Journal of Human Evolution 78 (2015) 114e121
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Journal of Human Evolution
journal homepage: www.elsevier.com/locate/jhevol
Early Pleistocene human hand phalanx from the Sima del Elefante (TE)
cave site in Sierra de Atapuerca (Spain)
n Pablos c, d, e, *, Jose
Miguel Carretero c, d, Rosa Huguet a, b, f,
Carlos Lorenzo a, b, c, Adria
n-Torres e, Juan Luis Arsuaga c, g, Eudald Carbonell a, b, f,
Josep Valverdú a, b, f, María Martino
e
María Bermúdez de Castro
Jose
a
ria, Universitat Rovira i Virgili, Avinguda Catalunya 35, 43002 Tarragona, Spain
Area de Prehisto
de Paleoecologia Humana i Evolucio
Social (IPHES), Marcel·lí Domingo s/n, 43007 Tarragona, Spain
Institut Catala
n sobre Evolucio
n y Comportamiento Humanos, c/Monforte de Lemos 5, 28029 Madrid, Spain
Centro Mixto UCM-ISCIII de Investigacio
d
n Humana (LEH), Dpto. de Ciencias Histo
ricas y Geografía, Universidad de Burgos, Edificio IþDþi, Plaza Misael Ban
~ uelos s/n,
Laboratorio de Evolucio
09001 Burgos, Spain
e
National Research Center on Human Evolution (CENIEH), Paseo Sierra de Atapuerca s/n, 09002 Burgos, Spain
f
Unit Associated to CSIC, Campus Sescelades URV, (Edifici W3) E3, 43007 Tarragona, Spain
g
Departamento de Paleontología, Universidad Complutense de Madrid, Avenida Complutense s/n, 28040 Madrid, Spain
b
c
a r t i c l e i n f o
a b s t r a c t
Article history:
Received 17 February 2014
Accepted 12 August 2014
Available online 8 September 2014
In this study, a new Early Pleistocene proximal hand phalanx (ATE9-2) from the Sima del Elefante cave
site (TE e Sierra de Atapuerca, Spain), ascribed to Homo sp., is presented and comparatively described in
the context of the evolution of the genus Homo. The ATE9-2 specimen is especially important because of
the paucity of hand bones in the human fossil record during the Early Pleistocene. The morphological and
metrical analyses of the phalanx ATE9-2 indicate that there are no essential differences between it and
comparator fossil specimens for the genus Homo after 1.3 Ma (millions of years ago). Similar to Sima de
los Huesos and Neandertal specimens, ATE9-2 is a robust proximal hand phalanx, probably reflecting
greater overall body robusticity in these populations or a higher gracility in modern humans. The age of
level TE9 from Sima del Elefante and morphological and metrical studies of ATE9-2 suggest that the
morphology of the proximal hand phalanges and, thus, the morphology of the hand could have remained
stable over the last 1.2e1.3 Ma. Taking into account the evidence recently provided by a metacarpal from
Kaitio (Kenya) from around 1.42 Ma, we argue that modern hand morphology is present in the genus
Homo subsequent to Homo habilis.
© 2014 Elsevier Ltd. All rights reserved.
Keywords:
Phalanges
Homo
Postcranial evolution
Western Europe
Introduction
Study of the hominin hand provides important information
about tool use (Susman, 1994; Ward et al., 2014 and references
therein) as well as phylogeny and taxonomy. Analysis of a third
metacarpal from Kaitio dated to around 1.42 Ma (millions of years
ago) indicates a styloid process morphology similar to that of
modern humans (Ward et al., 2014). This suggests that this early
Homo individual was potentially able to manufacture lithic tools, as
later Homo individuals did (Marzke and Marzke, 2000). However,
the scarcity of hand bones in the human fossil record complicates
investigation of the evolution of this anatomical region in the genus
* Corresponding author.
E-mail addresses: adrizaino@yahoo.es, apablos@isciii.es (A. Pablos).
http://dx.doi.org/10.1016/j.jhevol.2014.08.007
0047-2484/© 2014 Elsevier Ltd. All rights reserved.
Homo and consequently when modern-like hand morphology
appeared in the fossil record. In particular, little is known about the
morphology of hand phalanges in the genus Homo, especially from
the Early to Middle Pleistocene. Only a small number of Homo
proximal hand phalanges have been described (Leakey et al., 1964;
-Sola
et al., 2008).
Lorenzo et al., 1999; Susman et al., 2001; Moya
Furthermore, in the case of the Early Pleistocene African fossils, it
may be difficult to determine whether they belong to the genus
Homo, Australopithecus or Paranthropus (Susman et al., 2001; Moy
a et al., 2008).
