Science 2 October 2009: vol. 326 no. 5949 pp.

71-71e6
DOI: 10.1126/science.1175831
235 remains from, at least, 36 specimens,
most of wich belong to an adult female (4.4
Mya)
Ethiopia's harsh Afar desert
Is this species belonged to the branch of
bipedal hominins or left out along with the
Great Apes?
Fig. 1. Original and reconstructed os coxa of ARA-
VP-6/500. (Left) Anterolateral and anteromedial views
of the original, with a close-up of the AIIS. (Middle) CT
scans of same views, except the bottom view is a
close-up of the pubic symphyseal face, preserved in
its entirety but damaged at its inferior extremity.
Arrows indicate major areas of distortion corrected by
reconstruction. (Right) Anterolateral and anteromedial
views of the reconstructed os coxa and of the entire
pelvis using mirror reconstruction and a conjectural
sacrum.
Illium lateraly flared and has a
forward sweeping anterior
superior iliac spine
Anteriorly positioned anterior
inferior iliac spine is associated
wi t h a broad, short and
sagitally disposed iliac isthmus
Figure S4. Craniocaudal Pelvic Height and Pelvic Geometry in
Hominoids. Posterolateral views of the os coxae of hominoids: A) Pan;
B) Proconsul (KNM-MW 13142 D); C) Ar. ramidus; D) Au. afarensis (A.L.
288-1). Two chords are indicated on each specimen: one from the
acetabular center to the most caudal point on the auricular surface
(upper chord), and the other from the acetabular center to the ischial
spine (lower chord). Note that the angle between the two chords is most
obtuse in Proconsul and declines in the other taxa but for different
reasons.
(D) Minimum ischial length (from
inferior acetabular border to juncture
of the ischial body and tuberosity
surface) normalized by acetabular
di ameter. Two val ues for ARA-
VP-6/500 bracket possible extremes
for the acetabular diameter. Pan-
Homo di f f er ences ar e hi ghl y
significant.
(C) Angle made by two chords
connecting three landmarks on the
ilium. The third chord was measured
(auricular surface to the ischial spine),
and the angl e was determi ned
trigonometrically. All early hominids lie
within the human range because all
have a greatly shortened iliac isthmus.
Figure S5. Relative lateral Iliac Breadth in Anthropoids.
Natural log-log scatterplot of lateral iliac breadth (distance from
the most lateral point on the auricular surface to the ilium's lateral
margin taken at the location of maximum iliac breadth) and
acetabular height in anthropoids. A regression line (blue: RMA: y
= .485x + 1.573; r = .938; N = 107) has been fitted to the Old
World monkey data, and another of slope equal to one (red) has
been fitted through two points estimated as possible upper and
lower limits of Proconsul (based on KNM-MW 13142 D). Upper
and lower limits of possible acetabular height have been used for
ARA-VP-6/500.
Figure S7. Relative Maximum Iliac Height in Anthropoids. Natural log-
log scatterplot of maximum iliac length and acetabular height in anthropoids.
A regression line (blue: RMA: y = .802x + 2.130; r = .876; N = 145) has been
fitted through Old World monkey taxa and another of slope equal to one
(red) has been fitted through Proconsul (KNM-MW 13142 D). Four points
are provided for ARA-VP-6/500 reflecting upper and lower limits for both
dimensions (acetabular height and maximum iliac length). Note that the ilia
of African apes (but not Pongo) appear slightly elongated relative to
Proconsul (distinctly so in Pan), but that hominid ilia are dramatically
shortened. The strong shift made by modern humans largely reflects
enlargement of the acetabulum.
Figure S10. Correlates of the Trans-Iliac Space in
Hominoids. Viewed anteriorly, the posteromedial edges of
right and left ilia and cranial edge of the sacrum form a "U",
the trans-iliac space, indicated here by the horizontal
bracket. It contains and consequently confines the lower
lumbar vertebrae. African apes evidence unique ligaments
between the ilium and these lowest lumbar vertebrae.
