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Biology II Notes
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Human Evolution
       1. Describe primate origins and characteristics 
       2. Discuss the development of Hominids 
       3. Compare the two models for the origin of modern Homo Sapiens
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     Modern humans are in the primate family Hominidae.   We are the only living members of that family today.  In the past, australopithecines and other hominid genera (as well as other Homo species) have lived but are now extinct. 
Note: The evolutionary theory of human developement is perhaps one of the most volatile of the science world.  Every few years, new ideas and fossil evidence surfaces and causes changes in theories.  You can ask 5 leading paleoanthropologists of today to diagram the human family tree, and you'd probably get 5 different versions. 
     Australopithecines are thought to be the link between Ardipithecus ramidus and the Homo genus.  Some scientists put several larger species of australopithecenes into their own genus, Paranthropus.
     Up till now, ardipithecus and australopithecene fossils have only been found in Africa.  Although most have been found in the southern and eastern parts of the continent, some have recently been found in the north-central country of Chad. The first species in Homo, the  Homo habilis, has also been found only in Africa.  Some believe that the next step in the hominid tree splits in two.  Those that stayed in Africa were Homo ergaster, while Homo erectus is thought to be the first hominid to expand its range beyond Africa.
     Besides the major Australopithecenespecies detailed below, there are other where minor fossil evidence has been found.  The fossils found in Chad have been named Australopithecus bahrelghazali.  This species lived from 3.5 to 3 million years ago. Since the fossil evidence is limited, we don't know much about Australopithecus bahrelghazali. Also, some fossils found in east Ethiopia (dated about 2.5 million years old) have been called Australopithecus garhi.  Some say that a. garhi is a good candidate as an intermediary between australopithecenes and Homo habilis.  Fossils found within the last 5 years seemingly push back the hominid family tree even further, with Sahelanthropus
 tchadensis and Orrorin tugenensis dating back 6 million years.
    Generally speaking, the Australopithecenes were thought to be our primitive ancestors ... the first that started the switch to bipedalism and larger brains (among other characteristics).  In the human family tree, they are supposed to have been followed by the Homo species.  Homo habilis came first, followed by Homo erectus/ergaterHomo erectus/ergaster was followed by Homo sapiensHomo neanderthalensis is often thought to be a "cousin" to modern man.  Some think that Homo heidelbergensis may have been the common ancestor to both modern humans and Neanderthals.  Check the table at the bottom of the page for a summary of most hominid species.

Hominid characteristics: there are certain features that place a species in the Hominidae family.  The more advanced they appear, the closer the species is to modern humans. 
     - Bipedalism - walking upright; evidence for this includes a change in the femure/pelvis structure, the foramen magnum placed under the cranium, and shorter arms; early australopithecenes seem to show that they were starting to walk upright part of the time
     - Brain size - modern humans average about 1350cc in brain size; increasing brain size of hominids may indicate evolution towards H. sapiens; a more vertical forehead indicates the development of the frontal lobe of the brain
     - Speech - the ability to speak is a hominid characteristic; measuring brain casts tells us which hominids may have been able to speak; the first words may have begin with H. habilis
     - Tools - hominids are thought to be the first animal to make and use tools; the first tools were thought to have been made by H.habilis

     NOTE: both Sahelanthropus tchadensis and Orrorin tugenensis predate A. ramidus, but their discovery is so recent that there is not much written on how they may fit into the hominid family tree.
 

A. ramidus tooth Ardipithecus ramidus
     There is only one species in this genus, A. ramidus (sometimes A. ramidus kadabba).  Fossil evidence dates back to about 4.5 mya.  This, perhaps the earliest hominid, was identified from fossils found around Aramis, Ethiopia in the 1990s.  This individual was probably part of a radiation that included other hominids.  Because of its recent discovery, not much has been written about it.   If you take a look at the fossils recovered thus far [picture], you can see how difficult it might be to speculate as to it's skull characteristics. 
     The specimen’s hominid characteristics suggest that it was 'on the road ' to being bipedal.  The characteristics include a valgus knee angle, anteriorly placed foramen magnum and most importantly, a short, broad pelvis.

