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Carlos Lorenzo & Juan Luis ArsuagaDepartamento dePaleontologıa, Instituto deGeologıa Economica, Facultad de Ciencias Geologicas,Universidad Complutense,
28040 Madrid, Spain.E-mail:carlets@eucmax.sim.ucm.es
Jose MiguelCarreteroDepartamento de CienciasHistoricas y Geografıa, Facultad de Humanidades yEducacion, Universidad deBurgos, 09001 Burgos, Spain
Received 10 November1998Revision received 26 March1999and accepted 5 June 1999
Keywords: Early Pleistocene,hand, foot, Homo antecessor ,Gran Dolina, Atapuerca.
Hand and foot remains from the GranDolina Early Pleistocene site (Sierra deAtapuerca, Spain)
We report here the study of the 22 hand and foot remains from theEarly Pleistocene level TD6 of the Gran Dolina site at Sierra deAtapuerca (Burgos, Spain) recovered from 1994 to 1996. Theseremains are paratypes of Homo antecessor . All of the elements arebriefly described and compared with other fossil hominids. Thecapitate has a constricted neck, well developed head, strong attach-ment for the ligamentum interosseum trapezoid-capitate, a palmarlyplaced trapezoid facet with a distinctive small dorsal trapezoid facet,a highly curved and oblique orientation of the second metacarpalfacet, and a transversally oriented dorsodistal border. A hamate witha moderately projecting and lightly built hamulus; an inferredreduced styloid process on the third metacarpal base; a wide secondmetacarpal head; and middle phalanges with well marked insertionsfor the flexor digitorum superficialis muscle and wide heads. Themorphology and dimensions of the pedal remains from TD6 are very
similar to modern humans; but the base, proximal articular surfaceand shafts of the proximal hallucal phalanges are more rounded andthe midshaft of the proximal toe phalanx is wider.
1999 Academic Press
Journal of Human Evolution (1999) 37, 501–522Article No. jhev.1999.0341Available online at http://www.idealibrary.com on
Introduction
From 1994 to 1996 just under 80 human
fossil remains were recovered from level
TD6 of the Early Pleistocene cave site of
Gran Dolina at the Sierra de Atapuerca
(Burgos, Spain) (Carbonell et al ., 1995;
Pares & Perez-Gonzalez, 1995; Bermudez
de Castro et al ., 1997).
An inventory of the 36 human remains
found in the 1994 field season was publishedby Carbonell et al . (1995). These remains
include six hand and eight foot remains,
none of which was described or figured in
Carbonell et al . (1995).
Bermudez de Castro et al . (1997) added
38 new specimens recovered in 1995 and
1996 from TD6, which include six hand
and two foot remains. Based on cranial,
mandibular, and dental traits these remains
have been ascribed to a new species of
Homo, Homo antecessor , that may represent
the last common ancestor of Neandertals
and modern humans (Bermudez de Castro
et al ., 1997). The 22 hand and foot remains
described here, as well as the rest of the
postcranial remains (Carretero et al ., 1999),
are paratypes of H. antecessor , but none of
the postcranial features are included in the
traits defining this species (Bermudez de
Castro et al ., 1997). Some of thesemanual and pedal remains bear cut marks,
that have been interpreted as evidence of
defleshing (Fernandez-Jalvo et al ., 1996,
1999).
We report here the first study of the 22
hand and foot remains from the Early
Pleistocene level TD6 of the Gran Dolina
site at Sierra de Atapuerca (Burgos, Spain)
recovered from 1994 to 1996.
0047–2484/99/090501+22$30.00/0 1999 Academic Press
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Material and methods
Inventory of TD6 hand and foot remains
A complete inventory of the H. antecessor
postcranial remains can be found in
Carretero et al . (1999). The 12 handremains from level TD6 of Gran Dolina
include one complete capitate bone, one
fragment of a hamate bone, a second meta-
carpal, a metacarpal distal epiphysis, plus
four proximal and four middle phalanges.
The ten foot remains include a second meta-
tarsal, a proximal fragment of a metatarsal
base, plus three proximal, three middle and
two distal phalanges (Figures 1 and 2).
ATD6-23 [Figure 1(a)]. Palmar fragment
of a left hamate with the hamulus well
preserved. It articulates perfectly with the
capitate ATD6-24 and undoubtedly belongs
to the same adult individual. The majority of
the dorsal surface and the metacarpal facets
are lost but the hamulus is well preserved.
ATD6-24 [Figure 1(b)]. Complete left
capitate. There is a minor abrasion in the
dorsal border of the distal articular surface.
It belongs to the same adult individual as the
hamate ATD6-23.
ATD6-25 [Figure 2(b)]. Fragment of a base
of a left second or third metatarsal.
ATD6-26 [Figure 1(i)]. Distal fragment of
an adult second metacarpal. Although this
fossil is very fragmentary, the asymmetry of
the head in distal view and the presence
of an indentation in the dorsoradial border
of the articular surface indicate that it
most probably represents a second left
metacarpal.
ATD6-27 [Figure 1(d)]. Diaphyseal frag-
ment of a proximal hand phalanx.
ATD6-28 [Figure 1(j)]. Second or third
middle hand phalanx from an adult
individual.
ATD6-29 [Figure 1(e)]. Distal fragment of a
proximal hand phalanx.
ATD6-30 [Figure 2(c)]. Complete left
proximal hallucal phalanx from an adultindividual.
ATD6-31 [Figure 2(d)]. Complete left
proximal hallucal phalanx. This phalanx
presents scars of the epiphyseal line.
ATD6-32 [Figure 2(e)]. Distal half of a
proximal pedal phalanx. This phalanx is notfrom the first or fifth ray. A more precise
identification is not possible.
ATD6-33 [Figure 2(f)]. Complete middle
pedal phalanx from an adult individual.
ATD6-34 [Figure 2(g)]. Complete middle
pedal phalanx from an adult individual.
ATD6-35 [Figure 2(h)]. Complete fourth or
fifth middle pedal phalanx from an adult.
ATD6-36 [Figure 2(i)]. Apical tuberosity
fragment of a distal pedal phalanx. ATD6-44 [Figure 1(l)]. Second or fifth
middle hand phalanx from an adolescent. It
lacks the proximal epiphysis and prob-
ably belongs to the same individual as
ATD6-53.
ATD6-46 [Figure 1(f)]. Distal fragment of
the third or fourth middle hand phalanx
from an adult. It presents a longitudinal
fracture along the diaphysis that results in
moderate distortion of the diaphyseal
breadth.
ATD6-53 [Figure 1(k)]. Third or fourth
middle hand phalanx from an adolescent
individual. It lacks the proximal epiphysis
and probably belongs to the same individual
as ATD6-44.
ATD6-59 [Figure 1(c)]. A distal fragment
of a left second metacarpal from an adult
individual lacking the proximal base.
ATD6-67 [Figure 1(g)]. Eroded diaphysis of
a proximal hand phalanx. ATD6-68 [Figure 2(j)]. Complete distal
pedal phalanx from an adult individual.
ATD6-70+ 107 [Figure 2(a)]. Left second
metatarsal from an adult. This bone is
composed of three fragments, the base and
the diaphysis (ATD6-107) and the distal
epiphysis (ATD6-70), that were found very
close to one another during excavation. The
base of this metatarsal is eroded.
ATD6-82 [Figure 1(h)]. Distal half of a leftsecond proximal hand phalanx.
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Figure 1. (a) ATD6-23, proximal, dorsal, ulnar, radial, palmar and distal views. (b) ATD6-24, proximal,dorsal, ulnar, radial, palmar and distal views. (c) ATD6-59, dorsal and radial views. (d) ATD6-27, dorsaland palmar views. (e) ATD6-29, dorsal and palmar views. (f) ATD6-46, dorsal and palmar views. (g)
ATD6-67, dorsal and lateral views. (h) ATD6-82, dorsal and palmar views. (i) ATD6-26, distal andlateral views. (j) ATD6-28, dorsal and palmar views. (k) ATD6-53, dorsal and palmar views. (l)ATD6-44, dorsal and palmar views. Scale bar=2 cm.
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Figure 2. (a) ATD6-70+107, dorsal and medial views. (b) ATD6-25, lateral and medial views. (c)ATD6-30, dorsal, medial and proximal views. (d) ATD6-31, dorsal, medial and proximal views. (e)ATD6-32, dorsal and palmar views. (f) ATD6-33, dorsal and palmar views. (g) ATD6-34, dorsal and
palmar views. (h) ATD6-35, dorsal and palmar views. (i) ATD6-36, dorsal and palmar views. (j)ATD6-68, dorsal and palmar views. Scale bar=2 cm.
