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Journal of Human Evolution 59 (2010) 685e691

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Journal of Human Evolution

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News and Views Initial fossil discoveries from Hoogland, a new Pliocene primate-bearing karstic system in Gauteng Province, South Africa

J.W. Adams a,b,*, A.I.R. Herries c, J. Hemingway b, A.D.T. Kegley a,b, L. Kgasi d, P. Hopley e, H. Reade f, S. Potze d, J.F. Thackeray g a Biomedical Sciences Department, Grand Valley State University, 312 Padnos Hall, Allendale, MI 49401, USA b School of Anatomical Sciences, University of the Witwatersrand Medical School, 7 York Road, Parktown, Johannesburg 2193, Republic of South Africa c UNSW Archaeomagnetism Laboratory, Integrative Palaeoecological and Anthropological Studies, School of Medical Sciences, University of New South Wales, Kensington NSW 2052, Sydney, Australia d Department of Palaeontology and H.O.P.E., Ditsong National Museum of Natural History, Kruger Street, Pretoria 0001, Republic of South Africa e Department of Earth and Planetary Sciences, Birkbeck College, University of London, Malet Street, London WC1E 7HX, UK f Department of Archaeology, University of Cambridge, Cambridge CB2 3DZ, UK g Institute for Human Evolution, University of the Witwatersrand, Private Bag 3, Johannesburg 2050, Republic of South Africa

article info palaeocave, located in the Schurveberg (Farm Vlakplaats 354 JR; Gauteng Province; 2548048.3000S, 280020.4000E). Given that Article history: Hoogland lies on the former Hausleitner property and Broom’s Received 11 March 2010 descriptions of the locality, Hoogland likely represents a source for Accepted 1 July 2010 at least some of the ‘Schurveberg collection’ fossils.

Keywords: This announcement provides a primary description of the Pliocene geology and stratigraphy of the Hoogland cave system and prelim- Theropithecus inary data from paleomagnetic, isotopic and macromammalian Cave geology faunal analysis from ex situ sediments. The results of these analyses Paleomagnetism are then used to provide a provisional date for the deposits.

Site geology and paleomagnetism Introduction The Hoogland site includes an active cave system and a series of In 1936, Robert Broom explored fossil-bearing breccias from paleokarstic fossil-bearing deposits exposed during opencast spe- a previously mined cave system on the Hausleitner property near leothem mining. Until recently, the only entrances into the active the Hennops River, 22.5 km west of Pretoria in the Schurveberg system would have been a series of deep circular shafts running mountain region (Broom, 1936; Fig. 1). Broom collected a series of across the hillside above the fossil site. However, opencast lime fossil specimens from this site and other paleocaves in the region, mining of the paleokarstic deposit in the early 20th century broke including the type specimen of Papio (Dinopithecus) ingens (SB 7), through into a large chamber at the far reaches of this modern and a ‘Felis whitei’ mandible (TM 856; now attributed to Mega- system. The lime miners then followed the open cavities removing ntereon cultridens; Turner, 1987), that comprise the ‘Schurveberg any large quantities of speleothem. collection’ at the Ditsong (formerly Transvaal) Museum, Pretoria. A remnant of a similar circular shaft can be seen against the However, discoveries at Sterkfontein and later involvement at the northern wall of the mined cavity and indicates the deep vertical other Blaauwbank River Valley sites (e.g., Swartkrans and Krom- morphology of the entrance to the paleocave. The formation of draai) appears to have permanently forestalled Broom’s return to speleothems at the base of the ancient shaft suggests it was deep, or the Schurveberg region. In 2008, our team began the first in situ that an upper cave existed and was not subject to exterior conditions excavation and processing of ex situ deposits from the Hoogland that form more tufa-like deposits (see Herries et al., 2006b). Exog- enous material that fell down this shaft accumulated as a large talus slope at its base. This material was then periodically winnowed into * Corresponding author. a lateral cavern, depositing a series of interstratified siltstones, E-mail addresses: [email protected] (J.W. Adams), [email protected] gravels and flowstones. As a result, and in contrast with many South (A.I.R. Herries), [email protected] (J. Hemingway), kegleya@gvsu. African Plio-Pleistocene cave systems (e.g. Sterkfontein), the Hoog- edu (A.D.T. Kegley), lkgasi@nfi.museum (L. Kgasi), [email protected] (P. Hopley), [email protected] (H. Reade), potze@nfi.museum (J.F. Potze), francis.thackeray@ land deposits formed with an inclined ‘layer cake-like’ stratigraphy. wits.ac.za (J.F. Thackeray). Bone-rich clastic deposits are interstratified with flowstone lenses

