Sahelanthropus Tchadensis

Sahelanthropus tchadensis is an extinct hominid species that is dated to about 7 million years ago. Whether it can be regarded as part of the Hominina tree is unclear; there are arguments both supporting and rejecting it. Another complication in its classification is that it is older than the humanchimpanzee divergence (estimated to 6.3 to 5.4 million years ago) seen in genetic data,[2] and that there are few if any specimens other than the partial cranium known as Touma.

Fossils
Existing fossils a relatively small cranium nicknamed Touma ("hope of life" in the local Dazaga language of Chad in central Africa), five pieces of jaw and some teeth make up a head that has a mixture of derived and primitive features. The braincase, being only 320 cm to 380 cm in volume is similar to that of extant chimpanzees and is notably less than the approximate human volume of 1350 cm. The teeth, brow ridges, and facial structure differ markedly from those found in Homo sapiens. Cranial features show a flatter face, u-shaped dental arcade, small canines, an anterior foramen magnum, and heavy brow ridges. No postcranial remains have been recovered. The fossil suffered a large amount of distortion during the time of fossilisation and discovery. Since no postcranial remains (bones below the skull) have been discovered, it is as of yet unknown whether Sahelanthropus tchadensis was indeed bipedal or two-footed, although claims for an anteriorly placed foramen magnum suggests that this may have been the case, some paleontologists have disputed this interpretation of the basicranium. Its canine wear is similar to other Miocene apes.[3] Moreover, according to recent information, the femur of a hominid may have been discovered alongside the cranium but never published.[4]

Discovery
The fossils were discovered in the Djurab desert of Chad by a team of four led by Michel Brunet; three Chadians, Adoum Mahamat, Djimdoumalbaye Ahounta and Gongdib Fanon, and Frenchman, Alain Beauvilain et al.[5][6] All known material of Sahelanthropus were found between July 2001 to March 2002 at three sites (TM 247, TM 266 which yielded most of the material, and TM 292). The discoverers claimed that S. tchadensis is the oldest known human ancestor after the split of the human line from that of chimpanzees. The bones were found far from most previous hominin fossil finds, which are from Eastern and Southern Africa. However, an Australopithecus bahrelghazali mandible was found in Chad by Beauvilain A., Brunet M. and Moutaye A.H.E. as early as 1995.[7] With the sexual dimorphism known to have existed in early hominids, the difference between Ardipithecus and Sahelanthropus may not be large enough to warrant a separate species for the latter.[8]

Relationship to humans and chimpanzees

Sahelanthropus may represent a common ancestor of humans and chimpanzees; no consensus has been reached yet by the scientific community. The original placement of this species as a human ancestor but not a chimpanzee ancestor would complicate the picture of human phylogeny. In particular, if Touma is a direct human ancestor, then its facial features bring the status of Australopithecus into doubt because its thickened brow ridges were reported to be similar to those of some later fossil hominids (notably Homo erectus), whereas this morphology differs from that observed in all australopithecines, most fossil hominids and extant humans. Another possibility is that Touma is related to both humans and chimpanzees, but is the ancestor of neither. Brigitte Senut and Martin Pickford, the discoverers of Orrorin tugenensis, suggested that the features of S. tchadensis are consistent with a female proto-gorilla. Even if this claim is upheld, then the find would lose none of its significance, for at present precious few chimpanzee or gorilla ancestors have been found anywhere in Africa. Thus if S. tchadensis is an ancestral relative of the chimpanzees (or gorillas) then it represents the first known member of their lineage. Furthermore, S. tchadensis does indicate that the last common ancestor of humans and chimpanzees is unlikely to resemble chimpanzees very much, as had been previously supposed by some paleontologists.[9][10] Unfortunately, the exact age of the fossil is somewhat hard to determine. While molecular clocks are increasingly found to be far less reliable than initially believed,[11] sediment isotope analysis which yielded an age of about 7 million years is generally considered quite reliable. In this case however, the fossils were found exposed in loose sand; co-discoverer Beauvilain cautions that such sediment can be easily moved by the wind, unlike packed earth.[12] In fact, Touma was probably reburied in the recent past. Taphonomic analysis reveals the likelihood of one, perhaps two, burial(s) which seemingly occurred after the introduction of Islam in the region. Two other hominid fossils (a

left femur and a mandible) were in the same grave along with various mammal remains. The sediment surrounding the fossils might thus not be the material the bones were originally deposited in, making it necessary to corroborate the fossil's age by some other means.[13] The fauna found at the site namely the anthracotheriid Libycosaurus petrochii and the suid Nyanzachoerus syrticus suggests an age of more than 6 million years, as these species were probably extinct already by that time.[14]

Orrorin Tugenensis

Orrorin tugenensis was named in July 2001 on the basis of fossils discovered in the Lukeino Formation, near Lake Baringo in western Kenya (Senut et al. 2001). The fragmentary remains include portions of arm and thigh bones, lower jaws, and teeth. They date to between 6.1 and 5.8 mya and are therefore of Miocene age. They were discovered by a expedition led by Brigitte Senut and Martin Pickford of the Musum national d'Histoire naturelle in 2000. This hominid is the only member of the genus Orrorin. The scanty remains assigned to Orrorin tugenensis suggest it was bipedal (unlike Sahelanthropus tchadensis, which was once billed as the earliest hominid, but now considered a Miocene ape). The limb bones, about 50 percent longer than those of Lucy, suggest that Orrorin tugenensis was about the size of a chimpanzee. Its discoverers have claimed Orrorin tugenensis was adapted to both bipedality and tree climbing, and that it was a direct human ancestor, with the australopithecines as an extinct offshoot not ancestral to modern humans. Other scientists are skeptical of these claims, due to the highly fragmentary nature of the remains (Aiello and Collard 2001). But an additional paper (Galik et al. 2004) has provided further evidence of bipedality in this form. The Orrorin femur is more similar to that of H. sapiens than is Lucy's. The brain size of this hominid is unknown, since there is no skull material allowing cranial capacity to be measured. Fossils of other organisms from the Lukeino Formation show this hominid lived in a dry evergreen forest habitat, which suggests it probably had a diet similar to that of a modern ape. This contradicts the many theories depicting earliest humans as savanna hunters (Orrorin tugenensis remains long predate stone tools and the first use of fire). Thus, if the bipedality of O. tugenensis is confirmed, the only possible conclusion will be that human bipedalism actually arose in a forest-dwelling ancestor and not in the descendants of a quadrupedal form that moved out into the open savanna. Orrorin tugenensis is considered to be the second-oldest (after Sahelanthropus) known hominin ancestor that is possibly related to modern humans, and it is the only species classified in genus Orrorin. Orrorin is significant because it can be an early bipedal hominin. The name was given by the discoverers who found Orrorin fossils in the Tugen Hills of Kenya in 2000. By analysing radiometric decay (KAr dating), paleomagnetism, and biochronology the age of the specimen have been estimated to 6 to 5.8 million years (Ma). At present, 20 fossils have been found at four sites in the Lukeino Formation: of these, the fossils at Cheboit and Aragai are the oldest (6.1 Ma), while those in Kapsomin and Kapcheberek are found in the upper levels of the formation (5.7 Ma).

Fossils
The 20 specimen found this far include: the posterior part of mandible in two pieces; a symphysis and several isolated teeth; three fragments of femurs; a partial humerus; a proximal phalanx; and a distal thumb phalanx.

Orrorin had small teeth relative its body size. Its dentition differs from that found in Australopithecus' in that its cheek teeth are smaller and less elongated mesiodistally; and from Ardipithecus in that its enamel is thicker. The dentition differs from both these species in the presence of mesial groove on the upper canines. The canines are ape-like but reduced, like those found in Miocene apes and female chimpanzees. Orrorin had small post-canines and was microdont like modern humans, whereas robust Australopithecenes were megadont. In the femur, the head is spherical and rotated anteriorly; the neck is elongated and oval in section; and the lesser trochanter protrudes medially. While this suggest that Orrorin was bipedal, the rest of the postcranium indicates it climbed trees. while the proximal phalanx is curved, the distal pollical phalanx is of human proportions and have thus been associated with toolmaking, but should probably be associated with grasping abilities useful for tree-climbing in this context. After the fossils were found in 2000, they were held at the Kipsaraman village community museum, but the museum was subsequently closed. Since then, according to the Community Museums of Kenya chairman Eustace Kitonga, the fossils are stored at a secret bank vault in Nairobi.

Classification
If Orrorin proves to be a direct human ancestor, then australopithecines such as Australopithecus afarensis ("Lucy") may be considered a side branch of the hominid family tree: Orrorin is both earlier, by almost 3 million years, and more similar to modern humans than is A. afarensis. The main similarity is that the Orrorin femur is morphologically closer to that of H. sapiens than is Lucy's; there is, however, some debate over this point. Other fossils (leaves and many mammals) found in the Lukeino Formation show that Orrorin lived in dry evergreen forest environment, not the savanna assumed by many theories of human evolution.

Discovery
The team that found these fossils in 2000 was led by Brigitte Senut and Martin Pickford from the Musum national d'histoire naturelle. The discoverers conclude that Orrorin is a hominin on the basis of its bipedal locomotion and dental anatomy; based on this, they date the split between hominins and African great apes to at least 7 million years ago, in the Messinian. This date is markedly different from those derived using the molecular clock approach, but has found general acceptance among paleoanthropologists.

Ardipithecus
Ardipithecus is a fossil hominoid, described by its discoverers as a very early hominin genus. Two species are described in the literature: A. ramidus, which lived about 4.4 million years ago during the early Pliocene, and A. kadabba, dated to approximately 5.6 million years ago (late Miocene).

Ardipithecus ramidus
A. ramidus was named in September 1994. The first fossil find was dated to 4.4 million years ago on the basis of its stratigraphic position between two volcanic strata: the basal Gaala Tuff Complex (GATC) and the Daam Aatu Basaltic Tuff (DABT). The name Ardipithecus ramidus stems mostly from the Afar language, in which Ardi means "ground/floor" (borrowed from either the Semitic root in either Amharic or Arabic) and ramid means "root". The pithecus portion of the name is from the Greek word for "monkey". Like most primates, but unlike all previously recognized hominins, it had a grasping hallux or big toe adapted for locomotion in the trees. However, its describers claim that other features of its skeleton reflect adaptation to bipedalism on the ground as well. Like later hominins, Ardipithecus had reduced canine teeth. Its brain was small and comparable in size to that of the modern chimpanzee.

In 19921993 a research team headed by Tim White discovered the first A. ramidus fossilsseventeen fragments including skull, mandible, teeth and arm bonesfrom the Afar Depression in the Middle Awash river valley of Ethiopia. More fragments were recovered in 1994, amounting to 45% of the total skeleton. This fossil was originally described as a species of Australopithecus, but White and his colleagues later published a note in the same journal renaming the fossil under a new genus, Ardipithecus. Between 1999 and 2003, a multidisciplinary team led by Sileshi Semaw discovered bones and teeth of nine A. ramidus individuals at As Duma in the Gona Western Margin of Ethiopia's Afar Region.[5] The fossils were dated to between 4.32 and 4.51 million years old. Ardipithecus ramidus had a small brain, measuring between 300 and 350 cm3. This is about the same size as a modern bonobo or female common chimpanzee brain, but much smaller than the brain of australopithecines like Lucy (~400 to 550 cm3) and roughly 20% the size of the modern Homo sapiens brain. Like common chimpanzees, A. ramidus was much more prognathic than modern humans. The teeth of A. ramidus lacked the specialization of other apes, and suggest that it was a generalized omnivore and frugivore (fruit eater) with a diet that did not depend heavily on fibrous plants, ripe fruit or hard or abrasive food. The size of the upper canine tooth in A. ramidus males was not distinctly different from that of females. Their upper canines were less sharp than those of modern common chimpanzees in part because of this decreased upper canine size, as larger upper canines can be honed through wear against teeth in the lower mouth. The features of the upper canine in A. ramidus contrast with the sexual dimorphism observed in common chimpanzees, where males have significantly larger and sharper upper canine teeth than females. The less pronounced nature of the upper canine teeth in A. ramidus has been used to infer aspects of the social behavior of the species and more ancestral hominids. In particular, it has been used to suggest that the last common ancestor of hominids and African apes was characterized by relatively little aggression between males and between groups. This is markedly different from social patterns in common chimpanzees, among which intermale and intergroup aggression are typically high. Researchers in a 2009 study said that this condition "compromises the living chimpanzee as a behavioral model for the ancestral hominid condition." A. ramidus existed more recently than the most recent common ancestor of humans and chimpanzees (CLCA or Pan-Homo LCA), and thus is not fully representative of that common ancestor. Nevertheless, it is in some ways unlike chimpanzees, suggesting that the common ancestor differs from the modern chimpanzee. After the chimpanzee and human lineages diverged, both underwent substantial evolutionary change. Chimp feet are specialized for grasping trees; A. ramidus feet are better suited for walking. The canine teeth of A. ramidus are smaller, and equal in size between males and females, which suggests reduced male-to-male conflict, increased pair-bonding, and increased parental investment. "Thus, fundamental reproductive and social behavioral changes probably occurred in hominids long before they had enlarged brains and began to use stone tools," the research team concluded. Ardi On October 1, 2009, paleontologists formally announced the discovery of the relatively complete A. ramidus fossil skeleton first unearthed in 1994. The fossil is the remains of a small-brained 50-kilogram (110 lb) female, nicknamed "Ardi", and includes most of the skull and teeth, as well as the pelvis, hands, and feet. It was discovered in Ethiopia's harsh Afar desert at a site called Aramis in the Middle Awash region. Radiometric dating of the layers of volcanic ash encasing the deposits suggest that Ardi lived about 4.4 million years ago. This date, however, has been questioned by others. Fleagle and Kappelman suggest that the region in which Ardi was found is difficult to date radiometrically, and they argue that Ardi should be dated at 3.9 million years. The fossil is regarded by its describers as shedding light on a stage of human evolution about which little was known, more than a million years before Lucy (Australopithecus afarensis), the iconic early human ancestor candidate who lived 3.2 million years ago, and was discovered in 1974 just 74 km (46 mi) away from Ardi's discovery site. However, because the "Ardi" skeleton is no more than 200,000 years older than the earliest fossils of Australopithecus, and may in fact be younger than they are, some researchers doubt that it can represent a direct ancestor of Australopithecus. Some researchers infer from the form of her pelvis and limbs and the presence of her abductable hallux, that "Ardi" was a facultative biped: bipedal when moving on the ground, but quadrupedal when moving about in tree branches. A. ramidus had

a more primitive walking ability than later hominids, and could not walk or run for long distances. The teeth suggest omnivory, and are more generalised than those of modern apes.

Ardipithecus kadabba
Ardipithecus kadabba is "known only from teeth and bits and pieces of skeletal bones", and is dated to approximately 5.6 million years ago. It has been described as a "probable chronospecies" (i.e. ancestor) of A. ramidus. Although originally considered a subspecies of A. ramidus, in 2004 anthropologists Yohannes Haile-Selassie, Gen Suwa, and Tim D. White published an article elevating A. kadabba to species level on the basis of newly-discovered teeth from Ethiopia. These teeth show "primitive morphology and wear pattern" which demonstrate that A. kadabba is a distinct species from A. ramidus.[15] The specific name comes from the Afar word for "basal family ancestor".

Lifestyle
The toe and pelvic structure of A. ramidus suggest to some researchers that the creature walked upright. According to Scott Simpson, the Gona Project's physical anthropologist, the fossil evidence from the Middle Awash indicates that both A. kadabba and A. ramidus lived in "a mosaic of woodland and grasslands with lakes, swamps and springs nearby," but further research is needed to determine which habitat Ardipithecus at Gona preferred.

Contrary Views
The fossils of Ardipithecus have not yet been studied by researchers beyond the original (2009) group of describers, and the paleobiology and relationships of these creatures are the subject of controversy. Skeptics claim that many of the allegedly hominin-like features seen in the Ardipithecus material are found elsewhere among living and fossil primates, and that claims about its hominin status and locomotor habits are not adequately supported by the available evidence.

Australopithecus anamensis
Australopithecus anamensis (or Praeanthropus anamensis) is a stem-human species that lived approximately four million years ago. Nearly one hundred fossil specimens are known from Kenya and Ethiopia, representing over 20 individuals.

Discovery
The first fossilized specimen of the species, though not recognized as such at the time, was a single fragment of humerus (arm bone) found in Pliocene strata in the Kanapoi region of East Lake Turkana by a Harvard University research team in 1965. The specimen was tentatively assigned at the time to Australopithecus and dated about four million years old. Little additional information was uncovered until 1987, when Canadian archaeologist Allan Morton (with Harvard University's Koobi Fora Field School) discovered fragments of a specimen protruding from a partially eroded hillside east of Allia Bay, near Lake Turkana, Kenya.

In 1994, the London-born Kenyan paleoanthropologist Meave Leakey and archaeologist Alan Walker excavated the Allia Bay site and uncovered several additional fragments of the hominid, including one complete lower jaw bone which closely resembles that of a common chimpanzee (Pan troglodytes) but whose teeth bear a greater resemblance to those of a human. In 1995, Meave Leakey and her associates, taking note of differences between Australopithecus afarensis and the new finds, assigned them to a new species, A. anamensis, deriving its name from the Turkana word anam, meaning "lake". Leakey determined that this species was independent of many others. It does not represent an intermediate species of any type. Although the excavation team did not find hips, feet or legs, Meave Leakey believes that Australopithecus anamensis often climbed trees. Tree climbing was one behavior retained by early hominins until the appearance of the first Homo species about 2.5 million years ago. A. anamensis shares many traits with Australopithecus afarensis and may well be its direct predecessor. Fossil records for A. anamensis have been dated to between 4.2 and 3.9 million years ago, with recent findings from stratigraphic sequences dating to about 4.14.2million years ago. Specimens have been found between two layers of volcanic ash, dated to 4.17 and 4.12 million years, coincidentally when A. afarensis appears in the fossil record. The fossils (twenty one in total) include upper and lower jaws, cranial fragments, and the upper and lower parts of a leg bone (tibia). In addition to this, the aforementioned fragment of humerus found thirty years ago at the same site at Kanapoi has now been assigned to this species. In 2006, a new A. anamensis find was officially announced, extending the range of A. anamensis into north east Ethiopia. These new fossils, sampled from a woodland context, include the largest hominid canine tooth yet recovered and the earliest Australopithecus femur. The find was in an area known as Middle Awash, home to several other more modern Australopithecus finds and only six miles away from the discovery site of Ardipithecus ramidus, the most modern species of Ardipithecus yet discovered. Ardipithecus was a more primitive hominid, considered the next known step below Australopithecus on the evolutionary tree. The A. anamensis find is dated to about 4.2 million years ago, the Ar. ramidus find to 4.4 million years ago, placing only 200,000 years between the two species and filling in yet another blank in the preAustralopithecus hominid evolutionary timeline.