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Comparison samples
We have studied the originals of the Krapina
collection (Croatian Natural History
Museum, Zagreb), La Ferrassie 1, La
Ferrassie 2, La Chapelle-aux-Saints (Museede L’Homme, Paris), Kebara, Amud 1 (Tel
Aviv University), Tabun C1 (British
Museum of Natural History, London) and
the casts of Regourdou 1 (Natural History
Museum). We have used some raw data
published for La Ferrassie 1 and 2 (Heim,
1982), Kebara 2 (Vandermeersch, 1991),
Shanidar 3, 4, 5, 6 and 8 (Trinkaus, 1983a),
Tabun C1 (McCown & Keith, 1939),
Amud 1 (Endo & Kimura, 1970), Krapina
collection (Musgrave, 1977) and additional
data from other sources (Musgrave,
1973; Trinkaus, 1978; Villemeur, 1994;
Niewoehner et al ., 1997). For the South
African Australopithecus species we have
studied the original material housed in the
Transvaal Museum (Pretoria) and the
University of Witwatersrand Medical School
(Johannesburg). For East African Australo-
pithecus and the KNM-WT 15000 skeleton
we have studied the casts housed in theLaboratory for Human Evolutionary Studies
(University of California, Berkeley) and in
the Cleveland Museum of Natural History
(Cleveland, Ohio).
We include some original data of the
extensive fossil collection from the Sima de
los Huesos (SH) Middle Pleistocene site,
also at Sierra de Atapuerca (Burgos) with an
approximate age of 300,000 ka (Arsuaga
et al ., 1997a; Bischoff et al ., 1997). The SHfossils exhibit a number of primitive traits
absent in Late Pleistocene Neandertals, as
well as other traits transitional or close to the
Neandertal morphology. These features are
characteristic of H. heidelbergensis (Arsuaga
et al ., 1991, 1993, 1997b; Carretero et al .,
1997; Martınez & Arsuaga, 1997).
Finally two samples of modern humans,
Euroamericans and Afroamericans, from the
Hamann-Todd collection housed in theCleveland Museum of Natural History
(Cleveland, Ohio), have been used in the
comparative analysis. Also, some additional
comparative data of modern human samples
from literature has been used.
Variables
We define the hamate and capitate measure-
ments in Tables 1 and 2. We have measured
the metacarpals and hand phalanges follow-
ing Musgrave (1977), adding the proximal
articular height and proximal articular
breadth measurements (Trinkaus, 1983a).
For metatarsals and foot phalanges we fol-
low the same criteria used for the hand
remains.
Results and discussion
Minimum number of individuals
Based on teeth, six individuals could be
identified in the TD6 level of Gran Dolina
(Carbonell et al ., 1995; Bermudez de Castro
et al ., 1997, 1999): Hominid 1, an early
adolescent around 14 years old; Hominid 2,
a child between 3 and 4 years old; Hominid
3, a juvenile 10–11·5 years old; Hominid 4,a young adult around 20 years old; Hominid
5, another young adult also around 20 years
old; and Hominid 6, represented only by
ATD6-312, a left I2 germ, with an age at
death between 3 and 4 years.
The hand and foot remains from TD6
correspond to a minimum number of four
individuals: one juvenile, one adolescent
and two adults. The left hamate, ATD6-23,
and the left capitate, ATD6-24, both belongto the same adult individual. The hallucal
proximal phalanx ATD6-31 presents scars
of the epiphyseal line which, according to
modern human standards, suggests an age at
death between 13 and 15 years (Hoerr &
Pyle, 1962), and could be assigned to
Hominid 1. Using the modern human
patterns of hand development reported by
Greulich & Pyle (1959) the two middle
hand phalanges, ATD6-44 and ATD6-53,with their proximal epiphyses unfused,
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could belong to the same immature indi-
viduals, probably to Hominid 3 with an age
at death of around 10–11 years. ATD6-44
could be from the second or fifth ray and
ATD6-53 from the third or fourth ray.
ATD6-26 and ATD6-59 are both meta-
carpals from left second rays from adult
individuals without scars of the epiphyseal
line, and could represent Hominid 4 and
Hominid 5. The rest of the remains could
not be assigned securely to any of the adults
or adolescent individuals. None of the hand
and foot remains belong to either of the two
children (Hominids 2 and 6) identified byteeth (Bermudez de Castro et al ., 1999).
Carpals
Capitate. The overall dimensions of the
ATD6-24 capitate do not diff er substantially
when compared with modern humans,
Neandertals and SH (Table 1). Villemeur
(1994) has reported a slightly higher
maximum height and a lower breadth in the
Neandertal capitates relative to modern
human capitates. ATD6-24 presents a
maximum height/maximum length index
(77·7%) lower than the Neandertal values
(96·56·6, n=8; data from Villemeur,
1994), SH (85·63·0, n=6) and modern
humans (Euroamericans= 83·34·5%, n=20; Afroamericans= 83·45·5%, n=24).
Table 1 Dimensions of the ATD6-24 capitate (in mm and degrees)
ATD6-24 SH NeandertalsEuroamericans
(n=24)Afroamericans
(n=20)
Maximum length (M1) 24·1 22·81·5 22·31·9 22·61·9 22·41·6(n=6) (n=8)
Maximum breadth (M2) 14·1 13·11·1 12·31·1 13·91·1 13·61·2(n=6) (n=8)
Maximum height (M3) 18·8 19·61·8 21·41·6 18·81·5 18·61·2(n=6) (n=7)
Articular length (M4) 22·6 21·71·5 — 21·62·0 21·41·5(n=6)
Head breadth (M5) 13·2 12·81·0 11·31·1 12·31·4 11·91·2(n=6) (n=3)
Head height (M6) 12·3 11·50·9 12·10·9 12·11·1 12·30·9(n=6) (n=8)
MC3 facet height (CMC3Ht)* (15·7) 15·71·3 14·42·0 15·91·3 15·81·1
(n=6) (n=8)MC3 facet breadth (CMC3Br)* 6·1 7·91·1 8·91·4 — —
(n=6) (n=8)MC2+MC3 facet breadth (CapMxBr)* 10·8 10·21·0 10·61·3 — —
(n=6) (n=8)MC2 facet height (CMC2Ht)* 13·7 14·21·3 12·91·3 — —
(n=4) (n=8)MC2 facet breadth (CMC2Br)* 5·3 4·21·0 4·70·9 5·80·7 5·40·7
(n=5) (n=8)MC2 facet depth (CMC2Dp)* 2·3 1·80·2 1·20·3 — —
(n=3) (n=8)MC2 facet angle (CMC2A)* 48 552·2 6010·1 467·0† —
(n=4) (n=8) (n=41)
*Definition of variables and comparative data of Neandertals from Niewoehner et al. (1997).†Ameridian date from Niewoehner et al. (1997).M# refers to Martin & Saller (1957) measurements.Sima de los Huesos (SH) sample=AT-1008, AT-1009, AT-1305, AT-1309, AT-1319 and AT-1805.Neandertal sample=Amud 1, Chapelle, La Ferrassie 1, La Ferrassie 2, Kebara 2, Krapina 200, Shanidar 4 and
Tabun 1.
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ATD6-24 presents a maximum breadth/
maximum length index (58·5%) between
Neandertal values (55·43·0%, n=8; data
from Villemeur, 1994) and modern
humans (Euroamericans= 61·84·3%, n=
20; Afroamericans= 60·93·0%, n=24),but within the standard deviations of both.
Some authors have observed that
Neandertals have a more parasagittally
oriented capitate–metacarpal 2 facet
(CMC2), even though the articulations
between the capitate and the second and
third metacarpals in Neandertals are similar
to those of modern humans in their relative
dimensions (Riley & Trinkaus, 1989;
Trinkaus et al ., 1991). These authors con-cluded that Neandertals were not well
adapted for resisting oblique joint reaction
forces and inferred that the Neandertals
did not habitually employ tools which
required oblique power grips. In a more ex-
tensive analysis, Niewoehner et al . (1997)
demonstrate that Neandertal and recent
human capitate–metacarpal 2/3 articulations
have significant morphological diff erences,
but their behavioural correlates remainuncertain.
The ATD6-24 capitate presents an angle
between the capitate–metacarpal 2 facet
(CMC2) and the capitate–metacarpal 3
facet (CMC3) of 48 (CMC2A in
Table 1), a value intermediate between the
Neandertals and SH (6010·8 and55·32·2, respectively) and modern
humans (Puebloans=467·0, urbans=
398·9; from Niewoehner et al ., 1997).
Although the samples have overlapping
ranges of variation, Neandertals and SH
have a more parasagittally oriented CMC2
facet than modern humans and the Gran
Dolina capitate. Recently, Leakey et al .
(1998) reported an almost complete left
capitate (KNM-KP 31724) assigned to Australopithecus anamensis having a facet for
the second metacarpal that faces completely
laterally, a morphology more primitive than
both capitates of Australopithecus afarensis
from Hadar (Bush et al ., 1982; Johanson
et al ., 1982) and that of Australopithecus
africanus from Sterkfontein (McHenry,
1983).