686 J.W. Adams et al. / Journal of Human Evolution 59 (2010) 685e691

Figure 1. Topographic map of the Hoogland locality in reference to nearby Plio-Pleistocene fossil sites. Contour lines equal 20 m.

radiating from a series of superimposed, large stalagmite bosses formed at various periods on top of the talus cone. While some portions of the sequence have been partially obscured by mining rubble, there is continuity in exposed deposits between the upper layers at the base of the entrance shaft and the lower layers in the lateral chamber. The overall effect of this depositional history was to produce ‘flowstone bounded units’ (FBUs) as seen at Gladysvale (Pickering et al., 2007) and Buffalo Cave (Herries et al., 2006b). The stratigraphy is therefore described in terms of FBUs and sedimentologically distinct zones (siltstone, gravel, breccia, speleothem, etc.; Fig. 2). The deposits in the lower part of the sequence are horizontally bedded on the flat floor of what was a small chamber, and were deposited under alternating subaerial and subaqueous environments. The deposits of the upper part of the sequence are partly inclined to the south due to their deposition on the edge of the original talus cone running from the base of the entrance shaft in a subaerial environment. The base of the Hoogland paleocave sequence consists of 50 cm of exposed vuggy (inclusion-filled) speleothem, overlain by 60 cm of clean banded flowstone with interstratified layers rich in organic inclusions. This is termed the ‘southern basal speleothem’ (SBS). The top 25 cm of the SBS has occasional isolated clastic layers due to the formation and infill of small gour pools (pools of water surrounded by a rimstone dam) within the flowstone. Above the SBS is 30 cm of siltstone deposits with thin layers of subaqueous mammillary speleothem, suggesting formation in deeper pools. The lower half of this layer appears to represent a break in clastic deposition into the pools, Figure 2. Stratigraphy and magnetic polarity (Ogg and Smith, 2004) for the various ‘ ’ while the upper 15 cm represents dirty mammillary speleothem flowstone bounded units (FBUs) and sedimentological phases of deposition at the where clastic deposition had recommenced. This unit is termed the Hoogland fossil site. Author's personal copy

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Table 1 Table 1 (continued) Hoogland faunal assemblage Taxon NISP MNI Taxon NISP MNI Class Reptilia Class Mammalia Family Boidae Order Primates Subfamily Pythoninae Family Cercopithecidae Indet. 2 1 Subtribe Papioninae Theropithecus oswaldi oswaldi 11Total 1005 76 Indeterminate 1 1

Order Artiodactyla Family Bovidae ‘siltstone and subaqueous speleothem’ (STSAS). The lateral extent of Tribe Antilopini these deposits is unknown due to the infill of rubble to the north of Antidorcas bondi 21the site, but it appears that this layer formed synchronously as Antidorcas recki 10 4 fl Antidorcas sp. 6 2 subaerial owstone was forming towards the top of the talus slope. Antilopini indet. 3 1 This is the end of the basal zone of stratigraphy that is dominated by subaerial and subaqueous speleothem deposition, which is collec- Tribe Alcelaphini ‘ ’ Indeterminate (Class II) 17 6 tively termed the Basal Speleothem (BSP; Fig. 2). Indeterminate (Class III) 15 6 The BSP deposits are overlain by over 5 m of banded siltstone, conglomerate and breccia deposits that represent alternating Tribe Cephalophini cf. Sylvicapra sp. 1 1 periods of erosion, collapse, and periodic flooding events where Cephalophini indet. 2 1 clastic material that had entered the system through the vertical

