Primates evolved from these early forms. However, Paleocene deposits showed the presence

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of an early archaic primate group of the suborder Plesiadapiformes with unique auditory
region and dentition different from related insectivorous forms. By the Eocene, some such
groups evolved still further by developing bony orbital rings and digital nail. This split into
two families – Adapids (which gave rise to modern lemurs and lorises and – Omomyids (to
tarsiers).

The next evolutionary stage, leading to platyrrhine and catarrhine anthropoids is the subject of
considerable debate. Some primatologists believe that both anthropoid groups had a
monophyletic origin but controversies like whether they arose from adapids or omomyids or
whether the platyrrhine monkeys arrived in the New World from North America or Africa is
still raging. However, at present omomyid anthropoid ancestry is more acceptable but there is
no general agreement (Kay, et al., ).

In any case, both paleontological and molecular dating suggest that anthropoid lineage is
quite ancient (Paleocene epoch, 50 million years ago).

OLIGOCENE

This period is characterized by the divergence of old world monkeys from apes and man and
there were fossils of Proto-monkeys and Proto-Apes.

• Oligopithecus – This is one of the earliest fossils in the primate line

characterized by small size, mixed dental features with some showing cutting
first premolar, characteristic of anthropoids.
• Propliopithecus ~ 35 million years old found at Fayum in Egypt.

MIOCENE

a) Aegyptopithecus (belongs to early Miocene)

(“Dawn Ape”)

It was discovered from Fayum by Elwyn Simons in 1966, consisting of virtually complete
skull, headbones, upper and lower jaw and an almost full set of teeth. It’s the most primitive
ape yet discovered.

Characteristics: a) long canine teeth in the front of upper jaw fitting with the lower jaw allow
shearing like modern apes.

b) spaniel-sized

c) skull shaped like a monkey

d) long tail

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It probably diverged in early Miocene to a group of species now put under Dryopithecus.
There are 6 species known from East Africa (17-20 million years ago) and 1 from South East
Asia. It is believed that while one species. , earlier called Proconsul discovered on Rusinga
Island in Lake Victoria (Africa) gave rise to Pan/Chimpanzee, another to gorilla and still
another to hominoids.

b) Dryopithecus (middle Miocene, ~ 20 million years ago)

It was first discovered in Africa (although number of fossils were found in France earlier) and
was widespread in Europe, Asia and Africa about 14 million years ago. Post cranial skeleton is
scarce.

Characters – 1) blend of monkey and ape characteristics

2) various sized

3) jaws U-shaped ape-like

4) palate flat characters

5) arboreal life

6) poor brachiator (arms shorter than legs like man)

However, it seems to be the best contender for the position of hominid ancestor. It is probable
that the apes arose from moderate brachiators viz. Dryopithecus on one line of radiation and
man arose on another.

Man, in fact, resembles modern old-world monkeys more than the modern apes, leading some
to suggest that the human line diverges earlier (in Eocene) along with the old-world monkeys,
leaving Dryopithecus from direct lineage.

c) Ramapithecus (~ 9-15 million years ago)

This fossil (right upper jaw fragment with 4 attached teeth) was discovered by G.E. Lewis in
Siwalik Hills of India in 1932 with another fossil Bramapithecus (lower jaw).

Fossil evidence also came from Fort Ternan in Kenya (upper and lower jaw) by Louis &
Mary Leakey (Kenyapithecus), from Athens, Greece (European member of the genus
Ramapithecus) from Italy (Oreopithecus – more primitive (~ 10-12 million years) and
probably not in the hominid line) from Potwar plateau in Pakistan, from Candir & Pasalar in
Turkey in Rudabanya (Hungary) & from Lufeng County in Yunnan. This showed that
Ramapithecus was fairly widespread.

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 Miocene epoch, particularly mid-miocene was a key factor for hominid development as it
was characterized by the retraction of Tethys sea establishing contact between Africa and
Europe as well as Africa & Asia.

Characters: It was definitely a non-pongid & more of a hominid:

(1) mandibles relatively low & wide

(2) maxilla strongly buttressed, protruding
(3) relatively short face (prognathus)
(4) non-shearing premolars & molars
(5) wearing of teeth in older specimens
(6) tough herbivorous diet
(7) canines ape-like but didn’t project much beyond the teeth line (occulusal plane)
(8) very small diastema
(9) small canines & incisors – relatively short jaws.
(10) wide-curving jaw with arched palate. .

