Journal of Archaeological Science 40 (2013) 3674e3685

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Journal of Archaeological Science

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Identification of hairs in Parahyaena brunnea coprolites from Middle Pleistocene deposits at Gladysvale cave,

Phillip Taru a,*, Lucinda Backwell a,b a Bernard Price Institute for Palaeontological Research, School of Geosciences, University of the Witwatersrand, Private Bag 3, Wits 2050, Johannesburg, South Africa b Institute for Human Evolution, University of the Witwatersrand, Private Bag 3, Wits 2050, Johannesburg, South Africa article info abstract

Article history: This research focuses on scale pattern and cross sectional morphology of hair to identify an expanded Received 18 February 2013 sample of fossil hairs from Parahyaena brunnea coprolites from Gladysvale cave in the Sterkfontein Valley, Received in revised form South Africa. The coprolites are part of a brown hyaena latrine preserved in calcified cave sediment dated to 26 April 2013 the Middle Pleistocene (257e195 ka). Forty-eight fossil hairs were extracted from 12 coprolites using fine Accepted 29 April 2013 tweezers and a binocular microscope, and examined using a scanning electron microscope. Hair identi- fication was based on consultation of standard guides to hair identification and comparison with our own Keywords: collection of samples of guard hairs from 15 previously undocumented taxa of indigenous southern African Fossil hair Coprolites mammals. Samples were taken from the back of pelts curated at the Johannesburg Zoo and Ditsong Na- Brown hyaena tional Museum of Natural History (formerly Transvaal Museum, Pretoria). Based on the fossil hairs iden- Scale pattern tified here, this research has established that brown hyaenas shared the Sterkfontein Valley with hominins, Scanning electron microscopy warthog, impala, zebra and kudu. Apart from humans, these animals are associated with savanna grass- lands, much like the Highveld environment of today. These findings support the previous tentative identification of fossil human hair in the coprolites, provide a new source of information on the local Middle Pleistocene fossil mammal community, and insight into the environment in which archaic and emerging modern humans in the interior of the African subcontinent lived. Ó 2013 The Authors. Published by Elsevier Ltd. Open access under CC BY license.

1. Introduction Many chemicals and biological substances that accumulate in hair can be detected and measured and this makes hair samples Hair is a unique feature of mammals (Inagaki, 1986), the iden- good resource biomaterials in forensic science and physical an- tification of which provides a wealth of information to , thropology (Chang et al., 2005). Furthermore, the basic chemical epidemiology, ecology, criminology and forensic investigations. composition of hair is not affected by changes in blood chemistry or Hair morphology has been used to identify mammals to order, by exposure to chemicals after hair formation (Daniel et al., 2004). family or genus level since the beginning of this century (Hausman, Because of this, hair samples are often used for autopsy toxicology, 1920; Cole, 1924), and even though much is known about European including the detection of drug abuse, personal identification and mammalian hair (Brunner and Coman, 1974; Moore et al., 1974; the forensic genetic identification of relatives (Zaiats and Ivanov, Brothwell and Spearman, 1963; Brothwell, 1993; Teerink, 2003; 1997; Lebedeva et al., 2000). According to Wilson et al. (2007), Wilson et al., 2004, 2007; Wilson, 2005), relatively little research the hair shaft does not undergo any post-keratinization biogenic has been conducted on the morphology of fossil and modern change in contrast to bone and teeth, which are commonly ana- southern African animals. lysed human tissues in bioarchaeology. Hair is mostly made up of the fibrous protein keratin, which is extremely resistant to decomposition (Chang et al., 2005) and enzymatic digestion (Lubec et al., 1986, 1994), owing mainly to the presence of disulphide cross linkages of the amino acid cystine (Brothwell and Spearman, 1963; Taylor et al., 1995). Such an intensely cross-linked system is extremely resistant to decay (Brothwell, 1993), leading to the * Corresponding author. Tel.: þ27 11 717 6682; fax: þ27 11 717 6694. E-mail addresses: [email protected] (P. Taru), [email protected] preservation of hair. Hair proteins are hydrophobic in nature (Fraser (L. Backwell). et al., 1972) and this renders them insoluble in water, dilute acids