Tribe Neotragini shaft were washed from the top of the talus cone towards the back Oreotragus oreotragus 63wall of the cave. This sequence can be subdivided into three major cf. Raphicerus sp. 2 2 depositional zones based on sedimentological characteristics of the Neotragini indet. 7 4 deposits, or five flowstone bounded units (FBUs) that represent Tribe Reduncini discrete periods of clastic deposition covered by capping flow- Kobus sp. 1 1 stones. The lowest zone, ‘Basal Siltstone’ (BST) consists of alter- Redunca sp. 33 11 nating sequences of fine-grained conglomerate overlain by Reduncini indet. 5 2 siltstone deposits that represent alternating flooding events. These Tribe Tragelaphini deposits are seemingly entirely sterile of fossils. BST consists of Tragelaphus strepsiceros 21FBU1 and the lower part of FBU2. Towards the top of FBU2, the Tragelaphus sp. 2 2 Tragelaphini indet. 4 1 sequence begins to be dominated by coarser-grained conglomerate that represents the start of the next depositional zone, ‘Inter- Tribe Antilopini/Neotragini stratified Conglomerate and Flowstone’ (ICFS). ICFS consists of Indet. 6 1 conglomerates with large clasts of eroded, sub-rounded speleo- Tribe Cephalophini/Neotragini them. This phase contains almost all the fossil material so far e Indet. 1 fi fl Bovidae indet. craniodental 635 e identi ed in section at the site. A series of owstones, some capping Bovidae indet. postcranial 126 e the underlying deposits, others more ephemeral, are interstratified with the conglomerate and divide it into upper FBU2 through to Family Suidae fi Indet. 1 1 lower FBU5. This conglomerate rich phase is overlain by ne- grained silt deposits ‘the Upper Siltstone’ (UST), which contains Order Carnivora very little bone. The separation between the bone-rich ICFS and Family Canidae bone-poor UST is not well defined in some sections where there Indet. 1 1 appears to be a gradient from one to the other within FBU5. This Family Felidae density of fossils in ICFS suggests it is almost certainly the source of ? Acinonyx 11the current ex situ faunal sample. Family Hyaenidae Paleomagnetic analysis was undertaken on 44 samples across the ? Hyaena 11Hoogland sequence as per the procedures outlined in Dirks et al. e Carnivora indet. 5 (2010). This indicates the occurrence of a number of geomagnetic polarity transitions that suggest that the sequence covers an Order Perissiodactyla Family Equidae extremely long time period. Detailed work on the lower sections of Indet. 3 2 this sequence is ongoing, along with uranium-lead dating. The lower part of the dense bone-bearing zone ICFS has a reversed polarity at Order Hyracoidea its base in upper FBU2, intermediate polarity during the older fossil Family Procaviidae deposit of FBU3 and normal polarity for the remainder of ICFS, UST Procavia antiqua 52 Procavia transvaalensis 90 10 (Upper FBU5) then records a reversed polarity. Procavia sp. 5 3 Current faunal sample Order Lagomorpha Indet. 2 1 The current Hoogland faunal assemblage ( n ¼ 4327) is primarily Class Aves derived from nine fossiliferous ex situ blocks that correlate with the Indet. 1 1 FBU 5 in situ deposits. Despite the small number of blocks pro- cessed, 76 individuals are minimally represented across three Author's personal copy

