ORIGINAL ARTICLE The Oldowan and Early Acheulean Mammalian Fauna of Wonderwerk Cave (Northern Cape Province, South Africa) James Brink & Sharon Holt & Liora Kolska Horwitz # Springer Science+Business Media New York 2016 Abstract We describe and discuss the large vertebrates recovered from the basal layers (Strata 12 and 11) of Excavation 1 at Wonderwerk Cave, a site located in the Kuruman Hills, Northern Cape Province, South Africa. Stratum 12 is associated with a small core and flake Oldowan assemblage while Stratum 11 contains some Acheulean material. Based on palaeo-magnetism, the time span covered by these Strata is estimated to date to ca.1.8– 1.1 million years ago. Taxa identified include late Makapanian forms, such as Procavia transvaalensis, Procavia antiqua, a hipparionine and an unnamed species of large caprine, also found in the Makapan Limeworks deposits, confirming the antiquity of these layers. The bones are highly fragmented due to the action of multiple agencies, both pre- and postdepositional, which prevented diagnosis in many cases to lower levels of taxonomy. In support of other palaeo-environmental proxies from Strata 12 and 11, the large mammal remains reflect a semi-arid ecotone palaeo-environment, consisting of a mix of taxa associated with broken, montane habitat and semi-arid grassland-savanna plains habitat. Résumé Nous décrivons et discutons les grands mammifères qui sont retrouvés des niveaux inférieurs (Strates 12 et 11) de l’excavation de Wonderwerk Cave, situé dans les collines de Kuruman, Cap du Nord, Afrique de Sud. Strate 12 est associée avec l’assemblage de nucleus et éclat Oldowan alors que la Strate 11 contient quelques matériaux Acheuléens. Selon le paléomagnétisme, ces strates ont c. 1.8 – 1.1 mil- lions d’années. Les taxa identifiés incluent les formes Makapaniennes tardes, comme par exemple Procovia transvaalensis, P. antiqua, un hipparionin et une espèce sans nom de caprin grand, qui est aussi trouvée dans les dépôts de Limeworks à Makapan, qui confirme l’antiquité des niveaux. Les ossements sont très fragmentés à cause des agents taphonomiques avant et après déposition. Cela a empêché la détermination dans plusieurs cas aux niveaux inférieurs taxonomiques. Les restes des grandes mammifères soutiennent des autres indications paléo–environnementales des Strates 11 et 12, et ils reflétent un paléo – environnement écotone semi-aride. Les taxa sont associés avec un habitat quelque peu montagneux et un habitat de savanna-prairie semi-aride. Keywords Early Pleistocene . Mammals . Oldowan . Early Acheulean . South Africa Afr Archaeol Rev DOI 10.1007/s10437-016-9223-1 J. Brink (*) : S. Holt Florisbad Quaternary Research Department, National Museum, Bloemfontein, South Africa e-mail: [email protected]J. Brink Centre for Environmental Management, Faculty of Natural and Agricultural Science, University of the Free State, Bloemfontein, South Africa L. K. Horwitz National Natural History Collections, Faculty of Life Sciences, The Hebrew University, Jerusalem, Israel
Transcript
ORIGINAL ARTICLE
The Oldowan and Early Acheulean Mammalian Faunaof Wonderwerk Cave (Northern Cape Province, South Africa)
James Brink & Sharon Holt & Liora Kolska Horwitz
# Springer Science+Business Media New York 2016
Abstract We describe and discuss the large vertebratesrecovered from the basal layers (Strata 12 and 11) ofExcavation 1 at Wonderwerk Cave, a site located in theKuruman Hills, Northern Cape Province, South Africa.Stratum 12 is associated with a small core and flakeOldowan assemblage while Stratum 11 contains someAcheuleanmaterial. Based on palaeo-magnetism, the timespan covered by these Strata is estimated to date to ca.1.8–1.1 million years ago. Taxa identified include lateMakapanian forms, such as Procavia transvaalensis,Procavia antiqua, a hipparionine and an unnamed speciesof large caprine, also found in the Makapan Limeworksdeposits, confirming the antiquity of these layers. Thebones are highly fragmented due to the action of multipleagencies, both pre- and postdepositional, which preventeddiagnosis in many cases to lower levels of taxonomy. Insupport of other palaeo-environmental proxies from Strata12 and 11, the large mammal remains reflect a semi-arid
ecotone palaeo-environment, consisting of a mix of taxaassociated with broken, montane habitat and semi-aridgrassland-savanna plains habitat.
Résumé Nous décrivons et discutons les grandsmammifères qui sont retrouvés des niveaux inférieurs(Strates 12 et 11) de l’excavation de Wonderwerk Cave,situé dans les collines de Kuruman, Cap du Nord,Afrique de Sud. Strate 12 est associée avec l’assemblagede nucleus et éclat Oldowan alors que la Strate 11contient quelques matériaux Acheuléens. Selon lepaléomagnétisme, ces strates ont c. 1.8 – 1.1 mil-lions d’années. Les taxa identifiés incluent lesformes Makapaniennes tardes, comme par exempleProcovia transvaalensis, P. antiqua, un hipparioninet une espèce sans nom de caprin grand, qui estaussi trouvée dans les dépôts de Limeworks àMakapan, qui confirme l’antiquité des niveaux.Les ossements sont très fragmentés à cause desagents taphonomiques avant et après déposition.Cela a empêché la détermination dans plusieurscas aux niveaux inférieurs taxonomiques. Lesrestes des grandes mammifères soutiennent desautres indications paléo–environnementales desStrates 11 et 12, et ils reflétent un paléo–environnement écotone semi-aride. Les taxa sontassociés avec un habitat quelque peu montagneuxet un habitat de savanna-prairie semi-aride.
Keywords Early Pleistocene .Mammals . Oldowan .
Early Acheulean . South Africa
Afr Archaeol RevDOI 10.1007/s10437-016-9223-1
J. Brink (*) : S. HoltFlorisbad Quaternary Research Department, National Museum,Bloemfontein, South Africae-mail: [email protected]
J. BrinkCentre for Environmental Management, Faculty of Natural andAgricultural Science, University of the Free State, Bloemfontein,South Africa
L. K. HorwitzNational Natural History Collections, Faculty of Life Sciences,The Hebrew University, Jerusalem, Israel
There is a hiatus in our knowledge of the palaeo-ecologyand palaeo-environment of the arid interior of SouthAfrica during the Oldowan and Acheulean periods. Thisis because most sites in this time period represent opensites or isolated find localities that lack faunal or botanicalremains (e.g., several sites in Beaumont and Morris 1990;Butzer 1974; Walker et al. 2014), while others, like theVaal River gravels which may contain fauna, representsecondary depositional contexts (e.g., Cooke 1963;Klein 1984; Leader 2009; McNabb and Beaumont2011). An exception is the archaeological site ofWonderwerk Cave which has yielded well-preserved or-ganic remains that span approximately the entire 2millionyears of the archaeological record represented at the site.
Wonderwerk has been excavated since the 1940s byseveral different teams (for reviews, see Beaumont andVogel 2006; Horwitz and Chazan 2015). The most exten-sive excavations were undertaken by P.B. Beaumont overthe period 1978–1992. The current paper offers an in-depth description of the systematics of mammalian faunafrom the two lowest strata in the cave associated withOldowan and possibly an Early Acheulean lithic industry,respectively, and expands on a preliminary report on thismaterial published by Brink et al. (2015).