0305-4403 Ó 2013 The Authors. Published by Elsevier Ltd. Open access under CC BY license. http://dx.doi.org/10.1016/j.jas.2013.04.031 P. Taru, L. Backwell / Journal of Archaeological Science 40 (2013) 3674e3685 3675 and alkali, and various organic solvents at ambient temperatures study follows the recent discovery of possible human hair in a (Taylor et al., 1995). Because of this, hair is relatively durable in the single Parahyaena brunnea (brown hyaena) coprolite from Gla- natural environment, and unlike bone, does not fragment during dysvale cave, South Africa (Backwell et al., 2009). The coprolite is digestion (Keogh, 1979). While the properties of hair may enable it part of a brown hyaena latrine preserved in calcified cave sediment to survive for millennia, under unfavourable conditions, it can dated to the Middle Pleistocene (257e195 ka) (Pickering et al., degrade within a few weeks (Wilson, 2005; Wilson and Gilbert, 2007; Berger et al., 2009). Element analysis showed total replace- 2007). ment of keratin by calcium carbonate, confirming that the Gla- A review of the scientific literature reveals that relatively few dysvale hairs are high-resolution casts and do not preserve amino examples of prehistoric animal hair exist, mostly because of a lack acids (Backwell et al., 2009). Nonetheless, the exceptional preser- of suitable preservation conditions (Bonnichsen et al., 2001). The vation afforded by the dolomitic cave system, and potential for oldest fossil hairs are reported from China, from Early Cretaceous expanding the range of known fauna for this time period, stimu- (146 - 100 Ma) deposits in the Yixian Formation, where they are lated our interest in studying an enlarged sample of coprolites from preserved as carbonized filaments and impressions (Luo et al., the same deposit. Our aim was to enlarge the Gladysvale fossil hair 2003), and Late Palaeocene (w59e56 My) beds, where the hairs sample and discuss the implications of our findings for Middle are preserved as calcium carbonate filaments in carnivore faeces Pleistocene hyaena ecology and palaeoenvironment in the Sterk- (Meng and Wyss, 1997). Younger hairs are reported from Eocene fontein Valley, South Africa. (w50 Ma) deposits in the Messel Shales in Germany (Schaal and Ziegler, 1992; Schweitzer, 2011), Late Pleistocene (w50,000e 1.1. Archaeological context of Gladysvale cave 17,000 BP) permafrost deposits in Siberia (Gilbert et al., 2007) and Miocene (w20e15 My) amber from the Dominican Republic Gladysvale cave is located in the Eccles Formation of the Mal- (Poinar, 1988). Human hair samples over 8000 years old are asso- mani Subgroup of the Transvaal Supergroup (Berger et al., 2009) ciated with mummies from Peru (Benfer et al., 1978), and more and situated on the edge of mixed savanna (Scholes, 1997) and recent Roman hair has been found in a lead coffin burial from grassland (O’Connor and Bredenkamp, 1997) biomes. The site Dorchester (Green et al., 1981) and a Roman Ermine street site in (Fig. 1) is well known for yielding a rich Plio-Pleistocene fauna, Cambridgeshire, England (Brothwell, 1993). including specimens attributed to Australopithecus africanus Hairs found in archaeological sites are a unique resource for (Berger, 1993; Berger et al., 1993). A wealth of large vertebrate capturing a snapshot of life (Chang et al., 2005), and the (Berger, 1992; Lacruz et al., 2002) and micro-faunal remains morphology of hair such as scale pattern, medulla, cross-section (Avery, 1995), including diverse avian fauna (Stidham, 2004)have and colour patterns has provided significant information on spe- been reported from the site. Gladysvale is a complex cave system cies represented, for example, Brothwell and Grime (2003) showed made up of several underground chambers reaching a depth of that the hairs associated with the Neolithic ‘Iceman’ from the Alps about 65 m (Martini and Keyser, 1989; Schmid, 2002). The cave were that of a red deer, and that the fur armband on the Iron Age complex is made up of a roofed system of large underground caves Lindow bog body was made from fox (Budworth et al., 1986). referred to as the Gladysvale Internal Deposits (Pickering, 2005; Ancient mammalian hairs also provide insight into a site’s function, Pickering et al., 2007), and an outer de-roofed area known as the the nature of the environment, species evolution, and the relation Gladysvale External Deposits (Lacruz, 2002; Lacruz et al., 2002). between people and animals in the past (Davis et al., 2007). Ancient The internal deposits are clearly exposed, well preserved and human and animal hair can be an important data source for un- stratified and this makes them unusual for caves in the region. The derstanding palaeobiology, palaeoecology and palaeoanthropology strata consist of clastic sediments, ranging from coarse boulder (Bonnichsen et al., 2001), but unfortunately it is rarely preserved in piles, through to medium grained, sandy sediment to fine grained, the fossil record and researchers seldom attempt to find it. This often laminated muds (Pickering, 2005). Flowstones that act as