It was speculated earlier that hominid adaptation took place in Africa and then spread through
the land corridor to Europe & Asia or Dryopithecus like form migrated out of Africa whereas
Ramapithecus evolved in Arabian Gulf Area and then radiated to Europe, Asia & Africa. This
hypothesis seemed more probable because Ramapithecus was fairly widespread in all the
places simultaneously.

However, with the recent discovery of more complete ramapithecine fossils, researchers are of
the view that this group linked to a much earlier ape like lineage that may have been ancestral
to Orangutans (Wolpoff, 1982; Andrews, 1983; Pilbeam, 1984). African apes & humans may
therefore have diverged a long time after the ramapithecine radiations (supported by molecular
data, Ruvolo et al., 1994).

Fig. 20-5, Pg. 470, Strickberger + Fig. 26.2 Pg. 730, Futuyma.

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 d) Sivapithecus: It was an ape that developed someway along the same road as
Ramapithecus and was very similar to it but retained its long canine teeth and had
thick-enameled molar teeth. However, it never succeeded in fully adapting to open
habitats- the savannas – and their particular food resources and became extinct ~ 8
million years ago.

The specimen formerly classified as Ramapithecus are now usually included in the
genus Sivapithecus which in turn is thought to be a close relation of the ancestors of
modern orangutans.

Fig. 15.25b Ridley (3rd edn.)

e) Gigantopithecus – This open-habitat form was most successful. It was quite like a
gorilla, large-formed and survived as a successful quadraped in Asia for about 9
million years ago and became extinct only about 1 million years ago. But its extinction
has been questioned by those who believe in Yeti of the Himalayas or Sasquatch of
Northwest American Coast.

Characters – ground living savanna ape

• thick enamel Ramapithecus

• short canines characters
• immense molars (tough, fibrous vegetable diet)

It was probably a pongid which developed hominid dental characters as a parallel evolution
but became extinct when hominids invaded Asia. Last remains have been detected from S.E.
Asia, China and one species (G. bilaspurensis) from Siwaliks.

PLIOCENE

Australopithecines (Southern Ape). The earliest fossil hominids are called australopithecines.
Fig. 26.4 Pg.731, Futuyama.

Since Miocene apes were represented only by jaw fossils, they didn’t give us any clue about
bipedalism but somewhere in late Miocene and Pliocene, the adaptations of bipedalism made
their appearance in at least one kind of primate, which ultimately led to Homo.

(a) Australopithecus afarensis,: ( 3 – 3.9 million years ago)

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 These are a series of fossils found in East African sites at Laetoli (Tanzania)
(“footprints”) and Afar (Hadar) region of Ethiopia [extremely famous fairly complete
skeleton( 40%) of “Lucy”].

Characters :

• small brain (400 cc)

• heavy eye brow ridges
• low forehead
• prognathus jaw/face
• canines larger in males than female but sexual dimorphism less pronounced than
apes.
• thick enamel
• small, muscularly powerful body, 3.5 – 4 ft. high
• longer arms than man
• flat cranium
• curved bones in fingers and toes.(indicating they climbed trees)

1. but still fully bipedal.

(b) Australopithecus africanus (1.75 to 2 million years ago)

The first tangible evidence about a bipedal primate came again from Africa – from a place
called Transvaal region in S. Africa, discovered by Raymond Dart. (The famous “Taung
child”). It was named A. africanus which later came to be known as “gracile” form.
Australopithecines also found at Swartkrans.

Characters :

- brain capacity – 450 – 500 cm3

- foramen magnum situated near the centre of skull.
- face projecting but no muzzle (prognathus)

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 - small canines
- milk teeth human-like (multi-cusped) (anterior molar)
- shoulder adapted for arboreal locomotion.
- upright posture
- predominantly bipedal gait
- lunate sulcus (anterior border of visual area in the brain further back.

c) robustus was discovered from few km. from Sterktontein Kromdraii area by Dr. Robert
Broom. It was called as Paranthropus robustus (robust near-man) by him because it
was heavy jawed with extremely large molars, later renamed as A. robustus.

