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Brief communication Evidence bearing on the status of Homo habilis at Olduvai Gorge.

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AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 137:356–361 (2008)
Brief Communication: Evidence Bearing on the Status
of Homo habilis at Olduvai Gorge
Randall L. Susman*
Department of Anatomical Sciences, School of Medicine, Stony Brook University, Stony Brook, NY
KEY WORDS
early hominin postcrania; Homo habilis; Olduvai hominid 7; OH 8; OH 35
ABSTRACT
Students of the early hominin career
have debated the status of Homo habilis since its discovery in 1960. Today discussion centers on which specimens should be included in the species and what constitutes the holotype. Recent reviews of early Homo suggest
that the Olduvai Hominid 8 foot may sample Paranthropus while the OH 7 skull bones, mandible, and hand
sample H. habilis. Moreover, some suggest that while H.
habilis in Middle Bed I at Olduvai is craniodentally
Homo-like, the postcranial skeleton of H. habilis is more
like that of Australopithecus. Evidence presented here
indicates not only that OH 7 and OH 8 represent H.
habilis but also that they come from a single individual.
The association of OH 35 with OH 7 and OH 8 is less
certain. Morphological, pathological, and taphonomic evidence favors the inclusion of OH 35 in the holotype.
However, stratigraphic evidence suggests that OH 35
and OH 8 are not coterminous. With or without OH 35,
the holotype of H. habilis ranks as one of the most complete early hominin skeletons and the most complete and
functionally informative specimen of early Homo. Am J
Phys Anthropol 137:356–361, 2008. V 2008 Wiley-Liss, Inc.
Fossil evidence of Homo habilis was recovered from
Olduvai in 1960–1961 (Leakey, 1960, 1961a,b). When first
reported, a hand, foot, and clavicle, from site FLK NN
Level 3, and a leg from FLK ‘‘Zinj’’ (Level 22) represented
the first early hominid postcrania recovered in East
Africa. Together with a mandible and fragmentary skull
the fossils from FLK NN Level 3 were reported to be
those of a single juvenile whose age at death was 12
years (Leakey, 1961b).1 After the initial reports, Day and
Napier (1964) concluded that while the jaw, skull, and
hand were those of a ‘‘child,’’ the foot represented an
adult and thus signaled the presence at FLK NN 3 of a
second individual. Day and Napier’s conclusion was based
on the presence of what they considered to be ‘‘agerelated’’ arthritis in the midfoot and metatarsus (M.H.
Day, personal communication). The mandible, skull
bones, and hand, were designated OH 7. These fossils
became the holotype of the new species, Homo habilis
(Leakey et al., 1964). The foot was catalogued as OH 8,
and placed in the paratype along with other craniodental
and postcranial fossils by Leakey et al. (1964). In their diagnosis and description, Leakey, Tobias and Napier maintained that the holotype, OH 7, was a subadult while
other postcranial remains, including OH 8, OH 35 (a leg),
and OH 48 (a clavicle), were the bones of adults.
In addition to arthritis, Day also observed that the
OH 8 foot lacked evidence of unfused secondary growth
centers (epiphyses) such as he and Napier noted in the
OH 7 hand (Day, personal communication). According to
Day, unlike the OH 7 hand, the developmental keys in
the OH 8 foot, viz. the state of ossification of the metatarsals, did not indicate an age like that of the mandible
and hand of OH 7.
Other workers assented to Day and Napier’s ‘‘age’’
estimate for OH 8. As for arthritis, Oxnard and Lisowski
(1980) agreed that the unusually flattened transverse
arch of OH 8 was the possible result of age-related pathology. Some went even further and suggested that OH
8 not only did not belong with the youthful OH 7 ‘‘child,’’
but also that it did not even sample H. habilis. The latter group, represented by Day and Wood (1968 et seq.)
argued that OH 8 was morphologically closer to the second early hominin in Bed I at Olduvai, Australopithecus
boisei. In a multivariate analysis of the talus Day and
Wood identified features of OH 8 that were shared with
the Australopithecus (Paranthropus) robustus talus, TM
1517, from Kromdraai. Wood later extended his comparative set to include the derived KNM-ER 813 talus
assigned to Homo. Wood argued that OH 8 was on one
hand similar to A. robustus and on the other was unlike
the more derived, Homo-like, KNM-ER 813. This finding
provided support for the hypothesis that OH 8 belonged
in Australopithecus instead of Homo (Wood, 1974 et seq.)
