The morphology of the human shoulder seems well-suited to serving its role of providing a wide range of mobility for the upper limb. The dorsal position of a highly mobile scapula and the lateral orientation of the small, shallow glenoid fossa articulating with a medially directed, large round humeral head open up nearly a complete sphere of motion allowing the hand to be positioned almost anywhere around the body. For a creature that depends a great deal on its hands to manipulate objects in order to survive, such an ideal pairing of form and function would seem to be a wonderful example of the elegance of design in nature. Of course the human shoulder did not arrive at its present confi guration purely through natural selection for its current use. Humans share many features of upper limb morphology with extant apes suggesting that at least some aspects of human shoulder anatomy refl ect our phylogenetic heritage. In order to understand which features of the human shoulder have been inherited from a common hominin ancestor, including primitive retentions that continue to serve their original purpose and those that have been co-opted to serve a new function, and which are uniquely derived in humans, it is necessary to determine the course of evolutionary change in the hominin pectoral girdle and shoulder. This paper attempts to trace those changes, beginning with early hominins, but focusing on the characteristics of the shoulder region in early Homo erectus (see Larson, 2007, for a more detailed overview of hominin shoulder evolution).


Shoulder morphology scapula clavicle humerus Homo habilis Homo erectus Homo antecessor 


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.


  1. Alemseged, A., Spoor, F., Kimbel, W.H., Bobe, R. Geraads, D., Reed, D., Wynn, J.G., 2006. A juvenile early hominin skeleton from Dikika, Ethiopia. Nature 443, 296–301CrossRefGoogle Scholar
  2. Andrews, P., 1985. Family group systematics and evolution among catarrhine primates. In: Delson, E. (Ed.), Ancestors: The Hard Evidence. Alan R. Liss, New York, pp. 14–22Google Scholar
  3. Angel, J.L., Kelley, J.O., 1986. Description and comparison of the skeleton. In: Wendorf, F., Schild, R. (Eds.), The Wadi Kubbaniya Skeleton: A Late Paleolithic Burial from Southern Egypt. Southern Methodist University Press, Dallas, pp. 53–70Google Scholar
  4. Atwater, A.E., 1979. Biomechanics of overarm throwing movements and of throwing injuries. Exercise and Sport Sciences Reviews 7, 43–85CrossRefGoogle Scholar
  5. Beals, R.K., 2000. The short clavicle syndrome. Journal of Pediatric Orthopedics 30, 389–391Google Scholar
  6. Beals, R.K., Sauser, D.D., 2006. Nontraumatic disorders of the clavicle. Journal of the American Academy Orthopedic Surgery 14, 205–214Google Scholar
  7. Bermúdez de Castro, J.M., Arsuaga, J.L., Carbonell, E., Rosas, A., Martínez, I., Mosquera, M., 1997. A hominid from the lower Pleistocene of Atapuerca, Spain: possible ancestor to Neandertals and modern humans. Science 276, 1392–1395CrossRefGoogle Scholar
  8. Bingham, P.M., 1999. Human uniqueness: a general theory. Quarterly Review of Biology 74, 133–169CrossRefGoogle Scholar
  9. Bramble, D.M., Lieberman, D.E., 2004. Endurance running and the evolution of Homo. Nature 432, 345–352CrossRefGoogle Scholar
  10. Broom, R., Robinson, J.T., Schepers, G.W.H., 1950. Sterkfontein ape-men Plesianthropus. Transvaal Museum Memoir No. 4Google Scholar
  11. Campbell, B.G., 1966. Human Evolution. Heinemann Ed. Books, LondonGoogle Scholar
  12. Calvin, W.H., 1983. A stone's throw and its launch window: timing and precision and its implications for language and hominid brains. Journal of Theoretical Biology 104, 121–135CrossRefGoogle Scholar