Sola
Some morphological traits of the hominin hand seem to have
been in stasis for over a million years (Lorenzo et al., 1999; Ward
et al., 2014), although verifying evolutionary trends in hand phalanges through the late Early and early Middle Pleistocene is
thwarted by the paucity of fossil remains from early Homo and
Middle Pleistocene hominins. The ATE9-2 specimen (a hominin
C. Lorenzo et al. / Journal of Human Evolution 78 (2015) 114e121
adult left hand phalanx, probably of the fifth finger, Fig. 1) from
Sima del Elefante described and compared here thus represents an
important contribution in the understanding of the evolution of the
hand phalanges in the Early and Middle Pleistocene. The best evidence for the evolution of the Pleistocene hominin hand comes
from comparisons of Neandertals and modern humans. Although
there are no key differences in the morphology of the hand between these two taxa, the proximal hand phalanges of Neandertals
display broader diaphyses and trochleas (Musgrave, 1970; Trinkaus,
1983; Villemeur, 1994) compared with the more gracile form
evident in modern human populations. These traits observed in
Neandertals have been related to corporal robusticity (Trinkaus,
1983). Alternatively, they could represent the primitive morphological pattern, since relatively broad trochleas in proximal hand
phalanges are evident in Homo antecessor (Lorenzo et al., 1999;
Carretero et al., 2001). Nonetheless, the H. antecessor hand phalanges from the Early Pleistocene site of TD6, dated to around
900e950 ka (thousands of years ago) (Berger et al., 2008), are very
similar to modern humans and Neandertals (Lorenzo et al., 1999).
During the 2008 field season, the ATE9-2 specimen was recovered from square I-28 of the TE9c level of the Sima del Elefante cave
site (Sierra de Atapuerca, Burgos, northern Spain). It was found at
the same depth and less than 2 m from the hominin mandible
ATE9-1 (Carbonell et al., 2008) that has been referred to Homo sp.
(Bermúdez de Castro et al., 2011). The Sima del Elefante cave site
(TE) is located in the Railway Trench, 100 m from the entrance and
about 200 m away from the Gran Dolina cave site (Bermúdez de
Castro et al., 1999; Fig. 2). The TE site corresponds to a sedimentary karstic infilling stopping up the entrance to the so-called
‘Galería Baja’, which belongs to the Cueva Mayor-Cueva del Silo
complex where the Sima de los Huesos site is also located (Arsuaga
et al., 1997a; Fig. 2).
The Sima del Elefante cave is about 15 m wide and the railway
outcrop exposed a sedimentary thickness of about 25 m. The section is formed by 16 lithostratigraphic units mostly composed of
debris flow deposits and defined by major unconformities,
115
(TE7eTE21) (Rosas et al., 2006; Fig. 2). Starting in 1996, the entire
TE sequence was cleaned, and a vertical profile of the infilling has
been undertaken. Samples were taken mainly for biostratigraphic
and geochronological studies. Furthermore, starting in 2005, an
excavation of the TE9 level was conducted on an area of more than
30 m2. A complete and detailed biostratigraphical study was done
s et al. (2010, 2013) and Rofes and Cuenca-Besco
s
by Cuenca-Besco
(2013). The climatic conditions and environment at the time of
formation of the TE7eTE16 levels (the so-called Sima del Elefante
Lower Red Unit) was researched by Blain et al. (2010), whose
findings were based on the amphibian and squamate reptile
assemblages.
Paleomagnetic study of TE reveals that stratigraphic layers
s et al.,
TE7eTE16 have reverse magnetization directions only (Pare
2006; Fig. 1). These results are consistent with a Matuyama age of
the sediments (1.78e0.78 Ma) and with the mammal assemblage
s et al.,
(Carbonell et al., 2008; García et al., 2008; Cuenca-Besco
2013). Two dates based on the radioactive decay of cosmogenic
26
Al and 10Be obtained in the TE9 and TE7 (Fig. 1) suggest a range of
0.95e1.38 Ma for these levels (Carbonell et al., 2008). Thus, based
on a combination of paleomagnetism, cosmogenic nuclides, and
biostratigraphical data, the TE9 level has been dated to the Early
Pleistocene (about 1.2 Ma) or possibly even older (1.3 Ma).
The objective of this report is to present and comparatively
describe the ATE9-2 specimen, which together with the mandible
n deciduous
ATE9-1 (Carbonell et al., 2008) and the Barranco Leo
molar (Toro-Moyano et al., 2013), represents one of the oldest
hominin fossils found in Europe. We also carry out a study of
proximal hand phalanges in fossil humans in comparison with
ATE9-2 in order to test when the modern hand morphology
appeared and establish the polarity of the ATE9-2 morphology.
Material and methods
Our comparative sample comprised original fossils of proximal
hand phalanges of the fifth finger (PHP5) from Kebara 2, La
Figure 1. ATE9-2. Fifth proximal hand phalanx. Views: A) dorsal, B) lateral, C) plantar, D) medial, E) proximal, F) distal, G) 3D reconstruction from CT images. Scale in cm.