This idea persists at the present time (e.g., Gebo and
Schwartz, 2006; Wood and Constantino, 2007a,b).
The presence of early Homo in Bed I and the taxonomic validity of H. habilis (i.e. whether it merited separate status from H. erectus and A. africanus) were hotly
1
Louis and Mary Leakey applied a modern human schedule to
determine the subadult status of the Olduvai hominid. The use of a
human standard was based on the perceived hominid (hominin)
morphology of the dentition, hand, and foot. It is important to stress
that it is the relative age rather than the absolute age, of the Olduvai fossils from FLK NN Level 3 that is important. An ape developmental schedule will yield a younger absolute age for the OH 7 and
OH 8 but, as Smith has shown (1993), humans and (laboratory)
chimpanzees share the same approximate sequence of epiphyseal
fusion. Again, for determining the association of fossils remains at
FLK NN Level 3, relative age is the important variable.
C 2008
V
WILEY-LISS, INC.
C
*Correspondence to: Randall L. Susman, Department of Anatomical
Sciences, School of Medicine, Stony Brook University, Stony Brook,
NY 11794-8081, USA. E-mail: randall.susman@stonybrook.edu
Received 19 September 2007; accepted 4 June 2008
DOI 10.1002/ajpa.20896
Published online 25 July 2008 in Wiley InterScience
(www.interscience.wiley.com).
HOMO HABILIS POSTCRANIA BED I, OLDUVAI GORGE
debated in the years following its description. Despite
the different assessments of Leakey (1960) and Day and
Napier (1964) early on, questions regarding the relative
ontogenetic ages of the type and paratype received little
subsequent attention. Today, while the presence of early
Homo in Bed I is no longer at issue, debate continues on
the question of which fossils should be included in H.
habilis and, specifically, whether H. habilis as presently
constituted is an overly inclusive taxon containing more
than one species of early Homo (Wood, 1992; Wood and
Collard, 1999). Wood (1992) has suggested that postcranial fossils of H. habilis, including the holotype, more
closely resemble Australopithecus and Paranthropus
than Homo. The issue of what fossils should be included
in H. habilis has been argued by others as well
(e.g.,Walker and Leakey, 1978; Chamberlain and Wood,
1987; Stringer, 1989).
QUESTIONS
This article addresses two questions that bear on the
interpretation of H. habilis. The first is an examination
of the relative ontogenetic age of the Olduvai postcrania.
This issue was addressed before by Susman and Stern
(1982) but a survey of the current literature regarding
early Homo reveals a difference of opinion on this subject. We maintain that the key hominin remains at FLK
NN 3 are all of the same relative age and that they
belong to one, subadult, individual. This statement is
significant due to the importance of the FLK NN 3
remains for understanding the holotype of H. habilis
and, in turn, the taxonomy of early Homo. The foregoing
leads to the second question, viz. how we view and interpret the functional morphology of H. habilis and early
Homo. A proper understanding of the holotype is a necessary precursor to understanding adaptive differences
between H. habilis, Australopithecus and later members
of the genus Homo. At the very least, before we argue
that the postcranial skeleton of H. habilis is australopithecine-like (Wood, 1992, p 790), we need to understand
which fossils belong to H. habilis and which belong to
contemporary australopithecines.
MATERIALS
To address the above, measurements, plain X-rays,
and stereophotography were carried out on the original
fossils. A Wild dissecting scope was used for observation
of surface features and fine details of anatomy. The
above took place on three occasions. The first was at the
National Museums of Kenya in 1976 at which time I
studied the OH 7 hand and other hand fossils from East
Africa. The second was in 1982 at which time I studied
OH 8 and OH 35 at the KNM. The third study of the
Olduvai hominins was carried out at the National Museum of Tanzania in Dar es Salaam and at the KNM, in
2008.