The Site and Its Environment
Wonderwerk Cave (27° 50′ 45′′ S. 23° 33′ 19′′ E) lies onthe lower slopes of the Kuruman Hills, between thetowns of Danielskuil and Kuruman in the NorthernCape Province of South Africa. The cave falls withinthe Savanna biome but lies in close proximity to theGrassland and Nama Karoo biomes (Fig. 1; Mucina andRutherford 2006; Rutherford and Westfall 1986). TheSavanna biome covers the majority of northern SouthAfrica and extends into Namibia, Botswana and Zim-babwe. It comprises several vegetation complexes thatare variable in their vegetation cover and fauna theysupport. Thus, Wonderwerk Cave lies within theKuruman Mountain Bushveld complex (Mucina andRutherford 2006), which is composed of a wide spec-trum of grasses, a short-tree stratum and an open-to-closed woody shrubland component.
A botanical survey conducted by T. Anderson(1991a, b) for the McGregor Museum in the immediatevicinity of the cave, showed that the vegetation is amosaic of at least three different landscape types(Fig. 1; for detailed floral species lists, see Appendix1; the vegetation divisions follow Acocks 1988):
(i) On top of the Kuruman Hills, a sparse and openvegetation—dominated by Orange River BrokenVeld (Karroid Veld, type 9).
(ii) On the slopes of the low-banded ironstone hillssurrounding the cave, a mixture of TarchonanthusVeld (Shrubveld type 12) and False Orange RiverBroken Veld (Acocks Thornveld type 17) equiva-lent to Kuruman Thornveld SKv9 (Mucina andRutherford 2006), with a high species diversity oftrees, shrubs, flowering plants and ferns.
(iii) The plains of calcareous tufa on the Ghaap Plateaubelow the cave are covered by TarchonanthusVeld(Acocks Shrubveld type 12; equivalent toKuruman Vaalbosveld subdivision SVk8 ofMucina and Rutherford 2006), but with a highproportion of Tarchonanthus shrubs, a high diver-sity of grass species and in parts, parklands ofOlea.
A checklist compiled by C. Anderson (1991a, b) ofmammals and birds in the area around the cave,
Fig. 1 Map of the biomes of South Africa (after Mucina andRutherford 2006), showing the location of Wonderwerk Caveand some of the sites mentioned in the text
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indicates a high species diversity with representatives ofca. 24 mammalian families and 51 bird families.Mammalian taxa encountered were all typical of thesemi-arid Northern Cape (for detailed species list, seeAppendix 2). The species richness was attributed byAnderson to the different vegetation regimes and geo-graphic features covered by the area. However, the listedtaxa are all small-sized forms which contrast to historicalaccounts of early nineteenth-century travellers in the re-gion (e.g., Lichtenstein, Burchell, Methuen etc.) who not-ed the presence of lion, hyaena, jackal, zebra, quagga,white rhinoceros, giraffe, blue wildebeest, roan antelope,hartebeest, buffalo, eland and ostrich (Humphreys 1975;Skead 1980). Over-exploitation combined with habitatdestruction, especially farming, are the main contributingfactors to the to the depletion of wild game populations(Plug and Badenhorst 2001).
Wonderwerk Cave is a phreatic cavity that formedwithin Archaean dolomite and is ca. 140 m long and ca.2,400m2 in extent. Excavation 1, which is the focus of thispaper, is situated ca. 20 m in from the cave mouth. It hasyielded a long archaeological sequence spanning the LaterStone Age (ca. 2 m deep deposits), underlain by over 2 mofEarlier StoneAge (ESA)material, including anOldowanoccurrence at the base of the sequence (Beaumont 1990,2004; Beaumont and Vogel 2006; Chazan 2015; Chazanand Horwitz 2015; Chazan et al. 2008, 2012; Horwitz andChazan 2015; Matmon et al. 2012).
Stratum 12 is the basal archaeological deposit in thecave and to date has only been identified in Excavation 1.As described by Chazan et al. (2008, 2012), the sedimen-tary matrix of this layer comprises two LithostratigraphicUnits, 9 and 8, together ca. 1 m thick, made up of claysand reworked Kalahari sands interspersed with layers ofbanded ironstonemicro-gravels. Features characteristic ofwater deposition are evident in Stratum 12, but there is noevidence for high-energy water action that could accountfor the introduction of artefacts or bones into this layerfrom outside the cave (Goldberg et al. 2015). The overly-ing Stratum 11 (ca. 30 cm thick), is heterogeneous andcomprises three Lithostratigraphic Units (Units 7–5); theinterface betweenUnit 7 and the underlying topmost layerof Stratum 12 (Unit 8) reflects an episode of erosionalchannelling. The Middle Lithostratigraphic layer of Stra-tum11, Unit 6, represents a rock fall, while Unit 5 repre-sents a layer of calcite or phosphate nodules.
The Stratum 12 lithic industry (N=65 artefacts) ischaracterised by small flakes and cores and has beencharacterised as Oldowan (Chazan et al. 2012). The firstbiface appears in the overlying Stratum 11, implying anEarly Acheulean industry, but the lithic assemblagefrom this stratum is extremely small, such that no clearcultural attribution could be made (Chazan 2015;Chazan et al. 2008, 2012).
Stratum 12 is interpreted as dating to the end of theOlduvai subchron (1.96–1.78 Ma) and the beginning ofthe subsequent interval of reversed polarity (1.78–1.07 Ma), while Stratum 11 spans this period of reversepolarity (Chazan et al. 2012; Matmon et al. 2012).
Taphonomy
The taxonomic work has been hampered by the extremefragmentation of the faunal remains. All remains fromStrata 12 and 11 were separated into two groups—thosethat could be identified (comprising bones/teeth identi-fied either to skeletal element/family or species, thoughnot necessarily all of these categories) and those whichcould not be identified to even one of these levels(=unidentified bones/teeth). The maximum length,greatest breadth and depth of all material in both groupswere measured using a digital calliper to two places afterthe decimal point.
The histograms shown in Fig. 2, give the length andbreadth distributions of identified and unidentifiedbones and teeth from Strata 12 and 11. It is evident thatin both layers, the vast majority of the bones recoveredwere less than 5 cm in size (for both dimensions), with asignificant component that is even smaller. Furthermore,as illustrated in the triplots (Fig. 3), the size distributionsof both the identified and unidentified bone and toothremains from Strata 12 and 11 closely resembleeach other. In both layers, there is little differencethe size distributions of the identified and uniden-tified bones, but a marked difference can be ob-served between the identified and unidentifiedteeth, with the unidentified tooth fragments signif-icantly smaller. The latter result is probably due toa methodological effect; fragments of tooth enamelwere easily identified, but for an identification tothe genus or species level, a more complete tooth
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is needed. In contrast, most identified bone couldbe classified only to the level of skeletal elementand not to the level of genus or species.
Detailed taphonomic work on this assemblage iscontinuing, but it is evident that the high degree offragmentation is the end result of activities of severalagents, possibly including hominin processing activi-ties, in situ burning, sediment compaction, carnivoreand porcupine damage (Fig. 4), trampling, rock falls aswell as damage incurred during excavation and sieving(e.g., Lyman 1994; Outram 2001; Stiner et al. 1995).
Methods
Due to the poor preservation of the material, much effortwas expended in reconstructing specimens from fragmentswithin the same square (in spits above and below the one
being studied), as well as from adjoining squares of thesame spit. Importantly, no conjoins were found acrossstrata.