116
C. Lorenzo et al. / Journal of Human Evolution 78 (2015) 114e121
Figure 2. A) Location of the Sima del Elefante site in relation to other sites in the Railway Trench at Atapuerca (Burgos, Spain). B): Stratigraphic profile of the Sima del Elefante cave
site. Cosmogenic burial ages in TE7 and TE9 are also shown, with the standard error given at the 68% confidence interval.
rez-Martínez in Huguet (2007).
Modified from Pe
Ferrassie 1 and 2 and Middle Pleistocene material from Sima de los
Huesos (SH) in Atapuerca (Lorenzo et al., 2012), as well as previously published raw data from Shanidar 4 and 5 (Trinkaus, 1983), El
mez, 2000), Dolní Ve
stonice 3, 14, 15 and
n SDR-083 (Sierra Go
Sidro
16 (Trinkaus and Jelínek, 1997; Sl
adek et al., 2000), Qafzeh 7, 8 and 9
(Vandermeersch, 1981), Skhul 4 (McCown and Keith, 1939) Pavlov
31p (Trinkaus et al., 2010), and UW.88.121 (Australopithecus sediba)
from Malapa (Kivell et al., 2011). We also included our own
measurements of the Stw 28 cast (early Homo or Australopithecus)
from Sterkfontein Member 4. The modern human sample was
drawn from the Hamann-Todd collection (composed of 96 individuals consisting of 48 Euro-Americans and 48 Afro-Americans)
housed in the Cleveland Museum of Natural History (Ohio, USA), as
well as a further recent human sample of 38 Europeans (Musgrave,
1970). In addition, published phalangeal curvature data from PlioPleistocene southern African hominin specimens, and Pliocene
C. Lorenzo et al. / Journal of Human Evolution 78 (2015) 114e121
Hadar specimens (Susman et al., 1984, 2001; Susman, 1989) were
used in comparisons.
The morphological variables used in our analyses are as
€uer (1988), detailed in Appendix A, Supplementary
described in Bra
Online Material (SOM Table SI.1 and Figure SI.1). We used standard
anthropometric techniques and instruments to take all measurements (digital calipers to the nearest 0.1 mm). In order to describe
the size, relative proportions and articular dimensions of the
proximal hand phalanx ATE9-2, ten linear variables (total length,
articular length, proximal maximum height, proximal maximum
breadth, proximal articular height, proximal articular breadth,
midshaft height, midshaft breadth, distal height and distal breadth)
and five indices (Table 1) were used, along with qualitative
morphological descriptions. In order to test to which finger the
117
phalanx ATE9-2 belongs, a discriminant function analysis (DFA)
was carried out on our modern human sample of known rays, and
the fossil ATE9-2 classified into one of the phalanx groups.
Phalanx curvature was assessed through examining the
included angle (q). One of several methods proposed to quantify the
curvature of the phalanges (Stern et al., 1995; and references
therein), q has been demonstrated to be highly correlated with the
normalized curvature moment arm and can be obtained from
phalangeal landmarks and distances (Susman et al., 1984; Jungers
et al., 1997; see Table 2 for measurement protocol).
A comparative univariate analysis of all of the variables was
undertaken. To compare individual values from ATE9-2 with the
averages from the fossil and modern samples, Z-scores were
calculated, and a value of 1.96 was considered significant (p < 0.05,
Table 1