Australopithecenes

     Australopithecenes are considered to be the progenitors of the Homo genus.  There are two main types: gracile and robust. 
     Gracile australopithecenes: Their skeletons are less "bony."  For example, they do not have bony crests on their skulls to support large chewing muscles.  Their overall size was less than the robusts.  They started to appear about 4 million years ago and lasted till about 2 million years ago.  Gracile species include A. afarensis, A. africanus, and A. anamensis.
     Robust australopithecenes: An overall larger bone structure gives these species the robust title.  Their large molars and jawbones may mean that their diet required  tough chewing.    They started to appear about 4 million years ago and lasted till about 1 million years ago.  Robust species include A. aethiopicus, A. boisei and A. robustus.  Some scientists think that they are different enough from australopithecenes that they deserve their own genus, Paranthropus.


Australopithecus anamensis
     Australopithecus anamensis may have been the earliest australopithecine species.  They lived 4.2-3.9 million years ago in East Africa.  We know little about them because few fossils have been found.  It may have been partially bipedal.

Australopithecus afarensis
      Australopithecus afarensis lived 3.9-3.0 million years ago in East Africa (Ethiopia, Tanzania, Kenya.).  Their skeletons were still quite similar to early Pliocene apes.  For example, their legs were shorter than the later australopithecines.  They had large canines.  Some of the males had small sagittal crests. Afarensis had slender curved fingers like modern chimpanzees.  Because of these features, they may have been more adapted to tree climbing than bipedalism.  They were under 5 feet tall and weight about 100 pounds.  The females were significantly smaller (weighed 64% as much as males and were 70%  of the males' height).  Their slender structure also has led to the australopithecenes being known as gracile species. 

Homo

     The first member of the Homo genus is commonly thought to be Homo habilis (and/or Homo rudolfensis).  There is some disagreement as to how things evolved afterwards.  Homo ergaster and Homo erectus are thought to follow H. habilis. H. ergaster stayed in Africa, while H. erectus moved to the Asia about 2 million years ago.  While we know that modern humans are classified as Homo sapiens, the exact lineage is uncertain.  For a long time, it was assumed that we came from H. erectus.  Neanderthals (H. neanderthalensis) is thought to be a 'cousin' of H. sapiens.  Recent thought is that Homo heidelbergensis may be a common ancestor to H. sapiens and Neanderthals. 
     It is thought that H. habilis may have been the first tool makers. H. erectus advanced the art of toolmaking and was the first to utilize fire. 
 

Homo habilis / Homo rudolfensis
     Though many paleoanthropologists classify Homo habilis ("handy man") as a Homo sapiens ancestor, the exact bridge from one species to another has been debated since the Leakeys found the first specimen (OH 7) in 1960.
     Homo habilis is thought to be the first hominid to use simple tools. Its brain size is bigger than that of the australopithecenes.  Experts can't even agree on exactly which specimens should be definitely considered Homo habilis.
     Adding to the confusion over Homo habilis, some specimens have sufficient differences that another species has been proposed.  V.P. Alexeev, using thee KNM-ER 1470 as a type specimen, suggested the species Homo rudolfensis in 1986. Homo rudolfensis may have been the ancestor to the Homo habilis.  Perhaps they were two separate species.  Some even believe it should be classified as an australopithecene.

H. habilis


H. erectus
Homo erectus / Homo ergaster
     The Leakeys have long proposed that Homo erectus was the ancestor of modern Homo sapiens.  Other opinions do exist (see Homo sapiens below).  One theory is the Homo erectus left Africa to populate Asia and perhaps Europe, but Homo ergastor stayed in Africa and evolved into Homo sapiens who then replaced Homo erectus.  Of course, if you believe in the multiregional theory of H. sapiens origins, then you might say that H. ergaster evolved into a form of H. erectus and then into H. sapiens in Africa, while H. erectus evolved into H. sapiens in other parts of the world.
     Homo erectus is thought to have advanced the art of tool making.  They are also thought to be the first hominids to use fire.  Their brain capacity averaged over 1000cc (while H. habilis averaged 750cc).