688 J.W. Adams et al. / Journal of Human Evolution 59 (2010) 685e691 classes, including six orders of mammals (Table 1). In addition to Table 2 the taxonomically-identifiable sample presented in Table 1, the Measurements of the HLP 1600 lower molar and ranges of Theropithecus oswaldi subspecies.a current Hoogland fossil sample includes a single coprolite, 827 indeterminate craniodental specimens, 46 indeterminate post- BL (Mesial) BL (Distal) MD cranial specimens, and 2,448 indeterminate (primarily long-bone HLP 1600 9.83 10.19 15.57 (max)/13.29 (cej) diaphysis) fragments. m1 T. o. leakeyi 9.5e11.8 (4) 9.9e11.6 (4) 12.7e13.7 (5)b(max) T. o. oswaldi 9.4e12.8 (7) 9.6e12.4 (7) 10.8e16.5 (7) T. o. darti (Mak) 8.4e12.7 (8) 9.2e12.2 (8) 9.4e14.7 (8)b(max) e e e b Primates T. o. darti (Had) 7.3 11.8 (23) 7.0 11.6 (20) 8.5 14.1 (31) (max) m2 T. o. leakeyi 13.6e15.0 (4)b 12.6e14.6 (4)b 17.6e19.9 (5)b Two isolated primate specimens (HLP 1600: left m1, HLP 580: T. o. oswaldi 10.7 e12.8 (9)b 11.0e12.9 (9)b 13.1e17.0 (10) e b e e left lower canine) have been recovered from the Hoogland ex situ T. o. darti (Mak) 11.2 12.7 (7) 10.0 12.5 (9) 12.6 18.3 (9) T. o. darti (Had) 8.7e14.9 (24) 8.4e14.6 (26) 11.1e19.9 (30) breccia sample. The HLP 1600 molar (Fig. 3) is completely formed and exhibits the complex, derived crown morphology described MD, mesiodistal; BL, buccolingual; cej, measure taken at the cementoenamel junction; max, maximal measure of that dimension. for Theropithecus (Freedman, 1957; Jolly, 1972; Delson, 1975; Szalay a Theropithecus oswaldi darti samples from Makapansgat (Mak) and Hadar (Had) and Delson, 1979; Eck and Jablonski, 1984; Frost and Delson, have been listed separately because of their previously described morphological 2002). The high columnar cusps are separated by deep basins differences. b and a raised ridge of enamel lies between the two buccal cusps, The HLP 1600 specimen lies outside these described measurement ranges. yielding a high degree of occlusal relief. The mesial and distal buccal clefts are prominent, while the median buccal cleft extends Metric comparisons of HLP 1600 (following Freedman, 1957) into a pocket on the near-vertical buccal surface. The mesial and with the first and second mandibular molar measurements for T. distal foveas are deep and the lophids, together with the trigonid o. darti, T. o. oswaldi,andT. o. leakeyi obtained from the PRIMO basin, are angled mesio-lingually relative to the long axis of the database (http://primo.nycep.org) indicate that the tooth is tooth. mesiodistally expanded relative to what would be expected given Because of temporally-associated increases in body size, molar its buccolingual dimensions (Table 2). However, Theropithecus size, and crown complexity among specimens of Plio-Pleistocene molars generally become mesiodistally ‘shortened’ with occlusal Theropithecus, Leakey (1993) has treated Theropithecus oswaldi as wear because their maximum mesial and distal dimension occurs a chronospecies, comprised of three subspecies (T. o. darti, T. o. near the crown apex and does not extend to the cementoenamel leakeyi, T. o. oswaldi) spanning deposits from (tentatively) 3.9 Ma to junction (Szalay and Delson, 1979). As the comparative Ther- 0.4 Ma (Frost, 2001; Frost and Delson, 2002). Relative to these three opithecus molars include those with more significant wear, subspecies, the HLP 1600 specimen differs from early T. o. darti in a second mesiodistal measure of HLP 1600 at the cementoenamel having a more derived crown morphology; however, it lacks the junction (although an underestimate of the dimension of the degree of hypsodonty and enamel complexity characteristic of the tooth in normal wear) is within the established mesiodistal late-occurring T. o. leakeyi specimens. The HLP specimen does dimensions for the Theropithecus subspecies. The HLP specimen is possess both a ‘pocketed’ median buccal notch and deeply incised buccolingually narrower than Theropithecus second molars, with lingual notches extending below half the height of the protoconid. the exception of the T. o. darti from Hadar and Makapansgat These features have been used by Jolly (1972) and Jablonski and (which, as noted above, it is morphologically distinct from). Given Chaplin (2008), respectively, to distinguish specimens of T. o. that the overall dimensions of the HLP specimen is consistent oswaldi from those of T. o. darti. with lower first molar specimens, and the morphology of the