Modern and fossil osteological reference collec-tions were used as aids in the identification ofspecimens: the Florisbad Quaternary Research De-partment, National Museum (Bloemfontein); theDepartment of Mammalogy, National Museum(Bloemfontein), the Institute for Human Evolution(now part of the Evolutionary Studies Institute),University of the Witwatersrand (Johannesburg);and the Department of Palaeontology, DitsongNational Museum of Natural History (formerTransvaal Museum, Pretoria). The diagnosed spec-imens are listed by stratum (Stratum 12 or 11) andby unique catalogue number (WW#).
Abbreviationsused in the text and in the figuresareL forlength, B for breadth, C for canine, I for incisor, P for
Fig. 2 Length and breadth distribution histograms showing the extent of the fragmentation of theWonderwerk Cave large vertebrate sample
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premolar and M for molar. Upper teeth are designated bysuperscript, lower teethbysubscriptanddeciduous teethbya ‘d’ in lower case.Measurements are given inmillimetres.
The entire collection is housed and curated in theMcGregor Museum, Kimberley, South Africa.
Systematic Description
The faunal sample described here comprises 448 speci-mens that were taxonomically identified at the family levelor to lower levels of taxonomy: 260 from Stratum 12 and188 from Stratum 11. Despite the small number of iden-tified remains, the sample represents seven mam-malian Orders (Table 1) and is dominated by
Bovidae (67 %), Procaviidae (12 %) and Equidae(11 %) (Table 2). In addition, there are innumera-ble indeterminate remains (tooth and long bonefragments) from both strata. The vast majority ofremains could not be identified such that a largeproportion of the specimens, especially of bovids,were identified only to size classes. Present alsoare remains of micro-mammals (Avery 2007), rep-tiles and birds which are not dealt with here.
Order RodentiaFamily HystricidaeGenus HystrixHystrix sp. cf. H. africaeaustralisMaterialStratum 12: WW150, incisor fragment; WW232,
RP4; WW126, LM1; WW266, RM2; WW192, molar
Fig. 3 Size distribution triplots showing the nature of the fragmentation of the Wonderwerk Cave macro-faunal sample
Description and ComparisonsWe assign the Wonderwerk hystricid specimens ten-
tatively to the extant form, based on size comparisons.In occlusal dimensions, the three Stratum 12 teeth thatare sufficiently complete to be measured fall within thevariation of extant H. africaeaustralis (Table 3; Fig. 5).As noted byWinkler et al. (2010), attribution of isolatedteeth is extremely difficult, since the occlusal morphol-ogy changes with the attrition, especially in the firststages of abrasion (Van Weers 1990). As is typical inporcupine systematics, identifications of isolated teethto tooth type were based primarily on size and
angulation. In Excavation 1, porcupines occur only inStratum 12, although characteristic porcupine damage isevident on bones from both Strata 12 and 11.
WW232 RP4: The tooth is complete except forthe roots, which show recent breakage, and is inearly wear stage (Stage VI, ca. 24–30 months; VanAarde 1985; Table 3). The entoconid (Fig. 5,
Fig. 4 A carnivore tooth puncture on specimen WW385, medialphalanx of an antilopine (A) and porcupine damage on specimenWW314, an unidentified bone fragment (photos: by authors)
Table 1 Mammalian remains recovered fromWonderwerk Cave,Strata 11 and 12, according to number of identified specimens(NISP)
Stratum 11 Stratum 12 Total
Rodentia
Hystrix sp. cf.H. africaeaustralis
12 12
Pedetes sp. 1 6 7
Lagomorpha
Lepus sp. cf. L. capensis 6 6
Pronolagus sp. 1 6 7
Indet. 1 7 8
Primates
Cercopithecidae indet. 1 1
Primates indet. 1 1
Carnivora
Canidae indet. 1 1 2
Hyaenidae indet. 3 3
Felidae indet. 1 1
Carnivores indet. 5 5
Hyracoidea
Procavia transvaalensis 1 10 11
Procavia antiqua 4 10 14
Hyracoidea indet. 5 21 26
Perissodactyla
Eurygnathohippus sp. 1 3 4
Equidae indet. 14 29 43
Artiodactyla
Bovidae
Tragelaphini indet. 1 1
Caprini indet. (large) 3 3 6
Alcelaphini indet. 17 11 28
Gazella sp. 1 1
Antilopini indet. 6 1 7
Bovidae Indet. (large) 7 7
Bovidae Indet. (large-medium)
64 53 117
Bovidae Indet. (small-medium)
47 57 104
Indet. (small) 9 16 25
Total 188 260 448
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character 1) and the central conid (Fig. 5, charac-ter 2) (sensu; Sen 2001) are still unfused with therest of the tooth, reflecting its early wear stage.
WW126 LM1: The crown is complete, enamel issmooth. The roots are absent and appear to have beenburned. It is in a mid-late wear stage with some fosettesworn away (Stage VII, ca. 30 months; Van Aarde 1985;Table 3). In general, the enamel ridges are simple inoutline and not crenulated. The hypoflexus has straight,almost parallel sides (Fig. 5, character 3). Theparafosette forms an island due to the late wearstage of the tooth (Fig. 5, character 4). The anterior mesoflexus forms an island and is joined tothe walls of the paracone and the mesoloph for thesame reason (Fig. 5, character 5). The postfosetteis evenly rounded, and the enamel is not crenulated(Fig. 5, character 6).
WW266 RM2: The tooth is in early-mid wear (StageVI, ca. 24–30 months; Van Aarde 1985; Table 3). The
outer enamel surface is preserved and smooth, but theinterior of the tooth is damaged.
The three measured teeth represent three different in-dividuals, because their attrition levels suggest differentontogenetic ages and as they were found at differentdepths within Stratum 12 and in non-adjacent squares.
Of the family Hystricidae, the genus Hystrix hasthe broadest range from China to Southeast Asia,Indonesia, Indo-Pakistan, the Mediterranean regionof North Africa, eastern and sub-Saharan Africa(Kingdon 1974; Nowak 1991). Two members ofHystrix are found today in Africa: Hystrix cristata,Linnaeus in North Africa and H. africaeaustralis,Peters in southern Africa.