Characters - large size

- weight 150 pounds

- cranial capacity 530 cc.
- molars extremely large
- heavy jaws with large jaw muscles
- anterior teeth smaller than in gracile type
- canines ,skull with a bony ridge or crest

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-

Dia from Campbell, Fig. 6 -5 Pg.

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12 7.

(d) A. boisei (1.75 million years ago) was another fossil evidence (skull) discovered from
Olduvai Gorge (East Africa) at Koobi Fora, Kenya by the husband and wife team of Louis &
Mary Leakey
Characters:
• Morphologically closer to “robust” type charleCharles Boise, who financed Leakey’s
research). Later the skull was called A. boiesi.
(1) A. aethiopicus (2.5 m years ago) from deposits at Lake Turkana, Kenya.

• molars large
• shortening of face, more vertical
• massively built
They probably evolved separately but parallel to africanus – robustus lineage.
(2) Ardepithecus ramidus (4.3 – 4.5 million years ago) (mya)
- fossil evidence (teeth) is perhaps the most primitive hominid like fossils from
Ethiopia.
Characters :
• reduced sexual dimorphism for canine teeth
• forward positioning of foramen magnium
• Erect posture

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All these evidences, in any case indicate that Ramapithecus was probably an ape-like ancestor
and hominids diverged from apes about 15 million years ago while Australopithecines were
the first bipedal tropical apes evolved either from Ramapithecus or Ramapithecus – like forms.
Also, there is little question that A. afarensis stands at or very near the base of hominid
phylogeny. However, australopithecines definitely diverged along different lines. There are at
least three hypotheses of the phylogenetic relationships among fossil hominids, advocated by
different authors. Hypothesis ‘C’ is perhaps most widely accepted, though with little
uncertaintly about whether A. africanus is a separate lineage from Homo habilis or its direct
ancestor. (After Grine 1993 & Jones et al., 1992).

(A) (B) (C)

Fig.26.6. Pg.732 (Futuyma)

Origin & Evolution of Homo

A number of species of the genus Homo appeared in the evolutionary scene and became
extinct before the origin of modern man, Homo sapiens. These are called as pre-historic
species of man. Some of them evolved as direct ancestors of the modern man, while others
were merely off-shoots of the main line of human evolution.

The earliest fossils referred to as genus Homo range from 2.4 to 1.6 Mya (latest Pliocene &
early Pleistocene and came from Tanzania, Ethiopia, and South Africa).

a) Homo habilis

Early in 1960s, Jonathan Leakey (Louis Leakey’s Son) uncovered some teeth and bone
fragments from the same strata in Olduvai Gorge (East Africa) not far away from the region
where more heavily built A. boisei was discovered. It clearly represented a distinct species but
far closer to humankind than “gracile” A. africanus. It was described as Homo habilis, the
“handy” man (the tool maker).

Characters:

• Small brained ~ 650 cc (small for human but larger than A. africanus (450cc)
• 120 – 180cm tall (4.5 ft. tall)

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 • delicate lower jaw
• dentition modern man-like (short tooth row)
• omnivorous
• tool maker (patterned stone tools)
• Olduwan technology Oldowan Industry
• reduced prognathism

It’s the epitome of missing link no longer missing (Wood, 1992)

b) Homo erectus (1.6mya to 300,000 years).

H. habilis grade into later hominid fossils, referred to as H.erectus of mid-pleistocene era.

The fossil (Jaw bone, including several teeth, part of the skull and thigh bone-femur) was
unearthed by a Dutch anatomist, Eugene Dubois in 1891 from the bank of Solo river at
Trinil in Java and was named Pithecanthropus erectus initially (Java ape man).

Charactes:

• flat and rounded skull

• brain capacity ~800 – 1000cc.
• speech centre (Broca’ s area) well developed
• face steeper and less prognathous with no chin
• teeth smaller except larger canines of lower jaw
• beetling brows, prominent brow ridges
• heavy neck muscles
• brain deficient in pre-frontal and parietal areas (seat of higher mental faculties) and
heavy in rear part of skull (implying lack of delicate balance)
• Height – 5`8`` (150cm.)
• Weight – 154 pounds
• lips thick and protruding
• made more sophisticated tools of stones and bones including axes (Acheulean type)
• lived in small groups in caves
• Omnivorous and probably a cannibal
• first to make use of fire for hunting, defense and cooking.

