A comparative analysis of the hominin triquetrum (SKX 3498) from Swartkrans, South Africa

  • Tracy L. Kivell Department of Human Evolution, Max Planck Institute of Evolutionary Anthropology
Keywords: carpal, hominoid, wrist, Paranthropus, Homo


The SKX 3498 triquetrum from Member 2 at Swartkrans Cave, South Africa is the only hominin triquetrum uncovered (and published) thus far from the early Pleistocene hominin fossil record. Although SKX 3498 was found over two decades ago, its morphology has not been formally described or analysed, apart from the initial description. Furthermore, the taxonomic attribution of this fossil remains ambiguous as both Paranthropus and early Homo have been identified at Swartkrans. This analysis provides the first quantitative analysis of the SKX 3498 triquetrum, in comparison to those of extant hominids (humans and other great apes) and other fossil hominins. Although the initial description of the SKX 3498 triquetrum summarised the morphology as generally human-like, this analysis reveals that quantitatively it is often similar to the triquetra of all hominine taxa and not necessarily humans in particular. Shared hominid-like morphology between SKX 3498 and Neanderthals suggests that both may retain the symplesiomorphic hominin form, but that functional differences compared to modern humans may be subtle. Without knowledge of triquetrum morphology typical of earlier Pliocene hominins, the taxonomic affiliation of SKX 3498 remains unclear.

Author Biography

Tracy L. Kivell, Department of Human Evolution, Max Planck Institute of Evolutionary Anthropology
Junior Researcher


1. Brain CK. Structure and stratigraphy of the Swartkrans cave in light of the new excavations. In: Brain CK, editor. Swartkrans: A cave’s chronicle of early man. Pretoria: Transvaal Museum, 1993; p. 23–34.

2. Susman RL. New postcranial remains from Swartkrans and their bearing on the functional morphology and behaviour of Paranthropus robustus. In: Grine FE, editor. Evolutionary history of the ‘robust’ australopithecines. New York: Aldine de Gruyter, 1988; p. 149–172.

3. Susman RL. New hominid fossils from the Swartkrans formation (1979–1986 excavations): Postcranial specimens. Am J Phys Anthropol. 1989;79:451–474. doi:10.1002/ajpa.1330790403, PMid:2672829

4. Tocheri MW, Orr CM, Jacofsky MC, Marzke MW. The evolutionary history of the hominin hand since the last common ancestor of Pan and Homo. J Anat. 2008;212:544–562. doi:10.1111/j.1469-7580.2008.00865.x2672829, PMid:18380869, PMid:2409097

5. Susman RL, de Ruiter D, Brain CK. Recently identified postcranial remains of Paranthropus and early Homo from Swartkrans Cave, South Africa. J Hum Evol. 2001;41:607–629. doi:10.1006/jhev.2001.0510, PMid:11782111

6. Lovejoy CO, Simpson SW, White TD, Asfaw B, Suwa G. Careful climbing in the Miocene: The forelimbs of Ardipithecus ramidus and humans are primitive. Science. 2009;326:70e1–70e8.

7. Trinkaus E. The Shanidar Neandertals. New York: Academic Press; 1983.

8. Sládek V, Trinkaus E, Hillson SW, Holliday TW. The people of the Pavlovian: Skeletal catalogue and osteometrics of the Gravettian fossil hominids from Dolní Vĕstonice and Pavlov. Dolní Vĕstonice Studies 5. Brno: Akademie vĕd Česke republicky; 2000.

9. Jungers WL, Falsetti AB, Wall CE. Shape, relative size and size-adjustments in morphometrics. Yrbk Phys Anthropol. 1995;38:137–161. doi:10.1002/ajpa.1330380608

10. Mossiman JE. Size allometry: Size and shape variables with characterizations of the lognormal and generalized gamma distributions. J Am Stat Assoc. 1970;56:930–945.

11. Darroch JN, Mosimann JE. Canonical and principal components of shape. Biometrika. 1985;72:241–252. doi:10.1093/biomet/72.2.241

12. Coleman MN. What does geometric mean, mean geometrically? Assessing the utility of geometric mean and other size variables in studies of skull allometry. Am J Phys Anthropol. 2007;135:404–415. doi:10.1002/ajpa.20761, PMid:18067122

13. Quinn GP, Keough MJ. Experimental design and data analysis for biologists. Cambridge: Cambridge University Press; 2002.

14. Sanchez PM. The unequal group size problem in discriminant analysis. J Acad Marketing Sci. 1974;102:629–633. doi:10.1007/BF02729456

15. Taleisnik J. The wrist. New York: Churchill Livingstone; 1985.

16. Hamrick MW. Functional osteology of the primate carpus with special reference to Strepsirhini. Am J Phys Anthropol. 1997;104:105–116. doi:10.1002/(SICI)1096-8644(199709)104:1<105::AID-AJPA7>3.0.CO;2-Q

17. Bush ME, Lovejoy CO, Johanson DC, Coppens Y. Hominid carpal, metacarpal and phalangeal bones recovered from the Hadar Formation: 1974–1977 collections. Am J Phys Anthropol. 1982;57:651–677. doi:10.1002/ajpa.1330570410

18. Marzke MW. Joint functions and grips of the Australopithecus afarensis hand, with special reference to the region of the capitate. J Hum Evol. 1983;12:197–211. doi:10.1016/S0047-2484(83)80025-6

19. Feipel V, Rooze M, Louryan S, Lemort M. Functional anatomy of the carpus in flexion and extension and in radial and ulnar deviations: An in vivo two- and three-dimensional CT study. In: Schuind F, An K-N, Cooney WP III, Garcia-Elias M, editors. Advances in the biomechanics of the hand and wrist. New York: Plenum Press, 1994; p. 255–270.