  13. Carretero, J.M., Arsuaga, J.L., Lorenzo, C. 1997. Clavicles, scapulae and humeri from the Sima de los Huesos site (Sierra de Atapuerca, Spain). Journal of Human Evolution 33, 357–408CrossRefGoogle Scholar
  14. Carretero, J.M, Lorenzo, D., Arsuage, J.L., 1999. Axial and appendicu-lar skeleton of Homo antecessor. Journal of Human Evolution 37, 459–499CrossRefGoogle Scholar
  15. Carrier, D.R., 1984. The energetic paradox of human running and hom-inid evolution. Current Anthropology 25, 483–495CrossRefGoogle Scholar
  16. Ciochon, R.L., 1983. Hominoid cladistics and the ancestry of modern apes and humans. In: Corruccini, R.S., Ciochon, R.L. (Eds), New Interpretations of Ape and Human Ancestry. Academic, New York, pp. 783–843CrossRefGoogle Scholar
  17. Ciochon, R.L., Corruccini, R.S., 1976. Shoulder joint of Sterkfontein Australopithecus. South African Journal of Science 72, 80–82Google Scholar
  18. Churchill, S.E., 1994. Human upper body evolution in the Eurasian later Pleistocene. Ph.D. dissertation, University of New Mexico, AlbuquerqueGoogle Scholar
  19. Darlington, P.J., 1975. Group selection, altruism, reinforcement, and throwing in human evolution. Proceeding of the National Academy of Sciences 72, 3748–3752CrossRefGoogle Scholar
  20. Day, M.H., 1978. Hominid postcranial material from Bed I, Olduvai Gorge. In: Isaac, G.L., McCown, E.R. (Eds), Human Origins: Louis Leakey and the East African Evidence. W.A. Benjamin, Menlo Park, pp. 363–374Google Scholar
  21. Dunsworth, H., Challis, J.H., Walker, A., 2003. Throwing and bipedal-ism: a new look at an old idea. Courier Forschungsinstitut Senckenberg 243, 105–110Google Scholar
  22. Edelson G., 1999. Variations in the retroversion of the humeral head. Journal of Shoulder and Elbow Surgery 8, 142–145CrossRefGoogle Scholar
  23. Edelson, G., 2000. The development of humeral head retroversion. Journal of Shoulder and Elbow Surgery 9, 316–318CrossRefGoogle Scholar
  24. Evans, F.G., Krahl, V.E., 1945. The torsion of the humerus: a phylogenetic study from fi sh to man. American Journal of Anatomy 76, 303–337CrossRefGoogle Scholar
  25. Guidera, K.J., Grogan, D.P., Pugh, L., Ogden, J.A., 1991. Hypoplastic clavicles and lateral scapular redirection. Journal of Pediatric Orthopedics 11, 523–526CrossRefGoogle Scholar
  26. Harrison, T., 1986. A reassessment of the phylogenetic relationships of Oreopithecus bambolii Gervias. Journal of Human Evolution 15, 541–583CrossRefGoogle Scholar
  27. Harrison, T., 1987. The phylogenetic relationships of the early catarrhine primates: a review of the current evidence. Journal of Human Evolution 16, 41–80CrossRefGoogle Scholar
  28. Inman, V.T., Saunders, J.B. de C.M., Abbott, L.C., 1944. Observations on the function of the shoulder joint. Journal of Bone and Joint Surgery 26B, 1–30Google Scholar
  29. Inouye, S.E., Shea, B.T., 1997. What's your angle? Size correction and bar-glenoid orientation in “Lucy” (A.L. 288–1). International Journal of Primatology 18, 629–650CrossRefGoogle Scholar
  30. Jellema, L.M., Latimer, B., Walker, A., 1993. The rib cage. In: Leakey, R., Walker, A. (Eds), The Nariokotome Homo erectus Skeleton. Harvard University Press, Cambridge, pp. 294–325CrossRefGoogle Scholar
  31. Johanson, D.C., Lovejoy, C.O., Kimbel, W.H., White, T.D., Ward, S.C., Bush, M.E., Latimer, B.M., Coppens, Y., 1982. Morphology of the Pliocene partial hominid skeleton (A.L. 288–1) from the Hadar Formation, Ethiopia. American Journal of Physical Anthropology 57, 403–451CrossRefGoogle Scholar
  32. Jungers, W.L., 1994. Ape and hominid limb length. Nature 369, 194CrossRefGoogle Scholar