Comparisons of ATE9-2 fifth proximal hand phalanx measurements (in mm).
ATE9-2
Stw 28
A. sediba
(UW.88.121)
Total length (TL)
34.7
32.4
27.21,2,3
Articular length (AL)
32.3
30.1
e
Proximal maximum height (PMH)
10.3
11.1
8.52,4
Proximal maximum breadth (PMB)
15.1
12.7
10.81,2,4,5
Proximal articular height (PAH)
8.7
9.5
e
Proximal articular breadth (PAB)
11.9
11.7
e
Midshaft height (MdH)
6.52
6.32
4.31,2,4
Midshaft breadth (MdB)
8.9
8.8
7.7
Distal height (DH)
6.4
7.6
5.01,2,4,5
Distal breadth (DB)
9.8
9.6
7.71,2,5
Proximal index
68.2
87.41,4,5
78.71,5
Articular proximal index
73.1
81.2
e
Midshaft index
73.02,3
71.63
55.83,4,5
Distal index
65.32
79.21,2,4
65.92
Robusticity index
23.8
25.1
e
SH
Neandertals
LP H.sap
MHeHTH
MHeMus
33.7 ± 1.5
[31.8e35.0]
(n ¼ 5)
31.9 ± 1.3
[30.5e33.6]
(n ¼ 5)
9.92 ± 0.8
[9.2e11.1]
(n ¼ 6)
14.7 ± 1.5
[13.0e17.0]
(n ¼ 7)
8.6 ± 0.8
[7.7e9.5]
(n ¼ 5)
11.1 ± 1.0
[10.0e12.3]
(n ¼ 6)
6.1 ± 0.6
[5.3e6.8]
(n ¼ 7)
9.43,4,5 ± 1.0
[8.4e10.6]
(n ¼ 7)
6.5 ± 0.6
[5.7e7.3]
(n ¼ 8)
10.04 ± 0.9
[8.8e11.2]
(n ¼ 8)
69.03,4 ± 3.5
[63.9e72.1]
(n ¼ 6)
78.8 ± 3.5
[76.2e84.8]
(n ¼ 5)
64.5 ± 6.2
[53.9e73.3]
(n ¼ 7)
65.22,4,5 ± 1.4
[62.6e67.0]
(n ¼ 8)
23.95 ± 2.0
[22.6e27.4]
(n ¼ 5)
33.5 ± 2.6
[30.2e37.2]
(n ¼ 7)
31.8 ± 2.5
[28.9e35.4]
(n ¼ 7)
11.04,5 ± 0.7
[10.1e12.1]
(n ¼ 8)
14.8 ± 1.7
[12.9e16.7]
(n ¼ 8)
9.0 ± 0.9
[7.8e10.4]
(n ¼ 8)
11.2 ± 1.1
[9.7e12.7]
(n ¼ 8)
5.5 ± 0.4
[4.9e6.2]
(n ¼ 8)
9.54,5 ± 1.2
[7.7e10.8]
(n ¼ 8)
6.6 ± 0.7
[5.7e7.6]
(n ¼ 7)
10.74,5 ± 1.1
[9.5e12.2]
(n ¼ 7)
74.9 ± 6.6
[67.7e88.4]
(n ¼ 8)
80.4 ± 5.0
[73.6e90.5]
(n ¼ 8)
58.34,5 ± 7.4
[47.2e70.1]
(n ¼ 8)
62.04,5 ± 1.5
[60.0e63.5]
(n ¼ 7)
23.2 ± 1.3
[21.4e24.8]
(n ¼ 7)
35.4 ± 3.4
[29.8e38.4]
(n ¼ 5)
28.0
(n ¼ 1)
34.0 ± 2.7
[29.3e41.6]
(n ¼ 96)
32.1 ± 2.6
[27.5e39.5]
(n ¼ 96)
10.3 ± 0.9
[8.7e12.2]
(n ¼ 96)
14.1 ± 1.2
[11.7e17.1]
(n ¼ 96)
9.0 ± 0.9
[7.3e11.3]
(n ¼ 96)
11.0 ± 1.1
[8.5e13.6]
(n ¼ 96)
5.7 ± 0.7
[4.1e7.0]
(n ¼ 96)
8.4 ± 1.1
[5.9e10.6]
(n ¼ 96)
6.55 ± 0.7
[5.0e8.8]
(n ¼ 96)
9.35 ± 0.9
[7.3e11.5]
(n ¼ 96)
73.35 ± 3.2
[67.2e81.9]
(n ¼ 96)
81.8 ± 5.5
[69.9e97.1]
(n ¼ 96)
67.7 ± 5.3
[53.8e81.2]
(n ¼ 96)
70.55 ± 3.4
[60.0e78.5]
(n ¼ 96)
21.9 ± 2.2
[16.9e26.6]
(n ¼ 96)
e
10.1
[9.2e11.0]
(n ¼ 2)
13.8
[12.5e15.0]
(n ¼ 2)
e
e
5.3 ± 0.6
[4.4e6.0]
(n ¼ 6)
8.2 ± 1.0
6.9e9.6]
(n ¼ 6)
6.5 ± 0.8
[5.6e7.5]
(n ¼ 4)
10.3 ± 1.4
[8.3e11.1]
(n ¼ 4)
73.5
[73.3e73.6]
(n ¼ 2)
e
64.6 ± 1.6
[62.5e66.3]
(n ¼ 5)
65.1
[59.6e68.2]
(n ¼ 3)
20.2
(n ¼ 1)
31.8 ± 2.2
[26.2e36.0]
(n ¼ 38)
10.2 ± 0.9
[8.1e12.6]
(n ¼ 38)
14.4 ± 1.0
[11.7e16.8]
(n ¼ 38)
e
e
5.6 ± 0.7
[4.4e7.2]
(n ¼ 38)
8.3 ± 1.2
[5.6e11.0]
(n ¼ 38)
7.0 ± 0.8
[5.4e8.8]
(n ¼ 38)
9.7 ± 0.8
[8.2e11.9]
(n ¼ 38)
71.0 ± 3.4
[65.3e80.0]
(n ¼ 38)
e
68.2 ± 5.9
[55.4e83.9]
(n ¼ 38)
72.1 ± 3.8
[63.6e82.7]
(n ¼ 38)
21.9 ± 2.1
[17.4e26.5]
(n ¼ 38)
Mean ± standard deviation, range [ ] and sample size (n) are shown. Bold letters and superscript indicate significant differences with some of the samples (Z-score > 1.96 in
absolute terms and ManneWhitney test; p < 0.05); 1 ¼ SH, 2 ¼ Neandertals, 3 ¼ Late Pleistocene H. sapiens, 4 ¼ Hamann-Todd Osteological Collection (HTH), 5 ¼ Musgrave's
(1970) sample (Mus). Modern Humans (MHeHTH and MHeMus) are pooled sex samples. Data for Stw 28 from cast (AMNH); data for A. sediba (UW.88.121) from Kivell et al.