The proximo-ulnar concavity of the
CMC2 facet permits some degree of pronation/supination of the second ray, but
Table 2 Dimensions of the ATD6-23 hamate (in mm)
ATD6-23 SH NeandertalsEuroamericans
(n=25)Afroamericans
(n=21)
Articular length* (19·1) 15·70·7 17·60·7 18·42·5 18·71·7(n=4) (n=5)
Maximum height (M3) (23·5) 22·71·0 26·41·7 21·92·5 23·22·1
(n=3) (n=5)
Hamulus length* 10·8 11·60·3 11·91·5 10·01·7 10·51·4
(n=3) (n=8)
Hamulus thickness* 4·9 5·30·2 6·10·5 4·60·7 5·30·6
(n=3) (n=8)
Hamulus projection (M5) (9·0) 10·40·8 11·71·3 9·21·6 9·51·7(n=3) (n=5)
Hamulus area† 52·9 62·12·5 72·411·3 47·214·1 55·411·5
(n=3) (n=8)
*Definition of variables in Trinkaus (1983a).†Hamulus area=(hamulus lengthhamulus thickness).M# refers to Martin & Saller (1957) measurements.Sima de los Huesos (SH) sample=AT-939, AT-1310, AT-1311 and AT-1313.Neandertal sample=La Ferrassie 1, La Ferrassie 2, Amud 1, Kebara 2, Shanidar 3, 4, 5 and Tabun 1.
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Neandertals and recent humans do not
diff er in the CMC2 facet curvatures
(Niewoehner et al ., 1997). In ATD6-24 the
relative curvature of the CMC2 facet
[calculated as the ratio of the facet subtenseto its chord (CMC2 depth/CMC2
height100)] is 16·7, a value in the upper
range of variation seen in modern humans
(Puebloan=9·13·4, Urban=9·13·7;
from Niewoehner et al ., 1997) and
Neandertals (9·22·8; from Niewoehner
et al ., 1997).
In African apes a strong interosseous
MC2–MC3 ligament becomes continuous
with the interosseous ligament betweencapitate and trapezoid, dividing the CMC2
joint in two facets, and giving the CMC3
facet a constricted appearance in distal view
(Lewis, 1973; Marzke, 1983). Conversely,
modern humans present a single concave
CMC2 facet with a weak interosseous liga-
ment attachment. The ATD6-24 capitate
presents a strong attachment for the inter-
osseous ligament in the radial side, but the
groove dividing the CMC2 is absent. There-
fore the CMC2 facet of ATD6-24 is con-
tinuous and concave but the CMC3 facet
is somewhat constricted in distal view
[Figure 1(b)].
In dorsal and palmar view, the capitates of
the African apes have constricted necks
(Lewis, 1973). The ATD6-24 capitate
shares with A. anamensis, A. afarensis and A.
africanus a less constricted neck than the
apes, but more than most modern humans.
Also the ATD6-24 capitate shows a welldeveloped head [Figure 1(b)].
The capitate–metacarpal 3 joint (CMC3)
in modern humans is relatively immobile,
due to the flat articular surface and to the
presence of the styloid process in the third
metacarpal, which gives more stability to
this joint (Marzke & Marzke, 1987). The
capitates of A. anamensis and A. afarensis
present dorsal borders of the CMC3 more
perpendicularly oriented, and the styloidprocesses are absent in the third metacarpals
of A. afarensis (Bush et al ., 1982; Marzke,
1983; Leakey et al ., 1998). However Ricklan
(1987) reported a short styloid process in
the third metacarpal of A. africanus. The
orientation of the dorsodistal border of theATD6-24 capitate is more perpendicular to
the longitudinal axis and lacks the radial
bevelling for MC3 found in modern
humans. Thus, the ATD6-24 capitate
preserves the primitive morphology of the
dorsodistal border and, although no third
metacarpal was found in the TD6 level, we
infer a small styloid process on the MC3.
The dorsal border of the CMC3 of
Neandertals is oriented more perpendicularto the shaft and the styloid process is
absolutely and relatively shorter and ori-
ented more radially than in modern humans
(Villemeur, 1994; Riley & Trinkaus, 1989).
Hominid hand bones are scarce in the fossil
record during most of the period when stone
toolmaking and tool use developed. Never-
theless, Marzke & Marzke (1987) noted that
the earliest preserved evidence of a styloid
process is observed in the hands of the
Neandertals, and link the evolution of tool-
making capabilities with a developmental
process producing a styloid process in the
third metacarpal. However, we can observe
that approximately 1·5 million years after
the first appearance of the stone tools, the
ATD6-24 capitate, associated with more
than two hundred tools (Carbonell et al .,
1999), lacks a bevelled dorsodistal border to
accommodate the third metacarpal styloid
process. Therefore, the link between toolmaking and tool use and the presence of
the styloid process in the third metacarpal is
unclear.
The three capitates from A. afarensis
and A. africanus present dorsally placed
trapezoid facets (McHenry, 1983), contrary
to modern humans and Neandertals, which
exhibit a palmarly placed trapezoid facet.
ATD6-24 exhibits an intermediate mor-
phology with two trapezoid facets. The maintrapezoid facet (7·83·8 mm) is palmarly
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placed, and a very small trapezoid facet
(approximately 2 mm2) is dorsally placed,
which is reminiscent of the primitive loca-
tion. Thus, ATD6-24 is the most ancient
fossil with a morphology of the trapezoidfacet transitional between Australopithecus
and later Homo.
Hamate. One of the most striking features
of the Neandertal carpal bones is related to
the depth of the carpal tunnel and the
tendons that pass through (Trinkaus,
1983a). Well-developed scaphoid and
trapezium tubercles, as well as the hamulus,
are traits which reflect this feature because
they provide attachment to the transverse
carpal ligament, which bridges the tunnel
(Trinkaus, 1983a; Villemeur, 1994).
The hamulus of the left hamate ATD6-23
is well preserved and extends distally, pro-
viding the origin of the flexor digiti minimi
and opponens digiti minimi muscles. But
the overall dimensions (Table 2) and the
palmar projection of the ATD6-23 hamulus
are smaller than in Neandertals [see Figure
1(a)]. The cross-sectional area of theATD6-23 hamulus (hamulus length
hamulus thickness= 52·9 mm2) is closer to
modern humans (Euroamericans= 47·2
14·1 mm2; Afroamericans=55·411·5 mm2)
than to Neandertals (76·214·9 mm2,
n=7; data from Trinkaus, 1983a) or SH
(62·52·5 mm2, n=3).
On ATD6-23 the facet for the triquetral is
continuous, meeting the capitate facet at the
sharp proximal border, and we can notdiscern a lunatohamate contact [Figure
1(a)]. In the Viegas et al . (1990) study, 35%
of 165 modern humans lack lunatohamate
contact. Also, the A. afarensis hamate A.L.
333-50 does not seem to have a lunate facet
(Bush et al ., 1982; Marzke et al ., 1994).
The groove of the pisometacarpal liga-
ment for MC3 on the hamate is discernible
in ATD6-23. This ligament stabilizes and
buttresses the joints of the carpal bonesagainst forces associated with tool use
(Marzke, 1996). This groove is also
present in A.L. 333-50 (Marzke & Shackley,
1986).
Second metacarpal morphologyThe bilateral dorsal ridges of the ATD6-59
second metacarpal represent the dorsal
extent of the interosseous muscles [Figure
1(c)]. The ulnar ridge crosses the shaft
dorsally and joins with the radial ridge on
the dorsoradial border. The radial border of
the diaphysis in ATD-59 where the first
interosseous muscle inserts, presents several
oblique cutmarks.
The metacarpal heads of ATD6-29 and
ATD6-59 are asymmetrical and the dorsal
aspect of the phalangeal facets are narrow
and ulnarly placed. The latter presents two
strong tubercles for the collateral liga-
ments of the metacarpophalangeal joint.
Neandertals present relatively wide meta-
carpal heads and narrow metacarpal mid-
shafts (Musgrave, 1971). ATD6-59 also has
a wide head relative to the shaft breadth
and presents a head index of 50·7—closer
to Neandertals than to modern humans(Table 3).
Hand phalanges
The postcranium of A. afarensis retains
numerous primitive traits relative to
Homo sapiens, which include middle hand
phalanges with pronounced ridges lateral
to the insertion of the flexor digitorum
superficialis, and also strong impressions for
this muscle; proximal hand phalanges II-Vare slender, curved, and have strong flexor
sheath ridges (McHenry, 1994).