Australopithecus afarensis
Australopithecus afarensis is an extinct hominid that lived between 3.9 and 2.9 million years ago. A. afarensis was slenderly built, like the younger Australopithecus africanus. It is thought that A. afarensis was more closely related to the genus Homo (which includes the modern human species Homo sapiens), whether as a direct ancestor or a close relative of an unknown ancestor, than any other known primate from the same time.[2] The most famous fossil is the partial skeleton named Lucy (3.2 million years old) found by Donald Johanson and colleagues, who, in celebration of their find, played the Beatles song Lucy in the Sky with Diamonds over and over.[3][4][5](p234)

Localities
Australopithecus afarensis fossils have only been discovered within northern Africa. Despite Laetoli being the type locality for A. afarensis, the most extensive remains assigned to the species are found in Hadar, Afar Region of Ethiopia, including the above-mentioned "Lucy" partial skeleton and the "First Family" found at the AL 333 locality. Other localities bearing A. afarensis remains include Omo, Maka, Fejej and Belohdelie in Ethiopia, and Koobi Fora and Lothagam in Kenya.

Physical characteristics

Compared to the modern and extinct great apes, A. afarensis has reduced canines and molars, although they are still relatively larger than in modern humans. A. afarensis also has a relatively small brain size (~380430 cm3) and a prognathic face (i.e. a face with forward projecting jaws).

The image of a bipedal hominid with a small brain and primitive face was quite a revelation to the paleoanthropological world at the time. This was due to the earlier belief that an increase in brain size was the first major hominin adaptive shift. Before the discoveries of A. afarensis in the 1970s, it was widely thought that an increase in brain size preceded the shift to bipedal locomotion. This was mainly because the oldest known hominins at the time had relatively large brains (e.g. KNMER 1470, Homo rudolfensis, which was found just a few years before Lucy and had a cranial capacity of ~800 cm).

Skeletal morphology and locomotion

There is considerable debate regarding the locomotor behaviour of A. afarensis. Some believe that A. afarensis was almost exclusively bipedal, while others believe that the creatures were partly arboreal. The anatomy of the hands, feet and shoulder joints in many ways favour the latter interpretation. The curvature of the finger and toe bones (Phalanges) approaches that of modern-day apes, and is suggestive of their ability to efficiently grasp branches and climb. Alternatively, the loss of an abductable great toe and therefore the ability to grasp with the foot (a feature of all other primates) suggests that A. afarensis was no longer adapted to climbing. There are a number of traits in the A. afarensis skeleton which strongly reflect bipedalism, to the extent that some researchers have suggested that bipedality evolved long before A. afarensis. In overall anatomy, the pelvis is far more human-like than ape-like. The iliac blades are short and wide, the sacrum is wide and positioned directly behind the hip joint, and there is clear evidence of a strong attachment for the knee extensors. While the pelvis is not wholly human-like (being markedly wide, or flared, with laterally orientated iliac blades), these features point to a structure that can be considered radically remodeled to accommodate a significant degree of bipedalism in the animals' locomotor repertoire. Importantly, the femur also angles in toward the knee from the hip. This trait would have allowed the foot to have fallen closer to the midline of the body, and is a strong indication of habitual bipedal locomotion. Along with humans, present day orangutans and spider monkeys possess this same feature.[citation needed] The feet also feature adducted big toes, making it difficult if not impossible to grasp branches with the hindlimbs. The loss of a grasping hindlimb also increases the risk of an infant being dropped or falling, as primates typically hold onto their mothers while the mother goes about her daily business. Without the second set of grasping limbs, the infant cannot maintain as strong a grip, and likely had to be held with help from the mother. The problem of holding the infant would be multiplied if the mother also had to climb trees. Bones of the foot (such as the calcaneus) also indicate bipedality. Computer simulations using Dynamic modeling of the skeleton's inertial properties and kinematics suggest that A. afarensis was able to walk in the same way modern humans walk, with a normal erect gait or with bent hips and knees, but could not walk in the same way as chimpanzees. The upright gait would have been much more efficient than the bent knee and hip walking, which would have taken twice as much energy. It appears probable that A. afarensis was quite an efficient bipedal walker over short distances, and the spacing of the footprints at Laetoli indicates that they were walking at 1.0 m/s or above, which matches human small-town walking speeds. Yet, this can be questioned, as finds of Australopithecus foot bones indicate that the Laetoli footprints may not have been made by Australopithecus. Many scientists also doubt the suggestion of bipedalism, and argue that even if Australopithecus really did walk on two legs, it did not walk in the same way as a human. On the other hand, the presence of a wrist-locking mechanism might suggest that they engaged in knuckle-walking. (However, these conclusions have been questioned on the basis of close analysis of knuckle-walking and the comparison of wrist bones in different species of primates).[18] The shoulder joint is also oriented more cranially (i.e. towards the skull) than in modern humans. Combined with the relatively long arms A. afarensis are thought to have had, this is thought by many to be reflective of a heightened ability to use the arm above the head in climbing behaviour. Furthermore, scans of the skulls reveal a canal and bony labyrinth morphology, which is not supportive to proper bipedal locomotion. It is commonly thought that upright bipedal walking evolved from knuckle-walking with bent legs, in the manner used by chimpanzees and gorillas to move around on the ground, but fossils such as Orrorin tugenensis indicate bipedalism around 5 to 8 million years ago, in the same general period when genetic studies suggest the lineage of chimpanzees and humans diverged. Modern apes and their fossil ancestors show skeletal adaptations to an upright posture used in tree climbing, and it has been proposed that upright, straight-legged walking originally evolved as an adaptation to tree-dwelling. Studies of modern orangutans in Sumatra have shown these apes using four legs when walking on large stable branches and when swinging underneath slightly smaller branches, but are bipedal and maintain their legs very straight when using multiple small flexible branches under 4 cm in diameter while also using their arms for balance and additional support. This enables them to get nearer to the edge of the tree canopy to grasp fruit or cross to another tree.

Climate changes around 11 to 10 million years ago affected forests in East and Central Africa, establishing periods where openings prevented travel through the tree canopy, and during these times ancestral hominids could have adapted the upright walking behaviour for ground travel, while the ancestors of gorillas and chimpanzees became more specialised in climbing vertical tree trunks or lianas with a bent hip and bent knee posture, ultimately leading them to use the related knuckle-walking posture for ground travel. This would lead to A. afarensis usage of upright bipedalism for ground travel, while still having arms well adapted for climbing smaller trees. However, chimpanzees and gorillas are the closest living relatives to humans, and share anatomical features including a fused wrist bone which may also suggest knuckle-walking by human ancestors. Other studies suggest that an upright spine and a primarily vertical body plan in primates dates back to Morotopithecus bishopi in the Early Miocene of 21.6 million years ago. the earliest humanlike primates. Known from fossil remains found in Africa, australopithecines, or australopiths, represent the group from which the ancestors of modern humans emerged. As generally used, the term australopithecines covers all early human fossils dated from about 7 million to 2.5 million years ago, and some of those dated from 2.5 million to 1.4 million years ago. The group became extinct after that time.

Social characteristics
It is difficult to reconstruct the social behaviour of extinct fossil species. However, the social structure is likely to be comparable to that of modern apes, given the average difference in body size between males and females (known as sexual dimorphism). Although there is considerable debate over how large the degree of sexual dimorphism was between males and females of A. afarensis, it is likely that males were relatively larger than females. If observations on the relationship between sexual dimorphism and social group structure from modern great apes are applied to A. afarensis then these creatures most likely lived in small family groups containing a single dominant male and a number of breeding females. For a long time, there were no known stone tools associated with A. afarensis, and paleoanthropologists commonly thought stone artifacts only dated back to approximately 2.5 million years ago. However, a 2010 study published in Nature suggests the hominin species ate meat by carving animal carcasses with stone implements. This finding pushes back the earliest known use of stone tools among hominins to about 3.4 million years ago.

Notable fossils
Naming convention
The fossils found in the Afar region were named "AL" meaning "Afar Locality".

Type specimen
The type specimen for A. afarensis is LH 4, an adult mandible from the site of Laetoli, Tanzania.

AL 129-1
Main article: AL 129-1 The first A. afarensis knee joint was discovered in November 1973 by Donald Johanson as part of a team involving Maurice Taieb, Yves Coppens and Tim White in the Middle Awash of Ethiopia's Afar Depression.

AL 288-1 (Lucy)
Main article: Lucy (Australopithecus) The first A. afarensis skeleton was discovered on November 24, 1974 near Hadar in Ethiopia by Tom Gray in the company of Donald Johanson, as part of a team involving Maurice Taieb, Yves Coppens and Tim White in the Middle Awash of Ethiopia's Afar Depression.

AL 333
Main article: AL 333

In 1975, a year after the discovery of Lucy, Donald Johanson's team discovered another site in Hadar which included over 200 fossil specimens from at least 13 individuals, both adults and juveniles. This site, AL 333, is commonly referred to as the "First Family." The close alignment of the remains indicates that all the individuals died at the same time, a unique finding.

AL 444-2
This is the cranium of a male found at Hadar in 1992. By the time it was found, it was the first complete skull of A. afarensis. The rarity of relatively complete craniofacial remains in the A. afarensis samples before the discovery of A.L. 444-2 seriously hampered prior meaningful analysis of those samples' evolutionary significance. In light of A.L.444-2 and other new craniofacial remains, Kimbel, Rak and Johanson argue in favor of the taxonomic unity of A. afarensis.

Selam
Main article: Selam (Australopithecus) In 2006 a dig in Dikika, Ethiopia, a few miles from the place where Lucy was found recovered the skeleton of a 3-year-old A. afarensis girl which comprised almost the entire skull and torso, and most parts of the limbs. The features of the skeleton suggested adaptation to walking upright (bipedalism) as well as tree-climbing, features that match the skeletal features of Lucy and fall midway between human and humanoid ape anatomy. "Baby Lucy" has officially been named "Selam" (meaning peace in the Amharic/Ethiopian Language).

2011 Hadar finding

In February 2011, the discovery of a 3.2 million year-old bone in Hadar, Ethiopia at the AL 333 site was announced. The foot bone shows that the species had arches in its feet, which confirmed that the species walked upright for the majority of the time. The foot bone is one of 49 new bones discovered, and indicates that A. afarensis is "a lot more human-like than we had ever supposed before", according to the lead scientist on the study.

Others

AL 200-1 AL 444

Related work
Further findings at Afar, including the many hominin bones in site 333, produced more bones of concurrent date, and led to Johanson and White's eventual argument that the Koobi Fora hominins were concurrent with the Afar hominins. In other words, Lucy was not unique in evolving bipedalism and a flat face. Recently, an entirely new species has been discovered, called Kenyanthropus platyops, however the cranium KNM WT 40000 has a much distorted matrix making it hard to distinguish (however a flat face is present). This had many of the same characteristics as Lucy, but is possibly an entirely different genus. Another species, called Ardipithecus ramidus, was found by White and colleagues in the 1990s. This was fully bipedal, yet appears to have been contemporaneous with a woodland environment. Scientists have a rough estimation of the cranial capacity of Ar. ramidus at 300 to 350 cm, from the data released on the Ardipithecus specimen nicknamed Ardi in 2009.

Australopithecus bahrelghazali
Australopithecus bahrelghazali is a fossil hominin that was first discovered in 1993 by the paleontologist Michel Brunet in the Bahr el Ghazal valley near Koro Toro, in Chad, that Brunet named Abel. It was dated using Beryllium-based Radiometric dating as living circa 3.6 million years ago. The find consists of a mandibular fragment, a lower second incisor, both lower canines, and all four of its premolars, still affixed within the dental alveoli. The specimen's proper name is KT-12/H1; "Abel" is the informal name, a dedication to Brunet's deceased colleague Abel Brillanceau. The specimen located roughly 2,500 kilometers West from the East African Great Rift Valley. The mandible KT-12/H1 discovered has similar features to the dentition of Australopithecus afarensis; this has brought researchers like William Kimbel to argue that Abel is not an exemplar of a separate species, but "falls within the range of variation" of the Australopithecus afarensis. By 1996, Brunet and his team classified KT 12/H1 as the holotype specimen for Australopithecus bahrelghazali. This claim is difficult to substantiate, as the describers have kept KT 12/H1 locked away from the general paleoanthropological community, contrary to the International Code of Zoological Nomenclature 1999.[4] This species is a mystery to some as it is the only australopithecine fossil found in Central Africa. It is also of great importance as it was the first fossil to show that geographically there is a "a third window" of early hominid evolution.

Kenyanthropus Platyops

Kenyanthropus platyops is a 3.5 to 3.2 million year old (Pliocene) hominin fossil that was discovered in Lake Turkana, Kenya in 1999 by Justus Erus, who was part of Meave Leakey's team. Leakey (2001) proposes that the fossil represents an entirely new hominine genus, while others classify it as a separate species of Australopithecus, Australopithecus platyops, and yet others interpret it as an individual of Australopithecus afarensis.

Discovery and interpretations

The fossil found features a broad flat face with a toe bone that suggests it probably walked upright. Teeth are intermediate between typical human and typical ape forms. Kenyanthropus platyops, which means "Flat-faced man of Kenya" (a name given by Meave Leakey), is the only described species in the genus. However, if some paleoanthropologists are correct, Kenyanthropus may not even represent a valid taxon, as the specimen (KNM-WT 40000)[2] is so distorted by matrix-filled cracks that meaningful morphological characteristics are next to impossible to assess with confidence. It may simply be a specimen of Australopithecus afarensis, which is known from the same time period and geographic area, or its own species within Australopithicus, A. platyops. Other researchers speculate that the flatter face position of the rough cranium is similar to KNM ER 1470 (Homo rudolfensis), and suspect it to be closer to the genus Homo, perhaps being a direct ancestor. However the debate has not been concluded and the species remains an enigma. The bones discovered at the site included more than 30 skull and tooth fragments in a stratum dated to between 3.5 and 3.2 million years ago. Dr. Leakey believes that it belongs to an entirely new genus of ancestors, and is the oldest "reasonably complete" cranium found so far. Humans were once thought to have evolved only from Australopithecus afarensis, the species made famous by the fossil Lucy. But now it seems Lucy may have been sharing the woods and grass plains of prehistoric Africa with a rival. When learning of the discovery, Daniel Lieberman, an anthropologist at George Washington University expressed his opinion that between 3.5 and 2 million years ago there were several human-like species, each of which were well adapted to life in their particular environments. Also that, like that of many other mammalian groups, humans evolved through a series of complex radiations, known as "adaptive radiation".

The Kenyanthropus fossil has a small ear hole, like those of chimpanzees. It also shares many features of other primitive hominids, such as a small brain, but it also has striking differences, including high cheek bones, and a flat plane beneath its nose bone, which gives it a flat face.

Australopithecus africanus
Australopithecus africanus was an early hominid, an australopithecine, who lived between ~3.03 and 2.04 million years ago in the later Pliocene and early Pleistocene. In common with the older Australopithecus afarensis, Au. africanus was slenderly built, or gracile, and was thought to have been a direct ancestor of modern humans. Fossil remains indicate that Au. africanus was significantly more like modern humans than Au. afarensis, with a more human-like cranium permitting a larger brain and more humanoid facial features. Au. africanus has been found at only four sites in southern Africa Taung (1924), Sterkfontein (1935), Makapansgat (1948) and Gladysvale (1992).

Famous fossils
Taung Child Raymond Dart became interested in fossils found at the lime mine at Taung near Kimberley, South Africa in 1924. The most promising of these was a skull of an odd ape-creature sharing human traits such as eye orbits, teeth, and, most importantly, the hole at the base of the skull over the spinal column (the foramen magnum) indicating a human-like posture. Dart assigned the specimen the name Australopithecus africanus ("southern ape of Africa"). This was the first time the word Australopithecus was assigned to any hominid. Dart claimed that the skull must have been an intermediate species between ape and humans, but his claim about the Taung Child was rejected by the scientific community at the time due to the belief that a large cranial capacity must precede bipedal locomotion. This was exacerbated by the widespread acceptance of the Piltdown Man. Sir Arthur Keith, a fellow anatomist and anthropologist, suggested that the skull belonged to a young ape, most likely from an infant gorilla. It was not until 20 years later that the public accepted the new genus and that australopithecines were a true member of Homininae. Mrs. Ples Dart's theory was supported by Robert Broom.[5] In 1938 Broom classified an adult endocranial cast having a brain capacity of 485 cc, which had been found by G. W. Barlow, as Plesianthropus transvaalensis. On April 18, 1947, Broom and John T. Robinson discovered a skull belonging to a middle-aged female,[6] (catalogue number STS 5), while blasting at Sterkfontein. Broom classified it also as Plesianthropus transvaalensis, and it was dubbed Mrs. Ples by Broom's young coworkers (though the skull is now thought to have belonged to a young male). The lack of facial projection in comparison to apes was noted by Raymond Dart (including from Taung Child), a trait in common with more advanced hominines. Both fossils were later classified as Australopithecus africanus.

Morphology and interpretations

Like Au. afarensis, Au. africanus the South African counterpart was generally similar in many traits, a bipedal hominid with arms slightly larger than the legs (a physical trait also found in chimpanzees). Despite its slightly more human-like cranial features, seen for example in the crania Mrs. Ples and STS 71, other more primitive features including ape-like curved fingers for tree climbing are also present. Due to other more primitive features visible on Au. africanus, some researchers believe the hominin, instead of being a direct ancestor of more modern hominins, evolved into Paranthropus. One robust australopithecine seen as a descendent of Au. africanus is Paranthropus robustus. Both P. robustus and Au. africanus crania seem very alike despite the more heavily built features of P. robustus that are adaptations for heavy chewing like a gorilla. Au. africanus, on the other hand, had a cranium which quite closely resembled that of a chimp, yet both their brains measure about 400 cc to 500 cc and probably had an apelike intelligence.[5] Au. africanus had a pelvis that was built for slightly better bipedalism than that of Au. afarensis.

Sexual dimorphism Recent evidence regarding modern human sexual dimorphism (physical differences between men and women) in the lumbar spine has been seen in pre-modern primates such as Au. africanus. This dimorphism has been seen as an evolutionary adaptation of females to better bear lumbar load during pregnancy, an adaptation that non-bipedal primates would not need to make. A 2011 study using ratios of strontium isotopes in teeth suggested that Au. africanus and Paranthropus robustus groups in southern Africa were patrilocal: women tended to settle farther from their region of birth than men did.

Geochronology
Based on current data Au. africanus dates to between 3.03 and 2.04 million years based on a combination of palaeomagnetism (Andy Herries, La Trobe University, Australia, Uranium-lead (Robyn Pickering (U. Melbourne, Australia), electron spin resonance (Darren Curnoe, UNSW, Australia) and faunal dating. The Makapansgat fossils date to between 3.03 and 2.58 million years with fossils MLD37/38 likely dating close to 2.58 million years; Sterkfontein dates to between 2.58 and 2.04 million years with the Sts 5 Mrs Ples fossil dating to around 2.04 million years; and Gladysvale dates to between 2.4 and 2.0 million years. The age of the Taung child remains more difficult to determine and is the focus of a current project by Brian Kuhn (U. Witwatersrand, S. Africa), Phil Hopley (Birkbeck College, UK), Colin Menter (U. Johannesburg, S. Africa) and Andy Herries (La Trobe University, Australia).