20. Moritomo H, Goto A, Yoshinobu S, Sugamoto K, Murase T, Yoshikawa H. The triquetrum-hamate joint: An anatomic and in vivo three-dimensional kinematic study. J Hand Surg. 2003;28A(5):797–805.

21. Tuttle RH. Terrestrial trends in the hands of the Anthropoidea. Proceedings of the 2nd International Congress of Primatology; 1968; Atlanta, GA, USA. Basel: S. Karger; 1969. p. 192–200.

22. Ryu J, Palmer AK, Cooney WP III. Wrist joint motion. In: An K-N, Berger RA, Cooney WP III, editors. Biomechanics of the wrist joint. New York: Spinger-Verlag, 1991; p. 37–60.

23. Clarke RJ. Discovery of complete arm and hand of the 3.3 million-year-old Australopithecus skeleton from Sterkfontein. S Afr J Sci. 1999;95:477–480.

24. Clarke RJ. Latest information on Sterkfontein’s Australopithecus skeleton and a new look at Australopithecus. S Afr J Sci. 2008;104:443–449. doi:10.1590/S0038-23532008000600015

25. Pickering R, Kramers JD. Re-appraisal of the stratigraphy and determination of new U-Pb dates for the S terkfontein hominin site, South Africa. J Hum Evol. 2010;59:70–86. doi:10.1016/j.jhevol.2010.03.014, PMid:20605190

26. Berger LR, DeRuiter DJ, Churchill SE, et al. Australopithecus sediba: A new species of Homo-like australopith from South Africa. Science. 2010;328:195–204. doi:10.1126/science.1184944, PMid:20378811

27. Dirks PHGM, Kibii JM, Kuhn BF, et al. Geological setting and age of Australopithecus sediba from Southern Africa. Science. 2010;328:205–208. doi:10.1126/science.1184950, PMid:20378812

28. Niewoehner WA, Weaver AH, Trinkaus E. Neandertal capitate-metacarpal articular morphology. Am J Phys Anthropol. 1997;103:219–233. doi:10.1002/(SICI)1096-8644(199706)103:2<219::AID-AJPA7>3.0.CO;2-O

29. Ricklan DE. Functional anatomy of the hand of Australopithecus africanus. J Hum Evol. 1987;16:643–664. doi:10.1016/0047-2484(87)90018-2

30. Marzke MW. Precision grips, hand morphology and tools. Am J Phys Anthropol. 1997;102:91–110. doi:10.1002/(SICI)1096-8644(199701)102:1<91::AID-AJPA8>3.0.CO;2-G

31. Marzke MW, Wullsetin KL, Viegas SF. Evolution of the power (squeeze grip) and its morphological correlates in hominids. Am J Phys Anthropol. 1992;89:283–298. doi:10.1002/ajpa.1330890303, PMid:1485637

32. Marzke MW, Marzke RF. Evolution of the human hand: Approaches to acquiring, analysing and interpreting the anatomical evidence. J Anat. 2000;197:121–140. doi:10.1046/j.1469-7580.2000.19710121.x, PMid:10999274, PMid:1468111

33. Niewoehner WA. The functional anatomy of late Pleistocene and recent human carpometacarpal and metacarpophalangeal articulations. PhD thesis, Albuquerque, University of New Mexico, 2000.

34. Niewoehner WA. Behavioral inferences from the Skhul/Qafzeh early modern human hand remains. Proc Natl Acad Sci. 2001;98(6):2979–2984. doi:10.1073/pnas.041588898, PMid:11248017, PMid:30592

35. Tocheri MW, Marzke MW, Liu D, et al. Functional capabilities of modern an fossil hominid hands: Three-dimensional analysis of trapezia. Am J Phys Anthropol. 2003;122:101–112. doi:10.1002/ajpa.10235, PMid:12949830

36. Williams EM, Gordon AD, Richmond BG. Upper limb kinematics and the role of the wrist during stone tool production. Am J Phys Anthropol. 2010;143:134–145. doi:10.1002/ajpa.21302, PMid:20734439

37. Marzke MW, Toth N, Schick K, et al. EMG study of hand muscle recruitment during hard hammer percussion manufacture of Oldowan tools. Am J Phys Anthropol. 1998;105:315–332. doi:10.1002/(SICI)1096-8644(199803)105:3<315::AID-AJPA3>3.0.CO;2-Q

38. Willams EM, Richmond BG. Hand pressure during Oldowan stone tool production. Am J Phys Anthropol. 2010;141:243.

39. Solecki RS. Shanidar IV, a Neanderthal flower burial in northern Iraq. Science. 1975;190:880–881.

40. Sarmiento EE. Comment on the palaeobiology and classification of Ardipithecus ramidus. Science. 2010;328:1105b. doi:10.1126/science.1184148, PMid:20508113