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    Science 2 October 2009: vol. 326 no. 5949 pp. 71-71e6DOI: 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?

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    Fig. 1. Original and reconstructed os coxa of ARA-VP-6/500.(Left)Anterolateral and anteromedial viewsof the original, with a close-up of the AIIS. (Middle) CT

    scans of same views, except the bottom view is aclose-up of the pubic symphyseal face, preserved in

    its entirety but damaged at its inferior extremity.Arrows indicate major areas of distortion corrected byreconstruction. (Right)Anterolateral and anteromedial

    views of the reconstructed os coxa and of the entirepelvis 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

    with a broad, short and

    sagitally disposed iliac isthmus

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    Figure S4. Craniocaudal Pelvic Height and Pelvic Geometry inHominoids. 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 ischialspine (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.

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    (D) Minimum ischial length (frominferior acetabular border to juncture

    of the ischial body and tuberosity

    surface) normalized by acetabular

    diameter. Two values for ARA-

    VP-6/500 bracket possible extremesfor the acetabular diameter. Pan-

    Homo d i f ferences are h igh ly

    significant.

    (C) Angle made by two chordsconnecting three landmarks on the

    ilium. The third chord was measured

    (auricular surface to the ischial spine),

    and the angle was determined

    trigonometrically. All early hominids liewithin the human range because all

    have a greatly shortened iliac isthmus.

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    Figure S5. Relative lateral Iliac Breadth in Anthropoids.

    Natural log-log scatterplot of lateral iliac breadth (distance fromthe 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.

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    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, andanother of slope equal to one (red) has been fitted through Proconsul (KNM-MW 13142D). 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 climbinghas generated spatial compensation by further elongation of the iliac isthmus. Four pointsare provided forARA-VP-6/500 reflecting upper and lower limits for acetabular height andlower iliac height. These points plot close to Proconsul, suggesting maintenance or slightshortening of lower iliac height relative to its dimension in the GLCA. The demonstrableshift from the two early hominids A.L.288-1 and STS-14 to modern humans reflectsseveral effects on lower iliac height by later pelvic evolution, but especially elongation ofthe posterior ilium to increase the sagittal diameter of the birth canal.

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    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 inAu. 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.

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    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

    environments, l ikely taking the

    advantage of water-fed ecosystems

    developed in a narrow forest corridorbordered by more open woody

    grasslands

    Role of physical environment as a driver

    for hominin evolution, including the

    development of bipedalism

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    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.