Paranthropus aethiopicus
Paranthropus aethiopicus is an extinct species of hominid. The finding discovered in 1985 by Alan Walker in West Turkana, Kenya, KNM WT 17000 (known as the "Black Skull" due to the dark coloration of the bone, caused by high levels of manganese), is one of the earliest examples of robust pliocene hominids. The skull is dated to 2.5 million years ago, older than the later forms of robust australopithecines. Anthropologists suggest that P. aethiopicus lived between 2.7 and 2.5 million years ago. The features are quite primitive and share many traits with Australopithecus afarensis; thus P. aethiopicus is likely to be a direct descendant. With its face being as prognathic (projecting) as A. afarensis, its brain size was also quite small at 410 cc. P. aethiopicus was first proposed in 1967 to describe a toothless partial mandible (Omo 18) found in Ethiopia by French paleontologists. Lower jaw and teeth fragments have been uncovered. P. aethiopicus had a large sagittal crest and zygomatic arch adapted for heavy chewing (as in gorilla skulls). Not much is known about this species since the best evidence comes from the "Black Skull" and the jaw. There is not enough material to make an assessment to how tall they were, but they may have been as tall as Australopithecus afarensis. Not all anthropologists agree that P. aethiopicus gave rise to both Paranthropus boisei and Paranthropus robustus, since the skull more closely resembles that of A. afarensis. The one clue that makes P. aethiopicus a possible ancestor to both P. boisei and P. robustus is the similarity in jaw size. P. aethiopicus is known to have lived in mixed savanna and woodland. More evidence must be gathered about P. aethiopicus in order to accurately describe its physiology. The bizarre primitive shape of the "Black Skull" gives evidence that P. aethiopicus and the other australopithecines are on an evolutionary branch of the hominid tree, distinctly diverging from the Homo (human) lineage.

Paranthropus boisei
Paranthropus boisei (originally called Zinjanthropus boisei and then Australopithecus boisei until recently) was an early hominin and described as the largest of the Paranthropus species. It lived from about 2.6 until about 1.2 million years ago during the Pliocene and Pleistocene epochs in Eastern Africa.

Discovery
First discovered by anthropologist Mary Leakey on July 17, 1959, at Olduvai Gorge, Tanzania, the well-preserved cranium (nicknamed "Nutcracker Man") was dated to 1.75 million years old and had characteristics distinctive of the robust australopithecines. Mary and her husband Louis Leakey classified the find as Zinjanthropus boisei: "Zinj" for the medieval East African region of Zanj, "anthropus" meaning ape or ape-human, and "boisei" for Charles Boise (the anthropologists teams funder at the time). Paranthropus boisei (as the species was eventually categorized) proved to be a treasure especially when the anthropologists' son Richard Leakey considered it to be the first hominin species to use stone tools. Another skull was unearthed in 1969 by Richard at Koobi Fora near the Lake Turkana region, in Kenya.

Morphology and interpretations

The brain volume is quite small, about 500 and 550 cm, not much larger in comparison to Australopithecus afarensis and Australopithecus africanus. It had a skull highly specialized for heavy chewing and several traits seen in modern day gorillas. P. boisei inhabited savannah woodland territories. Males weighed 68 kg (150 lb) and stood 1.3 m (4 ft 3 in) tall, while females weighed 45 kg (99 lb) and stood 1.05 m (3 ft 5 in) tall. The average adult males were much larger than females (sexual dimorphism), as was the case in virtually all australopithecine species. The back molar teeth were relatively large, with an area over twice as great as is found in modern humans. The species is sometimes referred to as Nutcracker Man because it has the biggest, flattest cheek teeth and the thickest enamel of any known hominin. Some argue that the craniodental morphology of this taxon (e.g., large postcanine dentition, thick enamel, robust mandibles, sagittal cresting, flaring zygomatic region) are indicative of a diet of hard or tough foods such as ground tubers, nuts and seeds. However, research on the molar microwear of P. boisei found a microwear pattern very different than that observed for P. robustus in South Africa which is thought to have fed on hard foods as a "fallback resource. This work suggests that hard foods were an infrequent part of its diet. The carbon isotope ratios of P. boisei suggest that it had a diet dominated by C4 vegetation.

Fossils
In 1993, A. Amzaye found fossils of P. boisei at Konso, Ethiopia. The partial skull's designation is KGA10-525 and is dated to 1.4 million years old. It is the biggest skull specimen ever found of P. boisei. The oldest specimen of P. boisei was found in Omo, Ethiopia and dates to 2.3 million years old, classified as (L. 74a-21) while the youngest specimen from Olduvai Gorge dates 1.2 million years old, classified as OH 3 and OH 38. Other well preserved specimens

OH 5 Zinjanthropus or "Zinj" or "Nutcrackerman" is the first P. boisei found by Mary Leakey at Olduvai Gorge, Tanzania belonging to an adult male (circa 1.75 mya). KNM ER 406 is a small partial cranium discovered by H. Mutua and Richard Leakey in 1969 found at Koobi Fora, Kenya displays large zygomatic arches, a cranial capacity of 510 cm (circa 1.7 mya). KNM WT 17400 is a partial cranium with similar characteristics as KNM WT 17000 "Black skull" belonging to Paranthropus aethiopicus. The skull was found at West Turkana, Kenya (circa 1.7 mya). ER 406, found by Richard Leakey and H. Mutua in 1970 at Koobi Fora, Kenya, is a partial cranium most likely identified as belonging to a female (circa 1.7 mya).

Australopithecus garhi
Australopithecus garhi is a gracile australopithecine species whose fossils were discovered in 1996 by a research team led by Ethiopian paleontologist Berhane Asfaw and Tim White, an American paleontologist. The hominin remains are believed to be a human ancestor species and the final missing link between the Australopithecus genus and the human genus, Homo. Tim White was the scientist to find the first of the key A. garhi fossils in 1996 in the Bouri Formation, located in the Middle Awash of Ethiopia's Afar Depression. The species was confirmed and established as A. garhi on November 20, 1997 by the Ethiopian paleoanthropologist Yohannes Haile-Selassie. The species epithet "garhi" means "surprise" in the local Afar language.

Morphology and interpretations

The traits of A. garhi fossils such as BOU-VP-12/130 are somewhat distinctive from traits typically seen in Australopithecus afarensis and Australopithecus africanus. An example of the distinction can be seen when comparing the Hadar maxilla (A. afarensis) to the Bouri specimen of A. garhi. The cranial capacity of A. garhi measures 450cc, the same size as other australopithecines. The mandible classified as Asfaw et al. has a morphology generally believed to be compatible with the same species, yet it is possible that another hominin species may have been found within the same deposits. Studies made on the premolars and molar teeth have a few similarities with those of Paranthropus boisei since they are larger than any other gracile form of australopithecine. It has been suggested that if A. garhi is ancestral to Homo (i.e. Homo habilis) the maxillary morphology would have undergone a rapid evolutionary change in roughly 200,000 and 300,000 years. Earliest stone tools Few primitive shaped stone tool artifacts closely resembling Olduwan technology were discovered with the A. garhi fossils, dating back roughly 2.5 and 2.6 million years old. The tools are suggested to be older than those acquired by Homo habilis, which is thought to be a possible direct ancestor of more modern hominins. For a long time anthropologists assumed that only members of early genus Homo had the ability to produce sophisticated tools. However, the crude ancient tools lack several techniques that are generally seen in later forms Olduwan and Acheulean such as strong rock-outcroppings. In another site in Bouri, Ethiopia, roughly 3,000 stone artifacts had been found to be an estimated 2.5 million years old in age.

Homo habilis
Homo habilis is a species of the genus Homo, which lived from approximately 2.33 to 1.4 million years ago, during the Gelasian Pleistocene period. The discovery and description of this species is credited to both Mary and Louis Leakey, who found fossils in Tanzania, East Africa, between 1962 and 1964. Homo habilis (or possibly H. rudolfensis) was the earliest known species of the genus Homo until May 2010, when H. gautengensis was discovered, a species believed to be even older than H. habilis. In its appearance and morphology, H. habilis is thus the least similar to modern humans of all species in the genus (except possibly H. rudolfensis). H. habilis was short and had disproportionately long arms compared to modern humans; however, it had a less protruding face than the australopithecines from which it is thought to have descended. H. habilis had a cranial capacity slightly less than half of the size of modern humans. Despite the ape-like morphology of the bodies, H. habilis remains are often accompanied by primitive stone tools (e.g. Olduvai Gorge, Tanzania and Lake Turkana, Kenya). Homo habilis has often been thought to be the ancestor of the more gracile and sophisticated Homo ergaster, which in turn gave rise to the more human-appearing species, Homo erectus. Debates continue over whether H. habilis is a direct human ancestor, and whether all of the known fossils are properly attributed to the species. However, in 2007, new findings suggest that the two species coexisted and may be separate lineages from a common ancestor instead of H. erectus being descended from H. habilis.

Findings

One set of fossil remains (OH 62), discovered by Donald Johanson and Tim White in Olduvai Gorge in 1986, included the important upper and lower limbs. Their finding stimulated some debate at the time. An older (1963) finding from the Olduvai site found by N. Mbuika had included a lower jaw fragment, teeth and upper mandible possibly from a female dating 1.7 million years old. The remains from 3 skeletons stacked on top of each other demonstrated australopithecine-like body with a more human-like face and smaller teeth. Compared to australopithecines, H. habilis's brain capacity of 363 and 600 cm was on average 50% larger than australopithecines, but considerably smaller than the 1350 to 1450 cm range of modern Homo sapiens. These hominins were smaller than modern humans, on average standing no more than 1.3 m (4 ft 3 in) tall. The small size and rather primitive attributes have led some experts (Richard Leakey among them) to propose excluding H. habilis from the genus Homo, and renaming as "Australopithecus habilis". KNM ER 1813 KNM ER 1813 is a relatively complete cranium which dates 1.9 million years old, discovered at Koobi Fora, Kenya by Kamoya Kimeu in 1973. The brain capacity is 510 cm, not as impressive as other early specimen and forms of Homo habilis discovered. OH 7 OH 7 dates 1.75 million years old and was discovered by Mary and Louis Leakey on November 4, 1960 at Olduvai Gorge, Tanzania. It is a lower jaw complete with teeth and due to the size of the small teeth; researchers estimate this juvenile individual had a brain volume of 363 cm. Also found were more than 20 fragments of the left hand. Tobias and Napier assisted in classifying OH-7 as the type fossil. OH 24 OH 24 (AKA Twiggy) is a roughly deformed cranium dating 1.8 million years old discovered in October 1968 at Olduvai Gorge, Tanzania. The brain volume is just under 600 cm; also a reduction in a protruding face is present compared to members of more primitive Australopithecines. KNM ER 1805 KNM ER 1805 is a specimen of an adult H. habilis made of 3 pieces of cranium dating 1.74 million years old from Koobi Fora, Kenya. Previous assumptions were that this specimen belongs to H. erectus based on the degree of prognathism and overall cranial shape.

Interpretations
Homo habilis is thought to have mastered the Olduwan era (Lower Paleolithic) tool case which utilized stone flakes. These stone flakes were more advanced than any tools previously used, and gave H. habilis the edge it needed to prosper in hostile environments previously too formidable for primates. Whether H. habilis was the first hominid to master stone tool technology remains controversial, as Australopithecus garhi, dated to 2.6 million years ago, has been found along with stone tool implements at least 100,000 - 200,000 years older than H. habilis. Most experts assume the intelligence and social organization of H. habilis were more sophisticated than typical australopithecines or chimpanzees. Yet despite tool usage, H. habilis was not the master hunter that its sister species (or descendants) proved to be, as there is ample fossil evidence that H. habilis was a staple in the diet of large predatory animals such as Dinofelis, a large scimitar-toothed predatory cat the size of a jaguar. H. habilis used tools primarily for scavenging, such as cleaving meat off carrion, rather than defense or hunting. Homo habilis is thought to be the ancestor of the lankier and more sophisticated Homo ergaster, which in turn gave rise to the more human-appearing species Homo erectus. Debates continue over whether H. habilis is a direct human ancestor, and whether all of the known fossils are properly attributed to the species. Homo habilis co-existed with other Homo-like bipedal primates, such as Paranthropus boisei, some of which prospered for many millennia. However, H. habilis, possibly because of its early tool innovation and a less specialized diet, became the

precursor of an entire line of new species, whereas Paranthropus boisei and its robust relatives disappeared from the fossil record. Homo habilis may also have coexisted with Homo erectus in Africa for a period of 500,000 years.

Paranthropus robustus
Paranthropus robustus was originally discovered in Southern Africa in 1938. The development of P. robustus, namely in cranial features, seemed to be aimed in the direction of a "heavy-chewing complex". Because of the definitive traits that are associated with this robust line of australopithecine, anthropologist Robert Broom erected the genus Paranthropus and placed this species into it. Paranthropus robustus (considered for a time by the scientific community as Australopithecus robustus) is generally dated to have lived between 2.0 and 1.2 million years ago. P. robustus had large sagittal crests, jaws, jaw muscles, and post-canine teeth that were adapted to serve in the dry environment that they lived in. After Raymond Darts discovery of Australopithecus africanus, Broom had been in favour of Dart's claims about Australopithecus africanus being an ancestor of Homo sapiens. Broom was a Scottish doctor then working in South Africa who began making his own excavation in Southern Africa to find more specimens, which Dart had found earlier. In 1938, at 70 years old, Broom, excavating at Kromdraai, South Africa discovered pieces of a skull and teeth which resembled Dart's Australopithecus africanus find, but the skull had some "robust" characteristics. The fossils included parts of a skull and teeth; all dated to 2 million years old. Fossil sites found on Paranthropus robustus are found only in South Africa in Kromdraai, Swartkrans, Drimolen, Gondolin and Coopers. In the cave at Swartkrans, the remains of 130 individuals were discovered. The study made on the dentition of the hominins revealed that the average P. robustus rarely lived past 17 years of age. Paranthropus robustus became the first "robust" species of hominid ever uncovered well before P. boisei and P. aethiopicus. Broom's first discovery of P. robustus had been the first discovery of a robust australopithecine and the second australopithecine after Australopithecus africanus, which Dart discovered. Broom's work on the australopithecines showed that the evolution trail leading to Homo sapiens was not just a straight line, but was one of rich diversity.

Morphology
Typical of robust australopithecines, P. robustus had a head shaped a bit like a gorilla's with a more massive built jaw and teeth in comparison to hominins within the Homo lineage. The sagittal crest that runs from the top of the skull acts as an anchor for large chewing muscles. The DNH 7 skull of Paranthropus robustus, "Eurydice", was discovered in 1994 at the Drimolen Cave in Southern Africa by Andre Keyser, and is dated to 2.3 million years old, possibly belonging to a female. The teeth of these primates were larger and thicker than any gracile australopithecine found, due to the morphology differences Broom originally designated his find as Australopithecus robustus. On the skull, a bony ridge is located above from the front to back indicating where the jaw muscles joined. P. robustus males may have stood only 1.2m (4 feet) tall and weighed 54 kg (120 lb) while females stood just under 1 meter (3 feet 2 inches) tall and weighed only 40 kg (90 lb), indicating a large sexual dimorphism. The teeth found on P. robustus are almost as large as those of P. boisei. Broom analyzed his findings carefully and noted the differences in the molar teeth size which resembled a gorilla's a bit more than a human's. Other P. robustus remains have been found in Southern Africa. The average brain size of P. robustus measured to only 410 and 530 cc, about as large as a chimpanzee's. Some have argued that P robustus had a diet of hard gritty foods such as nuts and tubers since they lived in open woodland and savanna. More recent research suggests that this taxon was more of a dietary generalist, and others have argued that they principally consumed hard and gritty resources as fallback foods only during time of nutritional stress. A 2011 study using ratios of strontium isotopes in teeth suggested that Australopithecus africanus and P. robustus groups in southern Africa were patrilocal: females tended to settle farther from their region of birth than males did.

Homo erectus

Homo erectus (meaning "upright man," from the Latin rgre, "to put up, set upright") is an extinct species of hominid that lived from the end of the Pliocene epoch to the later Pleistocene, about 1.3 to 1.8 million years ago. The species originated in Africa and spread as far as India, China and Java. There is still disagreement on the subject of the classification, ancestry, and progeny of H. erectus, with two major alternative hypotheses: erectus may be another name for Homo ergaster, and therefore the direct ancestor of later hominids such as Homo heidelbergensis, Homo neanderthalensis, and Homo sapiens; or it may be an Asian species distinct from African ergaster.

Origin
The first theory is that H. erectus migrated from Africa during the Early Pleistocene, possibly as a result of the operation of the Saharan pump, around 2.0 million years ago, and dispersed throughout much of the Old World. Fossilized remains 1.8 to 1 million years old have been found in Africa (e.g., Lake Turkana[5] and Olduvai Gorge), Europe (Georgia and Spain), Indonesia (e.g., Sangiran and Trinil), Vietnam, China (e.g., Shaanxi) and India. The second theory is that H. erectus evolved in Asia and then migrated to Africa. The species occupied a West Asian site called Dmanisi, in Georgia, from 1.85 million to 1.77 million years ago, at the same time or slightly before the earliest evidence in Africa. Excavations found 73 stone tools for cutting and chopping and 34 bone fragments from unidentified creatures.