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 It appeared that H. erectus grade hominids had moved to the temperate West European
areas by 0.5 million years.

c) Homo heidelbergensis (Heidelberg man)

The fossil (toothed lower jaw) ~ a million years old was discovered from middle
Pleistocene sand beds at Mauer Heidelberg in Germany in 1907.

Characters:

• massive ape-like jaw

• dentition man-like, small
• chin sloping, area of muscular attachment powerful
• teeth with short divergent roots and large pulp cavity like the teeth of cattle (perhaps
an adaptation for harsh food).

These characters are not found anywhere except Neanderthal man, to which Homo
heidelbergensis may be an ancestor. Presumably it’s an offshoot of the main human
evolutionary line.

d) Homo erectus pekinensis (Peking man)

The story of human evolution from Java continued in China as fossil teeth of lower middle
Pleistocene of about 600,000 years old in Cave Choukoutien (Beijing) .It was discovered by
W. C.Pei working with Davidson Black (1924) and named as Sinanthropus Pekinenesis. It was
very much similar to Java ape man and the two are considered to be variants of the same type
and often placed in the same genus.

Character

• 155cm high, lighter and weaker than Java man

• skull very thick, cranial vault more rounded
• cranial capacity ~ 1075cc.
• forehead low, tori more pronounced
• inconspicuous chin
• canine teeth large
• omnivorous – cannibal
• used sharp chisel – shaped tools of bones, stones and quartz (Oldovian industry)
• fire for cooking
• lived in caves in small groups

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 • used non-syllabic and non-articulate language
• probably an ancestor of modern man goloids (Franz Weiden reich).

e) H. erectus mauri tanicus (Atlantic Man or ternifine man)

These fossils (three jaws and a parietal bone) were discovered from ~ 600,00 years
old Pleistocene deposits in Algeria (1975). Later, more fossils of this type were
discovered from Casablanca, Morocco.

Characters:

• most primitive ape man found so far in North Africa

• more advanced than Java or Peking man
• low cranial dome
• heavy projecting jaws
• no chin.

Thus, Homo erectus grade hominids were very widespread over Asia, Africa and Europe
between 1.5 and 0.5 million years ago. The first hominids were found in temperate regions,
even at altitudes of upto 2500 metres. By the end of Pleistocene era, they had refined hunting
techniques, certainly used stone tools and fire was also used at some sites for cooking.
However, it’s still controversial whether Homo habilis diverged into H. erectus & H. sapiens
or they were directly in the line of evolution of modern man.

Homo sapiens

Fossils of a form intermediate between H. erectus & H. sapiens are called as archaic modern
man or pre-sapiens and were mainly discovered from Britain and China.

Most hominid fossils from about 0.3 mya onward as well as some African fossils about 0.4 my
old are referred to as Homo sapiens. The best known populations of archaic H. sapiens are:

a) The Neanderthal/Neandertal (modern spelling) Man

The first Neandertal was discovered in 1856 in Dusseldorf (Germany) near the Neander
Valley (:. the name Neandertal) and was handed over to local science teacher, J.K.
Fuhlrott who consulted an expert, Hermann Schaaffhausen. It was named Homo
neanderthalensis by William King. The name however is now applied to all the fossil
material dating from the end of third inter-glacial and fourth glacial period.

Subsequently specimens have been discovered from Belgium, France, Spain, Romania
etc., thus giving Neandertals a European distribution. Later, however specimens were
also unearthed from Broken Hill in Zambia (1921) with the same characteristics. This
was named as “Rhodesian Man”.

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 In 1931-32, the specimens were discovered from Java, who however showed an early
evolutionary level than Neandertals (Solo Man) and in 1938, there were similar
discoveries from Central Russia (Teshik-Tash) ,Isreal (Mt: Caramel and Shanidar). So far
~300 Neandertals individuals are documented.

Thus, there are basically two types of Neandertals –

1. West European Type &

2. Non European Type

Characters of West European “Classic” Type

• Large cranium (could accomodate brain of modern size) ~ 1500cc.