The earliest record of hystricids in southern Africa isfrom Earliest Pliocene at Langebaanweg, but the mate-rials are not yet assigned to genus or species(Hendey 1981). Generally, the Plio-Pleistocene re-cord of Hystrix in southern Africa is disjunct(Winkler et al. 2010), but there is some fragmen-tary evidence for extinct and extant forms. Themajority of finds date to the Plio-Pleistocene andderive from the hominin-bearing breccia sites ofsouthern Africa. H. africaeaustralis has been re-corded at Sterkfontein Members 2 and 4–6(Maguire 1976; Ogola 2009), Taung (McKee1993), Swartkrans Members 1, 2 and 3 (Brain1981; De Ruiter 2003; Watson 2004), KromdraaiA and Makapan Limeworks Members 3, 4 and 5(Maguire 1976). However, two extinct fossil formshave also been identified at some of the samesites; Hystrix makapanensis, Greenwood (originallynamed Hystrix major), is a form that is about athird larger than the extant H. africaeaustralis, andhas been reported from Swartkrans Member 1,Makapan Limeworks Member 3 and 4 and
Table 2 Taxa identified at Wonderwerk Strata 11 and 12according to family
Family NISP %
Hystricidae 12 2.72
Pedetidae 7 1.59
Leporidae 21 4.76
Cercopithecidae 1 0.23
Canidae 2 0.45
Hyaenidae 3 0.68
Felidae 1 0.23
Procaviidae 51 11.56
Equidae 47 10.66
Bovidae 296 67.12
Total 441 100
Table 3 Measurements (mm) of Hystrix sp. dentition
Specimen Stratum ToothID
Attritiona Mesio-distallength (L)
Bucco-lingualbreadth (B)
Enamel height(H)b
RatioH/L
WW232 12 RP4 VI 6.8 5.5 11.4 1.68
WW126 12 LM1 VIII 7.8 6.9 10.4 1.39
WW266 12 RM2 VI 8.9 7.5 14.2 1.6
Length and width measurements correspond to the maximum mesio-distal length and bucco-lingual breadth of the teeth, while heightmeasurements correspond to enamel height (after Sen 2001)a Attritional stages according to Van Aarde (1985)b Height of enamel measured according to Van Weers (1990)
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Kromdraai A (Greenwood 1958; Maguire 1976,1978). Xenohystrix crassidens, Greenwood, whichis the largest Hystrix, but thought to have been aforest-dwelling form, based on its brachyodontdentition and assumed soft-diet, is also document-ed from Makapan Limeworks Members 3 and 4,dating to 3.7 and 2.5 Ma (Greenwood 1955, 1958;Herries et al. 2009; Maguire 1976, 1978).
In East African sites such as Laetoli, three species ofhystricids have been found: Hystrix leakeyi, H. cf.makapanensis and X. crassidens (Denys 1987).H. makapanensis is also recorded at the Pliocene sitesof Omo and at Olduvai Bed 1. X. crassidens also ap-pears at the Pliocene sites of Hadar (Denys 1987).
H. africaeaustralis is currently found in the in theenvirons of Wonderwerk Cave (Appendix 2) and was
Fig. 5 Porcupine specimens (a) and their size plots (b, c) in relation to extant Hystrix africaeaustralis (photos: by authors)
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also identified in the Holocene layers of the cave(Thackeray 2015), and there is clear evidence of itsactivity inside the cave in the form of characteristicallygnawed bones (Fig. 4).
Family PedetidaeGenus PedetesPedetes sp. cf. P. capensisMaterialStratum 12: WW683, RI1; WW705, I1 fragment;
WW727, I fragment; WW985, RM3; WW240, LM3
Stratum 11: WW590, I fragment; WW929, RM1
Description and ComparisonsAt Wonderwerk, scanty remains of springhare,
consisting only of isolated teeth, occur in Strata 12 and11. In morphology and in size, we cannot separate theWonderwerk specimens from modern comparative spec-imens of Pedetes capensis (Table 4; Fig. 6). However,given the paucity of the materials and their fragmentarystate, we assign the springhare specimens tentatively tothe extant species. We were able to determine the meristicposition of the teeth using Pickford and Mein (PickfordandMein 2011). In the upper jaw, the mesial loph is widerbucco-lingually than the distal loph, while the oppositeapplies to the lower premolars and molars. We couldattribute the lower third molar due to its angulation. Themeasurements follow Pickford and Mein (2011), with theaddition of measurements of both mesial and distal lophsand lophids at the occlusal surface.
In southern Africa, the genus Pedetes appears in theEarly Pliocene (Pickford and Mein 2011; Winkler et al.2010) and is widely recorded in the interior at sites, suchas Taung (Broom 1934), Swartkrans Members 1, 2 and 3
(De Ruiter 2003) and Cave of Hearths (Cooke 1963).Although the Florisbad Pedetes was described originallyas a distinct species, Pedetes hagenstadti (Dreyer andLyle 1931), it was later listed as Pedetes sp. (Brink1987). Springhare is found in the environs ofWonderwerk Cave still today (Appendix 2).
Order LagomorphaFamily LeporidaeGenus LepusLepus sp. cf. L. capensisMaterialStratum 12:WW875, Rmandible (P3–M1);WW290,
Rmandible (without teeth); WW748, Rmandible (with-out teeth); WW902, lower molar WW698, left pelviswith acetabulum; WW888, distal L metatarsal III.
Description and ComparisonsThe osteomorphology of southern African Leporidae,
and particularly the differentiation between hares, genusLepus, and the rabbits, genus Pronolagus (sensu Skinnerand Smithers 1990), is not well known. An additionalcomplication is the scarce riverine rabbit, Bunolagusmonticularis, which is in external morphology
Table 4 Measurements (mm) of Pedetes sp.
Specimen no. Stratum ToothID
Mesio-distallength (L)
Bucco-lingualwidth (B)
Mesialbucco-lingualdepth (Bm)
Distalbucco-lingualdepth (Bd)
WW683 12 I1 4.08
WW929 12 RM1 4.04 4.6 4.2
WW985 12 RM3 3.27 2.73 2.76
WW240 12 LM3 4.2 3.61 3.73
Pedetes capensisa
NMB-F9477 LM3 3.84 3.33 3.53
NMB-F842 LM3 4.08 3.35 3.41
NMB-F672 LM3 4.34 3.9 4.01
Length and breadth measurements are taken at the occlusal surface. In the premolars and molars, both the mesial (Bm) and distal (Bd) lobesare measureda Extant specimens from the Florisbad collections
Fig. 6 Pedetes cf. P. capensis: buccal, occlusal and lingual viewsof a LM3 WW240, mesial and distal views of a RI1, WW683(photos: by authors)
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intermediate between the hares and rabbits. Althoughknown as a ‘rabbit,’ it shares some characteristics withhares (Skinner and Smithers 1990). However, given thatthe Cape hare (Lepus capensis) and the scrub hare (Lepussaxatilis) are both larger than the members of the genusPronolagus, we could use size to assign the larger speci-mens to the genus Lepus and the smaller specimens to thegenus Pronolagus.
Based on the proportions of the mandibular depthand diastema length, we could demonstrate the likeli-hood of the presence of L. capensis (Table 5; Fig. 7). InL. capensis, there is a positive functional relationshipbetween mandibular depth and diastema length, but lessso in L. saxatilis, which reflects the more specialisedopen habitat and grazing niche of the former. In thisrespect specimenWW875 falls within the 95 % limits ofvariation of extant L. capensis, while specimenWW748plots just outside of these limits (Fig. 7). However, itshould be noted that the height of corpus of specimenWW748 is a conservative estimate (Table 5), since thespecimen is damaged in this region. The likelihood isthat the original height of this specimen would havebeen within the normal limits of variation of extantL. capensis. WW290 falls outside of this variation andits proportionally shallower corpus depth may point to it
being L. saxatilis. However, since we do not have agood understanding of evolutionary shifts that may haveaffected L. capensis since the Early Pleistocene in south-ern Africa, and because we do not have further evidencein support of L. saxatilis, we tentatively assign WW290also to the Cape hare.
Genus PronolagusPronolagus sp.MaterialStratum 12: WW124, glenoid of L scapula; WW799,
distal L humerus; WW90, distal R humerus; WW711,distal tibia; WW768, R calcaneus; WW144, Lcalcaneus.
Stratum 11: WW582 L calcaneusDescription and ComparisonsA small, very gracile lagomorph is represented by a
number of skeletal elements, including two distal hu-meri (WW799 and WW90) and a right and left calca-neus (WW768 and WW144; Table 5). These remainsare likely to represent a species of the rock rabbit, genusPronolagus. In Stratum 12, we identified both the gen-era Lepus and Pronolagus, while in Stratum 11 onlyPronolagus.