Figure 3. HLP 1600, Theropithecus sp. left m1. (a) Buccal view, (b) oblique lingual view, (c) occlusal view. Scale bar equals 1 cm. Author's personal copy

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Figure 4. HLP 580, Papionini indet. lower left canine. (a) Mesiolabial view, (b) distolabial view, (c) distolingual view. Scale bar equals 1 cm.

specimen corresponds with features diagnostic of T. o. oswaldi,we Acinonyx sp. (HLP 383; left I3) and Hyaena sp. (HLP 1; left i2), provisionally assign HLP 1600 as a first lower left molar of T. o. along with a canid terminal phalanx (HLP 2023) are the only oswaldi. carnivore specimens that can be confidently identified below the The HLP 580 lower canine is a complete unworn crown with the Order level. In addition, the current Hoogland assemblage has root broken just below the cementoenamel junction (Fig. 4). The three unattributable partial carnivore tooth fragments (HLP 496, overall dimensions of the specimen (MD: 5.6 mm, BL: 8.5 mm; 638, 942), a mandibular incisor (HLP 733), and an isolated inter- measurements following Swindler, 2002) fall within the ranges of mediate cuneiform (HLP 180) from a large-bodied felid or many fossil papionin species and are therefore not taxonomically hyaenid. diagnostic (PRIMO). Morphologically, HLP 580 has a moderate The largest single taxonomic group represented in the Hoogland distostylid and low bulbous crown, lacks cingulum development, sample thus far is the Procaviidae. Most of the well-preserved and has shallow mesial and distal lingual grooves. In comparison remains are directly attributable to the extinct Procavia trans- with South African papionin lower canine specimens, HLP 580 is vaalensis, with strongly developed lingual cingula and crenulated similar to the smaller, probable female Parapapio specimens from ectolophs in comparison with extinct Procavia antiqua and extant Swartkrans Member 1 (SK 571A: w2.0 Ma; Herries et al., 2009) and Procavia capensis (Churcher, 1956). All three relatively complete Sterkfontein Member 4 (STS 563; 2.6e2.0 Ma; Herries et al., 2010) isolated upper incisors (HLP 14, 34 and 404) exhibit more centrally assemblages, although the mesial lingual groove of HLP 580 is positioned labial ridges that are consistent with typical male hyrax shallower than observed in both SK 571A and STS 563. However, morphology (Churcher, 1956). given the diversity of Plio-Pleistocene South African papionins and Only three equid dental specimens have been recovered from that the specimen is an isolated canine, we can only confidently Hoogland. All are incisors from Block 9 (HLP 303; 304; 594). The attribute the specimen as an indeterminate papionin. isolated right i1 (HLP 303) is within the metric range of South African Hipparion and Equus but is too worn to attribute to a specific Bovidae taxon. The other specimens are upper left i2 (HLP 304) and upper left i3 (HLP 594) crown tips, likely from the same immature equid. There is significant taxonomic and trophic diversity among the These specimens also fall within the metric and morphological current bovid sample. Of particular note are mandibular third range of both equid genera and thus cannot be more specifically (n ¼ 6) and fourth (n ¼ 2) premolar specimens attributable to the attributed. genus Redunca. These specimens have widely-separated lingual cusps (paraconid, metaconid, and entoconid) and a gracile hypo- Discussion and summary conid. In these respects, the Hoogland specimens are analogous to Redunca darti from Makapansgat Member 3 (3.0e2.6 Ma; Hopley Our initial paleomagnetic results and faunal description et al., 2007a; Herries et al., 2010), and differ from the fused indicates deposition of FBU4 and 5 lower during a late Pliocene lingual cusps and projecting hypoconid of the Redunca sp. sample or early Pleistocene normal polarity period. Normal polarity from Gondolin GD 2 (w1.8 Ma; Adams and Conroy, 2005; Adams, periodsinthistimerangeincludetheendoftheGaussChron 2006; Herries et al., 2006a), extant Redunca arundinum, and (3.03e2.58 Ma), the Olduvai Subchron (1.95e1.78 Ma), or the Redunca fulvorufula. A detailed comparative analysis of the Hoog- Rèunion and Huckleberry Ridge events (2.2e2.0; Dirks et al., land reduncines is forthcoming. 2010). The normal polarity of the fossil deposits is unlikely to represent either of the short normal polarity events, given that Additional fauna thick, slowly deposited speleothem formed within this depositional zone. The occurrence of reduncines with primitive, Only a very small carnivore sample has been recovered thus Redunca darti-like premolars consistent with those from Maka- far. Two complete isolated incisors, provisionally attributed to pansgat Member 3 (3.0e2.6 Ma; Herries et al., 2010)makes Author's personal copy