Homo heidelbergensis
     Homo heidelbergensis fossils have been found in Africa and eastern & western Europe.  Their fossils date from about 600,000 to 200,000 years ago.  It's features are similar to Homo erectus, but its other features, the time frame, and location seem to indicate that it may have been the ancestor to Neanderthals and possibly H. sapiens

H. sapiens
Homo sapiens
     This is "us."  The modern traits of today's human started appearing about 100,000 years ago (though recent discoveries may push that back another 50,000 years).  The earliest fossil record of Homo sapiens is from Africa.  Sometimes early Homo sapiens is known as archaic or Cro-Magnon man. 
     Brain size is about 1350cc and we are fully bipedal.  Although we have hair over most of our bodies, at some point we lost the thick coat we once had. [see Why Humans and Their Fur Parted Ways]  Theories range from the idea that humans went through a semi-aquatic phase and lost it ... to the idea that it was lost through natural selection (more hair = more parasites & diseases).
Two models for the origin of the species, Homo sapiens
     Just as there is disagreement over the order of the entire hominid family tree, there are 2 different schools of thought on how our species came to be.  These 2 theories that developed in the 1980s are known as the 'Out of Africa' theory and the multiregional theory. [View Graphic]
     'Out of Africa' Theory:
          - This idea says that Homo sapiens evolved in Africa, and THEN spread out to the whole world. 
          - These Homo sapiens would replace any existing populations that were their (e.g. Neanderthals in Europe).
          - This is the leading theory because of the hominid fossils and fossil artifacts found in Africa.  Also, mitochondral DNA studies (1987 et. al) suggest a common ancestor in Africa (the "Eve" hypothesis). But recent DNA research is casting doubt on this idea. [see Is Out of Africa Going Out the Door?]
          - The different races are a more modern development.
     Multiregional Continuity theory:
          - This idea says that about 7 mya, Homo erectus left Africa and settled in different parts of the world. 
          - They (or their descendants) evolved into Homo sapiens in those different parts of the world. 
          - For example, Homo erectus evolved into Homo sapiens in Asia, Homo ergaster evolved into Homo sapiens in Africa, Homo neanderthalensis evolved into Homo sapiens in Europe, etc. 
          - Natural selection in those different areas led to the variations in Homo sapiens races.
         - One of the problems with this theory is that in some areas, Homo sapiens existed at the same time as their proposed ancestors.

These hominids currently thought NOT to be in our direct human lineage.
Australopithecus africanus
     Until 1925, it was thought that human ancestors would only be found in Europe.  But that year, Raymond Dart labeled his find (the Taung child) as a hominid because it was partially bipedal and created the genus australopithecus ("southern ape"). 
     Australopithecus africanus lived about 3.0-2.5 million years ago in East and South Africa.  Their bone structure was less ape-like than earlier species of australopithecines.  Their skeletons were still small and lightweight ... even smaller than a.afarensis.  Unlike a. afarensis, their canines did not project beyond the other teeth. 
A. africanus

A. robustus
Some scientists give the larger australopithecenes their own category (paranthropus), though some paleoanthropologists still call them australopithecenes.
     Paranthropus or A. aethiopicus - the earliest robust species; lived about 2.5 million years ago; found only in East Africa; smaller brain than the other robust species; large sagittal crest. 
     Paranthropus or A. boisei - the largest of the three Paranthropes; found in East Africa; lived about 2.3-1.4 million years ago; large canines and grinding teeth 
     Paranthropus or A. robustus - lived about 1.9-1.0 million years ago in South Africa; smaller version of the p. boisei 

Homo sapeins neanderthalensis or
Homo neanderthalensis
      Neanderthals started appearing about 130,000 years ago, and they seem to have died out about 30,000 years ago.  Their fossil evidence is mainly in western Europe.  Some people think that rather than becoming extinct, surviving Neanderthals mixed in with the Homo sapien population. 
H. neanderthalensis


Comparative Brain Sizes:
A. afarensis
H. habilis
H. erectus
H. sapiens neand.
H. sapiens
Chimp Gorilla
400cc
750cc
900-1200cc*
1400cc
1350cc
400cc
500cc
*900cc for early specimens (>1 mya), 1200cc for later ones (1/2 mya)

The Hominid Family Tree

     Here is one of the more commonly accepted views of the hominid family tree. 
     Other versions can be found at the Hominid Family Tree page.