Pronolagus and Lepus co-occur in the Pleistocenedeposits at the Cave of Hearths, Limpopo Province,
Table 5 Measurements (mm) of Leporidae cranial and post-cranial elements
Specimen no. Stratum Element Length of diastema Height of corpus at P3 L Bm Bd
Lepus capensis
WW875 12 R mandible 20.8 11.5
P3 2.5 2.8 3.3
P4 2.8 3.0 3.1
M1 2.5 2.86 2.93
WW290 12 R mandible 21.0 10.3
WW748 12 R mandible 18.9 9.9a
WW698 12 L acetabulum 8.74 (LA) 6.8 (BA)
WW888 12 Distal L metatarsal III 4.83 (Bd) 4.88 (Dd)
In the premolars and molars both the mesial (Bm) and distal (Bd) lobes are measuredaMeasurement may be an estimatebMeasurement may be affected by slight damage to the element
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South Africa (Cooke 1963). In the Plio-Pleistocenehominin-bearing breccia sites of southern Africa, mostlagomorph remains have not been identified to genus orspecies, e.g., Sterkfontein Member 5, Swartkrans Mem-ber 2 and the Channel Fill, and Kromdraai A. Brain(1981) hypothesised that at least two forms were repre-sented in these sites—Pronolagus randensis, the redrock hare found in rocky habitats with mixed densebush and grass (Kingdon 1974; Skinner and Smithers1990), and L. capensis, the Cape hare that inhabits awide range of generally open habitats (Kingdon 1974;Skinner and Smithers 1990). At Wonderwerk, we
provide positive evidence for the presence of L. capensisand a species of Pronolagus, which may suggestthat the co-existence of a larger and smaller-bodiedleporid within a given ecosystem may have beencommon and wide-spread during the Early Pleisto-cene in central southern Africa.
Leporidae indet.MaterialStratum 12: WW167, molar fragment; 654, M/P
Fig. 7 Leporidae: a right lowerjaw of Lepus capensis, WW875,and size plots of fossil and extantlower jaws of Lepus spp. (photos:by authors)
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Stratum 11: WW654, premolar/molar fragment.The Leporid remains from the Holocene levels
of the cave were not identified to species,(Thackeray 2015), but today three species co-exist in the semi-arid Northern Cape in the vicinityof the cave; L. capensis, L. saxatilis and Pronolagusrupestris (Appendix 2).
Order PrimatesFamily CercopithecidaeCercopithecidae indet.MaterialStratum 12: WW1055, anterior distal phalanx (I).Description and ComparisonsPrimate remains are rare in Wonderwerk Cave. An
anterior distal phalanx of the first ray (thumb) has beenassigned to an indeterminate cercopithecid (Fig. 8).
Primates indet.Another distal phalanx (WW148) has been attributed
to an indeterminate primate.The Chacma baboon (Papio ursinus) occurs in the
area surrounding Wonderwerk, and occasionally troopsenter the cave in search of food (Neels Lehule, caretakerof cave, personal communication 2011). Papio remainswere also identified in the Holocene levels of the cave(Thackeray 2015).
Order CarnivoraFamily CanidaeCanidae indet.MaterialStratum 12: WW652, distal radius;Stratum 11: WW459, proximal ulna.Description and ComparisonsCanids are present in both Stratum 11 and 12, but in
low numbers. The distal radius, WW652, and the prox-imal ulna, WW459 (Fig. 9) appear slightly larger than
Canis mesomelas, but the specimens are too fragmen-tary to allow detailed metric comparisons.
Today, three canid species occur in the caves vicinity;bat-eared fox (Otocyon megalotis), Black-backed jackal(Canis mesomelas) and Cape fox (Vulpes chama). Onlyjackal was positively identified in the Holocene strata(Thackeray 2015).
Family HyaenidaeHyaenidae indet.MaterialStratum 11: WW952, incisor fragment; WW1390,
incisor fragment; WW1391, left P2Description and ComparisonsThe hyaenid specimens derive only from Stra-
tum 11, but they are too fragmentary to be iden-tified to genus or species (Fig. 10). However, theavailable morphology of the incisor fragment,WW1390, differs from extant hyaena species inhaving a root that is more robust and extendedlabio-lingually. The premolar fragment is too frag-mentary to identify more specifically. These spec-imens may represent an extinct form, but we can-not confirm this as yet.
Family FelidaeFelidae indet.MaterialStratum 12: WW983, Proximal phalanx (III).Description and ComparisonsWe could not assign the proximal phalanx to species,
which is intermediate in size but smaller than compara-tive specimens of the leopard, Panthera pardus(Table 6) and so could possibly represent caracal (Feliscaracal).
Though remains of both leopard and African wild cat(Felis libyca) were noted in the Holocene layers of the
Fig. 9 Canidae indet.: a lateral (a) and medial view (b) of aproximal ulna (WW459). The scale bar is 10 mm (photos: byauthors)
Fig. 8 Cercopithecidae: a proximal view (a), dorsal view (b) andvolar view (c) of an anterior distal phalanx I (photos: by authors)
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cave, currently no leopards survive in the region. Felidslisted by C. Anderson (1991a, b) in the region werecaracal, African wild cat and small-spotted cat (Felisnigripes), the latter noted as scarce.
Description and ComparisonsThe fragmentary carnivore materials can be
grouped into a larger and a smaller size category.The larger size category includes WW914 andWW952, while the smaller category includesWW1042 and WW889. The latter appears to bean indeterminate carnivore that is smaller than thejackal, C. mesomelas.
Order HyracoideaFamily ProcaviidaeGenus ProcaviaProcavia transvaalensis (Shaw 1937)
MaterialStratum 12: WW33, mandible plus dP4; WW97, I2;
Description and ComparisonsAt Wonderwerk, dental and post-cranial remains of
hyrax are common in both Strata 12 and 11. A total of 42hyrax remains were identified in Stratum 12. Cranialand post-cranial elements are both well represented. Thecranial remains comprise skull fragments, maxillaryfragments (one from an immature specimen), mandiblesand loose teeth. In Stratum 11, there are eight hyrax
Fig. 10 Hyaenidae: a mesial and frontal views of an incisorfragment, WW1390 (a) and a lingual and occlusal view of a leftP2, WW1391 (b) (photos: by authors)
Table 6 Measurements (mm) ofFelis sp. proximal phalanx (III) Specimen Stratum Element GL Bp Dp SD DD Bd Bd
remains, representing both cranial and post-cranial ele-ments (Tables 7 and 8; Fig. 11).
Two size classes of Procavia were identified inStratum 12. The larger hyrax agrees in size withP. transvaalensis and the smaller form agrees withP. antiqua. In Stratum 11, only the smaller P. antiquais found. Overall, hyrax remains are more abundantin Stratum 12 than in Stratum 11, but the volume ofdeposit excavated in the two strata differs, a featurethat may affect the size of the faunal samples andhence richness of specific species.