Discovery and representative fossils

The Dutch anatomist Eugne Dubois, who was fascinated especially by Darwin's theory of evolution as applied to man, set out to Asia (the place accepted then, despite Darwin, as the cradle of human evolution), to find a human ancestor in 1886. In 1891, his team discovered a human fossil on the island of Java, Indonesia; he described the species as Pithecanthropus erectus (from the Greek , "ape", and , "man"), based on a calotte (skullcap) and a femur like that of H. sapiens found from the bank of the Solo River at Trinil, in East Java. (This species is now regarded as Homo erectus.) The find became known as Java Man. Thanks to Canadian anatomist Davidson Black's (1921) initial description of a lower molar, which was dubbed Sinanthropus pekinensis,[11] however, most of the early and spectacular discoveries of this taxon took place at Zhoukoudian in China. German anatomist Franz Weidenreich provided much of the detailed description of this material in several monographs published in the journal Palaeontologica Sinica (Series D). Nearly all of the original specimens were lost during World War II; however, authentic Weidenreichian casts do exist at the American Museum of Natural History in New York and at the Institute of Vertebrate Paleontology and Paleoanthropology in Beijing, and are considered to be reliable evidence. Throughout much of the 20th century, anthropologists debated the role of H. erectus in human evolution. Early in the century, however, due to discoveries on Java and at Zhoukoudian, it was believed that modern humans first evolved in Asia. A few naturalists (Charles Darwin most prominent among them) predicted that humans' earliest ancestors were African: he pointed out that chimpanzees and gorillas, who are human relatives, live only in Africa. From 1950s to 1970s, however, numerous fossil finds from East Africa yielded evidence that the oldest hominins originated there. It is now believed that H. erectus is a descendant of earlier genera such as Ardipithecus and Australopithecus, or early Homo-species such as H. habilis or H. ergaster. H. habilis and H. erectus coexisted for several thousand years, and may represent separate lineages of a common ancestor. The skull of Tchadanthropus uxoris, discovered in 1961 by Yves Coppens in Chad, is the earlier fossil human discovered in Africa. This fossil "had been so eroded by wind-blown sand that it mimicked the appearance of an australopith, a primitive type of hominid". Though some first considered it to be a specimen of H. habilis, it is no longer considered to be a valid taxon, and scholars rather consider it to represent H. erectus. Homo erectus georgicus Homo erectus georgicus (Georgian: ) is the subspecies name sometimes used to describe fossil skulls and jaws found in Dmanisi, Georgia, which although first proposed as a separate species is now

classified within H. erectus. A partial skeleton was discovered in 2001. The fossils are about 1.8 million years old. The remains were first discovered in 1991 by Georgian scientist, David Lordkipanidze, accompanied by an international team which unearthed the remains. Implements and animal bones were found alongside the ancient human remains. At first, scientists thought they had found mandibles and skulls belonging to Homo ergaster, but size differences led them to name a new species, Homo georgicus, which was posited as a descendant of Homo habilis and ancestor of Asian Homo erectus. This classification was not upheld and the fossil is now considered a divergent subgroup of of Homo erectus, sometimes called Homo erectus georgicus'. At around 600 cubic centimetres (37 cu in) brain volume, the skull D2700 is dated to 1.77 million years old and in good condition offering insights in comparison to the modern human cranial morphology. At the time of discovery the cranium was the smallest and most primitive Hominina skull ever discovered outside of Africa. However, in 2003 a significantly smaller brained hominid was found on the isle of Flroes H. erectus floresiensis. Homo erectus georgicus exhibits strong sexual dimorphism with males being significantly larger than females. Subsequently, four fossil skeletons were found, showing a species primitive in its skull and upper body but with relatively advanced spines and lower limbs, providing greater mobility. They are now thought not to be a separate species, but to represent a stage soon after the transition between Homo habilis and H. erectus, and have been dated at 1.8 million years before the present, according to the leader of the project, David Lordkipanidze. The assemblage includes one of the largest Pleistocene Homo mandibles (D2600), one of the smallest Lower Pleistocene mandibles (D211), a nearly complete sub adult (D2735), and a completely toothless specimen (D3900).

Classification and special distinction

Many paleoanthropologists still debate the definition of H. erectus and H. ergaster as separate species. Several scholars suggested to drop the taxon Homo erectus, and suggested instead to equate Homo erectus with archaic Homo sapiens Some call H. ergaster the direct African ancestor of H. erectus, proposing that it emigrated out of Africa and migrated into Asia, branching into a distinct species.[32] Most dispense with the species-name ergaster, making no distinction between such fossils as the Turkana Boy and Peking Man.[citation needed] Though "Homo ergaster" has gained some acceptance as a valid taxon, these two are still usually defined as distinct African and Asian populations of the larger species H. erectus. While some have argued (and insisted) that Ernst Mayr's biological species definition cannot be used here to test the above hypotheses, one can, however, examine the amount of morphological cranial variation within known H. erectus / H. ergaster specimens, and compare it to what one sees in disparate extant groups of primates with similar geographical distribution or close evolutionary relationship. Thus, if the amount of variation between H. erectus and H. ergaster is greater than what one sees within a species of, say, macaques, then H. erectus and H. ergaster may be considered two different species. The extant model of comparison is very important, and selecting appropriate species can be difficult. (For example, the morphological variation among the global population of H. sapiens is incredibly small and our own special diversity may not be a trustworthy comparison.) As an example, fossils found in Dmanisi in the Republic of Georgia were originally described as belonging to another closely related species, Homo georgicus, but subsequent examples showed their variation to be within the range of Homo erectus, and they are now classified as Homo erectus georgicus. H. erectus had a cranial capacity greater than that of Homo habilis (although the Dmanisi specimens have distinctively small crania): the earliest remains show a cranial capacity of 850 cm, while the latest Javan specimens measure up to 1100 cm, overlapping that of H. sapiens.; the frontal bone is less sloped and the dental arcade smaller than the australopithecines'; the face is more orthognatic (less protrusive) than either the australopithecines' or H. habilis's, with large brow-ridges and less prominent zygomata (cheekbones). These early hominins stood about 1.79 m (5 ft 10 in), and were more robust than modern humans. The sexual dimorphism between males and females was slightly greater than seen in H. sapiens, with males being about 25% larger than females. However, their dimorphism is drastically lesser than that of the earlier Australopithecus genus. The discovery of the skeleton KNM-WT 15000, "Turkana boy" (Homo ergaster), made near Lake Turkana, Kenya by Richard Leakey and Kamoya Kimeu in 1984, is one of the most complete hominid-skeletons discovered, and has contributed greatly to the interpretation of human physiological evolution.

Use of tools
Homo ergaster used more diverse and sophisticated stone tools than its predecessors: H. erectus, however, used comparatively primitive tools. This is possibly because H. ergaster first used tools of Oldowan technology and later progressed to the Acheulean: while the use of Acheulean tools began ca. 1.8 million years ago, the line of H. erectus diverged some 200,000 years before the general innovation of Acheulean technology. Thus the Asian migratory descendants of H. ergaster made no use of any Acheulean technology. In addition, it has been suggested that H. erectus may have been the first hominid to use rafts to travel over oceans. Use of fire Sites in Europe and Asia seem to indicate controlled use of fire by H. erectus, some dating back 1.5 million years ago. A presentation at the Paleoanthropology Society annual meeting in Montreal, Canada in March 2004 stated that there is evidence for controlled fires in excavations in northern Israel from about 690,000 to 790,000 years ago. A site called Terra Amata, located on the French Riviera, which lies on an ancient beach, seems to have been occupied by H. erectus; it contains the earliest evidence of controlled fire, dated at around 300,000 years ago. Excavations dating from approximately 790,000 years ago in Israel suggest that H. erectus not only controlled fire but could light fires. Despite these examples, some scholars continue to assert that the controlled use of fire was not typical of H. erectus, but only of later species of Homo, such as H. heidelbergensis, H. neanderthalensis, and H. sapiens).

Sociality
Homo erectus was probably the first hominid to live in a hunter-gatherer society, and anthropologists such as Richard Leakey believe that it was socially more like modern humans than the more Australopithecus-like species before it. Likewise, increased cranial capacity generally coincides with the more sophisticated tools occasionally found with fossils. The discovery of Turkana boy (H. ergaster) in 1984 gave evidence that, despite its Homo-sapiens-like anatomy, it may not have been capable of producing sounds comparable to modern human speech. Ergaster likely communicated in a protolanguage lacking the fully developed structure of modern human language but more developed than the non-verbal communication used by chimpanzees. Such inference has been challenged by the discovery of H. ergaster/erectus vertebrae some 150,000 years older than the Turkana Boy in Dmanisi, Georgia, that reflect vocal capabilities within the range of H. sapiens. Both brain-size and the presence of the Broca's area also support the use of articulate language. H. erectus was probably the first hominid to live in small, familiar band-societies similar to modern hunter-gatherer bandsocieties. H. erectus/ergaster is thought to be the first hominid to hunt in coordinated groups, use complex tools, and care for infirm or weak companions. There has been some debate as to whether H. erectus, and possibly the later Neanderthals may have interbred with anatomically modern humans in Europe and Asia. See Neanderthal admixture theory.

Descendants and subspecies

Homo erectus remains one of the most successful and long-lived species of Homo. As a distinct Asian species, however, no consensus has been reached as to whether it is ancestral to H. sapiens or any later hominids.

Homo erectus o Homo erectus erectus o Homo erectus yuanmouensis o Homo erectus lantianensis o Homo erectus wushanensis o Homo erectus nankinensis

o o o o o

Homo erectus pekinensis Homo erectus palaeojavanicus Homo erectus soloensis Homo erectus tautavelensis Homo erectus georgicus

Related species

Homo ergaster Homo floresiensis Homo antecessor Homo heidelbergensis Homo neanderthalensis Homo sapiens o Homo sapiens sapiens Homo rhodesiensis Homo cepranensis

The discovery of Homo floresiensis in 2003 and of the recentness of its extinction has raised the possibility that numerous descendant species of Homo erectus may have existed in the islands of Southeast Asia and await fossil discovery (see Orang Pendek). Homo erectus soloensis, who lived on Java at least as late as about 50,000 years ago, would be one of them. Some scientists are skeptical of the claim that Homo floresiensis is a descendant of Homo erectus. One explanation holds that the fossils are of a modern human with microcephaly, while another one holds that they are from a group of pygmies.

Individual fossils
Some of the major Homo erectus fossils:

Indonesia (island of Java): Trinil 2 (holotype), Sangiran collection, Sambungmachan collection, Ngandong collection China: Lantian (Gongwangling and Chenjiawo), Yunxian, Zhoukoudian, Nanjing, Hexian India: Narmada (taxonomic status debated!) Kenya: WT 15000 (Nariokotome), ER 3883, ER 3733 Tanzania: OH 9 Vietnam: Northern, Tham Khuyen, Hoa Binh Republic of Georgia: Dmanisi collection Ethiopia: Daka calvaria Eritrea: Buia cranium Denizli Province ,Turkey: Kocabas fossil

Homo rudolfensis
Homo rudolfensis is a fossil human species discovered by Bernard Ngeneo, a member of a team led by anthropologist Richard Leakey and zoologist Meave Leakey in 1972, at Koobi Fora on the east side of Lake Rudolf (now Lake Turkana) in Kenya. The scientific name Pithecanthropus rudolfensis was proposed in 1978[1] by V. P. Alekseyev later changed to Homo rudolfensis by Bernard Wood, for the specimen Skull 1470 (KNM ER 1470). Skull 1470 has an estimated age of 1.9 million years.

Originally thought to be a member of the species Homo habilis, the fossil was the center of much debate concerning its species. Assigned initially to Homo habilis, the skull was at first incorrectly dated at nearly three million years old. The differences in this skull, when compared to others of the Homo habilis species, are too pronounced, leading to the presumption of a Homo rudolfensis species, contemporary with Homo habilis. It is not certain if H. rudolfensis was ancestral to the later species in Homo, or if H. habilis was, or if some third species yet to be discovered was. In March 2007, a team led by Timothy Bromage, an anthropologist at New York University, reconstructed the skull of KNMER 1470. The new construction looked very ape-like (possibly due to an exaggerated rotation of the skull) and the cranial capacity based on the new construction was reported to be downsized from 752 cm to about 526 cm, although this seemed to be a matter of some controversy. Bromage published his results in 2008 where the cranial capacity was now estimated at 700 cm. Bromage said his teams reconstruction included biological principles not known at the time of the skulls discovery, which state that a mammals eyes, ears and mouth must be in precise relationships relative to one another.

Australopithecus sediba
Australopithecus sediba is a species of Australopithecus of the early Pleistocene, identified based on fossil remains dated to about 2 million years ago. The species is known from at least four partial skeletons discovered in the Malapa Fossil Site at the Cradle of Humankind World Heritage Site in South Africa, one a juvenile male (MH1, the holotype), an adult female (MH2), at least one other adult and an 18-month-old infant. The MH1 and MH2 fossils were buried together, and have been dated to between 1.977 and 1.980 million years ago. Over two hundred and twenty fragments from the species have been recovered to date. The partial skeletons were initially described in two papers in the journal Science by American and South African palaeo-anthropologist Lee R. Berger and colleagues as a newly discovered species of early human ancestor called Australopithecus sediba ("sediba" meaning "natural spring" or "well" in the Sotho language).

Discovery
The first specimen of A. sediba was found by paleoanthropologist Lee Berger's nine-year-old son, Matthew, on August 15, 2008.[6] While exploring near his father's dig site in the dolomitic hills north of Johannesburg, on the Malapa Nature Reserve, Matthew stumbled upon a fossilized bone. The boy alerted his father of the find, who could not believe what he saw, a hominid clavicle and upon turning the block over; "sticking out of the back of the rock was a mandible with a tooth, a canine, sticking out. And I almost died", he later recalled. The fossil turned out to belong to a 4 ft 2 in (1.27 m) juvenile male, the skull of which was discovered in March 2009 by Berger's team. The find was announced to the public on April 8, 2010. Also found at the Malapa archeological site were a variety of animal fossils, including saber-toothed cats, mongooses, and antelopes. Berger and geologist Paul Dirks speculated that the animals might have fallen into a deep, 100150-foot (3046 m) "death-trap", perhaps lured by the scent of water. The bodies may have then been swept into a pool of water rich with lime, and with sand at the bottom, making it possible for the remains to become fossilized.

Age estimates
The fossil was dated using a combination of palaeomagnetism and uranium-lead (U-Pb) dating by Andy Herries (La Trobe University, Australia), Robyn Pickering (U. Melbourne, Australia) and Jan Kramers (U. Johannesburg). U-Pb dating of the underlying flowstone indicates that the fossils are not older than ~2.0 Ma. The occurrence of species of animal that became extinct at ~1.5 Ma indicate the deposit is not younger than 1.5 Ma. The sediments have a 'normal' magnetic polarity and the only major period between 2.0 and 1.5 Ma when this occurred is the Olduvai sub-Chron between 1.95 and 1.78 Ma. As such, the fossils were originally dated to ~1.95 Ma. Recent dating of a capping flowstone illustrated this was not possible and the normal magnetic polarity sediments have since been correlated to the 3000 year long Pre-Olduvai event at ~1.977 Ma.

Morphology and interpretations

Because of the wide range of mosaic features exhibited in both cranial and post cranial morphology, the authors suggest that A. sediba may be a transitional species between the southern African A. africanus (the Taung Child, Mrs. Ples) and either Homo habilis or even the later H. erectus (Turkana boy, Java man, Peking man). The cranial capacity of MH1, which has been estimated to be at 95% of adult capacity (420 ml/cc), is at the higher end of the range for A. africanus and far from the lower range of early Homo (631 ml/cc), but the mandible and tooth size are quite gracile and similar to what one would expect to find in H. erectus, so similar that if found in isolation without other skeletal remains could be classified as Homo based on tooth and mandible size. However, the cusp spacing is more like Australopithecus. Regardless of whether Australopithecus sediba is an ancestor of early Homo or not, our understanding of the range of variation in early hominins has been greatly increased with the finding of these new specimens. A. sediba compared to its ancestor species A. africanus on the whole is described by Berger et al. as more derived towards Homo than A. garhi, especially showing a number of synapomorphies taken to anticipate the reorganization of the pelvis in H. erectus, associated with "more energetically efficient walking and running". The femur and tibia are fragmentary and the foot is more primitive. Its cranial capacity is estimated at around 420450 cm3 (2627 cu in), about one-third that of modern humans. A. sediba had a surprisingly modern hand, whose precision grip suggests it might have been another tool-making Australopithecus. As opposed to the authors of the initial description, who interpreted both fossils as a possible transitional species between Australopithecus and Homo, other palaeoanthropologists are reluctant to do so. In an accompanying news article published with the initial descriptions in 2010, detractors of the idea that A. sediba might be ancestral to the genus Homo (e.g. Tim White and Ron Clarke) suggest that the fossils could be a late southern African branch of Australopithecus, co-existing with already existing members of the Homo genus. This interpretation is based on the observation that the lower jaw, discovered by Friedemann Schrenk, of a 2.5 million year old fossil attributed to H. rudolfensis is the oldest known fossil ascribed to the Homo genus. This specimen is thus presumed to be older than the Australopithecus sediba fossils. These critics continue to ascribe to A. africanus the status of precursor of the genus Homo genus. Criticism has been raised in a news report on the find in Nature magazine that the authors of the initial description have failed to take the wealth of variation within A. africanus into account, prior to defining the finds as an independent species. Additionally, the basing of the description of the species largely on the skeleton of a juvenile specimen has been subject to criticism, given that there is no certain way of saying to what extent adults would differ from juvenile specimens. More recent reviews appear to have accepted the specific status of A. sediba as an independant species from A. africanus and other early hominins . Additionally, the use of juvenile specimens as Type specimens has been a common practice in palaeoanthropology, with even the type specimens of Homo habilis and Australopithecus africanus being juvenile specimens.

Homo georgicus

The remains of a new hominid, subsequently named Homo georgicus, were discovered in Dmanisi, Republic of Georgia in 1991 by a team led by Georgian paleoanthropologist David Lordkipanidze (Lordkipanidze et al. 2006; Vekua et al. 2002). Skulls and several near complete skeletons have been found, which have been reliably dated to 1.77 mya. Homo georgicus was bipedal, but was rather small. Males stood only about four feet (1.3 m) tall, and the females were even smaller. Cranial measurements of the well-preserved skull D2700 indicate the brain of H. georgicus was about half as large (around 600cc) as that of a modern human being the smallest of any adult hominid yet found outside Africa other than that of the far more recent Homo floresiensis. Tooth-wear patterns and remains found at the site show H. georgicus had an omnivorous diet, but there is no evidence of the use of fire. Crude stone tools have been found in association with the remains

(see picture >>). It has been suggested that Homo georgicus may represent a link between Homo habilis and Homo erectus. With an age of about 1.8 million years, the remains of H. georgicus do in fact date to a period when H. habilis and H. erectus overlapped in time. Moreover, georgicus individuals are about the same size as the few adult habilis specimens known, and their skulls are rather similar. However, H. habilis is known only from sub-Saharan Africa. Since these fossils were found on the eastern shore of the Black Sea, it's unclear though some have made the suggestion that H. georgicus was the first hominid to settle in Europe. The oldest human remains found on actual European soil are those from the Sima del Elefante in Spain, which date to 1.11.2 mya (Carbonell et al. 2008). For the present, about the only sure conclusion is that H. georgicus represents a new and interesting twig on the hominid bush. Homo georgicus is the subspecies name sometimes used to describe fossil skulls and jaws found in Dmanisi, Georgia, which although first proposed as a separate species is now classified within H. erectus. A partial skeleton was discovered in 2001. The fossils are about 1.8 million years old. The remains were first discovered in 1991 by Georgian scientist, David Lordkipanidze, accompanied by an international team which unearthed the remains. Implements and animal bones were found alongside the ancient human remains. At first, scientists thought they had found mandibles and skulls belonging to Homo ergaster, but size differences led them to name a new species, Homo georgicus, which was posited as a descendant of Homo habilis and ancestor of Asian Homo erectus. This classification was not upheld and the fossil is now considered a divergent subgroup of of Homo erectus, sometimes called Homo erectus georgicus'. At around 600 cubic centimetres (37 cu in) brain volume, the skull D2700 is dated to 1.77 million years old and in good condition offering insights in comparison to the modern human cranial morphology. At the time of discovery the cranium was the smallest and most primitive Hominina skull ever discovered outside of Africa. However, in 2003 a significantly smaller brained hominid was found on the isle of Flroes H. erectus floresiensis. Homo erectus georgicus exhibits strong sexual dimorphism with males being significantly larger than females. Subsequently, four fossil skeletons were found, showing a species primitive in its skull and upper body but with relatively advanced spines and lower limbs, providing greater mobility. They are now thought not to be a separate species, but to represent a stage soon after the transition between Homo habilis and H. erectus, and have been dated at 1.8 million years before the present, according to the leader of the project, David Lordkipanidze. The assemblage includes one of the largest Pleistocene Homo mandibles (D2600), one of the smallest Lower Pleistocene mandibles (D211), a nearly complete sub adult (D2735), and a completely toothless specimen (D3900).

