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C. O. Lovejoy, in Primate Morphology and Evolution, R. H. Tuttle, Ed. (Mouton, The Hague, 1975), pp. 291-326.
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Lovejoy, C.O.1
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4
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70349663832
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note
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The trait nomenclature system used here is taken from (47, 54) and is briefly as follows [for more complete explanations, see (2)]. Type 1: traits whose morphogenesis is the direct consequence of pattern formation; usually (but not always) subject to direct selection. Type 2: traits that are genetic but are pleiotropic to, or result from hitchhiking on, type 1 traits and are not themselves subject to selection [2A: parent type 1 is inferred to be under selection but its secondary effects are not; 2B: neither parent trait nor derivative is inferred to be under selection (rare)]. Type 3: resulting from a systemic growth factor. Type 4: cpigenctic consequence of osteochondral remodeling and/or response to environmental stimuli, i.e., not heritable but useful in interpreting behavior. Type 5: similar to type 4, but uninformative.
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5
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0002236234
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R. L. Ciochon, R. S. Corruccini, Eds. (Plenum, New York)
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A. C. Walker, M. Pickford, in New Interpretations of Ape and Human Ancestry, R. L. Ciochon, R. S. Corruccini, Eds. (Plenum, New York, 1983), pp. 325-413.
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(1983)
New Interpretations of Ape and Human Ancestry
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Walker, A.C.1
Pickford, M.2
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70349675302
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C. O. Lovejoy et al., Science 326, 71 (2009).
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Science
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Lovejoy, C.O.1
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70349693767
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note
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Schultz noted that the medial cuneiform-Mt1 joint is "directed more forward and less sidewise in the mountain gorilla than in the other apes" [(20), p. 395]. Our sample contained only "western" specimens (G. g. gorilla).
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22
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70349663833
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M. E. Baratz, A. D. Watson, J. E. Imbriglia, Eds. (Thieme, New York)
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J. T. Campbell, L. C. Schon, in Orthopaedic Surgery: The Essentials, M. E. Baratz, A. D. Watson, J. E. Imbriglia, Eds. (Thieme, New York, 1999), pp. 591-614.
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Orthopaedic Surgery: The Essentials
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Campbell, J.T.1
Schon, L.C.2
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70349693766
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thesis, University of Michigan
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J. M. DeSilva, thesis, University of Michigan (2008).
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(2008)
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DeSilva, J.M.1
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27
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0347296385
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E. Vereecke, K. D'Aout, D. De Clercq, L. Van Elsacker, P. Aerts, Am. J. Phys. Anthropol. 120, 373 (2003).
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Vereecke, E.1
D'aout, K.2
De Clercq, D.3
Van Elsacker, L.4
Aerts, P.5
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70349668594
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In two Pan specimens dissected for this paper, a small cartilage nodule (invisible upon x-ray) could be palpated within the tendon but had no effect on its caliber, nor was either accompanied by a facet. This accounts for reports by some earlier anatomists that the sesamoid is present in African apes (although it is clearly present in gibbons, which exhibit regular facets). A recent review (55) concluded an incidence of 2/11 for the os peroneum in apes from classical literature. However, this datum is likely unreliable, because many early authors did not inspect the tendon closely and typically reported its structure "as in humans," whereas "the few anatomists who have explicitly looked for the [os peroneum] have noted its absence in Gorilla..., Pan troglodytes..., and... Pongopygmaeus" [(30), p. 93]. Examination of the cuboid is the most reliable standard because a functional os peroneum cannot obtain without an underlying facet, just as the presence of a clearly functional facet assures the ossicle's presence, whether or not it was calcified. In rare cases, a facet-like discoloration of the bottom of the fibularis longus groove in Pan can be seen. However, the depth and cylindrical nature of the groove make such facets (if in fact they supported an os peroneum during life) largely nonfunctional, and we here report only clearly functional facets. One such human-like facet was found in a Pan specimen in our survey. Interestingly, its associated Mt4 and Mt5 facets suggested hypermobility at these joints far in excess of other Pan specimens. This supports the argument that relocation of the tendon, as in humans, does in fact reduce general midtarsal mobility.