• different shape with lower flatter crown bulging backwards to which neck muscles
are attached as a bun or chignon the Neandertal “bun”
• Receding Chin
• Large projecting Jaws & face
• Large teeth & palate
• Round orbits for eyes
• Extremely prominent brow ridges covering each eye and meeting across the bridge of
nose (beetle browed look)
• Short (5ft. tall) but powerfully built body
• Long dense bone robust and slightly curved giving Bandy-legged appearance. The
first specimen had curved bones leading people to think that Neandertals were pro-
simians but later specimens didn’t show this characteristic. It was later established
that the earlier specimen was an old man suffering with arthritis.
• Hands large, short stubby fingers
• Thick skull
• Use of variety of stone tools (Mousterian culture)
• Elaborate culture, skilful hunters
• probably practiced ritualized burial of the dead with flowers and ornaments (thus
probably believing in spiritual immortality).

Their remains date from an earlier age (about 110 Kya) in the Middle East than in Europe (~50
Kya).

Their “footprints” are also nicely preserved pointing to stocky proportions of feet (Picture).

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Characters of Non-European Neandertals

• Less massive
• Taller and more finely made
• legs and arms not as robust and curved
• skull more lofty
• face smaller, bony features like modern man.

Thus, Neandertals were a varied and widespread group. However, little is known about
the abrupt disappearance of Neandertal. Some of their populations may have died out,
whereas others may have merged into the new dominant forms. One hypothesis suggests
that they represent a separate offshoot of human line, differing from both the Homo
sapiens groups that preceded and followed it (Stringer & Gamble). In support of this
view are findings that recovered sequences of Neandertal mtDNA are outside the limits
of normal H. sapiens variability. According to Krings & Coworkers, this evidence
indicates that “Neandertal” mtDNA and the human ancestral mtDNA gene pool have
evolved as separate entities for substantial period of time and gives no support to the
notion that they should have contributed mtDNA to modern human gene pool. Rak
pointed out that extraordinarily large face was probably a biological innovation to exert
stronger biting forces on front teeth whereas heavy wearing of incisors and canines
indicated processing of tough food or hides or both. The expanded nasal chamber in
their face may have served as a radiator, warming and humidifying inspired air in the
dry, cold glacial climates that many European Neandertals inhabited. In other words, the
“classic” type characters were an adaptation to the environment (“cold”). Many
anthropologists, however feel that the Neandertals probably deserve a full membership
in H. sapiens as H. sapiens neanderthalensis.

Homo sapiens sapiens

The modern sapiens anatomically virtually indistinguishable from today’s humans, appeared
earlier in Africa (~170Kya) than elsewhere. It overlapped with Neandertals in the Middle East
for much of Neandertal’s history, but abruptly replaced them in Europe about 40Kya. By
12,000 years ago, and possibly earlier, modern humans had spread from northeastern Asia
across the Bering Land Bridge to north-western North America and then rapidly throughout
the Americas.

Evidence of early populations of Homo sapiens sapiens tends to be concentrated in Europe for
historical reasons, therefore, the tendency has been to use European samples to typify the
morphology of modern humans.

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Cro-Magnon

he fossils of modern man were discovered first from Dordogne in S. France (in 1868) in a
rock shelter called Cro-Magnon after a local harbit Magnon. Although they are probably the
best known evidence in the upper Paleolithic, material is now known from sites right across
Europe into Russia, from Kanjera and Kanam on the river Omo in (Africa), from Niah
(Malaysia) and Wadjak (East Java) in Asia from Lake Mungo and Kow Swamp in Australia
and from S. California (“Laguna skull”), Taber and Alberta in Canada in America.

Characters- height 6ft. (males) & 5ft.5” (females)

• lower leg longer than Neandertals (indicating swift footedness)

• skull with high forehead
• distinctive chin
• no heavy tori
• orthognathous
• teeth and jaw modern
• skull large but narrow
• broad face hence disharmonic
• brain large, high type
• Burial elaborate
• Use of stone tools and that made of bone (Aurignacian industry)
• More Asiatic in character, reached Europe as invaders from Asia
• Paleolithic art in caves (shows employment of artificial illumination in dark caves)

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The shape of the face changed because of:-

• Brace’s hypothesis – Neandertals used teeth to hold and cut as a plier but with the use
of tools, face suddenly straightened up.
• Pilbeam’s hypothesis – Use of speech particularly articulate speech necessitated
development of pharynx which in turn brought about
• changes in skull shape. This’s more plausible.