  33. Jungers, W.L., Hartman, S.E., 1988. Relative growth of the locomotor skeleton in orang-utans and other large-bodied hominoids. In: Schwartz, J.E. (Ed.), Orang-utan Biology. Oxford University Press, Oxford, pp. 3473. 59Google Scholar
  34. Jungers, W.L., Falsetti, A.B., Wall, C.E., 1995. Shape, relative size, and size-adjustments in morphometrics. Yearbook of Physical Anthropology 38, 137–161CrossRefGoogle Scholar
  35. Krahl, V.E., 1947. The torsion of the humerus: its localization, cause and duration in man. American Journal of Anatomy 80, 275–319CrossRefGoogle Scholar
  36. Krahl, V.E., Evans, F.G., 1945. Humeral torsion in man. American Journal of Physical Anthropology 3, 229–252CrossRefGoogle Scholar
  37. Larson, S.G., 1988. Subscapularis function in gibbons and chimpanzees: implications for interpretation of humeral head torsion in hom-inoids. American Journal of Physical Anthropology 76, 449–462CrossRefGoogle Scholar
  38. Larson, S.G., 1996. Estimating humeral torsion on incomplete fossil anthropoid humeri. Journal of Human Evolution 31, 239–257CrossRefGoogle Scholar
  39. Larson, S.G., 2007. Evolutionary transformation of the hominin shoulder. Evolutionary Anthropology 16, 172–187CrossRefGoogle Scholar
  40. Leakey, R.E., Walker, A., Ward, C.V., Grausz, H.M., 1989. A partial skeleton of a gracile hominid from the upper Burgi member of the Koobi Fora formation, East Lake Turkana, Kenya. In: Giacobini, G. (Ed.), Hominidae: Proceedings of the 2nd International Congress of Human Paleontology. Jaka Books, Milano, pp. 167–173Google Scholar
  41. Le Gros Clark, W.E., 1959. The Antecedents of Man. Edinburgh University Press, EdinburghGoogle Scholar
  42. Le Gros Clark, W.E., Thomas, D.P., 1951. Associated jaws and limb bones of Limnopithecus macinnesi. Fossil Mammals of Africa, No. 3. British Museum of Natural History, LondonGoogle Scholar
  43. Lordkipanidze, D., Jashashvili, T., Vekua, A., Ponce de León, M.S., Zollikofer, P.E., Rightmire, G.P., Pontzer, H., Ferring R., Oms, O., Tappen, M., Bukhsianidze, M., Agusti, J., Kahlke, R., Kiladze, G., Martinez-Navarro, B., Mouskhelishvili, A., Nioradze, M., Rook, L., 2007. Postcranial evidence form early Homo from Dmanisi, Georgia. Nature 449, 305–310CrossRefGoogle Scholar
  44. Lovejoy, C.O., Johanson, D.C., Coppens, Y., 1982. Hominid upper limb bones recovered from the Hadar formation: 1974–1977 collections. American Journal of Physical Anthropology 57, 637–649CrossRefGoogle Scholar
  45. Marquer, P., 1972. Novelle contribution a l'étude du squelette des pyg-mées occidentaux du Centre Africain comparé a celui des pygmées orientaux. Mémoires du Muséum National D'Histoire Naturelle, Nouvelle Série, Série A, Zoologie 72, 1–122Google Scholar
  46. Martin, L.B., 1986. Relationships among extant and extinct great apes and humans. In: Wood, B.A., Martin, L.B., Andrews, P. (Eds), Major Topics in Primate and Human Evolution. Cambridge University Press, Cambridge, pp. 161–187Google Scholar
  47. Martin, R., Saller, K., 1959. Lerbuch der Anthropologie. Gustav Fischer Verlag, StuttgartGoogle Scholar
  48. McCown, T.D., Keith, A., 1939. The Stone Age of Mount Carmel. The Fossil Human Remains from the Levalloiso-Mousterian. Oxford University Press, OxfordGoogle Scholar
  49. Milgram, J.E., 1942. Congenital forward shoulders (quadrupedal type) – treatment by clavicular osteotomy. Bulletin of the Hospital for Joint Disease 3, 93–96Google Scholar
  50. Mivart, St.G., 1868. On the appendicular skeleton of the primates. Philosophical Transactions of the Royal Society of London 157, 299–426CrossRefGoogle Scholar
  51. Mosimann, J.E., 1970. Size allometry: Size and shape variables with characterizations of the lognormal and generalized gamma distributions. Journal American of the Statistical Association 65, 930–945CrossRefGoogle Scholar