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70349684471
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In primates, a fascial sheet spans the plantar aspect of the foot from calcaneus to cuboid, lateral cuneiform, and lateral metatarsal bases, and underlies the fibularis longus tendon in its course to the first ray. In humans this sheet is described as two separate elements, the short and long plantar ligaments. However, in apes the fascial sheet bridges the cuboidal groove, transforming it into a tunnel confining the fibularis longus tendon (56, 57). Old World monkeys have a rough equivalent but retain an os peroneum lateral to the tunnel. In humans, the fibularis longus tendon lies outside (plantar to) the cuboidal groove, and the short plantar ligament terminates proximal to it. A second portion of the sheet, which lies plantar to the tendon, spans it and inserts distally on the lateral metatarsal bases. In humans this is distinguished as the long plantar ligament (57, 58). These human divisions of the plantar sheet are therefore likely derived (13).
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70349668593
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J. P. Schaeffer, Ed. (Blakiston, Philadelphia)
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R. J. Terry, in Morris' Human Anatomy, J. P. Schaeffer, Ed. (Blakiston, Philadelphia, 1942), pp. 266-376.
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(1942)
Morris' Human Anatomy
, pp. 266-376
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Terry, R.J.1
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70349668595
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note
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We base this simply on torsion of the bone's diaphysis because its head was not preserved. We have not provided a numerical value because estimating the articular head's dorsoplantar axis in near-adult (but still unfused) African ape Mt2 growth plates provides only a broadly reliable estimate of the exact angle between the proximal and distal joint axes when the (unfused) head is articulated on its growth plate.
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35348854422
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X. Chen, C. Macica, A. Nasiri, S. Judex, A. E. Broadus, Bone 41, 752 (2007).
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(2007)
Bone
, vol.41
, pp. 752
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Chen, X.1
Macica, C.2
Nasiri, A.3
Judex, S.4
Broadus, A.E.5
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38
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34249815809
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P. L. Reno, W. E. Horton Jr., R. M. Elsey, C. O. Lovejoy, J. Exp. Zool. B 308, 283 (2007).
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J. Exp. Zool. B
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Reno, P.L.1
Horton Jr., W.E.2
Elsey, R.M.3
Lovejoy, C.O.4
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70349655353
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note
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Considerable debate has centered around the importance of phalangeal shaft curvature in Au. afarensis, largely with respect to its etiology. If developmentally plastic [cartilage modeling; type 4 (4)], then curvature may imply active grasping history. However, if a direct product (or pleiotropic effect) of positional information [types 1 or 2 (4)], then it can be moot with respect to phalangeal function in rapidly evolving species. The fact that curvature increases during maturation does not resolve this issue because increasing curvature during growth is as explicable by positional information as it is by hypothetical strain regiments). Current evidence, such as the phalangeal curvature in great ape fetuses, supports a type 1 status. An important element that has long been ignored is phalangeal morphology in Au. afarensis itself (A.L. 333-115); see fig. S11.
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41
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0346397925
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M. Nakatsukasa, Y. Kunimatsu, Y. Nakano, T. Takano, H. Ishida, Primates 44, 371 (2003).
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(2003)
Primates
, vol.44
, pp. 371
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Nakatsukasa, M.1
Kunimatsu, Y.2
Nakano, Y.3
Takano, T.4
Ishida, H.5
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70349658844
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C. O. Lovejoy et al., Science 326, 70 (2009).
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(2009)
Science
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Lovejoy, C.O.1
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70349661542
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note
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STW-573 ("Little Foot") preserves a talus, navicular, medial cuneiform, and Mt1. Its first ray has been described as partially abducent (17,18). Replacement of its talus and/or navicular with those of cither a chimpanzee or human would have had no substantial effect on first-ray abducence; they are not informative. The higher primate medial cuneiform itself is not mobile and is therefore equally uninformative, save for its Mt1 joint surface. This surface faces distally, is virtually flat, and is therefore immobile in STW-573. The claim that "the first metatarsal facet overflows from distal to proximal, as in apes" [see footnote 18 of (17)] is incorrect, and the dotted line drawn to indicate the proximal extent of the facet in their figure 3B is exaggerated. The joint surface of STW-573 is virtually identical to that of OH-8 and unlike that of any ape. This alone falsifies the contention that the specimen's hallux was abducent. Indeed, the fibularis longus insertion of STW-573 is described as spanning both the medial cuneiform and Mt1 and therefore cannot have adducted the Mt1 even if it retained any mobility in the first tarsometatarsal joint.