The proximal hand phalanges of A.
africanus are less curved than those of A.
afarensis (although Stw 28 is strongly curved
according to Susman, 1988), and have less
strongly developed flexor sheath ridges
(Ricklan, 1987). Two proximal hand
phalanges from Member III of Swartkrans
(SKX 5018 and 22741) are less curved thanthose from Hadar, but a third one (SKX
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T a b l e 3
D i m e n s i o n s o f t h e A T D 6 - 5
9 s e c o n d m e t a c a r p a l ( i n m m )
S i d e
M i d s h a f t
b r e a d t h
M i d s h a f t
h e i g h t
D i s t a l
b r e a d t h
D i s t a l
h e i g h t
M i d s h a f t
i n d e x
D i s t a l
i n d e x
H e a d
i n d e x
A T D
6 - 5
9
L
7 · 7
8 · 8
1 5 · 2
1 5 · 0
8 7 · 5
1 0 1 · 3
5 0 · 7
A T - 1
2 7 2 ( S H )
L
7 · 1
8 · 8
1 3 · 2
1 3 · 6
8 0 · 7
9 7 · 1
5 3 · 8
A T - 1
3 6 2 ( S H )
R
7 · 4
8 · 9
1 3 · 7
—
8 3 · 2
—
5 4 · 0
N e a n
d e r t a l s
7 · 4
0 · 9
8 · 8
1 · 1
1 4 · 2
0 · 8
1 5 · 4
1 · 7
8 3 · 9
4 · 4
9 6 · 3
5 · 1
5 2 · 2
4 · 0
( n = 6 )
( n = 6 )
( n = 3 )
( n = 7 )
( n = 6 )
( n = 3 )
( n = 3 )
E u r o a m e r i c a n s ( n = 4 8 )
8 · 6
1 · 7
8 · 8
1 · 2
1 4 · 1
1 · 3
1 4 · 1
1 · 4
9 6 · 3
1 0 · 4
1 0 0 · 3
4 · 4
5 9 · 5
5 · 3
6 · 8 – 1
0 · 3
6 · 9 – 1
0 · 5
1 1 · 4 – 1
6 · 9
1 1 · 0 – 1
7 · 4
7 5 · 3 – 1
1 8 · 3
9 1 · 4 – 1
1 7 · 4
4 9 · 4 – 7
3 · 6
A f r o a
m e r i c a n s ( n = 4 8 )
8 · 6
1 · 0
9 · 4
1 · 0
1 4 · 3
1 · 2
1 4 · 3
1 · 2
9 1 · 7
7 · 7
9 9 · 8
4 · 6
6 0 · 4
4 · 4
7 · 0 – 1
1 · 0
7 · 7 – 1
1 · 3
1 2 · 0 – 1
7 · 2
1 1 · 5 – 1
6 · 3
7 2 · 9 – 1
1 6 · 7
8 8 · 7 – 1
1 1 · 0
5 0 · 7 – 6
9 · 6
M i d s h a f t i n d e x = ( m i d s h a f t b r e a d t h / m i d s h a f t h e i g h t ) 1 0 0 ; d i s t a l i n d e x = ( d i s t a l b r e a d t h / d i s t a l h e i g h
t ) 1 0 0 ; h e a d
i n d e x = ( m i d s h a f t
b r e a d t h / d i s t a l
b r e a d
t h ) 1 0 0 .
N e
a n d e r t a l s a m p l e = L a F e r r a s s i e 1 , L
a C h a p e l l e , R e g o u r d o u ,
K r a p i n a 2
0 1 . 1 ,
K e b a r a 2 ,
S h a n i d a r 3 ,
4 ,
6 a
n d T a b u n 1 .
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27431) is as curved as those from Hadar
(Susman, 1988). All of them are attributed
by McHenry (1994) to Paranthropus, but
Susman (1988) attributed SKX 27431 to
Homo even though it derives from Member3 of Swartkrans which contains only
‘‘robust’’ australopithecine craniodental
material (McHenry, 1994). Homo habilis
retains some australopithecine traits in con-
trast to later species of Homo, including
a hand with robust and curved middle
phalanges with well marked insertions for
flexor digitorum superficialis, and thick and
curved proximal phalanges (Susman &
Creel, 1979; McHenry, 1994).
The dimensions of the ATD6 hand
phalanges are very similar to modern
humans (Tables 4 and 5). Although frag-
mentary, the diaphysis of the proximal hand
phalanges from Gran Dolina (ATD6-27,
ATD6-67 and ATD6-82) seems to be non-
curved [Figure 1(g)]. The well marked
ridges along their shafts reflect the attach-
ment of the fibrous sheaths which hold the
flexor tendons towards the phalangeal shafts
and prevent bowstringing of the musclesduring flexion. The middle hand phalanges
also present marked insertions for the
flexor superficialis in their diaphyses [Figure
1(f,j,k)]. However, ATD6 fossils present
less developed flexor musculature than
the hand phalanges of living apes, Australo-
pithecus and H. habilis.
A relatively broad head on the manual
phalanges is a typical Neandertal feature
(Musgrave, 1973). In Table 6 we show thatthe trochlear index of hand middle
phalanges ATD6-28 (52·9) and ATD6-46
(52·3) exhibit closer values to Neandertals
and SH than to Euroamericans and Afro-
americans. A broad trochlea is also present
in the middle phalanx WT-15000BO
(Walker & Leakey, 1993) but not in A.
afarensis (Bush et al ., 1982) or Paranthropus
(Susman, 1988). Thus, broad phalangeal
heads seem to be the primitive morphologyfor Homo, ATD6, SH and Neandertals, but
modern humans show derived narrow
phalangeal heads.
Metatarsal
The overall dimensions and morphology of the ATD6-70+ 107 second metatarsal are
very similar to that of modern humans
(Table 7). In ATD6-70+107 the dor-
sopalmar diameter of the metatarsal head
includes the proximally projecting plantar
cornua, with the lateral cornua larger
[Figure 2(a)]. A transverse groove separates
the expanded superior surface of the head
from an elevated ridge connecting the two
tubercles for the collateral ligaments of the
metatarsophalangeal joint, with the medial
tubercle more distally located. The A.
afarensis metatarsals from Hadar [A.L. 333-
72, A.L. 333-115(B) and A.L. 333-115(C)]
present a similar metatarsal head mor-
phology. The dorsal surface of the shaft
presents a sharp ridge for the attachment of
the interosseus muscles. Neandertals
present wider diaphyses than modern
humans, but ATD6-70+ 107 shows a low
midshaft index (Table 7), below theEuroamerican and Afroamerican means.
Hallucal proximal phalanges
Both hallucal proximal phalanges from
Dolina (ATD6-30 and ATD6-31) present
prominent plantar tubercles for the attach-
ment of the collateral ligament. The meta-
tarsal facets are proximodorsally oriented
and the insertion of the extensor digitorum
brevis muscle on the proximodorsal surfaceof the diaphysis is smoothly marked [Figure
2(c) and (d)]. Although their dimensions do
not diff er much from that of the modern
human samples (Table 8), we can identify
some diff erences in the shafts, bases and
proximal articular surfaces. The shaft
breadth relative to shaft height of the Nean-
dertal hallucal proximal phalanges tends to
be larger than that of recent human samples
(Trinkaus & Hilton, 1996). However, theDolina hallucal phalanges exhibit a more
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T a b l e 4
M e a s u r e m e n t s o f t h e T D 6
h a n d a n d f o o t p h a l a n g e s ( i n m m )
L a b e l
F i n g e r
S i d e
M a x i m u m
l e n
g t h
A r t i c u l a r
l e n g t h
P r o x i m a l
b r e a d t h
P
r o x i m a l
h e i g h t
P r o x i m a l
a r t i c u l a r
b r e a d t h
P r o x i m a l
a r t i c u l a r
h e i g h t
M i d s h a f t
b r e a d t h
M i d s h a f t
h e i g h t
T r o c h l e a r
b r e a d t h
T r o c h l e a r
h e i g h t
H a n d
:
A T
D 6 - 2
7
P P
—
—
—
—
—
—
—
1 0 · 4
6 · 6
—
—
A T
D 6 - 2
8
M P 3 / 4
—
2 9
· 4
2 6 · 8
1 3 · 1
9 · 6
1 1 · 3
7 · 1
( 9 · 1
)
( 5 · 1
)
1 0 · 2
5 · 4
A T
D 6 - 2
9
P P
—
—
—
—
—
—
—
—
—
1 1 · 6
7 · 5
A T
D 6 - 4
4 *
M P 2 / 5
—
—
—
( 7 · 8
)
( 5 · 5
)
—
—
( 6 · 6
)
( 3 · 5
)
—
—
A T
D 6 - 4
6
M P
—
—
—
—
—
—
—
( 9 · 7
)
( 5 · 4
)
1 0 · 7
5 · 6
A T
D 6 - 5
3 *
M P 3 / 4
—
( 2 0
· 3 )
—
( 1 0 · 3
)
( 7 · 3
)
—
—
8 · 1
4 · 5
7 · 0
4 · 2
A T
D 6 - 6
7
P P
—
—
—
—
—
—
—
1 0 · 3
6 · 7
—
—
A T
D 6 - 8
2
P P 2 ?
L ?
—
—
—
—
—
—
( 9 · 8
)
( 5 · 3
)
9 · 5
6 · 5
F o o t :
A T
D 6 - 3
0
P P 1
L
3 6
· 6
3 1 · 0
1 8 · 0
1 6 · 7
1 5 · 6
1 4 · 3
1 1 · 8
1 0 · 5
1 5 · 0
1 0 · 7
A T
D 6 - 3
1
P P 1
L
3 4
· 3
2 9 · 1
1 8 · 3
1 7 · 0
1 6 · 9
1 4 · 4
1 1 · 0
9 · 9
1 6 · 1
1 0 · 8
A T
D 6 - 3
6
D P
—
—
—
—
—
—
—
—
—
( 8 · 0 )
( 5 · 4
)
A T
D 6 - 3
2
P P 2 / 4
R ?