  52. Napier, J.R., 1965. Comment on ‘New Discoveries in Tanganyika: Their Bearing on Hominid Evolution by P.V. Tobias.’ Current Anthropology 5, 402–403Google Scholar
  53. Ohman, J.C., 1986. The fi rst rib of hominoids. American Journal of Physical Anthropology 70, 209–229CrossRefGoogle Scholar
  54. Oxnard, C.E., 1968. A note on the fragmentary Sterkfontein scapula. American Journal of Physical Anthropology 28, 213–217CrossRefGoogle Scholar
  55. Oxnard, C.E., 1969. A note on the Olduvai clavicular fragment. American Journal of Physical Anthropology 29, 429–432CrossRefGoogle Scholar
  56. Partridge, T.C., Granger, D.E., Caffee, M.W., Clarke, R.J., 2003. Lower Pliocene hominid remains from Sterkfontein. Science 300, 607–612CrossRefGoogle Scholar
  57. Perry, J., 1983. Anatomy and biomechanics of the shoulder in throwing, swimming, gymnastics, and tennis. Clinical Sports Medicine 2, 247–270Google Scholar
  58. Pickford, M., Johanson, D.C., Lovejoy, C.O., White, T.D., Aronson, J.L., 1983. A hominoid humeral fragment from the Pliocene of Kenya. American Journal of Physical Anthropology 60, 337–346CrossRefGoogle Scholar
  59. Roberts, D., 1974. Structure and function of the primate scapula. In: Jenkins, F.A. (Ed.), Primate Locomotion. Academic, New York, pp. 171–200Google Scholar
  60. Robinson, J.T., 1972. Early Hominid Posture and Locomotion. University of Chicago Press, Chicago, ILGoogle Scholar
  61. Schmid, P., 1983. Eine Rekonstruktion des Skelettes von A.L. 288-1 (Hadar) und deren Konsequenzen. Folia Primatologica 40, 283–306CrossRefGoogle Scholar
  62. Schultz, A.H., 1930. The skeleton of the trunk and limbs of higher primates. Human Biology 2, 303–438Google Scholar
  63. Schultz, A.H., 1937. Proportions, variability and asymmetries of the long bones of the limbs and the clavicles in man and apes. Human Biology 9, 281–328Google Scholar
  64. Sládek, V., Trinkuas, E., Hillson, S.W., Holliday, T.W., 2000. The People of the Pavlovian. Skeletal Catalogue and Osteometrics of the Gravettian Fossil Hominids from Dolní Věstonice and Pavlov. Academy of Sciences of the Czech Republic, BrnoGoogle Scholar
  65. Smith, B.H., 1993. The physiological age of KNM-WT-15000. In: Leakey, R., Walker, A. (Eds), The Nariokotome Homo erectus Skeleton. Harvard University Press, Cambridge, MA, pp. 195–233CrossRefGoogle Scholar
  66. Stern, J.T., Susman, R.L., 1983. The locomotor anatomy of Australopithecus afarensis. American Journal of Physical Anthropology 60, 279–317CrossRefGoogle Scholar
  67. Trinkaus, E., 1983. The Shanidar Neandertals. Academic, New YorkGoogle Scholar
  68. Tullos, H.S., King, J.W., 1973. Throwing mechanisms in sports. Orthopedic Clinics of North America 4, 709–720Google Scholar
  69. Voisin, J.-L., 2006. Clavicle, a neglected bone: morphology and relation to arm movements and shoulder architecture in primates. Anatomical Record 288A, 944–953Google Scholar
  70. Vrba, E.S., 1979. A new study of the scapula of Australopithecus africanus from Sterkfontein. American Journal of Physical Anthropology 51, 117–130CrossRefGoogle Scholar
  71. Walker, A., Leakey, R., 1993. The postcranial bones. In: Leakey, R., Walker, A. (Eds), The Nariokotome Homo erectus Skeleton. Harvard University Press, Cambridge, MA, pp. 95–160CrossRefGoogle Scholar
  72. Zapfe, H., 1960. Die Primatenfunde aus der miozänen Spaltenfüllung von Neudorf an der March (Děvínská Nová Ves), Tschechoslowakei. Schweizerische Palaeontologische Abhandlungen 78, 1–293Google Scholar

Copyright information

© Springer-Verlag Berlin Heidelberg 2009

Authors and Affiliations

  • Susan G. Larson

There are no affiliations available

Personalised recommendations