690 J.W. Adams et al. / Journal of Human Evolution 59 (2010) 685e691 deposition within the Olduvai Subchron unlikely. As such, our Adams, J.W., Conroy, G.C., 2005. Plio-Pleistocene faunal remains from the Gondolin GD 2 preliminary interpretation is thatthisnormalpolarityperiodin in situ assemblage, North West Province, South Africa. In: Lieberman, D., Smith, R.J., Kelley, J. (Eds.), Interpreting The Past: Essays on Human, Primate and Mammal the Hoogland sequence represents the end of the normal Evolution in Honor of David Pilbeam. Brill Academic Publishers, Boston, pp. 243e261. polarity Gauss Chron (C2An.1n), with the FBU 5 fossils deposited Adams, J.W., Hemingway, J., Kegley, A., Thackeray, J.F., 2007a. Luleche, a new at the end of this period around 2.58 Ma, fossil layer FBU 4 dated paleontological site in the Cradle of Humankind, North-West Province, South Africa. J. Hum. Evol. 53, 751e754. closer to 3.03 Ma and fossils from FBU 3 dating to between 3.12 Adams, J.W., Herries, A., Conroy, G.C., Kuykendall, K., 2007b. Taphonomy of a South and 3.03 Ma. The ICFS fossil deposits are thus contemporary African cave: geological and hydrological influences on the GD 1 fossil assem- with fossils from Members 2e4 from the Makapansgat Lime- blage at Gondolin, a Plio-Pleistocene paleocave system in the Northwest e e Province, South Africa. Quaternary Sci. Rev. 26, 2526 2543. works (3.03 2.58 Ma; Latham et al., 1999, 2003; Herries et al., Broom, R., 1936. Letter to Mr. Hausleitner, Schruveberg (Hennops River), Pretoria. 2010) and represent one of only a handful of well documented 05/19/1936. Transvaal Museum Document #TM 17/35. Pliocene (5.33e2.59 Ma) fossil deposits in South Africa, the Churcher, C., 1956. The fossil Hyracoidea of the Transvaal and Taungs deposits. Ann. w Transv. Mus. 224, 477e501. others being Langebaanweg at 5Ma (Roberts, 2006)and Delson, E., 1975. Evolutionary history of the Cercopithecidae. In: Szalay, F. (Ed.), portions of the Bolt’sFarmcomplexatw4.5e4.0 Ma (Gommery Approaches to Primate Paleobiology. Karger, New York, pp. 167e217. et al., 2008). Dirks, P.H.D.M., Kibii, J.N., Kuhn, B.F., Steininger, C., Churchill, S.E., Kramers, J.D., Initial 18Oand13C isotopic analysis of the BSP deposits indi- Pickering, R., Farber, D.L., Mériaux, S.-A., Herries, A.I.R., King, G.C.P., Berger, L.R., 2010. Geological setting and age of Australopithecus sediba from southern Africa. cates a predominantly C3 vegetation in the lower layers followed Science 328, 205e208. by a transition to a mixed C3 and C4 vegetation towards the top of Eck, G.G., Jablonski, N.G., 1984. A reassessment of the taxonomic status and phyletic relationships of Papio baringensis and Papio quadratirostris (Primates: Cercopi- the sequence (Reade, 2009). The occurrence of grazers and e e thecidae). Am. J. Phys. Anthropol. 