Comparative skeletal materials of human and nonhuman primates were studied at the American Museum
of Natural History, the Cleveland Museum of Natural
History (CMNH), the U.S. National Museum of Natural
History, and the Department of Anatomical Sciences at
Stony Brook. Human skeletons from the Dart Collection
at the University of the Witwatersrand and the CMNH
were also studied. Radiographs of modern humans come
from the Departments of Radiology and Emergency Medicine at Stony Brook University Medical Center.
357
RESULTS
Age of the OH 8 foot
Olduvai Hominid 8 consists of a virtually complete set
of tarsals and metatarsals. The tarsus is complete save
for the posterior calcaneus and posterior extremity of the
talus. OH 8 represents the most complete and best preserved foot of any early hominin.
Susman and Stern (1982) offered evidence that OH 8
was the foot of a subadult, as originally suggested by
Leakey (1960, 1961a,b). Leakey was never explicit about
his reasoning regarding the age of OH 8, but evidence
he presented included the close proximity of the subadult OH 7 mandible, hand bones and the OH 8 foot at
FLK NN Level 3 (see also Leakey, 1971, Fig. 20). Susman and Stern (1982) offered morphological evidence for
the age of OH 8 in the form of mediolateral plain radiographs of the OH 8 metatarsals showing that while the
distal ends of metatarsals 1, 4, and 5 were broken, metatarsals 2 and 3 were intact save for their epiphyses. Figure 1, here, illustrates that the distal ends of metatarsals 2 and 3 are missing their epiphyses.
As for the subadult status of OH 8, the presence of a
fused basal epiphysis on metatarsal I together with
unfused distal epiphyses on metatarsals 2 and 3, indicates an age identical to that indicated by the mandible
and hand (9.0–12.0 years depending on whether the
comparison is based on a female or a male schedule
Hoerr et al., 1962).
Figure 2 is a dorsoplantar radiograph of the foot
including the metatarsus of a modern human female of
12 years (a) and the OH 8 metatarsals (b). In this view
it is again apparent that the ends of metatarsals 2 and 3
are intact save for their heads, and that they resemble
the metaphyses seen in subadult humans (Fig. 2a).
From radiographic evidence, the relative age of OH 8
comports with that of OH 7, including both the mandibular dentition (Tobias, 1971, 1991) and the hand (Day,
1976; Susman and Creel, 1979). The radiological evidence indicates that OH 7 and OH 8 are the same relative age. Parsimony dictates that OH 7 and OH 8 represent a single individual. Ancillary evidence of association
comes from the close proximity of hand bones, foot
bones, parietals, and mandible on the FLK NN Level 3
‘‘floor’’ (Leakey, 1971, Fig. 20).
Evidence indicates that Louis Leakey was correct in
his initial assessment of the FLK NN Level 3 postcrania
(Leakey, 1961a,b) and the holotype of H. habilis. The alternative to the above is that OH 7 and OH 8 are not
from the same individual, but instead represent the
improbable case of two different individuals of the same
age who succumbed, in proximity, at the same time, and
left complementary body parts, at FLK NN Level 3.
Olduvai Hominid 8 foot and OH 35 leg
The Olduvai tibia and fibula were recovered from the
‘‘Zinjanthropus’’ floor, FLK Level 22, in 1960. Initially,
Louis and Mary Leakey assigned the leg to ‘‘Zinjanthropus’’ (Leakey, 1961b, p 348). Davis (1964), in his preliminary description, refrained from assigning the tibia
and fibula to either Paranthropus (Zinjanthropus) boisei
or to H. habilis. Subsequently, some have assigned the
leg to Paranthropus (Day, 1976; Tobias, 1991) while
others placed it in Homo habilis (Leakey et al., 1964;
Leakey, 1971; Day, 1978; Susman and Stern, 1982;
Susman, 1983). The conundrum of an essentially
American Journal of Physical Anthropology
358
R.L. SUSMAN
Fig. 1. Lateral plain radiograph of the OH 8 metatarsals showing the unfused, missing heads (5 epiphyses) of metatarsals 2
and 3 and the broken and missing ends of metatarsals 1, 4, and 5. The opacity on the base of metatarsal V is an artifact resulting
from a small piece of plasticine used to orient the fossil.