LINKS:
     - Evolution[PBS]
          - Human Evolution
     - Early Hominid Evolution
     - Becoming Human  [ws]
     - Hominid Species
     - Archaeology Info
          - Human Ancestry
               - Hall of Skulls
          - Hominidae
               - Sahelanthropus tchadensis
               - Kenyanthropus
     - Human Evolution 
     - Human Origins Program
     - The human past up to 200,000 years ago [PDF]
     - Archaeology 2.5mya to 30 kya [PDF]
     - Human Evolution: The Fossil Evidence in 3D
     - Human Evolution Links
     - Human Evolution
     - The Evolution of the Human
     - Natural History and Ecology of Homo Sapiens
     - Early Modern Homo Sapiens
     - Oludvai Gorge
     - Introducing the Skeletons in Your Closet 
     - Paleoanthropology: A Journey Through Time
     - Paleoanthropology in the 1990's
          - The 'Replacement' or 'Out of Africa 2' Hypothesis
          - Multiregional Evolution
     - Hunting Hominids
     - Did Humans and Neandertals Battle for Control of the Middle East?
     - Did Our Species Mate With Other Human Species? 
     - African Bone Tools Dispute Key Idea About Human Evolution 
     - Human Fossil Adds Fuel to Evolution Debate
     - A Look at Modern Human Origins
     - The Physical Characteristics of Humans
     - Early Humans Definition of Homo Minimal List [PDF]
     - Genetic Study Revises Human Family Tree
     - Human Biology Web Site
          - A Hominid Family Tree 
          - A Hominid Time Line 
     - Origins of Modern Humans:  Multiregional or Out of Africa?
     - Is Out of Africa Going Out the Door?
     - African Eve or multiregional origins? Maybe both
     - Skulls lend credibility to 'out of Africa' theory
     - Evolution: Out of Africa and the Eve Hypothesis
     - DNA Tests Reveal Modern Man Is 'Out of Africa' 
     - Human Evolution: You Try It
     - Journey into the World of Cladistics
     - Evolution
          - History of Evolutionary Thought
     - Time-Space Chart of Hominid Fossils
     - 
Skulls

(A) Pan troglodytes, chimpanzee, modern 
                  (B) Australopithecus africanus, STS 5, 2.6 My 
                  (C) Australopithecus africanus, STS 71, 2.5 My 
                  (D) Homo habilis, KNM-ER 1813, 1.9 My 
                  (E) Homo habilis, OH24, 1.8 My 
                  (F) Homo rudolfensis, KNM-ER 1470, 1.8 My 
                  (G) Homo erectus, Dmanisi cranium D2700, 1.75 My 
                  (H) Homo ergaster (early H. erectus), KNM-ER 3733, 1.75 My 
                  (I) Homo heidelbergensis, "Rhodesia man," 300,000 - 125,000 y 
                  (J) Homo sapiens neanderthalensis, La Ferrassie 1, 70,000 y 
                  (K) Homo sapiens neanderthalensis, La Chappelle-aux-Saints, 60,000 y 
                  (L) Homo sapiens neanderthalensis, Le Moustier, 45,000 y 
                  (M) Homo sapiens sapiens, Cro-Magnon I, 30,000 y 
                  (N) Homo sapiens sapiens, modern 

Hominid Skulls

 
 