Classification and special distinction

Many paleoanthropologists still debate the definition of H. erectus and H. ergaster as separate species. Several scholars suggested to drop the taxon Homo erectus, and suggested instead to equate Homo erectus with archaic Homo sapiens Some call H. ergaster the direct African ancestor of H. erectus, proposing that it emigrated out of Africa and migrated into Asia, branching into a distinct species.[32] Most dispense with the species-name ergaster, making no distinction between such fossils as the Turkana Boy and Peking Man.[citation needed] Though "Homo ergaster" has gained some acceptance as a valid taxon, these two are still usually defined as distinct African and Asian populations of the larger species H. erectus. While some have argued (and insisted) that Ernst Mayr's biological species definition cannot be used here to test the above hypotheses, one can, however, examine the amount of morphological cranial variation within known H. erectus / H. ergaster specimens, and compare it to what one sees in disparate extant groups of primates with similar geographical distribution or

close evolutionary relationship. Thus, if the amount of variation between H. erectus and H. ergaster is greater than what one sees within a species of, say, macaques, then H. erectus and H. ergaster may be considered two different species. The extant model of comparison is very important, and selecting appropriate species can be difficult. (For example, the morphological variation among the global population of H. sapiens is incredibly small[33] and our own special diversity may not be a trustworthy comparison.) As an example, fossils found in Dmanisi in the Republic of Georgia were originally described as belonging to another closely related species, Homo georgicus, but subsequent examples showed their variation to be within the range of Homo erectus, and they are now classified as Homo erectus georgicus. H. erectus had a cranial capacity greater than that of Homo habilis (although the Dmanisi specimens have distinctively small crania): the earliest remains show a cranial capacity of 850 cm, while the latest Javan specimens measure up to 1100 cm,[34] overlapping that of H. sapiens.; the frontal bone is less sloped and the dental arcade smaller than the australopithecines'; the face is more orthognatic (less protrusive) than either the australopithecines' or H. habilis's, with large brow-ridges and less prominent zygomata (cheekbones). These early hominins stood about 1.79 m (5 ft 10 in), and were more robust than modern humans. The sexual dimorphism between males and females was slightly greater than seen in H. sapiens, with males being about 25% larger than females. However, their dimorphism is drastically lesser than that of the earlier Australopithecus genus. The discovery of the skeleton KNM-WT 15000, "Turkana boy" (Homo ergaster), made near Lake Turkana, Kenya by Richard Leakey and Kamoya Kimeu in 1984, is one of the most complete hominid-skeletons discovered, and has contributed greatly to the interpretation of human physiological evolution.

Homo ergaster
Homo ergaster is an extinct chronospecies of Homo that lived in eastern and southern Africa during the early Pleistocene, between 1.8 million and 1.3 million years ago. There is still disagreement on the subject of the classification, ancestry, and progeny of H. ergaster, but it is now widely accepted to be the direct ancestor of later hominids such as Homo heidelbergensis, Homo sapiens, and Homo neanderthalensis rather than Asian Homo erectus. It is one of the earliest members of the genus Homo, possibly descended from, or sharing a common ancestor with, Homo erectus. The binomial name was published in 1975 by Groves and Mazk. The second part, "ergaster", is derived from the Ancient Greek "workman", in reference to the comparatively advanced lithic technology developed by the species, introducing the Acheulean industry.

Discovery and representative fossils

The South African palaeontologist John T. Robinson first discovered a mandible of a new hominid in southern Africa in 1949; he named the species Telanthropus capensis, though it is now recognised as a member of Homo ergaster.[4] The name was first applied by Colin Groves and Vratislav Mazk to KNM-ER 992, a mandible discovered near Lake Rudolf (now Lake Turkana), Kenya in 1975, which became the type-specimen of the species. The most complete skeleton of H. ergaster (and one of the most complete extinct hominids to date), KNM-WT 15000, was discovered at Lake Turkana, Kenya, in 1984 by paleoanthropologists Kamoya Kimeu and Alan Walker. They nicknamed the 1.6-million-year-old specimen "Turkana Boy".

Classification and special distinction

Many paleoanthropologists still debate the definition of H. ergaster and H. erectus as separate species. Some call H. ergaster the direct African ancestor of H. erectus, proposing that H. ergaster emigrated out of Africa and into Asia, branching into a distinct species. Most dispense with the species-name ergaster, making no distinction between such fossils as the Turkana Boy and Peking Man. Though "Homo ergaster" has gained some acceptance as a valid taxon, H. ergaster and H. erectus are still usually defined as distinct African and Asian populations of the larger species H. erectus. (For the remainder of this article, the name "Homo ergaster" will be used to describe a distinct species for the convenience of continuity in reading.)

H. ergaster may be distinguished from H. erectus by its thinner skull-bones and lack of an obvious supraorbital foramen. It may be distinguished from Homo heidelbergensis by its thinner bones, more protrusive face, and lower forehead. Derived features separating it from earlier species include reduced sexual dimorphism, a smaller, more orthognathous (less protrusive) face, a smaller dental arcade, and a larger cranial capacity (700-900cm in earlier ergaster-specimens, and 900-1100 in later specimens). It is estimated that male H. ergaster stood 1.89 meters (6 ft 2 in) tall. Remains have been found in Tanzania, Ethiopia, Kenya, and South Africa.

Divergence
Homo habilis is generally accepted as the putative ancestor of the genus Homo,[4] and often of H. ergaster most directly. This taxon's status as a legitimate species within "Homo", however, is particularly contentious. H. habilis and H. ergaster coexisted for 200,000-300,000 years, possibly indicating that these species diverged from a common ancestor. It is unclear what genetic influence H. ergaster had on later hominids. Recent genetic analysis has generally supported the Out-of-Africa hypothesis, and this may designate H. ergaster the role of ancestor to all later hominids. Origin and extinction H. ergaster is believed to have diverged from the lineage of H. habilis between 1.9 and 1.8 million years ago; the lineage that emigrated from Africa and fathered H. erectus diverged from the lineage of H. ergaster almost immediately after this. These early descendants of H. habilis may have been discovered in Dmanisi, Georgia. H. ergaster remained stable for ca. 500,000 years in Africa before disappearing from the fossil record around 1.4 million years ago. No identifiable cause has been attributed to this disappearance; the later evolution of the similar H. heidelbergensis in Africa may indicate that this is simply a hole in the record, or that some intermediate species has not yet been discovered.

Use of tools
Homo ergaster used more diverse and sophisticated stone tools than its predecessor, Homo habilis. H. ergaster refined the inherited Oldowan developing the first Acheulean bifacial axes: while the use of Acheulean tools began ca. 1.6 million years ago, the line of H. erectus diverged some 200,000 years before the general innovation of Acheulean technology. Thus the Asian migratory descendants of H. ergaster made no use of any Acheulean technology.

Sociality
Sexual dimorphism in H. ergaster is greatly reduced from its australopithecine ancestors (around 20%), but still greater than dimorphism in modern humans. This diminished dimorphism is speculated to be a sign of reduced competition for mates between males, which may also correspond to the more modern social practices of ergaster. Not only was H. ergaster like modern humans in body, but also more in organisation and sociality than any earlier species. It is conceivable that H. ergaster was the first hominin to harness fire: whether as the containment of natural fire, or as the lighting of artificial fire, is still a matter of contention. It is now assumed, however, that erectus did have control of fire, as well as each other hominin sharing a common ancestor with ergaster. The social organisation of H. ergaster probably resembled that of modern hunter-gatherer societies. Unlike australopithecines, ergaster males presumably did not compete at all for females, which had themselves increased in size greatly in proportion to males. This reduced competition and dimorphism also coincided with an increase in brain size and efficiency of stone tools. Linguistic use Homo ergaster was probably the first hominid to "use a human voice", though its symbolic cognition was probably somewhat limited compared to modern humans. It was thought for a long time that H. ergaster was restricted in the physical ability to regulate breathing and produce complex sounds. This was based on Turkana Boy's cervical vertebrae, which were far narrower than in later humans. Discoveries of cervical vertebrae in Dmanisi, Georgia some 300,000 years older than those of Turkana Boy are well within the normal human range. It has been established, furthermore, that the Turkana Boy probably suffered from a disease of the spinal column that resulted in narrower cervical vertebrae than in modern humans (as well as the older Dmanisi finds). While the Dmanisi finds have not been established definitively as H. ergaster; they are older than Turkana Boy (the only definite ergaster vertebrae on record), and thereby suggest kinship to ergaster. Turkana Boy, therefore, may be an anomaly.

There is no archaeological evidence that Homo ergaster made use of symbolic thought (such as figurative art), but the wellevolved brain and physical capabilities (along with reconfiguration of ergaster's breathing-apparatus) suggest some form of linguistic or symbolic communication.

Homo antecessor
Homo antecessor is an extinct human species (or subspecies) dating from 1.2 million to 800,000 years ago, that was discovered by Eudald Carbonell, Juan Luis Arsuaga and J. M. Bermdez de Castro. H. antecessor is one of the earliest known human varieties in Europe. Various archaeologists and anthropologists have debated how H. antecessor related to other Homo species in Europe, with suggestions that it was an evolutionary link between H. ergaster and H. heidelbergensis, although Richard Klein believes that it was instead a separate species that evolved from H. ergaster. Others believe that H. antecessor is in fact the same species as H. heidelbergensis, who inhabited Europe from 600,000 to 250,000 years ago in the Pleistocene. The best-preserved fossil is a maxilla that belonged to a 10-year-old individual found in Spain. Based on palaeomagnetic measurements, it is thought to be older than 780857 ka. The average brain was 1,000 cm in volume. In 1994 and 1995, 80 fossils of six individuals that may have belonged to the species were found in Atapuerca, Spain. At the site were numerous examples of cuts where the flesh had been flensed from the bones, which indicates that H. antecessor may have practised cannibalism.

Physiology
H. antecessor was about 1.6-1.8 m (5-6 feet) tall, and males weighed roughly 90 kg (200 pounds). Their brain sizes were roughly 1,0001,150 cm, smaller than the 1,350 cm average of modern humans. Due to its scarcity, very little more is known about the physiology of H. antecessor, yet it was likely to have been more robust than H. heidelbergensis. According to Juan Luis Arsuaga, one of the co-directors of the excavation in Burgos, H. antecessor might have been right-handed, a trait that makes the species different from the other apes. This hypothesis is based on tomography techniques. Arsuaga also claims that the frequency range of audition is similar to H. sapiens, which makes him believe that H. antecessor used a symbolic language and was able to reason. Arsuaga's team is currently pursuing a DNA map of H. antecessor after elucidating that of a bear that lived in northern Spain some 500,000 years ago. Based on teeth eruption pattern, the researchers think that H. antecessor had the same development stages as H. sapiens, though probably at a faster pace. Other features acquired by the species are a protruding occipital bun, a low forehead and a lack of a strong chin. Some of the remains are almost indistinguishable from the fossil attributable to the 1.5 million year old Turkana Boy, belonging to H. ergaster.

Fossil sites
The only known fossils of H. antecessor are from two sites in the Sierra de Atapuerca region of northern Spain (Gran Dolina and Sima del Elefante). Gran Dolina Archaeologist Eudald Carbonell i Roura of the Universidad Rovira i Virgili in Tarragona, Spain and palaeoanthropologist Juan Luis Arsuaga Ferreras of the Complutense University of Madrid discovered Homo antecessor remains at the Gran Dolina site in the Sierra de Atapuerca, east of Burgos. The H. antecessor remains have been found in level 6 (TF6) of the Gran Dolina site. Over 80 bone fragments from six individuals were uncovered in 1994 and 1995. The site had also included roughly 200 stone tools and about 300 animal bones. Stone tools including a stone carved knife were found along with the

ancient hominin remains. All these remains were dated at least 780,000 years old. The best-preserved remains are a maxilla (upper jawbone) and a frontal bone of an individual who died at the age of 1011. Sima del Elefante On June 29th, 2007, Spanish researchers working at the Sima del Elefante site in the Atapuerca Mountains (Spain) announced that they had recovered a molar dated to 1.11.2 million years ago. The molar was described as "well worn" and from an individual between 20 and 25 years of age. Additional findings announced on 27 March 2008 included the discovery of a mandible fragment, stone flakes, and evidence of animal bone processing. Suffolk, England In 2005 flint tools and teeth from the water vole Mimomys savini, a key dating species, were found in the cliffs at Pakefield near Lowestoft in Suffolk. This suggests that hominins can be dated in England to 700,000 years ago, potentially a cross between Homo antecessor and Homo heidelbergensis. Norfolk, England In 2010 stone tools finds were reported in Happisburgh, Norfolk, England, believed to have been used by H. antecessor, suggesting that the early hominin species also lived in England about 950,000 years ago the earliest known population of the genus Homo in Northern Europe.

Homo heidelbergensis
Homo heidelbergensis ("Heidelberg Man", named after the University of Heidelberg) is an extinct species of the genus Homo which may be the direct ancestor of both Homo neanderthalensis in Europe and Homo sapiens. The best evidence found for these hominins date between 600,000 and 400,000 years ago. H. heidelbergensis stone tool technology was very close to that of the Acheulean tools used by Homo erectus.

Morphology and interpretations

Both H. antecessor and H. heidelbergensis are likely to be descended from the morphologically very similar Homo ergaster from Africa. But because H. heidelbergensis had a larger brain-case with a typical cranial volume of 11001400 cm overlapping the 1350 cm average of modern humans and had more advanced tools and behavior, it has been given a separate species classification. The species was tall, 1.8 m (6.0 ft) on average, and more muscular than modern humans. Males may have weighed 100 kg (220 lb). According to Professor Lee R. Berger of the University of Witwatersrand, numerous fossil bones indicate some populations of Heidelbergensis were "giants" routinely over 2.13 m (7 ft) tall and inhabited South Africa between 0.5 million and 300,000 years ago. Social behavior Recent findings in a pit in Atapuerca (Spain) of 28 human skeletons suggest that H. heidelbergensis may have been the first species of the Homo genus to bury their dead. Some experts believe that H. heidelbergensis, like its descendant H. neanderthalensis, acquired a primitive form of language. No forms of art or sophisticated artifacts other than stone tools have been uncovered, although red ochre, a mineral that can be used to create a red pigment which is useful as a paint, has been found at Terra Amata excavations in the south of France. Language The morphology of the outer and middle ear suggests they had an auditory sensitivity similar to modern humans and very different from chimpanzees. They were probably able to differentiate between many different sounds. Dental wear analysis suggests they were as likely to be right-handed as modern people.

H. heidelbergensis was a close relative (most probably a migratory descendant) of Homo ergaster. H. ergaster is thought to be the first hominin to vocalize and that as H. heidelbergensis developed more sophisticated culture proceeded from this point. Evidence of hunting A number of 400,000-year-old wooden projectile spears were found at Schningen in northern Germany. These are thought to have been made by H. erectus or H. heidelbergensis. Generally, projectile weapons are more commonly associated with H. sapiens. The lack of projectile weaponry is an indication of different sustenance methods, rather than inferior technology or abilities. The situation is identical to that of native New Zealand Mori, modern H. sapiens, who also rarely threw objects, but used spears and clubs instead.

Divergent evolution
Because of the radiation of H. heidelbergensis out of Africa and into Europe, the two populations were mostly isolated during the Wolstonian Stage and Ipswichian Stage, the last of the prolonged Quaternary glacial periods. Neanderthals diverged from H. heidelbergensis probably some 300,000 years ago in Europe, during the Wolstonian Stage; H. sapiens probably diverged between 200,000 and 100,000 years ago in Africa. Such fossils as the Atapuerca skull and the Kabwe skull bear witness to the two branches of the H. heidelbergensis tree. Homo neanderthalensis retained most of the features of H. heidelbergensis after its divergent evolution. Though shorter, Neanderthals were more robust, had large brow-ridges, a slightly protruding face and lack of prominent chin. With the possible exception of Cro-Magnon Man, they also had a larger brain than all other hominins. Homo sapiens, on the other hand, have the smallest brows of any known hominin, are tall and lanky, and have a flat face with a protruding chin. H. sapiens have a larger brain than H. heidelbergensis, and a smaller brain than H. neanderthalensis, on average. To date, H. sapiens is the only known hominin with a high forehead, flat face, and thin, flat brows. Some believe that H. heidelbergensis is a distinct species, and some that it is a cladistic ancestor to other Homo forms sometimes improperly linked to distinct species in terms of populational genetics. Some scenarios of survival include

H. heidelbergensis > H. neanderthalensis H. heidelbergensis > H. rhodesiensis > H. sapiens idaltu > H. sapiens sapiens

Those supporting a multiregional origin of modern humans envision fertile reproduction between many evolutionary stages and homo walking,[9] or gene transfer between adjacent populations due to gene passage and spreading in successive generations.

Discovery
The first fossil discovery of this species was made on October 21, 1907, and came from Mauer where the workman Daniel Hartmann spotted a jaw in a sandpit. The jaw (Mauer 1) was in good condition except for the missing premolar teeth, which were eventually found near the jaw. The workman gave it to Professor Otto Schoetensack from the University of Heidelberg, who identified and named the fossil. The next H. heidelbergensis remains were found in Steinheim an der Murr, Germany (the Steinheim Skull, 350kya); Arago, France (Arago 21); Petralona, Greece; and Ciampate del Diavolo, Italy. Boxgrove Man In 1994 British scientists unearthed a lower hominin tibia bone just a few miles away from the English Channel, along with hundreds of ancient hand axes, at the Boxgrove Quarry site. A partial leg bone is dated to between 478,000 and 524,000 years old. H. heidelbergensis was the early proto-human species that occupied both France and Great Britain at that time; both locales were connected by a landmass during that epoch. Prior to Gran Dolina, Boxgrove offered the earliest hominid occupants in Europe.

