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The Natural Endocast of Taung (Australopithecus Africanus): Insights from the Unpublished Papers of Raymond Arthur Dart Dean Falk*

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The Natural Endocast of Taung (Australopithecus Africanus): Insights from the Unpublished Papers of Raymond Arthur Dart Dean Falk* YEARBOOK OF PHYSICAL ANTHROPOLOGY 52:49–65 (2009) The Natural Endocast of Taung (Australopithecus africanus): Insights From the Unpublished Papers of Raymond Arthur Dart Dean Falk* Department of Anthropology, Florida State University, Tallahassee, FL 32306-7772 KEY WORDS Australopithecus africanus; Brain; Raymond Dart; Endocast; Arthur Keith; Lunate sulcus; Grafton Elliot Smith; Taung ABSTRACT Dart’s 1925 announcement of Australo- man (1931)), Schepers (Schepers: The endocranial casts pithecus africanus (Dart: Nature 115 [1925] 195–199) of the South African ape-men. In: Broom R, Schepers was highly controversial, partly because of an interpre- GWH, editors. The South African fossil ape-men; the tation of the Taung natural endocast that rested on an Australopithecinae [1946] p 155–272), and Falk (Falk: erroneous identification of the lambdoid suture as the Am J Phys Anthropol 53 [1980] 525–539), and the lunate sulcus. Unpublished materials from the Univer- thorny issue of the location of the lunate sulcus is sity of Witwatersrand Archives (Dart, unpublished revisited in light of new information. Archival materi- material) reveal that Dart reacted to the controversy als reveal that Dart believed that Taung’s brain was by: 1) describing and illustrating the entire sulcal pat- reorganized globally rather than in a mosaic manner, tern on the Taung endocast, in contrast to just two and that the shapes of certain cortical association sulcal identifications in 1925, 2) identifying a hypothet- areas showed that Australopithecus was closer to ical part of the lambdoid suture and revising his Pithecanthropus than to the living apes. Although a description of the lunate sulcus, and 3) bolstering his few of Dart’s hitherto-unpublished sulcal identifica- argument that Taung’s brain was advanced by detail- tions, including his revision for the lunate sulcus, were ing expansions in three significant cortical association questionable, his claim that the Taung endocast repro- areas. Four unpublished illustrations of Dart’s identifi- duced a shape that was advanced toward a human cations for sulci and sutures on the Taung endocast condition in its prefrontal cortex and caudally pro- are compared here with those published by Keith truded occipital lobe was correct. Yrbk Phys Anthropol (Keith: New discoveries relating to the antiquity of 52:49–65, 2009. VC 2009 Wiley-Liss, Inc. Raymond Dart’s description of the fossilized face, jaw, colleagues recognized that his conclusion rested heavily and associated natural endocast from Taungs (now on a questionable identification of the lunate sulcus Taung) and his naming of Australopithecus africanus (Keith, 1931; Clark et al., 1936). Although skeptics even- (Dart, 1925) were received with skepticism by British tually accepted the importance of Dart’s discovery, as anatomists, which contributed to the fact that it was well as the fact that certain shape features of Taung’s decades before the importance of his discovery was endocast were, indeed, derived, few of them ever accepted (Tobias, 1984, 1992, 1996, 2008) (see Fig. 1). accepted Dart’s specific identification of lb as L (Clark Dart had identified what he thought was a posteriorly et al., 1936; Clark, 1947). (caudally) positioned lunate sulcus (L) on the Taung In 1980, I published an independent analysis of the endocast (Fig. 2a). This deep crescent-shaped sulcus sulcal patterns reproduced on the Taung endocast and approximates the rostral boundary of primary visual six other australopithecine natural endocasts, and con- (striate) cortex (Brodmann’s area [BA] 17) in monkeys cluded that they appeared apelike rather than human- and apes, and is located much further forward (more ros- like (Falk, 1980) as others had suggested (Schepers, trally) on their brains than is the case for lunate sulci 1946; Holloway, 1975). Ralph Holloway critically exam- that have been identified for humans (Connolly, 1950). ined my identifications of sulci on Taung, and stated Because of this, it had long been thought that L was dis- that in his ‘‘estimation, almost none can be identified placed caudally as the brain’s parietotemporo-occipital with any certitude’’ (Holloway, 1981, p 51). Having association cortices enlarged during the course of championed Dart’s interpretation of a caudal position for hominin evolution (Smith, 1903, 1904a,b, 1927). Dart’s L in Taung (Holloway, 1975), Holloway took particular identification of L in a humanlike position on the Taung exception to my identification of a relatively rostral endocast was surprising because of its small apelike vol- depression on Taung’s endocast as the likely