—
—
—
—
—
—
( 5 · 3
)
( 5 · 0
)
7 · 8
5 · 6
A T
D 6 - 3
3
M P
—
1 4
· 9
1 3 · 1
9 · 6
8 · 5
9 · 1
6 · 2
7 · 3
5 · 6
9 · 5
5 · 1
A T
D 6 - 3
4
M P
—
1 2
· 7
1 1 · 7
8 · 3
7 · 4
7 · 9
4 · 9
4 · 6
3 · 9
8 · 0
4 · 6
A T
D 6 - 3
5
M P
—
1 0
· 6
9 · 5
7 · 4
6 · 6
7 · 1
4 · 9
5 · 7
4 · 2
7 · 7
3 · 9
A T
D 6 - 6
8
D P
—
1 1
· 1
1 0 · 4
9 · 1
6 · 3
8 · 7
5 · 4
5 · 3
5 · 0
6 · 6
5 · 0
P a r e n t h e s e s ( ) i n d i c a t e e s t i m a t e d v a l u e s . P P = p r o x i m a l p h a l a n x ; M P = m
i d d l e p h a l a n x ; D P = d i s t a l p h a l a n x
. * I m m a t u r e p h a l a n x w i t h o u t p r o x
i m a l e p i p h y s i s .
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T a b
l e 5
C o m p a r i s o n s o f t h e A T D 6
- 2 8 , A T D 6 - 4 6 a n d A T D 6 - 5 3 m i
d d l e h a n d p h a l a n g e s d i m e n s i o n
s ( i n m m )
M a x i m u m
l e n g t h
A r t i c u l a r
l e n g t h
P r o x i m a l
b r e a d t h
P r o x
i m a l
h e i g h t
P r o x i m a l
a r t i c u l a r
b r e a d t h
P r o x i m a l
a r t i c u l a r
h e i g h t
M i d s h a f t
b r e a d t h
M i d s h a f t
h e i g h t
T r o c h l e a
r
b r e a d t h
T r o c h l e a r
h e i g h t
A T D
6 - 2
8
2 9 · 4
2 6 · 8
1 3 · 1
9 · 6
1 1 · 3
7 · 1
( 9 · 1
)
( 5 · 1
)
1 0 · 2
5 · 4
A T D
6 - 4
6
—
—
—
—
—
—
( 9 · 7
)
( 5 · 4
)
1 0 · 7
5 · 6
A T D
6 - 5
3 *
( 2 0 · 3
)
—
( 1 0 · 3
)
( 7 · 3
)
—
—
8 · 1
4 · 5
7 · 0
4 · 2
S H
2 8 · 8
1 · 3
2 7 · 3
1 · 1
1 4 · 4
1 · 1
9 · 6
0 · 9
1 2 · 0
1 · 1
7 · 6
1 · 1
9 · 5
0 · 6
5 · 8
0 · 7
1 1 · 1 0 · 8
5 · 7
0 · 5
( n = 8 )
( n = 8 )
( n = 7 )
( n =
6 )
( n = 7 )
( n = 6 )
( n = 8 )
( n = 8 )
( n = 7 )
( n = 7 )
N e a n d e r t a l s
2 8 · 3
2 · 0
2 6 · 2
2 · 4
1 4 · 4
1 · 4
1 0 · 1
1 · 0
1 1 · 8
1 · 3
6 · 5
0 · 9
8 · 8
0 · 9
5 · 7
0 · 7
1 1 · 5 1 · 0
5 · 9
0 · 6
( n = 1 7 )
( n = 2 3 )
( n = 2 2 )
( n =
2 3 )
( n = 1 5 )
( n = 1 8 )
( n = 2 3 )
( n = 2 3 )
( n = 2 3 )
( n = 2 3 )
E a r l y m o d e r n h u m a n s
2 7 · 7
1 · 7
2 8 · 8
2 · 5
1 3 · 7
1 · 4
9 · 8
0 · 6
—
—
9 · 3
1 · 4
5 · 4
0 · 7
1 0 · 1 1 · 2
6 · 2
0 · 8
( n = 3 )
( n = 8 )
( n = 8 )
( n =
8 )
( n = 9 )
( n = 9 )
( n = 9 )
( n = 9 )
3 r d
a n d 4 t h r a y s :
E u r o a m e r i c a n s ( n = 9 6 )
2 9 · 2
2 · 3
2 7 · 0
2 · 1
1 3 · 3
1 · 3
9 · 8
0 · 9
1 1 · 2
1 · 1
6 · 8
0 · 8
8 · 6
1 · 1
5 · 3
0 · 7
1 0 · 1 1 · 0
6 · 0
0 · 7
2 3 · 9 – 3
5 · 2
2 2 · 0 – 3
2 · 7
1 1 · 0 – 1
6 · 8
7 · 9 –
1 2 · 8
9 · 2 – 1
4 · 3
5 · 1 – 9 · 3
6 · 4 – 1
1 · 4
3 · 8 – 7 · 1
8 · 3 – 1 2 ·
8
4 · 8 – 7 · 9
A f r o
a m e r i c a n s ( n = 9 6 )
3 0 · 6
2 · 8
2 8 · 5
2 · 5
1 3 · 5
1 · 2
9 · 9
0 · 9
1 1 · 3
1 · 1
7 · 2
0 · 8
8 · 8
1 · 1
5 · 4
0 · 8
1 0 · 2 0 · 9
6 · 0
0 · 7
2 4 · 7 – 3
7 · 6
2 3 · 9 – 3
5 · 2
1 0 · 8 – 1
6 · 4
7 · 7 –
1 2 · 0
9 · 3 – 1
3 · 5
5 · 4 – 9 · 4
6 · 4 – 1
1 · 7
3 · 9 – 7 · 0
8 · 4 – 1 2 ·
8
4 · 6 – 7 · 6
P a r e n t h e s e s ( ) i n d i c a t e e s t i m a t e d v a l u e s . * I m m a t u r e i n d i v i d u a l w i t h o u t e p i p h y s i s .
S i m a d e l o s H u e s o s ( S H ) s a m p l e i s c o m p o s e d o f a d u l t m i d d l e p h a l a n g e s
f r o m t h e t h i r d a n d f o u r t h r a y s A T - 9
4 ,
A T - 1
0 7 ,
A T - 2
6 3 ,
A T - 3
0 6 ,
A T
- 6 8 3 ,
A T - 6
8 4 ,
A T -
8 9 1 a n d A T - 1
0 1 7 .
N
e a n d e r t a l s a m p l e = L a F e r r a s s i e 1 , L a F e r r a s s i e 2 ,
K i i k - K o b a , T a b u n 1 ,
K e b a r a 2 ,
S h a n i d a r 3 ,
4 ,
5 ,
6 , K
r a p i n a 2 0 5 · 1 ,
2 0 5 · 2 ,
2 0 5 · 3 ,
2 0 5 · 4
, 2 0 5 · 5 ,
2 0 5 · 6 ,
2 0 5 · 8 ,
2 0 5 · 1
2 a n d A m u d 1 .
E a r l y m o d e r n h u m a n s a m p l e = S k h u l
I V ,
D o l n ı ´ V e s t o n i c e 3 ,
C o m b e - C a
p e l l e , A r e n e C a n d i d e , B a r m a G r a n
d e .
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rounded section and a taller shaft height
relative to the shaft breadth than those of
either the modern human samples or
Neandertals (Table 9). The base and the
proximal articular surfaces of both ATD6
hallucal proximal phalanges are very
rounded, as can be observed in the base and
articular indices (Table 9).
Also, the pedal hypertrophy of the
Neandertals is reflected in the musculoliga-
ments’ attachment areas on the bases of the
proximal phalanges, relative to the size of
the articular area for the metacarpal that
they surround (Trinkaus, 1983b). But this
diff erence is most clear in the lateral digits
and the proximal rugosity index of the Gran
Dolina hallucal proximal phalanges is very
similar to Neandertals, SH and modern
humans (Table 9). The hallucal proximal
phalanges of the Neandertals tend to be
slightly more robust, showing larger diaphy-seal diameters relative to length (Trinkaus,
1983b). Also, the Middle Pleistocene SH
hallucal proximal phalanges present large
robusticity indices, contrary to the robustic-
ity of the ATD6 hallucal proximal phalanges
that are more similar to the modern humans
(Table 9).
Toe phalanges
The proximal foot phalanges of A. afarensis
are long, curved, broad-based, narrow-
bodied in dorsal view, and have a medi-
olateral flare of the body for the flexor
sheaths and a more highly circumferential
trochlea; the middle foot phalanges are
relatively long also (McHenry, 1994).