The tibia had been gnawed by a large carnivore, suggesting that he had been killed by a lion or wolf or that his unburied corpse had been scavenged after death. Sima de los Huesos Beginning in 1992, a Spanish team has located more than 5,500 human bones dated to an age of at least 350,000 years in the Sima de los Huesos site in the Sierra de Atapuerca in northern Spain. The pit contains fossils of perhaps 28 individuals together with remains of Ursus deningeri and other carnivores and a biface called Excalibur. It is hypothesized that this Acheulean axe made of red quartzite was some kind of ritual offering for a funeral. Ninety percent of the known H. heidelbergensis remains have been obtained from this site. The fossil pit bones include:

A complete cranium (skull 5), nicknamed Migueln, and fragments of other crania, such as skull 4, nicknamed Agamenn and skull 6, nicknamed Rui (from El Cid, a local hero). A complete pelvis (pelvis 1), nicknamed Elvis, in remembrance of Elvis Presley. Mandibles, teeth, and many postcranial bones (femurs, hand and foot bones, vertebrae, ribs, etc.)

Indeed, nearby sites contain the only known and controversial Homo antecessor fossils. There is current debate among scholars whether Sima de los Huesos is H. heidelbergensis, or another hominin that doesn't fit in the direct line from H. antecessor to H. neanderthalensis. Suffolk, England In 2005 flint tools and teeth from the water vole Mimomys savini, a key dating species, were found in the cliffs at Pakefield near Lowestoft in Suffolk. This suggests that hominins can be dated in England to 700,000 years ago, potentially a cross between Homo antecessor and Homo heidelbergensis.

Homo rhodesiensis
Homo rhodesiensis (Rhodesian man) is a hominin species described from the fossil Kabwe skull. Other morphologicallycomparable remains have been found from the same, or earlier, time period in southern Africa (Hopefield or Saldanha), East Africa (Bodo, Ndutu, Eyasi, Ileret) and North Africa (Sal, Rabat, Dar-es-Soltane, Djbel Irhoud, Sidi Aberrahaman, Tighenif). These remains were dated between 300,000 and 125,000 years old.

Discovery
Kabwe skull or Kabwe cranium, or Broken Hill 1 is the type specimen. The cranium was found in an lead and zinc mine in Broken Hill, Northern Rhodesia (now Kabwe, Zambia) in 1921 by Tom Zwiglaar, a Swiss miner. In addition to the cranium, an upper jaw from another individual, a sacrum, a tibia, and two femur fragments were also found. The skull was dubbed Rhodesian Man at the time of the find, but is now commonly referred to as the Broken Hill Skull or the Kabwe Cranium. The association between the bones is unclear, but the tibia and femur fossils are usually associated with the skull. Rhodesian Man is dated to be between 125,000 and 300,000 years old. Cranial capacity of the Broken Hill skull has been estimated at 1,100 cm, which, when coupled with the more recent dating, makes any direct link to older skulls unlikely and negates the

1.75 to 2.5 million year earlier erroneous dating. Bada & al (1974) published direct date of 110 ka for this specimen measured by aspartic acid racemisation. The destruction of the paleoanthropological site has made layered dating impossible. The skull is from an extremely robust individual, and has the comparatively largest brow-ridges of any known hominid remains. It was described as having a broad face similar to Homo neanderthalensis (i.e. large nose and thick protruding brow ridges), and has been interpreted as an "African Neanderthal". However, when regarding the skulls extreme robustness, recent research has pointed to several features intermediate between modern Homo sapiens and Neanderthal. Most current experts believe Rhodesian Man to be within the group of Homo heidelbergensis though other designations such as Archaic Homo sapiens and Homo sapiens rhodesiensis have also been proposed. According to Tim White, it is probable that Rhodesian Man was the ancestor of Homo sapiens idaltu (Herto Man), which would be itself at the origin of Homo sapiens sapiens. No direct linkage of the species can so far be determined. The skull has cavities in ten of the upper teeth and is considered the oldest occurrence of cavities for a hominid. Pitting indicates significant infection before death and implies a cause of death attributable to dental infection or chronic ear infection. Another specimen "the hominid from Lake Ndutu" may approach 400,000 years old, and Clarke in 1976 classified it as Homo erectus. Undirect cranial capacity estimate is 1100 ml. Also supratoral sulcus morphology and presence of protuberance as suggested by Philip Rightmire : give the Nudutu occiput an appearance which is also unlike that of Homo Erectus but Stinger 1986 pointed that thickened iliac pillar is typical for Homo erectus.

Classification
In Africa, there is a distinct difference in the Acheulian tools made before and after 600,000 years ago with the older group being thicker and less symmetric and the younger being more extensively trimmed. This may be connected with the appearance (some 300,000 years later) of Homo rhodesiensis in the archaeological record at this time who may have contributed this more sophisticated approach. Rupert Murrill has studied the relations between Archanthropus skull of Petralona (Chalcidice, Greece) and Rhodesian Man. Most current experts believe Rhodesian Man to be within the group of Homo heidelbergensis though other designations such as Homo sapiens arcaicus and Homo sapiens rhodesiensis have also been proposed. According to Tim White, it is probable that Homo rhodesiensis was the ancestor of Homo sapiens idaltu (Herto Man), which would be itself at the origin of Homo sapiens sapiens. No direct linkage of the species can so far be determined.

Homo Neanderthalensis
The Neanderthal, sometimes spelled Neandertal, is an extinct member of the Homo genus known from Pleistocene specimens found in Europe and parts of western and central Asia. Neanderthals are classified either as a subspecies of Homo sapiens (Homo sapiens neanderthalensis) or as a separate human species (Homo neanderthalensis). Genetic evidence suggests interbreeding took place with anatomically modern humans between roughly 80,000 and 50,000 years ago in the Middle East, resulting in 14% of the genome of people from Eurasia having been contributed by Neanderthals. The first proto-Neanderthal traits appeared in Europe as early as 600,000350,000 years ago. Proto-Neanderthal traits are occasionally grouped with another phenetic 'species', Homo heidelbergensis, or a migrant form, Homo rhodesiensis. The youngest Neanderthal finds include Hyena Den (UK), considered older than 30,000 years ago, while the Vindija (Croatia) Neanderthals have been re-dated to between 33,000 and 32,000 years ago. No definite specimens younger than 30,000 years ago have been found; however, evidence of fire by Neanderthals at Gibraltar indicate they may have survived there until 24,000 years ago. Cro-Magnon or early modern human skeletal remains with 'Neanderthal traits' were found in Lagar Velho (Portugal), dated to 24,500 years ago and interpreted as indications of extensively admixed populations. Several cultural assemblages have been linked to the Neanderthals in Europe. The earliest, the Mousterian stone tool culture, dates to about 300,000 years ago. Later Mousterian culture is also seen in Asia, and in Africa dated after 150,000 years ago at

the Jebel Irhoud site located 620 km south of Gibraltar. Late Mousterian artifacts were found in Gorham's Cave on the remote south-facing coast of Gibraltar. Other tool cultures associated with Neanderthal include Chtelperronian, Aurignacian, and Gravettian, developed with gradual continuity not distributed by population change. Neanderthal cranial capacity is thought to have been as large as that of a Homo sapiens, perhaps larger, indicating their brain size may have been comparable, or larger, as well. In 2008, a group of scientists created a study using three-dimensional computer-assisted reconstructions of Neanderthal infants based on fossils found in Russia and Syria. The study showed Neanderthal and modern human brains were the same size at birth, but by adulthood, the Neandertal brain was larger than the modern human brain. They were much stronger than Homo sapiens, having particularly strong arms and hands. Males stood 164168 cm (6566 in) and females about 152156 cm (6061 in) tall. In 2010 a U.S. researcher reported finding cooked vegetable in the teeth of a Neanderthal skull, contradicting the earlier belief they were exclusively (or almost exclusively) carnivorous and apex predators.

Etymology
The species is named after the Neander valley, located about 12 km (7.5 mi) east of Dsseldorf, Germany. This ravine formed by the river Dssel, widened out by mining, was named Neanderthal in the early 19th century to honour the clergyman and hymn writer Joachim Neander: "Tal" (spelled "Thal" until the German spelling reform of 1901) is the German word for valley. The fossil discovered in the Feldhofer Cave in the limestone cliffs of the Neanderthal in 1856,[17] Neanderthal 1, was known as the "Neanderthal skull" or "Neanderthal cranium" in anthropological literature, and the individual reconstructed on the basis of the skull was occasionally called the "Neanderthal man".[18] The binomial name Homo neanderthalensis, extending the name "Neanderthal man" from the individual type specimen to the entire species, was proposed by the Anglo-Irish geologist William King in 1864. This had priority over the proposal put forward in 1866 by Ernst Haeckel, Homo stupidus.[17] The practice of referring to the members of the species simply as "the Neanderthals", singular "a Neanderthal", emerges in popular literature of the 1920s.[19] The spelling of the German word Thal ("dale, valley"), was changed to Tal in 1901, and the spelling of the valley was also changed accordingly to Neandertal. The former spelling is, however, often retained in English for the hominid. The spelling with th is in addition always used in scientific names throughout the world. In German, however, the modern spelling with t is used in referring to both the hominid and the valley. Neandertal is a widespread alternative spelling in English. The pronunciation of the German name Neandert(h)aler (regardless of spelling) is [neand tal]. American English speakers commonly pronounce it as // (th as in thin), but American scientists usually use /t/. British English speakers usually pronounce it as /t/ followed by a long a as in tar, matching the German pronunciation. The pronunciation /nindrl/ is very common in the United States and is often listed first in US dictionaries, for example American Heritage Dictionary and Random House Dictionaries. The UK pronunciation is /nindrt as shown in the Cambridge Advanced l/, Learner's Dictionary, and Oxford Advanced Learner's Dictionary.

Classification
For some time, scientists have debated whether Neanderthals should be classified as Homo neanderthalensis or "Homo sapiens neanderthalensis", the latter placing Neanderthals as a subspecies of Homo sapiens.Some morphological studies support the view that Homo neanderthalensis is a separate species and not a subspecies. Others, for example University of Cambridge Professor Paul Mellars, say "no evidence has been found of cultural interaction" and evidence from mitochondrial DNA studies have been interpreted as evidence Neanderthals were not a subspecies of H. sapiens, though more recent genomic evidence showed otherwise. And as a result of strong evidence of interbreeding between the two races in order to give fertile offspring, some scientists are being inclined more and more to classify the Neanderthal as a subspecies of H. sapiens, as species are defined by reproductive isolation. Neanderthals evolved from early Homo along a path similar to Homo sapiens, both deriving from a chimp-like ancestor between five and 10 million years ago. Like H. sapiens, Neanderthals are related to Australopithecus, Homo habilis, and

Homo ergaster; the exact descent remains uncertain. The last common ancestor between anatomically modern Homo sapiens and Neanderthals appears to be Homo rhodesiensis, named after an archaic Homo sapiens fossil, Broken hill 1 (Kabwe 1) discovered in the territory of Rhodesia in 1921. Homo rhodesiensis arose in Africa an estimated 0.7 to 1 million years ago. The earliest estimates for Homo rhodesiensis reaching Europe are approximately 800 thousand years ago when a type of human referred to as Homo antecessor or Homo cepranensis already inhabited the region. These two human types may be forerunners to European Homo heidelbergensis; however, stone tools dating from 1.2 to 1.56 million years ago of an unknown creator have been discovered in south-western Europe. The evidence at the Sima de los Huesos (in the Atapuerca cave system on the Iberian Peninsula) suggests Homo heidelbergensis was already in Europe by 600,000 years ago. Molecular phylogenetic analysis suggests Homo rhodesiensis and Homo heidelbergensis continued to intermix until 350,000 years ago, after which they were separate species, and sometime within the last 200,000 years Homo heidelbergensis evolved into Homo neanderthalensis, the classic Neanderthal human. It appears the original Neanderthal population was, in fact, more distantly related to today's human than is Homo heidelbergensis. However, recent evidence of successful interbreeding between Neanderthals and modern humans has made that issue moot, at least insofar as some Neanderthal populations were concerned.

Discovery
Neanderthal skulls were first discovered in Engis Caves (fr), in what is now Belgium (1829) by Philippe-Charles Schmerling and in Forbes' Quarry, Gibraltar (1848), both prior to the type specimen discovery in a limestone quarry of the Neander Valley in Erkrath near Dsseldorf in August 1856, three years before Charles Darwin's On the Origin of Species was published.[29] The type specimen, dubbed Neanderthal 1, consisted of a skull cap, two femora, three bones from the right arm, two from the left arm, part of the left ilium, fragments of a scapula, and ribs. The workers who recovered this material originally thought it to be the remains of a bear. They gave the material to amateur naturalist Johann Carl Fuhlrott, who turned the fossils over to anatomist Hermann Schaaffhausen. The discovery was jointly announced in 1857. The original Neanderthal discovery is now considered the beginning of paleoanthropology. These and other discoveries led to the idea these remains were from ancient Europeans who had played an important role in modern human origins. The bones of over 400 Neanderthals have been found since. Timeline

1829: Neanderthal skulls were discovered in Engis, in present-day Belgium. 1848: Neanderthal skull found in Forbes' Quarry, Gibraltar. Called "an ancient human" at the time. 1856: Johann Karl Fuhlrott first recognized the fossil called "Neanderthal man", discovered in Neanderthal, a valley near Mettmann in what is now North Rhine-Westphalia, Germany. 1880: The mandible of a Neanderthal child was found in a secure context and associated with cultural debris, including hearths, Mousterian tools, and bones of extinct animals. 1886: Two nearly perfect skeletons of a man and woman were found at Spy, Belgium at the depth of 16 ft with numerous Mousterian-type implements. 1899: Hundreds of Neanderthal bones were described in stratigraphic position in association with cultural remains and extinct animal bones. 1908: A nearly complete Neanderthal skeleton was discovered in association with Mousterian tools and bones of extinct animals. 1925: Francis Turville-Petre finds the 'Galilee Man' or 'Galilee Skull' in the Zuttiyeh Cave in Wadi Amud in Palestine (now Israel). 19531957: Ralph Solecki uncovered nine Neanderthal skeletons in Shanidar Cave in northern Iraq. 1975: Erik Trinkaus's study of Neanderthal feet confirmed they walked like modern humans.

1987: Thermoluminescence results from Israeli fossils date Neanderthals at Kebara to 60,000 BP and humans at Qafzeh to 90,000 BP. These dates were confirmed by electron spin resonance (ESR) dates for Qafzeh (90,000 BP) and Es Skhul (80,000 BP). 1991: ESR dates showed the Tabun Neanderthal was contemporaneous with modern humans from Skhul and Qafzeh. 1993: A 127.000 years old DNA is found on the child of Sclayn, found in Scladina (fr), Belgium. 1997: Matthias Krings et al. are the first to amplify Neanderthal mitochondrial DNA (mtDNA) using a specimen from Feldhofer grotto in the Neander valley. 2000: Igor Ovchinnikov, Kirsten Liden, William Goodman et al. retrieved DNA from a Late Neanderthal (29,000 BP) infant from Mezmaiskaya Cave in the Caucasus. 2005: The Max Planck Institute for Evolutionary Anthropology launched a project to reconstruct the Neanderthal genome. 2006: The Max Planck Institute for Evolutionary Anthropology announced it planned to work with Connecticut-based 454 Life Sciences to reconstruct the Neanderthal genome. 2009: The Max Planck Institute for Evolutionary Anthropology announced the "first draft" of a complete Neanderthal genome is completed. 2010: Comparison of Neanderthal genome with modern humans from Africa and Eurasia shows that 14% of modern non-African human genome might come from the Neanderthals. 2010: Discovery of Neanderthal tools far away from the influence of Homo sapiens indicate that the species might have been able to create and evolve tools on its own, and therefore be more intelligent than previously thought. Furthermore, it was proposed that the Neanderthals might be more closely related to Homo sapiens than previously thought and that may in fact be a sub species of it.[34] Evidence has more recently emerged that these artifacts are likely of Homo sapiens origin.[35] 2012: Charcoal found next to six paintings of seals in Nerja caves, Malaga, Spain, has been dated to between 42,300 and 43,500 years old. The paintings themselves will be dated in 2013, and if their pigment matches the date of the charcoal, they would be the oldest known cave paintings. Jos Luis Sanchidrin at the University of Cordoba, Spain believes the paintings are more likely to have been painted by Neanderthals than early homo sapiens.

Habitat and range

Early Neanderthals lived in the Last Glacial age for a span of about 100,000 years. Because of the damaging effects the glacial period had on the Neanderthal sites, not much is known about the early species. Countries where their remains are known include most of Europe south of the line of glaciation, roughly along the 50th parallel north, including most of Western Europe, including the south coast of Great Britain, Central Europe and the Balkans,[38] some sites in Ukraine and in western Russia and east of Europe in Siberia to the Altai Mountains and south through the Levant to Indus River. It is estimated that the total Neanderthal population across this habitat range numbered at around 70,000 at its peak. Neanderthal fossils have not been found to date in Africa, but there have been finds rather close to Africa, both at Gibraltar and in the Levant. At some Levantine sites, Neanderthal remains, in fact, date after the same sites were vacated by Homo sapiens. Mammal fossils of the same time period show cold-adapted animals were present alongside these Neanderthals in this region of the Eastern Mediterranean. This implies Neanderthals were better adapted biologically to cold weather than H. sapiens and at times displaced H. sapiens in parts of the Middle East when the climate got cold enough. Homo sapiens appears to have been the only human type in the Nile River Valley during these periods, and Neanderthals are not known to have ever lived south-west of modern Israel. When further climate change caused warmer temperatures, the Neanderthal range likewise retreated to the north along with the cold-adapted species of mammals. Apparently these weather-induced population shifts took place before "modern" people secured competitive advantages over the Neanderthal, as these shifts in range took place well over ten thousand years before "moderns" totally replaced the Neanderthal, despite the recent evidence of some successful interbreeding. There were separate developments in the human line, in other regions such as Southern Africa, that somewhat resembled the European and Western/Central Asian Neanderthals, but these people were not actually Neanderthals. One such example is Rhodesian Man (Homo rhodesiensis) who existed long before any classic European Neanderthals, but had a more modern set of teeth, and arguably some H. rhodesiensis populations were on the road to modern Homo sapiens sapiens.

To date, no intimate connection has been found between these similar people and the Western/Central Eurasian Neanderthals, at least during the same time as classic Eurasian Neanderthals, and H. rhodesiensis seems to have evolved separately and earlier than classic Neanderthals in a case of convergent evolution. It appears incorrect, based on present research and known fossil finds, to refer to any fossil outside Europe or Western and Central Asia as a true Neanderthal. True Neanderthals had a known range that possibly extended as far east as the Altai Mountains, but not farther to the east or south, and apparently not into Africa. At any rate, in Africa the land immediately south of the Neanderthal range was possessed by "modern" H. sap., since at least 160,000 years before the present. Classic Neanderthal fossils have been found over a large area, from northern Germany to Israel and Mediterranean countries like Spain and Italy in the south and from England and Portugal in the west to Uzbekistan in the east. This area probably was not occupied all at the same time. The northern border of their range, in particular, would have contracted frequently with the onset of cold periods. On the other hand, the northern border of their range as represented by fossils may not be the real northern border of the area they occupied, since Middle Palaeolithic-looking artifacts have been found even further north, up to 60 N, on the Russian plain. Recent evidence has extended the Neanderthal range by about 1,250 miles (2,010 km) east into southern Siberia's Altai Mountains.