Cro Magnon (upper Paleolithic age)

• stone implements
• cultural evolution
• use of agriculture, domestic animals.

Summary of Hominid Evolution

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 Table 20-4, pg. 482 (Strickberger), Fig. 20-15, pg.483 (Strickberger)

Origin of Homo sapiens

There are 2 main views of the origin of modern humans from a Homo erectus ancestor:

1. The Single-Origin Hypothesis also known as the “Out of Africa” or “Noah’s Ark” model
proposes that modern humans (coloured sections) originated in one locality (Africa) and
then migrated to other continents where they replaced relict Homo erectus populations
(grey sections) that had entered these continents one million or more years ago.

This can also be termed as replacement hypothesis, because it holds that after H. erectus
spread from Africa to Asia and Europe and evolved into archaic H. sapiens, modern sapiens
evolved from archaic sapiens in Africa, spread throughout the world in a second expansion,
and replaced the populations of archaic sapiens without interbreeding with them to any
substantial extent. (Vigilant et. al., 1991)

Diagram from Stickberger Box 20-1

That is, modern sapiens that evolved from archaic sapiens in Africa was reproductively
isolated from Eurasian populations of archaic sapiens- i.e. it was a distinct biological species.
Most of the world’s populations of archaic sapiens became extinct due to competition, and
most genes in contemporary populations are descended from these carried by the population of
modern sapiens that spread from Africa.

The support for this hypothesis are following facts:

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 1. All the non-African mtDNA sequences are variants of the African sequence.
2. Most mtDNA sequence variability occurs among African populations suggesting
that these are oldest mtDNA populations.
Interestingly, like “Mitochondrial Eve” the “Y-chromosome Adam” was also of African
origin but African populations also received Y-chromosome returning from Asia.

3. Nei & Roychaudhry also suggested a single African Origin for H. sapiens with
subsequent widespread geographic divergence (by calculating genetic distances
between 26 human populations for 29 different nuclear genes).
4. Mountain & coworkers presented similar findings by working out polymorphism
for Alu chromosomal DNA sequences specific to human’s (Batzer, et.al.).

Although the dispute continues, Cann and coworker’s general conclusion for a single African
origin of H. sapiens still seems generally favored.

Taking all the many hominid variations into account, it is clear that “Instead of a ladder with
humans at the pinnacle, there is bush with humans as one little twig” (Foley, 1995).

II. The Multiple-Origin Hypothesis/”Candelabra” Model (because of its shape)

Modern humans originated in different localities independently of other such groups.

However, if each population evolved independently, how did they become so similar? Modern
humans may differ in color and other minor attributes, but they all share basic H. sapiens
traits. In answer some proponents or paleontologists of multiple origins (the “Multiregional”
Model, Wolpoff, 1989) suggest that probably some genes exchanged between continental
populations via cross migration(gene flow),enabling all these various evolving groups to reach
the same H.sapiens grade.

Conclusion

Human beings developed language, apart from other functions, which allowed him to transmit
knowledge and information to successive generations, in contrast to other animals. Not only
this, a more developed brain allowed the information to be stored and language helped to build
upon the accumulated experience of earlier generations. This acclerated the pace of cultural
adaptation and humans were no longer dependent on biological evolution.

A Note on cultural adaptation

A highly specialized hand, and bipedalism (which led to “freeing” the specialized hand so that
it could be used for other purposes e.g. making tools) are of course, important pre-requisites
for making tools. However, the real breakthrough came about with a large brain (1000cc. and
beyond). Tool manufacture calls for a high degree of motor skills and coordination of various
parts of the body as well as enormous concentration as well as certain measure of
conceptualization.