Today, three hyrax genera inhabit Africa and South-west Asia—Procavia, Dendrohyrax and Heterohyrax(Gheerbrant et al. 2005). In southern Africa, four hyracoidgenera existed in the Neogene: Procavia, Heterohyrax,Gigantohyrax and Prohyrax, with two genera and threespecies preserved in the Plio-Pleistocene record—Gigantohyrax maguirei, P. transvaalensis and P. antiqua(Broom 1934; Churcher 1956; Hendey 1978; Kitching1965; McMahon and Thackeray 1994; Meyer 1978;Pickford 1986, 2003; Rasmussen 1989; Rasmussenet al. 1996; Rasmussen and Gutiérrez 2010; Schwartz1997; Shaw 1937; Stromer 1926). P. antiqua andP. transvaalensis co-occur in the Plio-Pleistocenehominin-bearing breccia sites of southern Africa
(Rasmussen and Gutiérrez 2010), while G. maguirei isfound in the Plio-Pleistocene of southern Africa and Ethi-opia (Kitching 1965; Rasmussen and Gutiérrez 2010).
P. transvaalensis was first described by Shaw (Shaw1937) based on part of a skull and a right mandible fromSterkfontein. Compared with modern P. capensis, Shawnoted that in the fossil the teeth were considerablylarger, the lower incisors (I1 and I2) were almost equalin size, and the fossil retained the P1. The significantlysmaller P. antiqua was first described by Broom(1934) based on a partial cranium, posterior partof palate and a hemi-mandible from BuxtonLimeworks near Taung (Lectotype 4194A—Schwartz 1997). Compared with the extantP. capensis, this fossil form is of similar size orslightly smaller, less hypsodont, the M3 exhibits awell-developed hypocone and hypostyle, the M2
exhibits strong mesostyles, it has a bunodontprotocone, the I1 has a rounded posterior borderwhile in the upper molars the ectoloph length isrelatively shorter on the lingual border than in themodern form (Broom 1934; Churcher 1956;Schwartz 1997). At Wonderwerk, the smaller form,P. antiqua, is better represented in both Strata 12and 11 than P. transvaalensis.
Table 7 Measurements (mm) of Procavia transvaalensis cranial and post-cranial elements
Specimen no. Layer Element Mesio-distal length (L) Bucco-lingual breadth (B)
aMeasurement may be an estimatebMeasurement may be affected by slight damage to the element
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Procavia capensis remains were found in theHolocene strata of Wonderwerk Cave (Thackeray2015) and is a species listed as occurring in thecave’s vicinity today (Appendix 2)
Order PerissodactylaFamily Equidae
Genus Eurygnathohippus, (Van Hoepen 1930)Eurygnathohippus sp.MaterialStratum 12: WW223, LdI1 and LdI2; WW981, LM3;
WW116, lower R molarStratum 11: WW936, upper P fragment.
Table 8 Measurements (mm) of Procavia antiqua cranial and post-cranial elements
Specimen Layer Element Mesio-distal length (L) Bucco-lingual breadth (B)
WW592 11 M2 7.3 5.1
WW718 12 M3 5.5 3.6a
WW929 11 P4 4.0 4.7
WW790 12 Mandiblea
P2–4 13.2 –
P4 – 4.6a
WW332 11 Radius 9.2 (Bp) 4.7 (Dp)
WW780 12 Ulna 10.8 (DPA)
WW66 12 Tibia 10.7 (Bp) 11.4 (Dp)
WW73 12 Calcaneus 4.65 (Bd) 7.3 (Dd)
aWithout teeth
Fig. 11 Procaviidae: occlusaland buccal views of a LM2,WW267 (a) and a plot of lengthand breadth measurements offossil and modern Procaviaspecimens (b) (photos: byauthors)
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Description and ComparisonsWe follow Bernor et al. (2010) in referring African
hipparionines (three-toed equids) to Van Hoepen’s genus,Eurygnathohippus, which was first named from Cornelia-Uitzoek(VanHoepen1930).HipparionhasbeenpositivelyidentifiedinStratum12atWonderwerkandlesscertainly inStratum 11, using the dental criteria of Eisenmann et al.(1988) and Bernor et al. (2010). An upper third molar,WW981,has an isolatedprotocone,while a set of fragmen-tary lower deciduous incisors, WW223, have markedlycrenulated enamel, as seen inhipparionines.A lowermolaror premolar fragment, WW116, has a caballoid doubleknot. WW936 appears to have an isolated protocone, butis too fragmentary, so there is an element of uncertainty inthe diagnosis of this specimen.
Remains of hipparionines have been reported fromSwartkrans Members 1–3, Kromdraai A, and possiblyalso Gladysvale older deposits (Berger 1993; Brain1981; Churcher 1970). The last record of hipparioninesin southern Africa comes from Cornelia-Uitzoek and isdated to ca. 1 Ma (Brink et al. 2012).
The genus Equus first appeared in East Africa<2.3 Ma and co-existed there with hipparionines untilthe last appearance of hipparionines in East Africa atOlduvai Bed 4 (>0.78 Ma) and Bodo (0.6 Ma) (Cooke1983; Eisenmann 1979, 1983, 1992; O’Regan et al.2005; Tamrat et al. 1995). Current data indicate thatEquus capensis first appeared in the South Africanrecord ca. 1.0 Ma years ago at sites such as Cornelia-Uitzoek (Brink et al. 2012) and a little later atElandsfontein (Eisenmann 2000; Klein et al. 2007).However, E. capensis has been recorded at Bothaville(Broom and Le Riche 1937), Sterkfontein Member 4,Swartkrans Members 1–3, Kromdraai A, Minnaar’sCave, Gladysvale and Coopers D (Berger 1993;Berger et al. 2003; Brain 1981; Churcher 1970; DeRuiter 2009), where it may occur earlier than 1.0 Ma.It may be possible that early forms of Equus quagga(often referred to in the literature as ‘Equus burchellii’)also appear earlier than 1.0 Ma at sites, such asSterkfontein Members 4 and 5, Swartkrans Members1–3, Kromdraai A, Gladysvale older deposits and Coo-pers D (Berger 1993; Berger et al. 2003; Brain 1981;Churcher 1970; De Ruiter 2009). However, equids arenotoriously difficult to diagnose to species level, andwhere mainly size is used for diagnosis there may besome uncertainly.
Equidae indet.Material
Stratum 12: We could identify 29 equid dental re-mains to an unspecified equid category. It was notpossible to identify the genus Equus, although it is verylikely to be represented, given the time range of Strata12 and 11. At least seven teeth are represented; twodeciduous incisors, two deciduous lower premolars, adP4, an upper deciduous premolar and an upperpremolar.
Stratum 11: Fragments of 14 large-sized equid teethwere identified, which represent at least five differentteeth; two incisors, at least one lower deciduous premo-lar, a lower premolar and an upper premolar
Description and ComparisonsThe equid dental remains are poorly preserved
and very fragmentary in Strata 12 and 11, due inpart to the taphonomic effects of burning and thestructural nature of equid teeth. Equid teeth areabundant in both strata but especially in Stratum12. The majority of specimens are broken intosmall splinters such that their generic attribution,and in some cases even tooth type, is uncertain. Ina few cases we were able to refit some of thesplinters into incomplete teeth, but even these werenot identifiable to genus.
As noted by Thackeray (2015), Wonderwerkcave lies close to the biogeographic border be-tween E. quagga burchellii (the southwestern limitof this subspecies) and of E. quagga quagga (itsnorthern-most limit). In the Holocene layers, re-mains of at least two equid species were noted:E. capensis and remains of a smaller zebra, eitherE. quagga quagga or E. quagga burchellii, be-cause the dental measurements on the latter spec-imens were intermediate between these two taxa(Thackeray 1988, 2015). mtDNA analyses ofE. quagga burchellii and E. quagga quagga haveconfirmed that they are conspecific (Leonard et al.2005; Orlando et al. 2009). Today, there are nonatural populations of wild equids in the regionaround Wonderwerk Cave (Appendix 2).