Fig. 2. Dorsoplantar plain radiographs of a 12-year-old modern human female (a) and H. habilis (OH 8) (b). Damage to the distal ends of metatarsals 1, 4, and 5 is due breakage at the time of death. Note the similar chevron-shaped appearance of the distal
ends of metatarsals 2 and 3 in both the human and in OH 8. Unlike the lateral toes, the epiphysis of the big toe is located at its
base. Here the base is fused in both (a) and (b). In the human foot (a) fusion is not complete. Nonetheless, the big toe has attained
its adult length at this point in both (a) and (b). The combination of a fused epiphysis of the big toe together with unfused epiphyses
on toes 2 and 3 yields a skeletal age of 9–12 years for the OH 8 foot if we apply a modern human standard (Hoerr et al., 1962).
human-like leg in proximity to the skull of Paranthropus
boisei, OH 5, on the FLK ‘‘Zinj’’ floor is the reason that
OH 35 has remained in taxonomic limbo for the past 47
years.
American Journal of Physical Anthropology
Morphological evidence suggests that the OH 35 leg
belongs to holotype, OH 7/8. This evidence includes 1)
carnivore bites marks on both the OH 8 talus and on the
distal end of the OH 35 tibia (Figs. 3 and 4), 2) abutting
HOMO HABILIS POSTCRANIA BED I, OLDUVAI GORGE
Fig. 3. Dorsal view, OH 8 foot. The heterotopic ossification
of the lateral intermetatarsal joints 3 and 4 is observed in the
form of lipping osteophytes surrounding the joints (arrows).
Note the broken ends of the metatarsals 1, 4, and 5 missing epiphyses of toes 2 and 3. The ossification observed in toes 1 and 2
(together with a fused basal epiphysis on the hallux) signals a
developmental age for the OH 8 foot like that of the OH 7 mandible and hand.
surfaces of postmortem or perimortem, damage on the surfaces of the lateral talus and the fibular malleolus and, 3)
the derived, human-like morphology of both the foot (Day
and Napier, 1964; Archibald et al., 1972; Susman and
Stern, 1982) and the leg (Davis, 1964; Lovejoy, 1978).
Other evidence argues against the association of OH
35 and OH 7/8. Studies by Aiello et al. (1998) and Wood
et al. (1998) on the reciprocal talar and tibial surfaces
indicate that the contour of the OH 8 talus does not
match the inferior surface of OH 35 tibia. Aiello et al.
state that not only are OH 35 and OH 8 not from the
same individual, but they may not even come from the
same species. However, the findings reported by Aiello
359
et al. on the subject of the association of OH 35 and OH 8
are mixed. While some data falsify the hypothesis of
association, others do not. The authors state that OH 8
and OH 35 do not come from the same individual (1998,
p 70), but they also curiously note that ‘‘. . . the OH 35/8
pairing shows a degree of congruency that would be comparable with associated modern human (ankle) joints,’’
(Aiello et al., 1998, p 68). Some of their data are size
corrected and some are not. Not surprisingly the fit is
better when size corrected data are analyzed. The findings of Aiello et al. (1998) and Wood et al. (1998) do not
account for the natural joint space created by the chondral surfaces in the living ankle or the range of ‘‘fit’’
observed from the close-packed to the loose-packed position of the talocrural joint in both humans and apes.
Finally, the above studies of the talus and tibia did not
include the fibula, a not insignificant participant in the
talocrural joint.
The geological evidence from FLK NN Level 3 and
FLK ‘‘Zinj’’ (Level 22) is also mixed regarding the hypothesis of an association of OH 8 and 35. While lithologically the two sites bear a close resemblance to one
another, stratigraphy indicates that FLK NN Level 3
and FLK Level 22 sample different times. With regard
to the lithology of the two sites, Leakey (1971) notes that
OH 35 was recovered from a clay paleosol at FLK ‘‘Zinj’’
and that the H. habilis floor at FLK NN, where OH 8
was recovered, is a similar surface. Leakey states, ‘‘(FLK
‘‘Zinj’’) consisted of a grey-green silty clay, 1 ft. thick.
The upper surface of the clay was slightly uneven, with
the top few inches noticeably more friable than the lower
part, suggesting that it had been weathered into a paleosol similar to the clay of Levels 1 and 3 at FLK NN.’’