It is further demonstrated that Neandertal
proximal pedal phalanges are short relative
to foot length, and exhibit wide diaphyses
compared to that of recent humans
(Trinkaus, 1983a; Trinkaus et al ., 1991;
Trinkaus & Hilton, 1996). We compare thevalues of the proximal pedal phalanx
Table 6 Comparisons of the ATD6-28 and ATD6-46 middle phalanges indices (in mm)
Baseindex
Articularindex
Midshaftindex
Trochlearindex
Robusticityindex
ATD6-28 73·3 62·8 (56·0) 52·9 26·5ATD6-46 — — (55·7) 52·3 — SH 66·94·0 63·811·0 60·44·5 51·83·1 28·01·7
(n=6) (n=6) (n=8) (n=7) (n=8)Neandertals 71·14·0 55·36·2 65·17·7 51·43·0 27·83·2
(n=22) (n=15) (n=23) (n=23) (n=23)Early modern humans 71·63·7 — 59·15·1 61·04·3 26·32·1
(n=8) (n=9) (n=9) (n=8)3rd and 4th ray:Euroamericans (n=96) 73·93·9 60·94·9 61·54·3 59·03·3 25·82·2
66·7–86·3 49·2–73·2 47·5–70·8 51·5–66·7 20·8–31·3Afroamericans (n=96) 73·72·9 63·35·2 61·45·2 58·94·0 24·82·5
68·4–82·3 51·1–78·8 50·4–71·4 49·1–68·2 19·4–31·1
Parentheses ( ) indicate estimated values.Base index=(proximal height/proximal breadth)100; articular index= (proximal articular height/proximal
articular breadth)100; midshaft index=(midshaft height/midshaft breadth)100; trochlear index=(trochlearheight/trochlear breadth)100; robusticity index=0·5(midshaft breadth+midshaft height)/articular length100.
Sima de los Huesos sample is composed of adult middle phalanges from the third and fourth rays AT-94,AT-107, AT-263, AT-306, AT-683, AT-684, AT-891 and AT-1017.
Neandertal sample=La Ferrassie 1, La Ferrassie 2, Kiik-Koba, Tabun 1, Kebara 2, Shanidar 3, 4, 5, 6, Krapina205·1, 205·2, 205·3, 205·4, 205·5, 205·6, 205·8, 205·12 and Amud 1.
Early modern human sample= Dolnı Vestonice 3, Combe-Capelle, Arene Candide, Barma Grande.
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T a b
l e 7
D i m e n s i o n s o f t h e A T D 6 -
7 0 + 1 0 7 s e c o n d m e t a t a r s a l ( i n m
m )
M a x i m u m
l e n g t h
A r t i c u l a r
l e n g t h
M i d s h a f t
b r e a d t h
M i d s h a f
t
h e i g h t
D i s t a l
b r e a d t h
D i s t a l
h e i g h t
E p i c
o n d y l a r
b r
e a d t h
M i d s h a f t
i n d e x
D i s t a l
i n d e x
R o b u s t i c i t y
i n d e x
A T D
6 - 7
0 + 1 0 7
( 7 9 )
7 6 · 3
7 · 3
1 0 · 0
1 1 · 7
1 7 · 1
( 1 2 · 1
)
7 3 · 0
6 8 · 4
1 1 · 3
A T D
6 - 9
9 2 + 1 1 3 8 ( S H )
7 7 · 7
7 4 · 1
8 · 2
9 · 4
1 1 · 3
1 6 · 4
1 2 · 8
8 7 · 2
6 8 · 9
1 1 · 9
N e a n d e r t a l s
7 6 · 4
4 · 9
6 8 · 5
4 · 5
8 · 4
0 · 9
8 · 9 1 · 2
1 1 · 0
1 · 3
1 5 · 0
—
9 4 · 0
6 · 4
6 5 · 3
1 2 · 5
0 · 2
( n = 5 )
( n = 2 )
( n = 8 )
( n = 8 )
( n = 2 )
( n = 1 )
( n = 8 )
( n = 1 )
( n = 2 )
E u r o a m e r i c a n s ( n = 2 2 )
7 4 · 7
4 · 7
7 1 · 3
4 · 5
7 · 3
0 · 9
9 · 0 1 · 0
1 0 · 7
1 · 0
1 5 · 6
1 · 5
1 1 · 4
1 · 5
8 1 · 4
1 0 · 5
6 8 · 7 7 · 2
1 1 · 4
0 · 9
6 4 · 8 – 8
3 · 4
6 1 · 8 – 7
9 · 8
5 · 6 – 9 · 1
7 · 5 – 1 0 ·
6
8 · 5 – 1
2 · 3
1 1 · 6 – 1
8 · 7
8 · 6
– 1 5 · 7
6 2 · 9 – 1
0 3 · 4
5 9 · 0 – 9 7 · 4
9 · 5 – 1
3 · 0
A f r o
a m e r i c a n s ( n = 2 5 )
7 8 · 7
5 · 0
7 5 · 7
4 · 8
7 · 7
1 · 1
9 · 3 1 · 1
1 0 · 7
1 · 4
1 5 · 6
2 · 0
1 1 · 6
1 · 2
8 3 · 0
8 · 9
6 8 · 8 7 · 0
1 1 · 2
0 · 8
7 0 · 4 – 8
6 · 1
6 7 · 7 – 8
3 · 0
6 · 2 – 1
1 · 1
7 · 9 – 1 1 ·
6
8 · 5 – 1
4 · 0
9 · 5 – 1
9 · 2
9 · 6
– 1 5 · 4
6 0 · 6 – 9
7 · 9
5 7 · 1 – 9 2 · 6
9 · 7 – 1
4 · 0
P a r e n t h e s e s ( ) i n d i c a t e e s t i m a t e d v a l u e s .
M
i d s h a f t
i n d e x = ( m i d s h a f t
b r e a d t h / m i d s h a f t
h e i g h t ) 1 0 0 ;
d i s t a l
i n d e x = ( d i s t a l b r e a d t h / d i s t a l h e i g h t ) 1 0 0 ;
r o b u s t i c i t y
i n d e x = 0 · 5
( m i d s h a f t
b r e a
d t h + m i d s h a f t h e i g h t ) / a r t i c u l a r l e n g t h
1 0 0 .
N
e a n d e r t a l s a m p l e = L a F e r r a s s i e 1 , L a F e r r a s s i e 2 ,
K i i k - K o b a , A m u d 1
, T a b u n 1 ,
S h a n i d a r 1 ,
3 ,
4 ,
6 a n d
S p y .
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T a b l e 8
C o m p a r i s o n s o f t h e A T D 6 - 3 0 a n d A T D 6 - 3 1 h a l l u c a l p r o x i m a l p h a l a n g e s d i m e n s i o n s ( i n m m )
M a x i m u m
l e n g t h
A r t i c u l a r
l e n g t h
P r o x i m a l
b r e a d t h
P r o x i m
a l
h e i g h
t
P r o x i m a l
a r t i c u l a r
b r e a d t h
P r o x i m a l
a r t i c u l a r
h e i g h t
M
i d s h a f t
b r e a d t h
M i d s h a f t
h e i g h t
T r o c h l e a
r
b r e a d t h
T r o c h l e a r
h e i g h t
A T D
6 - 3
0
3 6 · 6
3 1 · 0
1 8 · 0
1 6 · 7
1 5 · 6
1 4 · 3
1 1 · 8
1 0 · 5
1 5 · 0
1 0 · 7
A T D
6 - 3
1
3 4 · 3
2 9 · 1
1 8 · 3
1 7 · 0
1 6 · 9
1 4 · 4
1 1 · 0
9 · 9
1 6 · 1
1 0 · 8
S H
3 4 · 6
1 · 3
2 8 · 3
1 · 4
2 1 · 1
2 · 0
1 7 · 3
1 · 1
1 8 · 3
1 · 5
1 4 · 1
1 · 1
1 4 · 8
1 · 3
1 2 · 0
0 · 7
1 8 · 4 0 ·
8
1 0 · 3
0 · 4
( n = 4 )
( n = 4 )
( n = 5 )
( n = 5 )
( n = 5 )
( n = 4 )
( n = 4 )
( n = 4 )
( n = 4 )
( n = 4 )
N e a n
d e r t a l s
3 0 · 4
2 · 4
2 6 · 5
2 · 2
1 7 · 2
1 · 5
1 7 · 1
1 · 5
1 7 · 4
2 · 0
1 3 · 8
1 · 6
1 2 · 9
1 · 3
9 · 7
0 · 9
1 6 · 9 2 ·
1
9 · 5
1 · 3
( n = 5 )
( n = 1 2 )
( n = 1 2 )
( n = 1 2
)
( n = 1 4 )
( n = 1 3 )
( n = 1 2 )
( n = 1 2 )
( n = 1 1 )
( n = 1 1 )
P a n (
n = 1 9 )
3 0 · 7
1 · 9
2 7 · 7
1 · 7
1 5 · 9
1 · 1
1 2 · 5
0 · 7
1 2 · 4
0 · 7
1 0 · 5
0 · 9
9 · 8
0 · 9
7 · 9
0 · 7
1 2 · 1 0 ·
9
9 · 1
0 · 6
2 7 · 2 – 3
3 · 7
2 4 · 5 – 3
0 · 4
1 3 · 8 – 1
7 · 9
1 1 · 3 – 1
4 · 1
1 1 · 1 – 1
3 · 8
9 · 3 – 1
2 · 6
8 · 2 – 1
1 · 5
6 · 8 – 9 · 2
1 0 · 5 – 1 4 ·
1
7 · 8 – 1
0 · 3
G o r i l l a ( n = 2 4 )
3 0 · 9
3 · 8
2 7 · 0
3 · 5
2 0 · 5
2 · 6
1 5 · 0
1 · 6
1 5 · 8
2 · 0
1 2 · 3
1 · 3
1 2 · 7
2 · 0
9 · 0
1 · 0
1 4 · 9 1 ·
8
9 · 7
1 · 3
2 3 · 1 – 3
6 · 0
2 0 · 1 – 3
2 · 3
1 6 · 0 – 2
4 · 6
1 2 · 5 – 1
7 · 8
1 2 · 7 – 1
9 · 9
9 · 6 – 1
4 · 6
9 · 2 – 1
5 · 4
7 · 0 – 1
1 · 0
1 1 · 3 – 1 7 ·
5
6 · 8 – 1
1 · 8
E u r o a m e r i c a n s ( n = 8 0 )
3 4 · 1
2 · 5
2 8 · 5
2 · 2
1 9 · 8
1 · 7
1 6 · 3
1 · 6
1 7 · 6
1 · 7
1 3 · 6
1 · 5
1 2 · 0
1 · 7
9 · 7
1 · 2
1 6 · 3 1 ·
6
1 0 · 0
0 · 9
2 9 · 4 – 3
9 · 6
2 4 · 1 – 3
4 · 5
1 7 · 1 – 2
7 · 7
1 2 · 9 – 2
2 · 2
1 4 · 6 – 2
3 · 9
1 0 · 9 – 1
9 · 7
8 · 7 – 1
6 · 9
7 · 2 – 1
3 · 0
1 3 · 0 – 2 1 ·
6
8 · 2 – 1
2 · 8
A f r o a
m e r i c a n s ( n = 7 9 )
3 5 · 1
2 · 6
2 9 · 5
2 · 5
2 0 · 1
1 · 7
1 6 · 9
1 · 5
1 7 · 8
1 · 7
1 3 · 8
1 · 6
1 1 · 8
1 · 5
9 · 9
1 · 2
1 6 · 4 1 ·
5
1 0 · 2
1 · 1
2 9 · 9 – 4
0 · 7
2 3 · 5 – 3
5 · 9
1 5 · 5 – 2
3 · 7
1 3 · 3 – 2
1 · 4
1 3 · 8 – 2
2 · 5
1 0 · 8 – 1
9 · 0
8 · 5 – 1
4 · 7
7 · 3 – 1
3 · 2
1 3 · 0 – 1 9 ·
5
7 · 9 – 1
2 · 8
S i m
a d e l o s H u e s o s ( S H ) s a m p l e i s c o m p o s e d o f A T - 9
6 ,
A T - 6
8 7 ,
A T - 7
7 2 ,
A T - 8
9 8 a n d A T - 8
9 9 .