(2011).
Neandertal sample includes: Shanidar 4 (right and left), Shanidar 5 (right), Kebara 2 (left), La Ferrassie 1 and 2 (right and left), SDR-083 (left). Sima de los Huesos (SH) sample
includes: AT-95, AT-515, AT-1326, AT-1327, AT-1390, AT-1486, AT-1827, AT-2479, AT-2831 and AT-4478. Late Pleistocene H. sapiens (LP H.sap) sample includes: Dolní
stonice-DV 3 (right), DV 14 (right?), DV 15 (right?), DV 16, Qafzeh 7 (right); Qafzeh 8 (right), Qafzeh 9 (right); Skhul 4 (left) and Pavlov 31p (right?).
Ve
Proximal index ¼ Prox. max. height/Prox. max. breadth 100. Articular proximal index ¼ Prox. artic. height/Prox. artic. breadth 100. Midshaft index ¼ Midshaft height/
Midshaft breadth 100. Distal index ¼ Dist. height/Dist. breadth 100. Robusticity index ¼ (Midshaft breadth þ Midshaft height) 100/2 (Articular length).
118
C. Lorenzo et al. / Journal of Human Evolution 78 (2015) 114e121
Table 2
Included angle of manual proximal phalanges.
Specimen
Phalangeal dorsal height (H)
Included angle
ATE9-2
5.3
27.0
Present study
Source
AL288-1
AL333-19
AL333-57
AL333-62
AL333-63
AL333-93
AL333-w4
SKX 5018
SKX 15468
SKX 16699
SKX 22741
SKX 27431
Stw 355
5.5
6.3
6
6.4
6.9
5.6
6.8
e
5.5
e
e
e
e
40.5
39.7
40.0
46.5
44.0
34.6
32.4
27.0
30.7
22.0
27.0
34.0
36.0
Bush et al., 1982; Susman
Bush et al., 1982; Susman
Bush et al., 1982; Susman
Bush et al., 1982; Susman
Bush et al., 1982; Susman
Bush et al., 1982; Susman
Bush et al., 1982; Susman
Susman, 1989
Susman et al., 2001
Susman et al., 2001
Susman, 1989
Susman, 1989
Susman et al., 2001
H. sapiens mean1
Chimp mean1
e
e
26.0
42.0
Susman, 1989
Susman, 1989
et
et
et
et
et
et
et
al.,
al.,
al.,
al.,
al.,
al.,
al.,
1984
1984
1984
1984
1984
1984
1984
Measurements in mm, included angle in degrees. The phalangeal dorsal height (H) is used to calculate the included angle following the method proposed by Stern et al. (1995)
[q ¼ ((H MdH/2)2 þ (TL/2)2)/2(H MdH/2)]. 1 ¼ The mean values for modern humans (H. sapiens) and chimps are provided (Susman, 1989).
Sokal and Rohlf, 2003). Z-scores were only calculated in cases
where the ‘n’ of the comparative samples was equal to or greater
than five. To better understand the position of ATE9-2 in the
context of human evolution, and also to characterize the comparative samples, we compared the values of the different samples/
populations with a ManneWhitney U-test (Mann and Whitney,
1947). For statistical analysis, we used STATISTICA 8.0 (StatSoft,
2007).
Results
Description and identification
The bone is complete and its dimensions are shown in Table 1.
The proximal epiphysis is fused, and therefore this bone belonged
to an adult individual over 16 years of age according to modern
human development patterns (Scheuer and Black, 2000; Cardoso
and Severino, 2010). The fossil displays a flat palmar surface and
a transversally convex dorsal surface. Bilateral flexor sheath ridges
are present on the palmar surface; the left ridge is slightly more
developed and more projected palmarly. The proximal articular
surface for the metacarpal is oval, concave and its dorsal border is
slightly indented, which facilitates the hyperextension of the digit.
In dorsal view, the base presents a prominent lateral tubercle for
the insertion of the abductor digiti minimi muscle on the left side
and the trochlea shows a right deviation. All of these traits suggest
identification as a proximal hand phalanx of a left fifth finger. The
distinction of proximal hand phalanges from rays 2e5 is difficult,
and often can be distinguished only when all of the phalanges of
the same hand are present. Given that ATE9-2 is an isolated
element, to corroborate the finger assignment we performed a DFA
to classify the ray of this proximal phalanx. The DFA (SOM Table SI.2
and Fig. SI.2) carried out with the proximal phalanges from ray 2e5
correctly classifies 82.6% of the modern phalanges, but specifically
for the fifth finger the correct classification is much higher (97.9%).