Leakey notes further that ‘‘Both the nature of the clay
and the mode of occurrence of the remains on the occupation floor (at FLK ‘‘Zinj’’) bear a close resemblance to
the condition pertaining to Level 3 at FLK NN,’’ (Leakey,
1971:49). As for the artifacts and bones on the FLK
‘‘Zinj’’ floor, Leakey states that ‘‘. . . the finds were dispersed at random and it is not possible to discern any
pattern in the distribution, although the scatter resembles that at FLK NN Level 3,’’ (Leakey, 1971, p 50).
Notwithstanding the above, the strongest evidence
against the association of OH 8 and OH 35 comes from
stratigraphy. Hay, 1976; personal communication) and
Njau (personal communication) observe that FLK NN
Level 3 and FLK ‘‘Zinj’’ do not sample the same time period. A recently completed study indicates that the two
surfaces at FLK NN and FLK may differ in age by
6,000 years (Njau, personal communication).
The evidence for the association of OH 35 and OH 8 is
thus equivocal. The leg and foot are a good match and
those who have studied the leg agree that it is derived
and human-like in proportion to the foot (Susman,
1983), in its morphology (Davis, 1964; Lovejoy, 1975;
Susman and Stern, 1982). Taphonomic evidence of carnivore damage to leg and foot may also be suggestive of
association. Finally, evidence of the depositional environment indicates similarities between FLK ‘‘Zinj’’ and FLK
NN Level 3, while the stratigraphy argues strongly
against the bones being coterminous.
Pathology: OH 8 and OH 35
Similar pathology is present in both OH 8 and OH 35.
While Day and Napier determined that bony changes in
American Journal of Physical Anthropology
360
R.L. SUSMAN
Fig. 4. Anterior oblique view of the ankle joint: (a), OH 35 leg with the OH 8 talus, (b) modern human with ossification of the
tibiofibular syndesmosis following injury and, (c) a healthy modern human ankle. The heterotopic ossification of the distal tibia and
fibula in both (a) and (b) is evidence of a ‘‘high ankle’’ sprain (Marder and Lian, 1997). Also seen in OH 35 and OH 8 are bite marks
left by at least two different predators, likely crocodile and leopard (Njau and Blumenschine, 2007). Scale bar in (c) 5 3 cm.
the OH 8 foot signaled the presence of ‘‘age-related’’ arthritis, the lesions on the tarsus and metatarsus are
more likely indicative of trauma not age-related pathology. Although osteoarthritic changes can be seen in both
old and young individuals, the osteophytes in OH 8 and
in OH 35 present as heterotopic ossification (HTO) in
both the midfoot of OH 8 and in the tibiofibular syndesmosis of OH 35.
Figures 3 and 4 reveal heterotopic ossification that
commonly follows trauma. The heterotopic changes in
OH 8 are manifest as osteophytes invading the adjoining
soft tissues and capsules at the margins rather than on
the joint surfaces (see Fig. 3). These changes are not specifically age-related, but may occur at any age, following
injury. Lipping osteophytes, a sign of HTO (Gamble and
Yale, 1975), are particularly evident on the bases of
metatarsals III, IV, and V of OH 8 (see Fig. 3).
In addition to the osteophytes seen in the foot, the distal tibiofibular syndesmosis of OH 35 reveals HTO of the
anterior inferior tibiofibular ligament and inferior end of
the interosseous membrane (Marder and Lian, 1997;
Fig. 4). Injuries similar to the one indicated in OH 35 are
known as ‘‘syndesmosis injuries’’ or ‘‘high ankle sprains.’’