Anatomy
Neanderthal anatomy was more robust than anatomically modern humans and they had less neotenized skulls.[46]

Behavior
Main article: Neanderthal behavior Neanderthals were largely carnivorous and apex predators; however, new studies do indicate that they had cooked vegetables in their diet. They made advanced tools, had a language (the nature of which is debated) and lived in complex social groups. The Molodova archaeological site in eastern Ukraine suggests some Neanderthals built dwellings using animal bones. A building was made of mammoth skulls, jaws, tusks and leg bones, and had 25 hearths inside.

Genome
Further information: Neanderthal genome project Early investigations concentrated on mitochondrial DNA (mtDNA), which, owing to strictly matrilineal inheritance and subsequent vulnerability to genetic drift, is of limited value in evaluating the possibility of interbreeding of Neanderthals with Cro-Magnon people. In 1997, geneticists were able to extract a short sequence of DNA from Neanderthal bones from 30,000 years ago.[51] The extraction of mtDNA from a second specimen was reported in 2000, and showed no sign of modern human descent from Neanderthals. In July 2006, the Max Planck Institute for Evolutionary Anthropology and 454 Life Sciences announced that they would sequence the Neanderthal genome over the next two years. This genome is expected to be roughly the size of the human genome, three-billion base pairs, and share most of its genes. It was hoped the comparison would expand understanding of Neanderthals, as well as the evolution of humans and human brains.

Svante Pbo has tested more than 70 Neanderthal specimens. Preliminary DNA sequencing from a 38,000-year-old bone fragment of a femur found at Vindija Cave, Croatia, in 1980 showed Homo neanderthalensis and Homo sapiens share about 99.5% of their DNA. From mtDNA analysis estimates, the two species shared a common ancestor about 500,000 years ago. An article appearing in the journal Nature has calculated the species diverged about 516,000 years ago, whereas fossil records show a time of about 400,000 years ago. A 2007 study pushes the point of divergence back to around 800,000 years ago. Edward Rubin of the Lawrence Berkeley National Laboratory in Berkeley, California, states recent genome testing of Neanderthals suggests human and Neanderthal DNA are some 99.5% to nearly 99.9% identical. On 16 November 2006, Lawrence Berkeley National Laboratory issued a press release suggesting Neanderthals and ancient humans probably did not interbreed. Edward M. Rubin, director of the U.S. Department of Energy's Lawrence Berkeley National Laboratory and the Joint Genome Institute (JGI), sequenced a fraction (0.00002) of genomic nuclear DNA (nDNA) from a 38,000-year-old Vindia Neanderthal femur. They calculated the common ancestor to be about 353,000 years ago, and a complete separation of the ancestors of the species about 188,000 years ago. Their results show the genomes of modern humans and Neanderthals are at least 99.5% identical, but despite this genetic similarity, and despite the two species having coexisted in the same geographic region for thousands of years, Rubin and his team did not find any evidence of any significant crossbreeding between the two. Rubin said, "While unable to definitively conclude that interbreeding between the two species of humans did not occur, analysis of the nuclear DNA from the Neanderthal suggests the low likelihood of it having occurred at any appreciable level." In 2008 Richard E. Green et al. from Max Planck Institute for Evolutionary Anthropology in Leipzig, Germany published the full sequence of Neanderthal mitochondrial DNA (mtDNA) and suggested "Neanderthals had a long-term effective population size smaller than that of modern humans." Writing in Nature about Green et al.'s findings, James Morgan asserted the mtDNA sequence contained clues that Neanderthals lived in "small and isolated populations, and probably did not interbreed with their human neighbours. In the same publication, it was disclosed by Svante Pbo that in the previous work at the Max Planck Institute that "Contamination was indeed an issue," and they eventually realized that 11% of their sample was modern human DNA. Since then, more of the preparation work has been done in clean areas and 4-base pair 'tags' have been added to the DNA as soon as it is extracted so the Neanderthal DNA can be identified. With 3 billion nucleotides sequenced, analysis of about 1/3 showed no sign of admixture between modern humans and Neanderthals, according to Pbo. This concurred with the work of Noonan from two years earlier. The variant of microcephalin common outside Africa, which was suggested to be of Neanderthal origin and responsible for rapid brain growth in humans, was not found in Neanderthals. Nor was the MAPT variant, a very old variant found primarily in Europeans. However, an analysis of a first draft of the Neanderthal genome by the same team released in May 2010 indicates interbreeding may have occurred. "Those of us who live outside Africa carry a little Neanderthal DNA in us," said Pbo, who led the study. "The proportion of Neanderthal-inherited genetic material is about 1 to 4 percent. It is a small but very real proportion of ancestry in non-Africans today," says Dr. David Reich of Harvard Medical School in Boston, who worked on the study. This research compared the genome of the Neanderthals to five modern humans from China, France, sub-Saharan Africa, and Papua New Guinea. The finding is that about 1 to 4 percent of the genes of the non-Africans came from Neanderthals, compared to the baseline defined by the two Africans. This indicates a gene flow from Neanderthals to modern humans, i.e., interbreeding between the two populations. Since the three non-African genomes show a similar proportion of Neanderthal sequences, the interbreeding must have occurred early in the migration of modern humans out of Africa, perhaps in the Middle East. No evidence for gene flow in the direction from modern humans to Neanderthals was found. The latter result would not be unexpected if contact occurred between a small colonizing population of modern humans and a much larger resident population of Neanderthals. A very limited amount of interbreeding could explain the findings, if it occurred early enough in the colonization process. While interbreeding is viewed as the most parsimonious interpretation of the genetic discoveries, the authors point out they cannot conclusively rule out an alternative scenario, in which the source population of non-African modern humans was already more closely related to Neanderthals than other Africans were, due to ancient genetic divisions within Africa.

Among the genes shown to differ between present-day humans and Neanderthals were RPTN, SPAG17, CAN15, TTF1 and PCD16.

Extinction hypotheses
Main article: Neanderthal extinction hypotheses The Neanderthals disappear from the fossil record after about 25,000 years ago. The last traces of Mousterian culture (without human specimens) have been found in Gorham's Cave on the remote south-facing coast of Gibraltar, dated 30,000 to 24,500 years ago. Possible scenarios are: 1. Neanderthals were a separate species from modern humans, and became extinct (due to climate change or interaction with humans) and were replaced by H. sapiens moving into its habitat beginning around 80,000 years ago.[66] Competition with H. sapiens probably contributed to Neanderthal extinction. Jared Diamond has suggested a scenario of violent conflict and displacement. 2. Neanderthals were a contemporary subspecies that bred with Homo sapiens and disappeared through absorption (interbreeding theory). 3. A Campanian ignimbrite volcanic super-eruption around 40,000 years ago, followed by a second one a few thousand years later, has been hypothesised as having contributed to the demise of the Neanderthal, based on evidence from Mezmaiskaya cave in the Caucasus Mountains of southern Russia Mitochondrial DNA (mtDNA) analysis of a specimen from Mezmaiskaya Cave is radiocarbon dated to be about 29,000 years BP and therefore from one of the latest living Neanderthal individuals. The sequence shows 3.48% divergence from the Feldhofer Neanderthal4. Phylogenetic analysis places the two Neanderthals from the Caucasus and western Germany together in a clade that is distinct from modern humans, suggesting that their mtDNA types have not contributed to the modern human mtDNA pool. As Paul Jordan notes: "A natural sympathy for the underdog and the disadvantaged lends a sad poignancy to the fate of the Neanderthal folk, however it came about." Jordan, though, does say that there was perhaps interbreeding to some extent, but that populations that remained totally Neanderthal were likely out-competed and marginalized to extinction by the Aurignacians.

Climate change About 55,000 years ago, the weather began to fluctuate wildly from extreme cold conditions to mild cold and back in a matter of a few decades. Neanderthal bodies were well suited for survival in a cold climatetheir barrel chests and stocky limbs stored body heat better than the Cro-Magnons. However, the rapid fluctuations of weather caused ecological changes to which the Neanderthals could not adapt; familiar plants and animals would be replaced by completely different ones within a lifetime. Neanderthal's ambush techniques would have failed as grasslands replaced trees. A large number of Neanderthals would have died during these fluctuations, which peaked about 30,000 years ago. Studies on Neanderthal body structures have shown that they needed more energy to survive than any other species. Their energy needs were up to 100350 calories more per day comparing to projected anatomically modern human males weighing 68.5 kg and females 59.2 kg. When food became scarce, this difference may have played a major role in the Neanderthals' extinction. Coexistence with H. sapiens There is no longer certainty regarding the identity of the humans who produced the Aurignacian culture, even though the presumed westward spread of anatomically modern humans (AMHs) across Europe is still based on the controversial first

dates of the Aurignacian. Currently, the oldest European anatomically modern Homo sapiens is represented by a robust modern-human mandible discovered at Petera cu Oase (southwest Romania), dated to 34,00036,000 years ago. Human skeletal remains from the German site of Vogelherd, so far regarded as the best association between anatomically modern Homo sapiens and Aurignacian culture, were revealed to represent intrusive Neolithic burials into the Aurignacian levels and subsequently all the key Vogelherd fossils are now dated to 3,9005,000 years ago instead. As for now, the expansion of the first anatomically modern humans into Europe cannot be located by diagnostic and well-dated AMH fossils "west of the Iron Gates of the Danube" before 32,000 years ago. Consequently, the exact nature of biological and cultural interaction between Neanderthals and other human groups between 50,000 and 30,000 years ago is currently hotly contested. A new proposal strives to resolve the issue by proposing the Gravettians rather than the Aurignacians as the anatomically modern humans who contributed to the Eurasian genetic pool after 30,000 years ago. Correspondingly, the human skull fragment found at the Elbe River bank at Hahnfersand near Hamburg was once radiocarbon-dated to 36,000 years ago and seen as possible evidence for the intermixing of Neanderthals and anatomically modern humans. It is now dated to the more recent Mesolithic.

Interbreeding hypotheses
An alternative to extinction is that Neanderthals were absorbed into the Cro-Magnon population by interbreeding. This would be counter to strict versions of the Recent African Origin, since it would imply that at least part of the genome of Europeans would descend from Neanderthals. The most vocal proponent of the hybridization hypothesis is Erik Trinkaus of Washington University.[79] Trinkaus claims various fossils as hybrid individuals, including the "child of Lagar Velho", a skeleton found at Lagar Velho in Portugal dated to about 24,000 years ago. In a 2006 publication co-authored by Trinkaus, the fossils found in 1952 in the cave of Petera Muierii, Romania, are likewise claimed as hybrids. Genetic research has confirmed that some admixture took place. The genomes of non-Africans include portions that are of Neanderthal origin, due to interbreeding between Neanderthals and the ancestors of Eurasians in Northern Africa or the Middle East prior to their spread. Rather than absorption of the Neanderthal population, this gene flow appears to have been of limited duration and limited extent. An estimated 1 to 4 percent of the DNA in Europeans and Asians (i.e. French, Chinese and Papua probands) is non-modern, and shared with ancient Neanderthal DNA rather than with Sub-Saharan Africans (i.e. Yoruba and San probands). Nonetheless, more recent genetic studies seem to suggest that modern humans may have mated with "at least two groups" of ancient humans: Neanderthals and Denisovans.

Specimens

Neanderthal 1: Initial Neanderthal specimen found during an archaeological dig in August 1856. Discovered in a limestone quarry at the Feldhofer grotto in Neanderthal, Germany. The find consisted of a skull cap, two femora, the three right arm bones, two of the left arm bones, ilium, and fragments of a scapula and ribs. La Chapelle-aux-Saints 1: Called the Old Man, a fossilized skull discovered in La Chapelle-aux-Saints, France, by A. and J. Bouyssonie, and L. Bardon in 1908. Characteristics include a low vaulted cranium and large browridge typical of Neanderthals. Estimated to be about 60,000 years old, the specimen was severely arthritic and had lost all his teeth, with evidence of healing. For him to have lived on would have required that someone process his food for him, one of the earliest examples of Neanderthal altruism (similar to Shanidar I.) La Ferrassie 1: A fossilized skull discovered in La Ferrassie, France, by R. Capitan in 1909. It is estimated to be 70,000 years old. Its characteristics include a large occipital bun, low-vaulted cranium and heavily worn teeth. Le Moustier: A fossilized skull, discovered in 1909, at the archaeological site in Peyzac-le-Moustier, Dordogne, France. The Mousterian tool culture is named after Le Moustier. The skull, estimated to be less than 45,000 years old, includes a large nasal cavity and a somewhat less developed brow ridge and occipital bun as might be expected in a juvenile. Shanidar 1: Found in the Zagros Mountains in northern Iraq; a total of nine skeletons found believed to have lived in the Middle Paleolithic. One of the nine remains was missing part of its right arm; theorized to have been broken off or

amputated. The find is also significant because it shows that stone tools were present among this tribe's culture. One was buried with flowers, showing that some type of burial ceremony may have occurred.

Chronology
Bones with Neanderthal traits in chronological order. (Sorted by time) Mixed with H. heidelbergensis traits

> 350 ka: Sima de los Huesos c. 500:350 ka ago 350200 ka: Pontnewydd 225 ka ago. 200135 ka: Atapuerca,[88] Vrtesszllos, Ehringsdorf, Casal de'Pazzi, Biache, La Chaise, Montmaurin, Prince, Lazaret, Fontchevade

Typical H. neanderthalensis traits

13545 ka: Krapina, Saccopastore, Malarnaud, Altamura, Gnovce, Denisova, Okladnikov Altai, Pech de l'Az, Tabun 120 ka 1005 ka, Qafzeh9 100, Shanidar 1 to 9 8060 ka, La Ferrassie 1 70 ka, Kebara 60 ka, Rgourdou, Mt. Circeo, Combe Grenal, Erd 50 ka, La Chapelle-aux Saints 1 60 ka, Amud I 538 ka, Teshik-Tash. 4535 ka: Le Moustier 45 ka, Feldhofer 42 ka, La Quina, l'Horus, Hortus, Kulna, ipka, Saint Csaire, Bacho Kiro, El Castillo, Bnolas, Arcy-sur-Cure. < 35 ka: Chtelperron, Figueria Brava, Zafaraya 30 ka, Vogelherd 3?, Vindija 32,400 800 14C B.P. (Vi-208 31,390 220, Vi-207 32,400 1,800 14C B.P.), Velika Peina,

Mixed with AMH traits

< 35 Pestera cu Oase 35 ka, Mlade 31 ka, Pestera Muierii 30 ka (n/s), Lagar Velho 24.5 ka.

Neanderthal reconstructions
Early artistic reconstructions mostly presented Neanderthals as beastly creatures, emphasizing hairiness and rough, dark complexion. More recent reconstructions acknowledge that due to the lineage evolution in European latitude there is reason to believe that Neanderthals were fair-skinned and probably with no more facial hair than modern man. Also, archaeological evidence exists indicating that they may have communicated by speech, used tools and engaged in artistic endeavours. Reconstructions of Neanderthal men, women and children have become much more intelligent-looking and pleasing to the modern eye.

Popular culture
Main article: Neanderthals in popular culture Neanderthals often appear in popular culture, often in unflattering and inaccurate light, much in the same way as "dinosaur" is also used.

Homo floresiensis
Homo floresiensis ("Flores Man", nicknamed "hobbit" and "Flo") is a possible species, now extinct, in the genus Homo. The remains were discovered in 2003 on the island of Flores in Indonesia. Partial skeletons of nine individuals have been recovered, including one complete cranium (skull). These remains have been the subject of intense research to determine whether they represent a species distinct from modern humans, and the progress of this scientific controversy has been closely followed by the news media at large. This hominin is remarkable for its small body and brain and for its survival until

relatively recent times (possibly as recently as 12,000 years ago). Recovered alongside the skeletal remains were stone tools from archaeological horizons ranging from 94,000 to 13,000 years ago. The discoverers (archaeologist Mike Morwood and colleagues) proposed that a variety of features, both primitive and derived, identify these individuals as belonging to a new species, H. floresiensis, within the taxonomic tribe of Hominini. Hominini currently comprises the extant species human (the only living member of the genus Homo), bonobo (genus Pan), and chimpanzee (genus Pan); their ancestors; and the extinct lineages of their common ancestor. The discoverers also proposed that H. floresiensis lived contemporaneously with modern humans (Homo sapiens) on Flores. Doubts that the remains constitute a new species were soon voiced by the Indonesian anthropologist Teuku Jacob, who suggested that the skull of LB1 was a microcephalic modern human. Two studies by paleoneurologist Dean Falk and her colleagues (2005, 2007) rejected this possibility. Falk et al. (2005) has been rejected by Martin et al. (2006) and Jacob et al. (2006) and defended by Morwood (2005) and Argue, Donlon et al. (2006). Two orthopedic researches published in 2007 both reported evidence to support species status for H. floresiensis. A study of three tokens of carpal (wrist) bones concluded there were similarities to the carpal bones of a chimpanzee or an early hominin such as Australopithecus and also differences from the bones of modern humans. A study of the bones and joints of the arm, shoulder, and lower limbs also concluded that H. floresiensis was more similar to early humans and apes than modern humans. In 2009, the publication of a cladistic analysis and a study of comparative body measurements provided further support for the hypothesis that H. floresiensis and Homo sapiens are separate species. Critics of the claim for species status continue to believe that these individuals are Homo sapiens possessing pathologies of anatomy and physiology. A second hypothesis in this category is that the individuals were born without a functioning thyroid, resulting in a type of endemic cretinism (myxoedematous, ME).

Discovery
The specimens were discovered on the Indonesian island of Flores in 2003 by a joint Australian-Indonesian team of archaeologists looking for evidence of the original human migration of Homo sapiens from Asia to Australia.[1][3] They were not expecting to find a new species, and were surprised at the recovery of a nearly complete skeleton of a hominin they dubbed LB1 because it was unearthed inside the Liang Bua Cave. Subsequent excavations recovered seven additional skeletons, dating from 38,000 to 13,000 years ago. An arm bone provisionally assigned to H. floresiensis is about 74,000 years old. The specimens are not fossilized and have been described as having "the consistency of wet blotting paper"; once exposed, the bones had to be left to dry before they could be dug up. Researchers hope to find preserved mitochondrial DNA to compare with samples from similarly unfossilized specimens of Homo neanderthalensis and H. sapiens. Sophisticated stone implements of a size considered appropriate to the 1-meter-tall human are also widely present in the cave. The implements are at horizons from 95,000 to 13,000 years ago and are associated with (found in the same stratigraphic layer as) an elephant of the extinct genus Stegodon (which was widespread throughout Asia during the Quaternary), presumably the prey of LB1. They also shared the island with giant rats, Komodo dragons, and even larger species of lizards. Homo sapiens reached the region by around 45,000 years ago. Homo floresiensis was unveiled on 28 October 2004, and was swiftly nicknamed the "Hobbit", after the main character of J. R. R. Tolkien's book The Hobbit, and a proposed scientific name for the species was Homo hobbitus. It was initially placed in its own genus, Sundanthropus floresianus ("Sunda man from Flores"), but reviewers of the article felt that the cranium, despite its size, belonged in the genus Homo.