65, 109 134. browsers in the FBU3 5 deposits suggests that C4 and C3 vege- Freedman, L., 1957. Fossil Cercopithecoidea of South Africa. Ann. Transv. Mus. 23, tation was present in the Schurveberg mountain region during 121e262. the depositional phases covered by the Hoogland faunal Frost, S.R., 2001. Fossil Cercopithecidae from the Afar Depression, Ethiopia: species systematics and comparison to the Turkana Basin. Ph.D. dissertation, City sequence. It is not yet apparent whether the fauna was deposited University of New York. in a stable mixed C3/C4 environment or experienced a range of Frost, S.R., Delson, E., 2002. Fossil Cercopithecidae from the Hadar Formation and vegetation regimes due to orbitally-forced climate change surrounding areas of the Afar Depression, Ethiopia. J. Hum. Evol. 43, 687e748. (Hopley and Maslin, 2010). Further faunal and isotopic work will Gommery, D., Thackeray, J., Senegas, F., Potze, S., Kgasi, L., 2008. The earliest primate focus on resolving the palaeoenvironmental reconstruction of this (Parapapio sp.) from the Cradle of Humankind World Heritage site (Waypoint locality. 160, Bolt’s Farm, South Africa). S. Afr. J. Sci. 104, 405e408. Herries, A., Adams, J., Kuykendall, K., Shaw, J., 2006a. Speleology and magneto- These initial results from Hoogland, along with interdisci- biostratigraphic chronology of the Gondolin hominin palaeocave, S. Africa. plinary studies of sites outside the Blaauwbank River Valley J. Hum. Evol. 51, 617e631. (Menter et al., 1999; Keyser et al., 2000; Adams, 2006; Herries Herries, A., Reed, K.E., Kuykendall, K., Latham, A., 2006b. Speleology and magne- et al., 2006a,b; Hopley et al., 2007a,b; Adams et al., 2007a,b) tobiostratigraphic chronology of the Buffalo Cave fossil site, Makapansgat, South Africa. Quatern. Res. 662, 233e245. have highlighted the importance of expanded excavation and Herries, A.I.R., Curnoe, D., Adams, J.W., 2009. A multi-disciplinary seriation of early analysis in establishing more accurate ranges of variation among Homo and Paranthropus bearing palaeocaves in southern Africa. Quatern. Int. e Plio-Pleistocene mammal lineages, paleocommunity composition, 202, 14 28. Herries, A.I.R., Adams, J.W., Curnoe, D., Warr, G., Latham, A.G., Shaw, J., 2010. A and the geological and taphonomic modes of fossil record multi-disciplinary perspective on the age of Australopithecus in southern formation. Ongoing fossil preparation, in situ excavations at Africa. In: Leakey, R., Reed, K. (Eds.), Paleoecology of Australopithecus. Verte- Hoogland, and continued geologic sampling for isotopes, paleo- brate Paleobiology and Paleoanthropology Series. Hopley, P.J., Marshall, J.D., Weedon, G.P., Latham, A.G., Herries, A.I.R., magnetism and uranium-lead dating will continue to expand our Kuykendall, K.L., 2007a. Orbital forcing and the spread of C4 grasses in the late understanding of the paleolandscape of southern Africa during Neogene: stable isotope evidence from South African speleothems. J. Hum. Evol. the terminal Pliocene; a period in which only one hominin 53, 620e634. Hopley, P.J., Weedon, G.P., Marshall, J.D., Herries, A.I.R., Latham, A.G., bearing site (Makapansgat) occurs in the region and as such Kuykendall, K.L., 2007b. High- and low-latitude orbital forcing of early hominin potentially marks the period of colonization of southern Africa by habitats. Earth