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47
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0036785121
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C. O. Lovejoy, R. S. Meindl, J. C. Ohman, K. G. Heiple, T. D. White, Am. J. Phys. Anthropol. 119, 97 (2002).
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Lovejoy, C.O.1
Meindl, R.S.2
Ohman, J.C.3
Heiple, K.G.4
White, T.D.5
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50
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70349672963
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note
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The plantaris is "normally present in prosimians, monkeys, and man, but is lacking in gibbons and gorillas, nearly all orang-utans, and a considerable percentage of chimpanzees" (53).
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52
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70349653138
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Interpretation of structure frequency requires constant attention to the selective mechanism in play. Narrowly defined (i.e., named) anatomical structures (e.g., long and short plantar ligaments) emerge from mesenchymal fields, which are manifestations of their parent positional information. It is therefore field configuration that is the target of selection. Constant structures within a species (e.g., the Achilles' tendon) suggest low field variance and intense stabilizing selection. Conversely, substantially reduced frequencies of a "named" tissue mass within a species (e.g., the quadratus plantae in Pan) signal the occurrence of underlying field shifts. These may emanate from selective encouragement of changes in target structures that share field commonality with the reduced structure, simple relaxation of selection, selection against the structure, or some combination of these.
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53
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0004064948
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Williams and Wilkins, Baltimore, ed.
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C. G. Hartman, W. L. Straus, The Anatomy of the Rhesus Monkey (Williams and Wilkins, Baltimore, ed. 1, 1933), pp. 1-383.
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The Anatomy of the Rhesus Monkey
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Hartman, C.G.1
Straus, W.L.2
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V. K. Sarin, G. M. Erickson, N. J. Giori, A. G. Bergman, D. R. Carter, Anat. Rec. 257, 174 (1999).
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Anat. Rec.
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Sarin, V.K.1
Erickson, G.M.2
Giori, N.J.3
Bergman, A.G.4
Carter, D.R.5
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58
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70349678374
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Lea and Febiger, Philadelphia, ed.
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H. Gray, Gray's Anatomy (Lea and Febiger, Philadelphia, ed. 26, 2008).
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Gray's Anatomy
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Gray, H.1
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59
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70349659307
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note
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Supported by NSF grants 8210897, 9318698, 9512534, 9632389, 9729060, 9910344, and 0321893 HOMINIDRHOI, and by the Japan Society for the Promotion of Science. We thank the Ministry of Tourism and Culture, the Authority for Research and Conservation of the Cultural Heritage, and the National Museum of Ethiopia for permissions and facilitation; the Afar Regional Government, the Afar people of the Middle Awash, and many other field workers for contributing directly to the data; the National Museum of Ethiopia, National Museums of Kenya, Transvaal Museum South Africa, Cleveland Museum of Natural History, Royal Museum of Central Africa Tervuren for access to comparative materials; L Spurlock and P. L Reno for assistance with dissections and histology preparations; D. Kubo and H. Fukase for assistance in computed tomography (CT) scanning; C. Hernandez for calculation of section modulus data for Mohr's Circle analyses; M. Brunet, C. V. Ward, and J. DeSilva for cooperation with comparative data; R. Meindl for statistical advice and assistance; J. DeSilva, P. L. Reno, M. A. Serrât, M. A. McCollum, M. Selby, A. Ruth, L. Jellema, S. W. Simpson, and B. A. Rosenman for aid in data collection and exceptionally helpful discussions; and H. Gilbert, J. Carlson, and K. Brudvik for figure preparation.
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