Order ArtiodactylaFamily BovidaeTribe Tragelaphinicf. Tragelaphini indet.MaterialStratum11: WW1136, an unerupted upper premolar.Description and ComparisonsWe tentatively assign the specimen to the tribe
Tragelaphini, because of its simple occlusal enamel
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outline and being low crowned. It is the only specimenin the large-medium size class of bovids in the presentstudy sample that we could assign to this tribe.
molar fragments (in wear); WW892, upper dP fragments.Stratum 11: WW401, R mandible with dP2, dP3 and
dP4 (all unworn); WW1065, R mandibular symphysiswith RdC and dental fragments; WW629, dP4.
Description and ComparisonsMost of the caprine materials in Strata 12 and 11 are
from very young individuals, with only WW848representing fragments of an upper molar in early wear.
The best preserved specimen is from Stratum 11, ajuvenile right mandible with unworn dP2, dP3 and dP4(WW401). It was reconstructed from a number of small-er fragments, including WW404, 423 and WW423(Fig. 12). The corpus of the mandible appears moder-ately low and the dentition is similar in morphology toMakapania broomi from Makapan Limeworks Mem-bers 3 and 4 (Brink 1999; Gentry 1970; Wells andCooke 1957; Table 9; Fig. 12), but it is larger and similarin size to an unnamed large caprine from the Limeworksdeposits (JSB, personal observation). As illustrated inBrink (1999),M. broomi has well-developed mesial anddistal basal pillars in the dP4, which is a common featureof underived caprines. The Wonderwerk Cave caprinealso has mesial and distal basal pillars. However, it isless hypsodont thanM. broomi, judging from the depthof the mandible at the dP4/M1 junction. It is not yetpossible to identify the specimen positively to genus orspecies, but it may represent a large-bodied caprine,such as ‘Bos makapani,’ which occurs at Buffalo Cavein the Makapan Valley (Herries et al. 2006) and isconsidered to be related to the Bhutan Takin, genusBudorcas (Gentry 1996). It is noteworthy that otherfragmented specimens identified to this taxon all appearto belong to very young individuals, possibly of similarbiological age as specimen WW401.
There is a sparse record of Plio-Pleistocene andPleistocene caprines in sub-Saharan Africa. M. broomiis abundantly represented in the Makapan Limeworksdeposits (Gentry 1980; Reed 1996) and a large-bodiedcaprine, possibly M. broomi, also occur in small num-bers in the lower levels of the Olduvai sequence (Gentryand Gentry 1978; JSB, personal observation). InMiddleand Late Pleistocene deposits in southern Africa, a
smaller-bodied, as yet unnamed caprine replacesM. broomi (Brink 1999; Lacruz et al. 2003). TheWonderwerk caprine is larger than M. broomi and issimilar in size to the Makapan Limeworks unnamedcaprine (Fig. 12), which would be in accordance withthe Early Pleistocene age for Strata 12 and 11.
Tribe AlcelaphiniAlcelaphini indet.MaterialAlcelaphini are well represented in Stratum 11 (17
specimens) and slightly less so in Stratum 12 (11 spec-imens). Extreme fragmentation, as in the equids, doesnot allow attribution to genus or species.
Description and ComparisonsIt is noteworthy that alcelaphines are relatively
more abundant in Stratum 11 than in Stratum 12.Given the sparseness of fossil presence in Stratum11, this may have significance in reflecting semi-arid conditions, as suggested by the phytoliths(see, Rossouw 2016; this issue).
Tribe AntilopiniGenus GazellaGazella sp.MaterialStratum 11: WW 643, horncore fragments.Description and ComparisonsAntilopine horncore fragments show a solid horn
base, which is atypical for springbok, genus Antidorcas,which have inflated horn bases. Due to the fragmentarystate of the material, it is not possible to identify thespecies.
Gazelles occur in both southern and East Africa intimes predating ca. 1.0 Ma, with the only exceptionbeing the Elandsfontein main site (Klein et al. 2007),where gazelles are found probably as biogeographicremnants in a time younger than 1.0 Ma (Brink 2016).
Description and ComparisonsSpecimen WW385 falls within the range of
both Gazella and Antidorcas (Table 9). However,no remains of springbok have been positivelyidentified in either stratum.
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Bovid species common in the area of the cave at thetime C. Anderson wrote his report (Appendix 2) were
the common duiker (Sylvicapra grimmia), steenbok(Raphicerus campestris) and mountain reedbuck
Fig. 12 Caprini: a juvenile rightmandible,WW401,with unworn dP2,dP3 and dP4. The specimen was reconstructed from smaller fragments(a) to produce a virtually complete specimen (b). The bivariate plots
show that it is larger thanMakapaniabroomi fromMakapanLimeworksand similar in size to an unnamed caprine (M.6449) from theLimeworks (c). Thearrows (b) indicate basal pillars (photos: by authors)
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(Redunca fulvorufula). Together with Alcelaphines,these taxa are commonly found in the Holocenelayers of the site, as were springbok, eland, roanantelope, amongst others (Thackeray 2015). Asnoted before, the impoverished spectrum of bovidsfound in the region today is due both to huntingas well as the expansion of farming areas at theexpense of natural habitats.
Discussion
Despite its small size, the diagnostic faunal com-ponent from Strata 12 and 11 at Wonderwerk Cavefills an important hiatus in our knowledge of thearid interior of South Africa ca. 2 Ma years ago.Bovids make up the bulk of the large-mediummammalian fauna at Wonderwerk, being morecommon than equids and other forms. Most bovidsbelong to the tribe Alcelaphini, and only a fewspecimens are referred to caprines and antilopines.It is significant that no specimens were identifiedto the tribes Reduncini or Hippotragini. This andthe absence of suids, may be seen as further sup-port for the overall semi-arid character of thefaunas from both Strata 12 and 11 at WonderwerkCave.
The phytolith (see Rossouw 2016, this issue) se-quence shows a distinct shift from a warm Savanna/Nama Karoo C4 grassland environment at the bottomof Stratum 12, to more arid and cooler conditionsanalogous to a Succulent Karoo (C3-dominated)
environment at the top of Stratum 12 through Stratum11 (Chazan et al. 2012). Stable isotope values forostrich eggshell from these layers support this trend(Ecker et al. 2015). Further support is found in themicro-mammal assemblage. St ra tum 12 ischaracterised by higher proportions of Gerbillinae/(gerbils) to Murinae (mice), reflecting a drier andmore open environment, while Stratum 11 containedhigher frequencies of Soricidae (Fernandez-Jalvo andAvery 2015). Despite the small sample size of iden-tified material, this shift may be expressed in themacro-faunal spectrum, as alcelaphine bovids are rel-atively more abundant in Stratum 11 than in Stratum12.