medial end ume, and it was the main reason he concluded that Taung’s brain had been derived toward a humanlike *Correspondence to: Dean Falk, Department of Anthropology, condition (Dart, 1925). Florida State University, Tallahassee, FL 32306-7772, USA. Dart’s description of Taung’s endocast as appearing E-mail: [email protected] humanlike despite its small apelike size contributed to the initial skepticism of scientists, including Arthur DOI 10.1002/ajpa.21184 Keith (1925, 1931), Grafton Elliot Smith (1925), and Published online in Wiley InterScience Arthur Smith Woodward (1925). Some of Dart’s (www.interscience.wiley.com). VC 2009 WILEY-LISS, INC. 50 D. FALK Fig. 1. Phillip Tobias elucidated the academic politics between the British scientific establishment and Raymond Dart that were one of the reasons it took decades for the importance of Australopithecus africanus to become recognized (Tobias, 1984, 1996). Here, Tobias is shown displaying the original Taung fossil in the Wits Anatomy Department. Photographs courtesy of Harry Jerison. [Color figure can be viewed in the online issue, which is available at www.interscience.wiley.com.] Fig. 2. The Taung fossil. (a) Dart’s 1925 illustration of the Taung endocast, in which he identified only two sulci: the superior temporal (parallel) sulcus; and a caudally positioned lunate sulcus that was actually the lambdoid suture (lb). Modified from Dart, 1925). (b) A copy of the Taung endocast that includes its right temporal pole (tp), which was restored by Ron Clarke by attaching a cast made from its fossilized impression in the back of Taung’s face (Falk and Clarke, 2007). The lambdoid (lb) and coronal (co) sutures, and an adhering bony chip are labeled. Notice, also, damaged (rough) areas and meningeal blood vessel (m). The chip, sutures, and vessels are superficial to other features. [Color figure can be viewed in the online issue, which is available at www. interscience.wiley.com.] of that sulcus (Holloway, 1981). Subsequently, we enlargement (Falk, 1985b). As will become clear, the dif- debated the position of L on a number of australopithe- ference between these two views has profound implica- cine endocasts, and neither of us changed our general tions for unraveling the evolutionary dynamics between views about australopithecine sulcal patterns (Holloway, changes in brain shape and altered sulcal patterns. 1983, 1984, 1985, 1988, 1991; Holloway and Kimbel, Since Dart’s initial description of the Taung endocast, 1986; Holloway et al., 2001, 2004; Falk, 1983, 1985a,b, new information has come to light including identifica- 1986, 1989, 1991; Falk et al., 1989). tion of its enlarged occipital-marginal (O/M) sinus The reader may wonder why so much attention has (Tobias and Falk, 1988), restoration of its missing right been given to the lunate sulcus. Determining the loca- temporal pole (Fig. 2b) (Falk and Clarke, 2007), identifi- tion of L for Taung and other small-brained early homi- cation of bony morphology adhering to its basal surface nins is highly relevant for understanding human brain (Falk and Clarke, 2007; Fig. 1), and a revised (reduced) evolution. On the one hand, a caudally located L would cranial capacity estimate of 382 cm3 with an estimated provide fossil evidence for the long-held view that as adult value of 406 cm3 (Falk and Clarke, 2007) in place brains evolved, the lunate sulcus was displaced caudally of earlier juvenile and adult estimates of 405 cm3 due to expansion of the bordering parieto-occipital asso- and440 cm3 (Holloway, 1970). ciation cortex (Smith, 1904a). It would also imply that In July–August, 2008, I visited the University of Wit- changes in the arrangement of sulci preceded brain size watersrand Archives in Johannesburg, South Africa, and increase (Holloway, 1983). On the other hand, a rostral gained access to a long unpublished manuscript of Dart’s apelike position of L in australopithecines would suggest that was completed in 1929, in which he provided exten- that changes in sulcal patterns did not precede brain sive details about Taung’s endocast (Dart, unpublished Yearbook of Physical Anthropology NATURAL ENDOCAST OF TAUNG 51 material). Tobias (1993) had reported on his recovery Taung’s endocast was, ‘‘You refer to the ‘failure of the and rescue of this manuscript in 1951. I also studied endocranial cast to meet the middle line in the cerebellar unpublished drafts of other manuscripts, personal corre- region.’ This has been criticized both as a statement of spondence, and notes written by Dart, and obtained fact and on the matter of style; on the matter of fact copies of four unpublished illustrations of the Taung because less than half of the cerebellar region is present endocast that were prepared under his direction, and on so that the statement with reference to the midline is which he labeled numerous sulci. Because Dart had not valid, and you attribute an active volition to the identified only
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