The results of the DFA indicate that ATE9-2 falls comfortably within
the range of variation of ray 5 but outside the range of variation of
rays 3 and 4. Although it falls just within the 95% equiprobability
ellipse for ray 2, the results of our DFA classified ATE9-2 as a fifth
phalanx with a posterior probability of 96.6%. Hence, we considered
that ATE9-2 is a proximal hand phalanx from the fifth finger. Due to
shape variability between rays, in our fossil comparative sample
and further analyses we thus only used phalanges that are known
to belong to the fifth finger.
Metrical comparisons
Table 1 shows the values of the different variables measured for
the phalanx ATE9-2, as well as for the rest of the fossil specimens,
and the main statistical parameters of the comparative samples
(mean, standard deviation, range and sample size). For nearly all of
the variables, ATE9-2 does not display significant differences from
the comparative samples. However, ATE9-2 and Stw 28 both show a
diaphysis that is absolutely and relatively higher than those of the
Neandertals, as evidenced by the midshaft height (Fig. 3A) and the
midshaft index (midshaft index ¼ midshaft height/midshaft
breadth 100), with the latter significantly higher in ATE9-2
relative to Neandertals and Late Pleistocene Homo sapiens samples (Fig. 3D). Moreover, the distal index (distal index ¼ dist. height/
dist. breadth 100) is also significantly higher in ATE9-2, Stw 28
and UW 88.121 compared with Neandertals (Fig. 3C). This is likely
due to the broad distal articulations of the fifth proximal hand
phalanges present in Neandertals (Trinkaus, 1983; Villemeur, 1994).
Although Stw 28 is similar to ATE9-2 in having a high diaphysis,
Stw 28 displays high proximal and distal indices due to the small
breadth of the proximal epiphysis and the high distal trochlea
(Table 1). The fifth proximal hand phalanx of A. sediba (UW.88.121)
is smaller than nearly all of the studied samples. However, its
proximal index is higher than that of the SH population and
modern humans. The midshaft index is lower than that of Late
Pleistocene H. sapiens and modern humans and the distal index is
higher than that of Neandertals.
The fifth proximal hand phalanges of Neandertals display
absolutely thick and wide diaphyses (Fig. 3B) and broader distal
articulations relative to the other samples (Musgrave, 1973;
Trinkaus, 1983), except SH (Lorenzo et al., 2012). Another trait
that the Neandertal and SH samples share is the low distal index
compared with modern humans. ATE9-2 shares a relatively broad
trochlea with SH, Neandertals and TD6, and it is different to the
earlier Stw 28 phalanx (Lorenzo et al., 1999 and Fig. 3C). Moreover,
the Neandertals exhibit a high base and a low midshaft index, due
mainly to the broad diaphyses of their hand proximal phalanges,
including the fifth finger.
The phalanx curvature value obtained for the ATE9-2 phalanx
(Table 2) is identical to the value for SKX 5018 and SKX 22741, close
to the modern human mean and above the value of SKX 16699
(Susman et al., 1984, 2001; Susman, 1989 and Table 2). The value of
ATE9-2 is lower than that obtained for SKX 15468, SKX 27431, Stw
355 and all the Hadar fossils, which represent the highest values
C. Lorenzo et al. / Journal of Human Evolution 78 (2015) 114e121
119
Figure 3. Univariate analysis of ATE9-2 and the comparative samples. HTH ¼ Modern Humans (MH) e Hamann-Todd Osteological collection, Mus ¼ Modern humans from
Musgrave (1970), SH ¼ Sima de los Huesos, Nea ¼ Neandertals, LP Hsap ¼ Late Pleistocene H. sapiens, Aust-E.Homo ¼ Australopithecus or early Homo, A. sediba ¼ UW.88.121 of
A. sediba. SD ¼ standard deviation. A) Midshaft height (in mm), B) Midshaft breadth (in mm), C) Distal index, D) Midshaft index.
among early hominins. Other early hominin phalanges show higher
values, which are closer to or above the chimpanzee mean (Susman
et al., 1984, 2001; Susman, 1989).
Discussion and conclusions
Our comparative analysis of ATE9-2 supports previous assertions that the morphology of the hand, at least as far as we can
ascertain from study of the fifth proximal phalanges, appears to
have remained quite stable during the Early, Middle and Late
Pleistocene (Vandermeersch, 1981; Trinkaus, 1983; Lorenzo et al.,
1999; Ward et al., 2014). Relatively broad phalangeal trochleas
seem to be the primitive morphology for Homo, ATE9-2, the TD6
sample, SH and Neandertals, although modern humans show
derived relatively narrow phalangeal trochleas (Lorenzo et al.,
1999; Carretero et al., 2001). In common with Neandertals, SH
and TD6 populations, ATE9-2 displays a robust proximal phalanx,
probably related to a greater overall bodily robusticity. The only
differences seen in the available fossil record for the fifth proximal
hand phalanx are the broad diaphyses and distal articulations
shared by Neandertals (Musgrave, 1973; Trinkaus, 1983) and SH
samples.