The archaic character of the large mammalian faunasof Strata 12 and 11 is reflected by the presence of twospec i e s o f ex t i nc t hy r ax (P. an t i qua andP. transvaalensis) known from several early and mid-Pleistocene sites in the Cradle of Humankind, aswell as a large-bodied caprine, similar to an un-named large caprine from Makapan Limeworks. Inbody size, these caprines would have been close tobovines, such as Simatherium kohllarseni(Fig. 12). The presence of large caprines is acommon feature of the geologically older assem-b l age s f r om Makapan L imework s , f r omSterkfontein and from the basal levels atSwartkrans. At Swartkrans large-bodied caprinesmake place for smaller-bodied forms more typicalof the Middle Pleistocene (JSB, personal observa-tion). The palaeo-magnetic and cosmogenic burial ageestimates for Stratum 12, which suggest an age
Table 9 Measurements (mm) of Bovidae cranial and post-cranial elements
of around the Olduvai Subchron, would be inagreement with the archaic character suggested bythe large-bodied caprine. It is now possible toplace the large mammal faunas of Strata 12 and11 in a revised biochronological position withinthe Makapanian Land Mammal age.
The salient ecological character of the fauna is that oftaxa adapted to a semi-arid, savanna conditions. This isattested to by the remarkable continuity of several small-bodiedmammalian species present in Stratum 12 as wellas in the region today; H. africaeaustralis, the presenceof both Lepus and Pronolagus spp. and the presence ofPedetes cf. P. capensis. This is not surprising as almostall of the micro-mammal species identified in Strata 12and 11 still inhabit the Northern Cape, though in somecase at some distance from the site (Fernandez-Jalvo andAvery 2015).
It is further noteworthy that the mammal faunasfrom Strata 12 and 11 reflect two distinctecozones; the plains further to the east of the caveand the broken and montane habitat of the moun-tainous area to the west of the cave (see also,Brink et al. 2015). The abundance of hyrax andthe presence of the large caprine clearly reflect thelocal montane habitat, while equids, alcelaphinesand antilopines are more typical inhabitants of thesemi-arid grassland-savanna habitat, of the plainsto the east of the cave. The basal layers reflect agenerally semi-arid palaeo-environment, given theprominence of equids and the absence of wetlandindicators, such as hippopotamuses suids, hippotraginesand reduncines. This is corroborated by the microfaunal and phytolith records from WonderwerkCave (Chazan et al. 2012; Rossouw 2016; this issue)and is in contrast with the evidence for generally moremesic conditions evident in the Middle Pleistocenemammal faunas of the interior of southern Africa(Brink 2016). However, it is likely that the mammalianfauna from Strata 12 and 11 are of low resolution indiscerning subtler palaeo-environmental changes—suchas the shift to more mesic conditions at the top ofStratum 12 identified in the phytolith record at the site(Rossouw 2016; this issue), given that the samples aretime averaged, and that the Stratum 11 assemblage isextremely small.
An important factor in the reduced level oftaxonomic resolution in the large mammal remains
from Strata 12 and 11 is the extent to which thematerials were burned in situ (vide; Berna et al.2012). This, as well as other taphonomic agentsnoted above, caused substantial in situ crackingand fracturing, which subsequently was exacerbat-ed by excavation without in situ consolidation ofthe specimens. In such conditions of preservation,it is essential that in situ consolidation of verte-brate materials should take place and if this hadbeen done, the level of taxonomic resolutionwould have been substantially improved. As it is,we have a reasonably clear impression of the tax-onomic representation of Strata 12 and 11, but it istantalisingly clear that much more would havebeen gleaned from the materials if in situ consol-idation had been undertaken. It serves as a remind-er for further work at the cave and at other similarsites.
Conclusions
The importance of the vertebrate remains fromStrata 12 and 11 at Wonderwerk cannot beoveremphasised, since at the present they are theonly samples in the interior of southern Africadating to the Early Pleistocene outside of thehominin-bearing breccias of Taung, Gauteng andMakapan. The relative ages of Stratum 12 and11 at Wonderwerk Cave are known due to lithictechnology (Chazan 2015; Chazan et al. 2008) andradiometric dating (Matmon et al. 2012). Some ofthe taxa l i s t ed in Tab le 1 have l imi tedbiochronological significance since they are ubiqui-tous over long time spans. However, the hyraxesand large caprine accords with an early to Mid-Makapanian age.
By and large the bovids, in particular alcelaphines,comprise the bulk of the sample. Together with theequids and in the absence of reduncines, hippotraginesand suids, they indicate semi-arid palaeo-climatic con-ditions during the early part of the Early Pleistocene,in support of the phytolith evidence and isotope evi-dence. On a local scale, the equids and alcelaphinesreflect taxa of the plains habitat to the east of the cave,while hyraxes and the caprine reflect the broken andmontane habitat of the Kuruman Hills to the west.
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Acknowledgements We wish to express our special thanksfor their assistance in various aspects of this project toProf. Michael Chazan, co-director of the Wonderwerk pro-ject; Mr. Colin Fortune (now retired, Director), Dr. DavidMorris (Head of Archaeology Department), Dr. LeonJacobson (now retired, Assistant Director) and the staff ofthe McGregor Museum; Mr. Rick Nuttall (Director), Dr.Nico Avenant (Head of the Mammalogy Department) andthe staff of the National Museum, Bloemfontein; Dr.Berhard Zipfel, Institute for Human Evolution, Universityof the Witwatersrand, for access to the MakapanLimeworks fossil assemblage; Ms. Stephany Potze andMr. Lazarus Kgasi, Department of Palaeontology, DitsongNational Museum of Natural History (previously the Trans-vaal Museum), for access to the Swartkrans fossil assem-blages; and Mr. Willem Carel Brink for translating the
abstract into French. JSB would like to thank the NRFfor financial support (grant no. 82603) and Prof. AndyHerries, La Trobe University, Australia, for fruitful discus-sions on geochronology. The authors thank two anonymousreviewers for their comments. The authors would like toacknowledge the contribution of the excavator of theWonderwerk faunal assemblage, Mr. Peter Beaumont.
Compliance with Ethical Standards
Conflict of Interest The authors declare that they have noconflict of interest.
Appendix 1: Checklist of Veld Types Found in a Field Survey in the Immediate Area Surrounding Wonderwerk Cave
From: Anderson, T. 1991. Wonderwerk Cave as a Botanical Reserve. UnpublishedReport. McGregor Museum Archive.
A. Karroid Veld – Orange River Broken Veld (9)
The vegetation is sparse and open, but contains a rich flora.
C. Shrubveld- Tarchonanthus veld of the Ghaap Plateau
Consists of essentially the same species as in Category B but has: far more Tarchonanthus camphorates shrubs, a few more grass species and fewer species of trees, and in places with parklands of Olea europaea subsp. africana.
* exotic species
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Appendix 2: Checklist of Mammalian Species Inhabiting the Semi-arid Northern Cape Region in the Area Surrounding
Wonderwerk Cave
From: Anderson, C. 1991. The Birds and Mammals of the Wonderwerk Cave Area.Unpublished Report. McGregor Museum Archive.
Acocks, J. P. H. (1988). Veld types of South Africa.Memoirs of theBotanical Survey of South Africa, 57, 1–146.
Anderson, C. (1991). The birds and mammals of the WonderwerkCave area. Unpublished Report. McGregor Museum Archive
Anderson, T. (1991). Wonderwerk Cave as a botanical reserve.Unpublished Report. McGregor Museum Archive
Avery, D. M. (2007). Pleistocene micromammals fromWonderwerk Cave, South Africa: Practical issues. Journalof Archaeological Science, 34, 613–625.
Beaumont, P. B. (1990). Wonderwerk Cave. In P. Beaumont & D.Morris (Eds.), Guide to archaeological sites in the NorthernCape (pp. 101–134). Kimberly: McGregor Museum.
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