Hominid Jaws



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Species
Date Found
Where Found
Dated At
Fossils Found
Notes
Specimens (Type specimen)
Sahelanthropus tchadensis
2001
Chad
6-7 myo
nearly complete cranium, lower jaw fragments & teeth 350cc brain size TM 266-01-060-1
Orrorin tugenensis
2001
Kenya
6 myo
arm & thigh bone fragments, cranium fragment, lower jaws & teeth    BAR 1000'00
Ardipithecus ramidus
1994
Ethiopia
4.4-5.8 myo
skull fragments, teeth; a 45% complete skeleton has been recently found   17 individuals at ARA-VP-6/1 & ARA-VP-7/2
Australopithecus anamensis
1965
Kenya
3.9-4.2 myo
tibia fragment, skull fragments, jawbones, & teeth the 12 fossils had been found in 1988; named 1994 KP 271, KP 29281, KP 29285
Australopithecus afarensis
1973
Ethiopia, Tanzania
3-3.9 myo
fragments from throughout the skeleton 375-550cc brain size; Lucy is a 40% complete skeleton; the Laetoli footprints may be from A. afarensis AL 129-1, AL 288-1 (Lucy), AL 333 Site (First Family); AL 444-2
Kenyanthropus platyops
1999
Kenya
3.5 myo
skull fragments, teeth   KNM WT 40000
Australopithecus africanus
1924
South Africa
2-3 myo
skull fragments, teeth, vertebrae, pelvis, rib & femur fragments 420-500cc brain size "Taung child", TM 1512. Sts 5 (Mrs Ples), Sts 14, Stw 573 ? (Little Foot)
Australopithecus garhi
1997
Ethiopia
2.5 myo
skull fragments, teeth   BOU-VP-12/130
Australopithecus aethiopicus
1985
Kenya
2.3-2.6 myo
almost complete skull 410cc brain size; 1 major specimen (the Black Skull); some classify as a Paranthropus Omo 18, KNM-WT17000 (A. walkeri)
Australopithecus robustus
1938
South Africa
1.5-2 myo
skull fragments, teeth, a few skeletal fragments 530cc brain size;some classify as a Paranthropus TM 1517, SK 48, DNH 7 (Eurydice), DNH 8 (Orpheus)
Australopithecus boisei
1959
Tanzania, Kenya, Ethiopia
1.1-2.1 myo
skull fragments, teeth 530cc brain size; some classify as a Paranthropus OH 5 (Nutcracker man), KNM-ER 406, KNM-ER 732, KGA10-525
Homo habilis
1960
Tanzania
1.5-2.4 myo
skull fragments, teeth 650cc brain size; a partial skeleton was recently found OH 7 (Jonny's child), OH 8, OH 13 (Cindy), OH 16 (George), OH 24 (Twiggy), OH 62 (Dik-dik hominid), OH 65; ***
Homo rudolfensis           KNM-ER 1470
Homo georgicus
2002
Georgia
1.8 myo
skull fragments, teeth 600cc brain size D2700, D2280, D2282
Homo erectus
1891
Africa, Asia, Europe
300,000-1.8 myo
 skull fragments, teeth, skeletal bones 1000cc brain size Trinil 2 (Java Man), Peking Man Site, Sangiran 2, OH 9 (Chellean Man), OH 12 (Pinhead), Sangiran 17, KNM-ER 3733?, KNM-WT 15000?
Homo ergaster         some say the H.erectus fossils in Africa should be H.ergaster KNM-ER 3733?, KNM-WT 15000? (Turkana boy), KNM-ER 992
Homo antecessor
1977
Spain
800,000 kyo
facial fragments 10-11 yr old child ATD6-69
Homo heidelbergensis
1907
Europe
500,000-? kyo
fairly complete skeletons 1200cc brain size Mauer 1 (Heidelberg Man), Arago XXI (Tautavel Man)
Homo neanderthalensis
1856
Europe, Middle East
30,000-230,000 kyo
fairly complete skeletons 1450cc brain size Feldhofer, Spy 1 and 2, Krapina Site, La-Chapelle-aux-Saints, Shanidar Site, Saint-Cesaire, 
Homo sapiens (archaic)     ???-150,000 kyo fairly complete skeletons 1350cc brain size Rhodesian Man, Petralona 1, Atapuerca 5
Homo sapiens (modern)
1868
Europe
present-30,000 kyo complete skeletons 1350cc brain size Cro-Magnon Site, many more
kyo = thousand years ago; myo = million years old

*** Several fossils may be H. habilis or H. rudolfensis or ???;  these include: KNM-ER 1470 (Kenya), KNM-ER 1805 (Mystery Skull; Kenya), KNM-ER 1813 (Kenya), Stw 53 (S.Africa)

Other species may include: A. bahrelghazali ? (KT 12/H1); A. crassidens (SK 6), A. praegens (KNM-T1 13150), Homo louisleakeyi (OH 9), Homo microcranous (KNM-ER 1813)

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