The expansion of the distal articulation in the fifth proximal
hand phalanges in SH and in Neandertals could be an adaptation for
an increased stress level at the fingertip (Trinkaus, 1983). It could
also reflect the evolutionary relationship between these two populations (Arsuaga et al., 1997b, 2014; Carretero et al., 1997; Martínez
mez-Olivencia et al., 2007; Bonmatí et al.,
and Arsuaga, 1997; Go
n-Torres et al., 2013; Pablos et al., 2013, 2014) or
2010; Martino
that both the SH population and Neandertals exhibit a broad and
robust body size. The body robusticity of these hominins is indicated by the robust and very broad pelves with very long superior
pubic rami, long femoral necks, absolutely and relatively long
clavicles, mediolaterally broad axes, long transversal processes in
the lumbar vertebrae, large thoraxes, large tali with broad lateral
malleolar facets and robust calcanei with broad sustentaculum tali
mez-Olivencia
(Carretero et al., 1997, 2004; Arsuaga et al., 1999; Go
et al., 2007, 2009; Bonmatí et al., 2010; Pablos et al., 2012, 2013,
2014). As was previously proposed, this robust morphotype is
probably the primitive condition within the genus Homo from
which modern humans departed (Arsuaga et al., 1999; Bonmatí
et al., 2010).
Compared with raw measurements of later specimens, Stw 28
only differs significantly in midshaft height, but because of this the
proximal and distal indices of Stw 28 are significantly different
from all of the comparative samples including ATE9-2. The phalanx
of A. sediba is smaller and with different proportions than all of the
later fifth proximal phalanges. Australopithecus fossils usually
120
C. Lorenzo et al. / Journal of Human Evolution 78 (2015) 114e121
display curved proximal hand phalanges (Susman et al., 1984,
2001). In contrast, it is accepted that phalanges from the genus
Homo are straighter, except those of OH 7 (Susman and Creel, 1979;
Susman et al., 1984) and other early Homo or Paranthropus specimens from Swartkrans. Our study indicates that the diaphysis of
ATE9-2 is as straight as other phalanges of later members of the
genus Homo, some South African fossils and modern humans, but is
different from the curved phalanges of Australopithecus afarensis
and OH 7.
Hand morphology has been linked to tool manufacture in a
number of studies (Susman, 1994; Marzke, 1997; Marzke and
Marzke, 2000; Susman et al., 2001), and on the basis of the
straightness of the diaphyses in the proximal phalanges from
Swartkrans it has been proposed that tools were likely used by all
early hominins from two million years ago (Susman, 1994, 1998).
Our analyses indicate some differences in fifth proximal phalanx
morphology between African Australopithecus/early Homo and
Eurasian Pleistocene samples, but given the apparent stability in
hand morphology over the past 1.4 Ma, future studies should
examine in more detail the discrepancy between this stability and
the radical changes in tool technology and manufacturing processes over the same time period. It is possible that either the
morphology of the hand is not related to the advances in lithic
technology or that the morphological features needed for
manufacturing complex technologies arose early in the evolution of
the genus Homo. Ideally, more fossil hand bones, particularly from
the Early and Middle Pleistocene are needed to test this hypothesis,
but further detailed morphological and biomechanical research is
possible on the proximal phalanges, such as ATE9-2, already
existing in the fossil record, which to date have received very little
attention.
Acknowledgments
The authors are grateful to all members of the Atapuerca
Research Team for their effort spanning decades recovering information from the Sierra de Atapuerca sites and their superb research
mez-Merino, A. Lombera and
work. Thanks to X.P. Rodríguez, G. Go
M. Terradillos and the rest of TE team, whose field work led to the
discovery of the findings presented here. We wish to thank B.
Latimer, Y. Haile-Selassie and L. Jellema (Cleveland Museum of
Natural History) for providing access to the Hamann-Todd Collece de l’Homme), D. Grimaudtion. Thanks also to P. Mennecier (Muse
(Musee National d'Histoire Naturelle), Y. Rak (Tel Aviv UniHerve
versity) and E. Delson and I. Tattersall (American Museum of Natural History) for providing access to human fossil remains and
skeletal collections in their care. We appreciate the constructive
and fruitful discussion provided by R. Quam. Lauren Ames kindly
reviewed a previous English version.
n General de
This research has been funded by the ‘Direccio
n’ of the Spanish Ministry of ‘Economía y ComInvestigacio
petitividad’ (Project numbers CGL2012-38434-C03-01/02/03), the
AGAUR 2014-SGR-899 project, the ‘Consejería de Cultura y
n’, the European Social Fund
Turismo de la Junta de Castilla y Leo
n Atapuerca’. R.Huguet, J.