Conversely, either broad sacral alae and/or reduction of iliac
height evacuates the trans-iliac space, freeing the lowest
lumbars and enhancing their mobility for lordosis. Specimen
shown here is a female Pan (CMNH-B1758). Gorilla also
exhibits this trait.
Figure S8. Relative Lower Pelvic Height in Anthropoids. Natural log-log scatterplot of
lower iliac height and acetabular height in anthropoids. A regression line (blue: RMA: y =
1.052x + .631; r = .791; N = 145) has been fitted through Old World monkey taxa, and
another of slope equal to one (red) has been fitted through Proconsul (KNM-MW 13142
D). While Gorilla specimens fall around this line, all of those from Pan fall above it,
suggesting that shortening of the back for suspensory locomotion and vertical climbing
has generated spatial compensation by further elongation of the iliac isthmus. Four points
are provided for ARA-VP-6/500 reflecting upper and lower limits for acetabular height and
lower iliac height. These points plot close to Proconsul, suggesting maintenance or slight
shortening of lower iliac height relative to its dimension in the GLCA. The demonstrable
shift from the two early hominids A.L.288-1 and STS-14 to modern humans reflects
several effects on lower iliac height by later pelvic evolution, but especially elongation of
the posterior ilium to increase the sagittal diameter of the birth canal.
Fig. 4. Lateral and posterior CT scan surface renders of (A)
MAK-VP-1/1 (Au. afarensis, cast) and (B) ARA- VP-1/701 (Ar.
ramidus, original). The preserved portion of the ARA-VP-1/701 shaft
is sufficient to demonstrate the absence of a lateral spiral pilaster and
the presence of a distinct rugose insertion area for the gluteus
maximus that is homologous to the true hypotrochanteric fossa
present in MAK-VP-1/1 (arrows indicate the third trochanter; brackets
indicate hypotrochanteric fossae). In ARA-VP-1/701, this area is
more laterally placed, as in other early hominid femora, including
BAR-1002'00. The more posterior position of this insertion in MAK-
VP-1/1 is almost certainly associated with decreased sagittal iliac
orientation in Au. afarensis, a consequence of further posterior pelvic
broadening and increased lateral iliac flare. None of these early
hominid specimens shows evidence of a lateral spiral pilaster, which
is restricted to African ape femora.
Fig. 2. CT comparison of ARA-VP-6/500 (left) os coxa reconstruction
and A.L. 288-1 (right) restoration. The broad sacral alae of ARA-
VP-6/500 are probably because African ape sacra have almost certainly
been narrowed since the GLCA. A novel ossification center for the AIIS and
a substantial reduction in the height of the iliac isthmus are derived
characters present in both hominids. Spatial orientation was made on an
assumption of vertical alignment of the pubic tubercle and ASIS, although
the primitive form of this pelvis suggests that lumbar lordosis during
terrestrial bipedality was partially situational and that the superior pelvis
may have been angled less anteriorly during arboreal clambering. There is
a dramatic reduction of lower pelvic length and robusticity in A.L. 288-1.
Figure 1 | Location map. Satellite (AsTER) image
showing study area at Aramis surrounded by modern
Jara and Awash rivers in Afar, Ethiopia, where
Ardipithecus ramidus skeleton was discovered.
Mixed vegetation (woody plants and
grasses) in floodplain environment
Early hominins inhabited riparian
envi r onment s, l i kel y t aki ng t he
advantage of water-fed ecosystems
developed in a narrow forest corridor
bor der ed by mor e open woody
grasslands
Role of physical environment as a driver
for hominin evolution, including the
development of bipedalism
Conclusions
Pelvis and femur of Ar. ramidus establish that adaptations
to upright walking were found by 4.4 Mya.
Evidence on the locomotor transition from arboreal life to
habitual terrestrial bipedality.