Planet. Sci. Lett. 256, 419e432. Australopithecus. Hopley, P.J., Maslin, M.A., 2010. Climate-averaging of terrestrial faunas: an example from the Plio-Pleistocene of South Africa. Paleobiology 36, 32e50. Jablonski, N.G., Chaplin, G., 2008. Natural language descriptions and keys of the Acknowledgements Koobi Fora monkey fossil species using Delta. In: Jablonski, N.G., Leakey, M.G. (Eds.), Koobi Fora Research Project. The Fossil Monkeys, vol. 6. California Academy of Sciences, San Francisco, pp. 301e336. We wish to thank the Kruger family for their assistance and Jolly, C.J., 1972. The classification and natural history of Theropithecus (Simopithecus) access to the Hoogland locality. Funding for research at Hoogland (Andrews, 1916) baboons of the African Plio-Pleistocene. Bull. British Museum e was provided by a grant by the Ford Foundation and by Faculty of (Natural History), Geol. 22, 1 122. Keyser, A.W., Menter, C.G., Moggi-Cecchi, J., Pickering, T.R., Berger, L.R., 2000. Health Sciences, University of the Witwatersrand. Preliminary Drimolen: a new hominid-bearing site in Gauteng, South Africa. S. Afr. J. Sci. paleomagnetic analysis was funded by a UNSW Faculty of Medicine 964, 193e197. research grant to A.I.R.H. and was additionally supported by ARC Latham, A., Herries, A., Kuykendall, K., 2003. The formation and sedimentary infilling of the Limeworks Cave, Makapansgat, South Africa. Palaeontologia Discovery Grant DP0877603. Thanks to Florian Stark (Geomagne- Africana 39, 69e82. tism Laboratory, University of Liverpool) for assistance with Latham, A., Herries, A., Quinney, P., Sinclair, A., Kuykendall, K., 1999. The Maka- paleomagnetic and isotope sampling, Ginette Warr for sample pansgat australopithecine site from a speleological perspective. In: Pollard, A. (Ed.), Geoarchaeology: Exploration, Environments, Resources. Geological preparation, and to Eric Delson and an anonymous reviewer for Society of London, London, pp. 61e77. comments on an early draft of this manuscript. Finally, we grate- Leakey, M., 1993. Evolution of Theropithecus in the Turkana Basin. In: Jablonski, N. fully acknowledge the encouragement and support of Charles (Ed.), Theropithecus: The Rise and Fall of a Primate Genus. Cambridge Univer- e fi sity Press, Cambridge, pp. 85 123. Lockwood for our Schurveberg eldwork as the Director-elect of Menter, C.G., Kuykendall, K.L., Keyser, A.W., Conroy, G.C., 1999. First record of the Institute for Human Evolution, University of the Witwatersrand. hominid teeth from the Plio-Pleistocene site of Gondolin, South Africa. J. Hum. Evol. 37, 299e307. Ogg, J.G., Smith, A.G., 2004. The geomagnetic polarity timescale. In: Gradstein, F., References Ogg, J., Smith, A. (Eds.), A Geologic Time Scale. Cambridge University Press, pp. 63e86. Adams, J.W., 2006. Taphonomy and paleoecology of the Gondolin Plio-Pleistocene Pickering, R., Hancox, P., Lee-Thorp, J., Grun, G., Mortimer, M., McCulloch, M., cave site, South Africa. Ph.D. Dissertation, Washington University in St. Louis. Berger, L.R., 2007. 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