N e
a n d e r t a l s a m p l e = L a F e r r a s s i e 2 , K
i i k - K o b a , S p y 2 5 F & G ,
S h a n i d a r 1 ,
3 ,
4 ,
6 ,
8 ,
K r a p i n a 2 5 0 · 1 ,
2 5 0 · 2
, 2 5 0 · 3 ,
2 5 0 · 4 ,
2 5 0 · 5 ,
2 5 3 · 3 a n d T
a b u n 1 .
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T a b
l e 9
C o m p a r i s o n s o f t h e A T D 6
- 3 0 a n d A T D 6 - 3 1 h a l l u c a l p r o x
i m a l p h a l a n g e s i n d i c e s ( i n m m )
B a s e
i n d e x
A r t i c u l a r
i n d e x
M i d s h a f t
i n d e x
T r o c h l e a r
i n d e x
R o b u s t i c i t y
i n d e x
P r o x i m a l
r u g o s i t y i n d e x
A T D
6 - 3
0
9 2 · 8
9 1 · 7
8 9 · 0
7 1 · 3
3 6 · 0
7 4 · 2
A T D
6 - 3
1
9 2 · 9
8 5 · 2
9 0 · 0
6 7 · 1
3 5 · 9
7 8 · 2
S H
8 1 · 5
6 · 2
7 7 · 1
2 · 2
8 1 · 1
4 · 5
5 6 · 0 2 · 8
4 7 · 6
5 · 4
7 0 · 2
3 · 4
( n = 5 )
( n = 4 )
( n = 4 )
( n = 4 )
( n = 4 )
( n = 4 )
N e a n d e r t a l s
8 5 · 8
4 · 5
7 8 · 9
5 · 0
7 5 · 4
5 · 9
5 6 · 4 4 · 8
4 2 · 6
3 · 0
7 4 · 7
7 · 2
( n =
1 1 )
( n = 1 2 )
( n = 1 2 )
( n = 1 0 )
( n = 1 2 )
( n = 1 1 )
P a n
( n = 1 9 )
7 8 · 3
4 · 2
8 4 · 1
6 · 3
8 0 · 7
7 · 4
7 4 · 8 4 · 8
3 1 · 9
1 · 9
6 5 · 8
7 · 3
6 8 · 2
– 8 5 · 5
6 7 · 4 – 9
5 · 0
6 6 · 1 – 9
1 · 5
6 1 · 0 – 8 1 · 3
2 8 · 4 – 3
6 · 2
5 5 · 5 – 8
1 · 5
G o r i l l a ( n = 2 4 )
7 3 · 3
4 · 3
7 8 · 6
8 · 2
7 1 · 8
5 · 9
6 5 · 2 3 · 6
4 0 · 4
4 · 0
6 4 · 1
8 · 1
6 5 · 9
– 7 9 · 2
6 5 · 1 – 9
4 · 7
6 3 · 0 – 8
8 · 0
5 5 · 3 – 7 0 · 5
3 2 · 4 – 5
1 · 2
3 7 · 5 – 7
4 · 9
E u r o a m e r i c a n s ( n = 8 0 )
8 2 · 4
5 · 6
7 7 · 7
5 · 9
8 1 · 4
7 · 1
6 1 · 5 4 · 5
3 8 · 3
5 · 0
7 4 · 1
5 · 4
7 0 · 1
– 9 2 · 4
6 2 · 3 – 9
1 · 4
6 7 · 3 – 1
0 0 · 0
5 0 · 0 – 7 2 · 8
2 5 · 7 – 5
0 · 2
5 8 · 4 – 8
8 · 0
A f r o
a m e r i c a n s ( n = 7 9 )
8 4 · 3
5 · 1
7 7 · 8
5 · 3
8 4 · 3
6 · 0
6 2 · 4 5 · 8
3 7 · 0
4 · 6
7 2 · 5
6 · 7
7 0 · 5
– 9 8 · 6
6 5 · 3 – 9
3 · 2
7 1 · 1 – 9
7 · 9
5 0 · 6 – 8 8 · 9
3 0 · 0 – 5
4 · 4
5 4 · 8 – 8
7 · 3
B a s e i n d e x = ( p r o x i m a l h e i g h t / p r o x i m a l b r e a d t h ) 1 0 0 ; a r t i c u l a r i n d e x = ( p r o x i m a l a r t i c u l a r h e i g h t / p r o x i m a l a r t i c u l a r b r e a d t h ) 1 0 0 ; m i d s h a f t i n d e x =
( m i d
s h a f t h e i g h t / m i d s h a f t b r e a d t h ) 1
0 0 ; t r o c h l e a r i n d e x = ( t r o c h l e a r h e
i g h t / t r o c h l e a r b r e a d t h ) 1 0 0 ; r o b u s t i c i t y i n d e x = 0 · 5
( m i d s h a f t b r e
a d t h + m i d s h a f t
h e i g h t ) / a r t i c u l a r l e n g t h
1 0 0 ; p r o x i m a l r u g o s i t y i n d e x = ( p r o x i m a l a r t i c u l a r h e i g h t / p r o x i m a l a r t i c u l a r b r e a d t h
) / ( p r o x i m a l m a x i m u m h e i g h t / p r o x i m a l m a x i m u m
b r e a
d t h ) 1 0 0 .
S i m a d e l o s H u e s o s ( S H ) s a m p l e = A T - 9
6 ,
A T - 6
8 7 ,
A T - 7
7 2 ,
A T - 8
9 8 a
n d A T - 8
9 9 .
N
e a n d e r t a l s a m p l e = L a F e r r a s s i e 2 , K i i k - K o b a , S p y 2 5 F & G ,
S h a n i d a r
1 ,
3 ,
4 ,
6 ,
8 ,
K r a p i n a 2 5 0 · 1 ,
2 5 0 · 2
, 2 5 0 · 3 ,
2 5 0 · 4 a n d T a b u n 1 .
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ATD6-32 with means of phalanges from the
second, third and fourth ray. The foot proxi-
mal phalanx ATD6-32 presents a larger
midshaft breadth than midshaft height and
has a midshaft index (midshaft breadth/midshaft height100=106·0) that is inter-
mediate between Neandertal (114·87·1,
n=11) and modern human means (Euro-
americans=87·413·6, n= 99; Afro-
americans=86·410·9, n=105). The foot
proximal phalanx ATD6-32 also presents a
higher midshaft index when we compare it
to two prehistoric Amerindian samples, who
probably wore little more than protective
footgear (Libben=95·0, n=120; Pecos=95·6, n=136; mean of proximal pedal
phalanges 2, 3 and 4 calculated using mean
values from Trinkaus & Hilton, 1996), but
is very similar to Early modern humans
(104·6, n=23; calculated as previously from
Trinkaus & Hilton, 1996).