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I. Oldowan [Paleolithic (early Characterized by Tools were crude, rudimentary
stone age) 1.5 – 2.0 culture] found hunting & plant small-sized made from pebbles
in East Africa gathering used mainly for cutting plant foods,
breaking nuts, digging roots,
scrapping wood & obtaining honey.
Also used for skinning the meat of
animals and extracting bone narrow

II. Acheulian [named after a site Characterized by a Most important tool was
in North France (St. Acheul)] predetermined “handaxe”. These were pear-shaped
found in East, South and West “design” on the or tear drop shaped pointed at one
Africa, in Spain, France and stone. end and broad at the other. Tools
several other parts of Europe and much larger and remarkably
in West Asia, India and some symmetrical. Hand axe was a
other regions of South Asia “core” tool (made from larger
block or “core”). The other type
was “flake tools” fashioned from
small bits of flakes which come off
a block of stone on hitting.

III. Clactonian* & 700,000 years ago Characterized by turtle shaped* or

Levalloisian** cultures (lower round shaped** made of wood and
“Paleolithic” cultures) found in bone
some parts of North Europe and
far East

IV. Mousterian Culture (middle A product of Very specialized tools. Sixty

Paleolithic) (named after a Neanderthals varieties identified careful disposal
French site of Le Moustier) found Quarternary of dead bodies
in West Asia and Africa period P*and
independently H**
*Pleistocene
epoch (1.64mya –
12,000ya)
**Holocene
(12,000ya to
present)

V. Aurignacian Upper 35,000 years ago Tools increasingly specialized

made for specific purposes, highly
Solutrean Paleolithic - 30,000 years
ago refined. A typical tool was “burin”
Magdalenian (a kind of chisel). Tools were small
Cultures blades with parallel and extremely
sharp edges, sharp “leaf point”,

27
Discovered in France harpoons used for fishing. Tools
made of wood, bone, ivory and
(South) and Spain
antler. Composite tools composed
*developed/evolved in Eastern of several parts viz. leaf points
22,000 – 18,000
Europe (Hungary, Yugoslavia) joined to wooden rods making
years ago
and South West Asia, reached a spear-like weapon. Arrows often
peak in Western Europe poisoned. Tools also made for
a) No Aurignacian paintings but crude clothing (from hides), for
other beautifully carved objects chiseling wood, bone, ivory and
viz – ivory beads used for antler for decorative purposes
ornamentation and flute like culture also known for earliest
18,000 – 11,000 examples *of art (development of
wind instrument from bone
years ago thought and imagination) Cave
b) Solutrean Culture – cave paintings (on the roof of a cave in
paintings major art form colors Altamira) discovered by Don
used were natural pigments – Marcelino de Sautuola (1879) Heri
expert at carving figures on walls Breuil researched on 200
of caves (technique known as paleolithic cave painting. Other art
“bas-relief” or low – relief) objects also found viz. – handles of
c) Magdalenian Culture tools, statuettes and clay figures
with animal or geometric designs.
- Cave paintings reached fullest
development
- paintings deep inside caves
using animal fat lamps depicting
fantastic animals, humans
engaged in hunting and some
geometric patterns
[V. Gordon Childe refers to the
Magdalenian as the most
brilliant culture created by
hunter- gathers]

Throughout the Paleolithic period, one feature of social organization stood out – Collective
effort and sharing. However, the concepts began to undergo changes with the beginning of
food production lead to individual control.

Table 11.10: Brief inferences about adaptions, behavior, and ecological factors in
hominid evolution

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Reference:
1. Campbell, Bernard, G. (1982), Humankind Emerging, Third Edition, Little Brown
& Campany, Boston/Toronto.
2. Farooqui, Amar (2001), Early Social Formations, Second Edition, Manak
Publications Pvt. Ltd., New Delhi.
3. Futuyma, Douglas, J. (1997), Evolutionary Biology, Third Edition, Sinauer
Associates, Inc. Publishers, Sunderland, Massachusetts.
4. Lull, R.S. (1977), Organic Evolution, MacMillan Co., New York.
5. Moody, P.A. (1978), Introduction to Evolution, Third Edition, Kalyani Publishers,
New Delhi.
6. Strickberger, M.W., (2000), Evolution, Third Edition, Jones & Barlett Publ. Int.

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7. Wood, B.A. (1978), Outline Studies in Biology – Human Evolution, Chapman &
Hall (London), John Wiley & Sons, New York.

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