Anatomy
The most important and obvious identifying features of H. floresiensis are its small body and small cranial capacity. Brown and Morwood also identified a number of additional, less obvious features that might distinguish LB1 from modern H. sapiens, including the form of the teeth, the absence of a chin, and the lesser angle in the head of the humerus (upper arm bone). Each of these putative distinguishing features has been heavily scrutinized by the scientific community, with different

research groups reaching differing conclusions as to whether these features support the original designation of a new species, or whether they identify LB1 as a severely pathological H. sapiens. The discovery of additional partial skeletons has verified the existence of some features found in LB1, such as the lack of a chin, but Jacob and other research teams argue that these features do not distinguish LB1 from local H. sapiens morphology. Lyras et al. have asserted, based on 3D-morphometrics, that the skull of LB1 differs significantly from all H. sapiens skulls, including those of small-bodied individuals and microcephalics, and is similar to the skull of Homo erectus alone. Small bodies The first set of remains to have been found, LB1, was chosen as the type specimen for the proposed species. LB1 is a fairly complete skeleton, including a nearly complete cranium (skull), determined to be that of a 30-year-old female. LB1 has been nicknamed the Little Lady of Flores or "Flo". LB1's height has been estimated at about 1.06 m (3 ft 6 in). The height of a second skeleton, LB8, has been estimated at 1.09 m (3 ft 7 in) based on measurements of its tibia. These estimates are outside the range of normal modern human height and considerably shorter than the average adult height of even the smallest modern humans, such as the Mbenga and Mbuti (< 1.5 m (4 ft 11 in)), Twa, Semang (1.37 m (4 ft 6 in) for adult women) of the Malay Peninsula, or the Andamanese (1.37 m (4 ft 6 in) for adult women). By body mass, differences between modern pygmies and Homo floresiensis are even greater. LB1's body mass has been estimated at 25 kg (55 lb). This is smaller than that of not only modern H. sapiens, but also H. erectus, which Brown and colleagues have suggested is the immediate ancestor of H. floresiensis. LB1 and LB8 are also somewhat smaller than the australopithecines from three million years ago, not previously thought to have expanded beyond Africa. Thus, LB1 and LB8 may be the shortest and smallest members of the extended human family discovered thus far. Aside from smaller body size, the specimens seem otherwise to resemble H. erectus, a species known to have been living in Southeast Asia at times coincident with earlier finds purported to be of H. floresiensis. These observed similarities form the basis for the suggested phylogenetic relationship. Controversially, the same team has reported finding material evidence (stone tools) on Flores of a H. erectus occupation dating back 840,000 years ago, but not remains of H. erectus itself or transitional forms. To explain the small stature of H. floresiensis, Brown et al. have suggested that in the limited food environment on Flores, H. erectus evolved a smaller body size via insular dwarfism,[1] a form of speciation which has been observed in other species on Flores also including several species of the proboscidean genus Stegodon. (A dwarf stegodont species of Flores, Stegodon sondaari, went extinct by about 850,000 years ago and was replaced by another species of normal size, Stegodon florensis, which then also evolved into a dwarf form, Stegodon florensis insularis, which disappeared about 12,000 years ago.) This hypothesis has been criticized by Teuku Jacob and colleagues who argue that LB1 is similar to the midget humans who populate a Flores village, Rampasasa, and who point out that size can vary substantially in pygmy populations. Contradictory evidence has emerged. Small brains

In addition to a small body size, H. floresiensis had a remarkably small brain. The brain of the holotype LB1 is estimated to have had a volume of 380 cm3 (23 cu in), placing it at the lower range of chimpanzees or the extinct australopithecines. LB1's brain size is half that of its presumed immediate ancestor, H. erectus (980 cm3 (60 cu in)). The brain to body mass ratio of LB1 lies between that of H. erectus and the great apes. Insular dwarfism has been posited to explain the brain size reduction. Scientists at the Natural History Museum in London have found that the reduction in brain size of extinct pygmy hippopotamuses in Madagascar compared with their living relatives is greater than the reduction in body size, and similar to the reduction in brain size of H. floresiensis compared with H. erectus. An indicator of intelligence is the size of Brodmann's area 10, the dorsomedial prefrontal cortex, an area of the brain associated with higher cognition. LB1's region 10 is about the same size as that of modern humans, despite the much smaller overall size of the brain. Notwithstanding the small brain of H. floresiensis, the discoverers have associated it with advanced behaviors. Their cave shows evidence of the use of fire for cooking, and Stegodon bones associated with the hominins have cut marks. The hominin

specimens have also been associated with stone tools of the sophisticated Upper Paleolithic tradition typically associated with modern humans, who have nearly quadruple the brain volume (1,3101,475 cm3 (8090.0 cu in)) and 2.6 times greater body mass. Some of these tools were apparently used in the necessarily cooperative hunting of Stegodon by these hominins.[2] This is an interesting paradox: there has been a gradual increase in brain volume as we progressed along the Human timeline of evolution (see Homininae), starting from about 600 cm3 (37 cu in) in Homo habilis up to 1,500 cm3 (92 cu in) in Homo sapiens neanderthalensis. Thus, in general, there is a correlation between brain volume and intelligence. However, modern Homo sapiens have a brain volume slightly smaller (1,400 cm3 (85 cu in)) than neanderthals, women have a brain volume slightly smaller than men and Homo floresiensis with a cranial capacity of about 380 cm3 (23 cu in), considered small for a chimpanzee) and about a third that of H. erectus, apparently used fire and made tools at least as sophisticated as those of their ancestor H. erectus. In this case, it seems that for intelligence, the structure of the brain is more important than its volume. Additional features Additional features used to argue that the finds come from a population of previously unidentified hominids include the absence of a chin, the relatively low twist of the arm bones, and the thickness of the leg bones. The presence of each of these features has been confirmed by independent investigators but their significance has been disputed. The forearm and pectoral girdle of H. floresiensis have been examined by Larson et al. (2007). Modern humans have the top of the bone twisted between 145 to 165 degrees to the plane of the elbow joint. For LB1, the twist was initially reported to be 110 degrees. Larson later revised this measurement to 120 degrees. This could be an advantage when arm-swinging, but it complicates activities associated with modern people, such as tool-making. As for the pectoral girdle of H. floresiensis, they studied a broken clavicle of LB1 and a shoulder blade of LB6. The clavicle was relatively short, which in combination with the shape of the shoulder blade and the low twist of the arm bone resulted in the shoulder being moved slightly forward, as if it was shrugged. Thus H. floresiensis could bend the elbow in the way modern people do and Larson concluded that it was able to make tools. Tocheri et al. (2007) examined three carpal bones believed to belong to LB1. The shapes of these bones were claimed to differ significantly from the bones of the modern human wrist and to resemble the wrist of great African apes or Australopithecus. The feet of H. floresiensis were unusually flat and unusually long in relation with the rest of the body. As a result, when walking, it would have to bend its knees further back than modern people do. This forced the gait to be high stepped and the creature was not able to walk very fast. The toes had an unusual shape and the big toe was very short.

Recent survival
The species is thought to have survived on Flores at least until 12,000 years before present, making it the longest lasting nonmodern human, surviving long past the Neanderthals (H. neanderthalensis), which became extinct about 24,000 years ago. Because of a deep neighboring strait, Flores remained isolated during the Wisconsin glaciation (the most recent glacial period), despite the low sea levels that united Sundaland. This has led the discoverers of H. floresiensis to conclude that the species, or its ancestors, could only have reached the isolated island by water transport, perhaps arriving in bamboo rafts around 100,000 years ago (or, if they are H. erectus, then about 1 million years ago). This idea of H. floresiensis using advanced technology and cooperation on a modern human level has prompted the discoverers to hypothesize that H. floresiensis almost certainly had language.[39] Local geology suggests that a volcanic eruption on Flores approximately 12,000 years ago was responsible for the demise of H. floresiensis, along with other local fauna, including the elephant Stegodon. Gregory Forth hypothesized that H. floresiensis may have survived longer in other parts of Flores to become the source of the Ebu Gogo stories told among the Nage people of Flores. The Ebu Gogo are said to have been small, hairy, language-poor cave dwellers on the scale of this species. Believed to be present at the time of the arrival of the first Portuguese ships during the 16th century, these creatures are claimed to have existed as recently as the late 19th century. Gerd van den Bergh, a paleontologist working with the fossils, reported hearing of the Ebu Gogo a decade before the fossil discovery. On the island of Sumatra, there are reports of a 11.5 m (3 ft 3 in4 ft 10 in) tall humanoid, the Orang Pendek which might be related to H. floresiensis. Henry Gee, senior

editor at Nature magazine, speculates that species like H. floresiensis might still exist in the unexplored tropical forest of Indonesia.

Scandal over specimen damage

In early December 2004, Teuku Jacob removed most of the remains from their repository, Jakarta's National Research Centre of Archaeology, with the permission of only one of the project team's directors and kept them for three months. Some scientists expressed the fear that important scientific evidence would be sequestered by a small group of scientists who neither allowed access by other scientists nor published their own research. Jacob returned the remains on February 23, 2005 with portions severely damaged and missing two leg bones to the worldwide consternation of his peers. Reports noted the condition of the returned remains; "[including] long, deep cuts marking the lower edge of the Hobbit's jaw on both sides, said to be caused by a knife used to cut away the rubber mould"; "the chin of a second Hobbit jaw was snapped off and glued back together. Whoever was responsible misaligned the pieces and put them at an incorrect angle"; and, "The pelvis was smashed, destroying details that reveal body shape, gait and evolutionary history" and causing the discovery team leader Morwood to remark "It's sickening, Jacob was greedy and acted totally irresponsibly". Jacob, however, denied any wrongdoing. He stated that the damages occurred during transport from Yogyakarta back to Jakarta despite the physical evidence to the contrary that the jawbone had been broken while making a mold of bones. In 2005 Indonesian officials forbade access to the cave. Some news media, such as the BBC, expressed the opinion that the reason for the restriction was to protect Jacob, who was considered "Indonesia's king of palaeoanthropology", from being proven to be wrong. Scientists were allowed to return to the cave in 2007 shortly after the death of Jacob.

Microcephaly hypothesis
Prior to Jacob's removal of the fossils, a CT scan was taken of the skull and a virtual endocast of the skull (i.e., a computergenerated model of the skull's interior) of H. floresiensis was produced and analyzed by Dean Falk et al. This team concluded that the brainpan was not that of a pygmy nor an individual with a malformed skull and brain. In response, Weber et al. conducted a survey the same year comparing the computer model of LB1's skull with a sample of microcephalic human skulls, concluding that the skull size of LB1 falls in the middle of the size range of the human samples and is not inconsistent with microcephaly. Next to dispute the finding of Falk et al. (2005) were Martin et al. (2006), who objected to the failure to compare the model of LB1's skull with a typical example of adult microcephaly. Martin and his coauthors concluded that the skull was probably microcephalic, arguing that the brain is far too small to be a separate dwarf species; if it were, the 400-cubic-centimeter brain would indicate a creature only one foot in height, one-third the size of the discovered skeleton. Shortly thereafter, a group of scientists from Indonesia, Australia, and the United States came to the same conclusion by examining bone and skull structure (Jacob (2006)). Brown and Morwood countered by claiming that the skeptics had drawn incorrect conclusions about bone and skull structure and mistakenly attributed the height of H. floresiensis to microcephaly. Falk's team replied to the critics of their study (Falk et al. (2006)). Morphologist Jungers examined the skull and concluded that the skeleton displays "no trace of disease". Argue, Donlon, et al. (2006) rejects microcephaly and concludes that the finds are indeed a new species. Falk et al. (2007) offered further evidence that the claims of a microcephalic H. sapiens were not credible. Virtual endocasts of an additional nine microcephalic brains and ten normal human brains were examined, and it was found that the floresiensis skulls are similar in shape to normal human brains, yet have unique features which are consistent with what one would expect in a new species. The frontal and temporal lobes of the floresiensis brain were found to be highly developed, in strong contrast to the microcephalic brain, and advanced in ways different from modern human brains. This finding also answered past criticisms that the floresiensis brain was simply too small to be capable of the intelligence required for the members of H. floresiensis to create the tools found in their proximity. Falk et al. (2007) conclude that the onus is now upon the critics that continue to claim microcephaly to produce a brain of a microcephalic that bears resemblance to the floresiensis brain. Falk's argument was supported by Lyras et al. (2008) in that 3D-morphometric features of the skulls of microcephalic H. sapiens indeed fall within the range of normal H. sapiens and that the LB1 skull falls well outside this range. This was interpreted as proving that LB1 cannot, on the basis of either brain or skull morphology, be classified as a microcephalic H. sapiens.

In 2009, a study by Jungers et al. presented a statistical analysis of skull shapes of healthy modern humans, microcephalic humans, and several ancient human species, as well as H. floresiensis. They showed that the three grouped separately, with H. floresiensis among the ancient humans, providing evidence that H. floresiensis is a separate species instead of a diseased modern human.

Laron syndrome hypothesis

The anatomist Gary D. Richards introduced a new skeptical hypothesis in June 2006: that the skeletons from Flores might be the remains of people who suffered from Laron syndrome, a genetic disorder first reported in 1966. The next year, a team including Laron himself published a paper arguing that the morphological features of H. floresiensis are essentially indistinguishable from those of Laron syndrome. The team said that to determine whether the H. floresiensis individuals had Laron syndrome would require testing their DNA for the presence of the defective genes, if samples of that DNA ever become available. Critics of the hypothesis have however pointed out that despite the low stature, people suffering from Laron syndrome look nothing like the Homo floresiensis remains, particularly in the anatomy of the cranial vault.[60]

Endemic cretinism hypothesis

In 2008 Australian researchers Peter J. Obendorf, Charles E. Oxnard, and Ben J. Kefford suggested that LB1 and LB6 suffered from myxoedematous (ME) endemic cretinism resulting from congenital hypothyroidism and that they were part of an affected population of H. sapiens on the island. This disease, caused by various environmental factors including iodine deficiency, is a form of dwarfism which can still be found among the local Indonesian population. Affected people, who were born without a functioning thyroid, have both small bodies and reduced brain size but their mental retardation and motor disability is not as severe as with neurological endemic cretins. According to the authors of the study, the critical environment could have been present on Flores approximately 18,000 years ago, the period to which the LB fossils are dated. They wrote that various features found on the fossils, such as enlarged pituitary fossa, unusually straight and untwisted tops of the upper arm bone and relatively thick limbs, are signs of this diagnosis. The double rooted lower premolar and primitive wrist morphology can be explained in this way as well. The oral stories about strange human-like creatures may also be a record of cretinism. Falk challenged the premise of Oberndorf et al. Studying computer tomography scans of LB1's pituitary fossa, she came to the conclusion that it is not larger than usual. Peter Brown declared that the remains of the pituitary fossa were very poorly preserved and no meaningful measurement was possible. In a paper delivered to the Australasian Society for Human Biology in 2009, Colin Groves and Catharine FitzGerald compared the Flores bones with those of ten people who had had cretinism, focusing on anatomical features which are typical of the disease. They found no overlap, and stated that they had put the claim to rest. However, an article by Oxnard, Obendorf and Kefford rejects Groves and FitzGerald's argument and revives the cretinism hypothesis. Oxnard and colleagues also criticise the cladistic analysis of Argue et al. (2009), stating that it is not logically possible for the analysis to conclude that the Liang Bua remains represent a separate species and not a pathology because the cladistics analysis assumes that they do not represent a pathology. Brown (2012) compared skeletal and dental morphology in H. floresiensis with the clinical and osteological indicators of cretinism, and the traits that have been argued to be associated with ME cretinism in LB1 and LB6. He concludes that LB1 and LB6 Homo floresiensis are not modern human (Homo sapiens) cretins.

Bone structure
The bone structure of H. floresiensis shoulders, arms and wrists have been described as very different from modern humans', much closer to the bone structure of chimpanzees or an early hominin. This adds support to the idea that H. floresiensis is a separate species of early human rather than a modern human with a physical disorder. Susan G. Larson et al. analyzed the upper limb of LB1. They found that in LB1 the angle of humeral torsion is much less than in modern humans. This had been previously studied by Richards et al., who declared that it is a sign of modern pygmy

populations, and T. Jacob et al., who pointed out that muscle attachments on the bone suggest LB1 had weak muscles which resulted in little development of humeral torsion. Larson et al. rejected Richards conclusion, arguing that the humeral torsion of pygmy populations is usually similar to that of peoples of average stature. They argued that Richards et al. cited a 1972 paper which had studied a sample of six female Eastern Central African pygmies and this sample was too small to represent the whole population. Larson et al. also failed to find signs of microcephaly on the studied bones. Larson et al. also studied the relatively short clavicle and the unusual formation of the pectoral girdle. They compared their finding with the skeleton of Nariokotome Boy (variously classified as H. ergaster or H. erectus), and suggested that the pectoral girdle of H. floresiensis was a transitional stage in human shoulder evolution. While some specialists, including paleoanthropologist Russell Ciochon of the University of Iowa, supported the conclusion, others, including Eric Delson of Lehman College, City University of New York, pointed out that the recent sample of H. floresiensis individuals is too small and that Larson's research was based just on one shoulder bone. Tocheri et al. (2007) (including Morwood, Larson, and Jungers), compared three carpal bones believed to belong to LB1 with carpal bones of modern humans, some earlier hominids and African apes. They concluded that the carpals from the Liang Bua cave resembled ape carpal bones and were significantly different from the bones of H. sapiens, Homo neanderthalensis or even Homo antecessor, and that they were comparable to carpal bones of Australopithecus. The carpal bones of H. floresiensis were found to lack features that evolved with ancestors of modern humans at least about 800,000 years ago. These features are already formed during embryogenesis and therefore Tocheri et al. argue that it is improbable that the shape of H. floresiensis wrist bones could be a result of a developmental disease. This conclusion was challenged by Robert Martin, since Jacob's death the leading proponent of the microcephaly hypothesis, and Alan Thorne. Martin noted that no research has been done on wrists of microcephalic people. Thorne maintained that the differences were small and that similar variation could occur with living modern humans. He also pointed out that the carpal bones had been found scattered in the cave and it was not certain that they all belonged to the same individual. Project leader Morwood countered that there were also other features, such as the stature, body proportions, brain size, shoulder, pelvis, jaw, and teeth which suggested that H. floresiensis is a separate species that evolved in isolation on the island.

DNA extraction
In around 2006, two teams attempted to extract DNA from a tooth discovered in 2003 - both teams were unsuccessful. It has been suggested that this happened because the dentine was targeted, whereas new research suggests that the cementum has higher concentrations of DNA. Also, the heat generated by the high speed of the drill bit may have denatured the DNA.