The three pedal middle phalanges of Gran
Dolina (ATD6-33, ATD6-34 and ATD6-
35) and the two distal phalanges (ATD6-26
and ATD6-68) have the same morphology
and dimensions of those of modern humans,
Neandertals and SH (Table 10).
Stature estimation
In spite of the problems of using metatarsal
length to estimate stature, we have used it
for this purpose because it is the only bone
available. Byers et al . (1989) provide some
regression formulae using two samples of
Euroamericans and Afroamericans from
the Terry collection. Using the metatarsal
articular length we estimate the stature of
the ATD6-70+ 107 individual. The corre-
lation coefficient between the metatarsal
articular length and stature ranges between
r =0·66 and r =0·75 (Byers et al ., 1989).
Applying the combined formula for both
samples and both sexes we obtain a stature
of 169·76·5 cm (r =0·78). Some authors
prefer to apply formulae derived from
European people to estimate the stature of Neandertals, due to their similar body pro-
portions (Vandermeersch & Trinkaus, 1995;
Holliday, 1997; Holliday & Ruff , 1997).
Using the Euroamerican male and female
samples we obtain values of 173·07·0 cm
and 168·95·2 cm respectively, with amean estimate of 170·9 cm. This figure is
very close to a stature of 172·5 cm obtained
using the radius ATD6-43 from eight
diff erent equations (Carretero et al ., 1999).
Summary and conclusions
In spite of the scarcity and the fragmentary
nature of the Gran Dolina fossils we can
observe some characteristics in the handremains: a capitate with a constricted neck,
well developed head, strong attachment for
the ligamentum interosseum trapezoid-
capitate, a palmarly placed trapezoid facet
with a small dorsal trapezoid facet, remi-
niscent of the primitive condition highly
curved and oblique orientation of the second
metacarpal facet, and a transversally ori-
ented dorsodistal border; a hamate with a
less projecting and lightly built hamulus; an
inferred reduced styloid process on the third
metacarpal base; a wide second metacarpal
head; and well marked insertions of the
flexor digitorum superficialis in the middle
phalanges. The morphology and dimensions
of the pedal remains from TD6 fits within
the modern human range of variation
(including an elongated second meta-
tarsal), but the base, proximal articular sur-
face and shaft of the hallucal proximal
phalanges are more rounded; and the mid-
shaft of the proximal toe phalanx is wider.
From a phylogenetic perspective, the
cranial and postcranial evidence from the
SH supports the view that H. heidelbergensis
was an exclusively European species
ancestral to H. neanderthalensis (Arsuaga
et al ., 1991, 1993, 1997b; Carretero et al .,
1997; Martınez & Arsuaga, 1997), and
according to Bermudez de Castro et al .
(1997) and Arsuaga et al . (1999) H.antecessor represents the last common
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T a b
l e 1 0
C o m p a r i s o n s o f t h e A T D
6 - 3 3 , A T D 6 - 3 4 a n d A T D 6 - 3 5 m
i d d l e f o o t p h a l a n x d i m e n s i o n s ( i n m m )
M a x i m u m
l e n g t h
A
r t i c u l a r
l e n g t h
P r o x i m a l
b r e a d t h
P r o x i m a l
h e i g h t
P r o x i m a l
a r t i c u l a r
b r e a d t h
P r o x i m a l
a r t i c u l a r
h e i g h t
M
i d s h a f t
b
r e a d t h
M i d s h a f t
h e i g h t
T r o c h l e a r
b r e a d t h
T r o c h l e a r
h e i g h t
A T D
6 - 3
3
1 4 · 9
1
3 · 1
9 · 6
8 · 5
9 · 1
6 · 2
7 · 3
5 · 6
9 · 5
5 · 1
A T D
6 - 3
4
1 2 · 7
1
1 · 7
8 · 3
7 · 4
7 · 9
4 · 9
4 · 6
3 · 9
8 · 0
4 · 6
A T D
6 - 3
5
1 0 · 6
9 · 5
7 · 4
6 · 6
7 · 1
4 · 9
5 · 7
4 · 2
7 · 7
3 · 9
S H ( n = 1 7 )
1 0 · 1
2 · 7
8 · 5
2 · 4
9 · 4
1 · 3
7 · 7 1 · 0
8 · 7
0 · 8
5 · 7
0 · 7
7 · 5
1 · 1
4 · 3
0 · 8
8 · 6
1 · 0
4 · 4
0 · 5
N e a n d e r t a l s
—
8 · 8
1 · 5
1 0 · 2
0 · 4
8 · 5 0 · 6
9 · 4
0 · 3
6 · 3
0 · 3
7 · 9
0 · 7
4 · 7
0 · 4
9 · 6
0 · 4
5 · 6
0 · 9
( n = 5 )
( n = 6 )
( n = 6 )
( n = 5 )
( n = 5 )
( n = 6 )
( n = 6 )
( n = 6 )
( n = 6 )
2 n d
a n d 5 t h r a y s :
E u r o a m e r i c a n s
1 1 · 0
3 · 7
9 · 7
3 · 3
9 · 2
0 · 9
7 · 5 0 · 8
7 · 7
1 · 1
4 · 9
0 · 6
6 · 7
1 · 3
4 · 3
0 · 7
8 · 1
0 · 8
4 · 9
0 · 7
( n = 1 3 0 )
( n = 1 3 0 )
( n = 1 3 0 )
( n = 1 3 0
)
( n = 1 3 0 )
( n = 1 3 0 )
(
n = 8 5 )
( n = 8 5 )
( n = 1 3 0 )
( n = 1 3 0 )
A f r o
a m e r i c a n s
1 0 · 8
3 · 4
9 · 7
3 · 1
9 · 1
0 · 9
7 · 5 0 · 9
8 · 0
0 · 9
5 · 1
0 · 7
6 · 0
1 · 2
4 · 0
0 · 6
8 · 3
0 · 9
4 · 8
0 · 6
( n = 1 4 0 )
( n = 1 3 8 )
( n = 1 4 0 )
( n = 1 4 0
)
( n = 1 4 0 )
( n = 1 4 0 )
( n = 1 1 1 )
( n = 1 1 0 )
( n = 1 3 6 )
( n = 1 3 7 )
S i m a d e l o s H u e s o s ( S H ) s a m p l e i s
c o m p o s e d o f a d u l t m i d d l e f o o t p h a l a n g e s f r o m t h e s e c o n d a n d fi f t h r a y s A T - 8
8 ,
A T - 1
0 9 ,
A T - 1
1 1 ,
A T
- 1 1 5 ,
A T - 2
1 4 ,
A T -
2 6 5 ,
A T - 5
1 4 ,
A T - 5
1 6 ,
A T - 5
2 3 , A
T - 5
2 4 ,
A T - 9
0 6 ,
A T - 1
2 8 5 ,
A T - 1 3
4 9 ,
A T - 1
4 3 3 ,
A T - 1
5 1 0 ,
A T - 1
5 1 1
a n d A T - 1
7 6 7 .
N
e a n d e r t a l s a m p l e = S h a n i d a r 4 ,
8 a n
d T a b u n 1 .
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ancestor for H. sapiens (modern humans)
and H. neanderthalensis.
The H. antecessor hand and foot remains
display a set of morphological traits that are
more similar to modern humans than tothe later Middle and Late Pleistocene
European hominids. Our results do not
contradict the previous phylogenetic analy-
sis, i.e., that H. antecessor represents the
last common ancestor for H. sapiens
(modern humans) and H. neanderthalensis
(Neandertals).
Acknowledgements
We thank especially the rest of the
Atapuerca team involved in the excavation
and in the preparation of the remains studied
here. Thanks also to I. Martınez, A. Gracia,
N. Garcıa, L. Lopez-Polın, J. M. Bermudez
de Castro and R. Quam for their discussion,
as well as useful and constructive comments.
M. Marzke and E. Trinkaus provided help-
ful suggestions on this manuscript. We also
thank Leslie Aiello and the editorial staff of
JHE for their invaluable help with the editing
of the text. We thank B. Latimer and L.
Jellema (Cleveland Museum of Natural
History) for access to the Hamann-Todd
collection; P. Andrews, R. Clarke, F. C.
Howell, R. Kruszynski, M. Laranjeira
Rodrigues de Areia, A. Langaney, H. de
Lumley, J. Radovcic, Y. Rak, C. Stringer, F.
Thackeray, P. Tobias and T. D. White for
access to human fossil remains and skeletal
collections under their care. Help in the field
from the Grupo Espeleologico Edelweiss of
Burgos was also essential. The first author
received a grant from the Ayuntamiento de
Madrid in the Residencia de Estudiantes.
Field work at the Atapuerca sites is sup-
ported by the Junta de Castilla y Leon, and
this research was funded by the Direccion
General de Ensenanza Superior (PB96-
1026-C03-03), by the Comunidad de
Madrid (06/0037/1997) and by the UnidadAsociada (CSIC-UCM).
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