CONCLUSIONS
The OH 8 foot and the OH 7 mandible and hand (and
likely also the parietals) all represent an individual of
the same age. It is much more likely that OH 7 and OH
8 represent the same subadult individual than they do
twin adolescents who died at the same time at FLK NN
Level 3 and left complementary body parts. Even more
unlikely are the persistent claims that OH 8 represents
Paranthropus boisei. The significance of determining the
association of the Olduvai hominid remains is more than
just accounting. The subadult individual representing
the revised holotype offers a perspective on Homo habilis
that clarifies its membership in the genus Homo. The
OH 7/8 holotype is relatively well-represented as African
Plio-Pleistocene hominins go. Functionally, OH 7/8 indicates a hominin with a hand and a foot that share key
derived features with those of modern humans. OH 8
has a suite of derived features including: 1) an
American Journal of Physical Anthropology
‘‘elongate’’ distal tarsal row that provides for a proportionately longer midfoot, 2) an adducted hallux, 3) longitudinal and transverse arches and, 4) and an inferior navicular angle that suggests a human-like inferior plantar
calcaneonavicular (‘‘spring’’) ligament (Day and Napier,
1964; Archibald et al., 1972; Susman and Stern, 1982;
Susman, 1983). The prevailing view of the functional
morphology of the Olduvai hand, OH 7, suggests it had
derived adaptations for precision grasping seen in the
trapezium and distal phalanges, Napier, 1962; Susman
and Creel, 1979) together with primitive, apelike features of the fingers seen in the morphology of the proximal and middle phalanges (Napier, 1962; Susman and
Creel, 1979). The latter were related to power grasping
(Susman and Creel, 1979; Susman and Stern, 1979) and
climbing, although given the total morphological pattern
of OH 7/8 and the morphology of OH 35, H. habilis did
not climb like an African ape.
As for taxonomic affinities, the OH 7/8 individual indicates, as Tobias suggested, ‘‘. . . a total morphological pattern for H. habilis which is outside the range for the
Australopithecinae and closer to, if not within, the range
of the Homininae,’’ (Tobias, 1964, p 4). These results call
into question the proposal to remove H. habilis from the
genus Homo and align it with the australopithecines
(Wood and Collard, 1999). The hand bones of OH 7 indicate adaptations for climbing beyond that which we see
in larger bodied later Homo, and certainly in modern
humans, (see Susman and Creel, 1979; Susman and
Stern, 1979) but not ape-like climbing (Wood and Collard, 1999).
The arboreality practiced by some early Homo was
probably influenced by selective pressures of predator
avoidance, feeding, and sleeping in relatively small-bodied hominins with only an incipient capacity for tool
behavior. The OH 8 foot and the OH 35 leg, on the other
hand, show a resolute adaptation to human-like bipedality. The suggestion of ape-like, arboreal adaptations in
the OH 8 foot including among them an abductable hallux, (Lewis, 1980; Oxnard and Lisowski, 1980; Kidd et
al., 1996) is at odds with the morphology (and total morphological pattern) of OH 8. A cruropedal index, if the
leg and foot come from the same individual (see Fig. 6,
HOMO HABILIS POSTCRANIA BED I, OLDUVAI GORGE
Susman, 1983), is human-like indicating, at minimum, a
human-like swing phase during walking and running,
different from the pattern that would have attended a
more primitive, ape-like foot of A. afarensis (Stern and
Susman, 1983) or A. africanus (Clarke and Tobias,
1995). The inferred presence of longitudinal and transverse arches and a spring ligament in OH 7 are suggestive of a foot that was adapted to bipedal support. The
evidence of a ‘‘high ankle sprain,’’ seen in the tibiofibular
syndesmosis of OH 35 may also bear witness to humanlike positional behavior of H. habilis.
The combined evidence of OH 7, 8, and 35 does not
support the statement that ‘‘Whereas H. habilis sensu
stricto is hominine with respect to its masticatory complex, it retains an essentially australopithecine postcranial skeleton,’’ (Wood, 1992, p 790). Instead, the fossils
provide a cohesive picture of the craniodental and the
postcranial evidence of H. habilis at Olduvai Gorge.
Both the craniodental and the postcranial evidence distinguish H. habilis from the australopithecines and align
it with later members of the genus Homo.
ACKNOWLEDGMENTS
I thank the National Museums of Kenya and Dr.
Emma Mbua for facilitating this research and the
National Museum in Dar es Salaam for permission to
study the fossil hominids from Olduvai. Dr. Devorah Balsam and Dr. Solomon Spector (Department of Radiology,
Stony Brook University Medical Center) offered valuable
comments on the radiographs of the Olduvai fossils.
Jack Stern and Bill Jungers provided helpful comments
on the manuscript and Lucille Betti assembled Figures
1, 2, and 3 from original radiographs and photographs.
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