(Fondo Social Europeo) and the ‘Fundacio
Vallverdú and E. Carbonell belong to a unit associated with the
CSIC.
Further thanks go to our colleagues at the ‘Centro Mixto UCMn sobre Evolucio
n y Comportamiento
ISCIII de Investigacio
n Humana (LEH)’
Humanos’ and from the ‘Laboratorio de Evolucio
at the University of Burgos. Thanks also go to the BBP and R3
groups.
Finally, we appreciate the helpful comments and suggestions
provided by the editor (Sarah Elton), the associate editor, and the
three anonymous reviewers that improved the manuscript.
Appendix A. Supplementary online material
Supplementary data related to this article can be found at http://
dx.doi.org/10.1016/j.jhevol.2014.08.007
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Supplementary Online Material
Table SI.1.
Metric variables studied in the fifth proximal hand phalanges. In order to study the
ATE9-2 fifth proximal hand phalanx, the standard anthropometric variables were used
(Martin and Saller, 1957; Bräuer, 1988) (Figure SI-1).
Name
Abbrev.
Description
Fig.
Maximum distance from the most proximal point of the
Total length
TL
SI-1a
proximal epiphysis to the most distal point of the trochlea.
Distance from the midpoint of the proximal articular facet to
Articular length
AL
SI-1b
the center point of the trochlea.
Proximal
PMH
Maximum dorso-palmar diameter of the base.
SI-1b
PMB
Maximum medio-lateral diameter of the base.
SI-1a
maximum height
Proximal
maximum breadth
Proximal articular
Maximum dorso-palmar diameter of the proximal articular
SI-1c
PAH
height
facet.
Maximum medio-lateral diameter of the proximal articular
Proximal articular
PAB
breadth
SI-1c
facet.
Dorso-palmar diameter at midshaft. This measurement does
Midshaft height
MdH
SI-1b
not include the flexor sheath ridge.
Midshaft breadth
MdB
Medio-lateral diameter at midshaft.
SI-1a
Distal height
DH
Maximum dorso-palmar diameter of the trochlea.
SI-1b
Distal breadth
DB
Maximum medio-lateral diameter of the trochlea.
SI-1a
Abbrev. = Abbreviation. Fig. = Figure in the SOM.
Figure SI.1. Metric variables studied in the fifth proximal hand phalanx ATE9-2.
Views: (a) dorsal, (b) lateral, (c) proximal. Abbreviations as in Table SI.1: TL = Total
length, AL = Articular length, PMH = Proximal maximum height, PMB = Proximal
maximum breadth, PAH = Proximal articular height, PAB = Proximal articular breadth,
MdH = Midshaft height, MdB = Midshaft breadth, DH = Distal height, DB = Distal
breadth.
Table SI.2. Discriminant function analysis (DFA) of the 10 variables of the
proximal hand phalanges.
Function 1
Function 2
Function 3
Wilks'
Lambda
Eigenvalue
4.118
1.059
0.126
Cumulative proportion
0.776
0.976
1.000
Unstandardized coefficient
TL - Total length
0.083
-0.833
-0.303
0.090
AL - Articular length
0.234
0.770
0.349
0.090
PMB - Prox. max. breadth
-0.958
-1.492
0.196
0.132
PMH - Prox. max. height
0.428
0.640
0.554
0.089
PAB - Prox. artic. breadth
0.048
0.419
0.539
0.087
PAH - Prox. artic. height
-0.640
0.079
0.563
0.094
MdB - Midshaft breadth
-0.071
0.463
-0.666
0.089
MdH - Midshaft height
0.271
0.646
1.060
0.089
DB - Distal breadth
1.212
-0.182
-0.024
0.105
DH - Distal height
-0.448
-0.206
-2.124
0.090
Constant
-8.278
8.954
-6.502
Figure SI.2. Scatter diagram based on discriminant function analysis (DFA) for the
finger assignation. The dashed lines indicate the 90% equiprobability ellipses of modern
humans for the corresponding finger. The solid lines indicate the 95% equiprobability
ellipses of modern humans. PP2-PP5 = proximal hand phalanges from the second to
fifth rays. Note that the fossil ATE9-2 is outside the 90% equiprobability ellipse for the
second finger and well inside that of the fifth finger. The phalanx ATE9-2 is classified
as belonging to a fifth finger with a posterior probability of 96.6%.
Supplementary references
Bräuer, G., 1988. Osteometrie. In: Martin, R., Knussman, R. (Eds.), Anthropologie.
Handbuch der vergleichenden Biologie des Menschen. Fisher, Stuttgart, pp. 160–232.
Martin, R., Saller, K., 1957. Lehrbuch der Anthropologie, Third edition. Gustav Fisher,
Stuttgart.