Fig. 1. Map of southern Africa showing Gladysvale cave (a), and geological section of Gladysvale deposits showing the position of the latrine within flowstone bounded unit 14 (b). Scale bar in b ¼ 30 cm (modified after Backwell et al., 2009). 3676 P. Taru, L. Backwell / Journal of Archaeological Science 40 (2013) 3674e3685 chronostratigraphic markers are interbedded within the strata and Table 1 are used to divide the sediments into flowstone bounded units Coprolites and hairs from Gladysvale cave analysed in this study. (FBUs) (Pickering et al., 2007). The fossil hyaena latrine is preserved Coprolite Diameter Shape No. of hairs Hair specimen Fig. no. in FBU 14 (Fig. 1), which has well developed lower flowstones and a no. (mm) extracted no. small stalagmite at the base of the unit, constraining the age of the C1 21 Ovoid 4 F1 7a,b latrine to between 195 and 257 ka (Pickering et al., 2007). The C2 19 Ovoid 5 ee position of the latrine is directly under the major palaeo-drip C3 8 Ovoid 2 F2 7c source of the cave, providing calcium carbonate-rich water, which C4 11 Spherical 4 F3 and F5 7d, 8a C5 10 Spherical 3 ee aided in the fossilization and preservation of the latrine and sur- C6 15 Spherical 5 F7 8c,d rounding sediment (Berger et al., 2009). C7 20 Spherical 4 ee

C8 32 Spherical 7 F8, F9, F11 8e, 9a,c C9 19 Spherical 4 F10 9b 2. Materials and methods C10 18 Spherical 4 ee

C11 10 Ovoid 3 F6 8b In this research, a block of the calcified latrine (Fig. 2) was taken C12 8 Spherical 3 F4 7e from Gladysvale cave for laboratory analysis. From this block, 12 C1eC12 refers to coprolite specimens, and F1 e F11 refers to fossil hair specimens. e coprolites were studied (Table 1). The coprolites varied in size, with indicates that the fossil hairs had extremely degraded and unidentifiable scale larger scats approximately 32 mm in diameter. The number of patterns, which are not shown here. coprolites analysed in this study were deemed sufficient to deter- mine the diet of Parahyaena brunnea considering that previous Guard hairs were selected because they give the most characteristic dietary descriptions were successfully achieved with 10 scats and distinct scale patterns and cross section shapes among (Pontier et al., 2002; Sinclair and Zeppelin, 2002; Zabala and different mammals (Keogh, 1979; Kondo, 2000), and because these Zubergogoitia, 2003). Forty-eight fossil hairs were extracted from are the hair types presented by other researchers (e.g. Williams, fi fi the coprolites using ne tweezers and a low magni cation binoc- 1938; Stoves, 1942; Mayer, 1952; Stains, 1958; Keogh, 1979, 1983; ular microscope. Thereafter, they were ultrasonically cleaned with Teerink, 2003; Backwell et al., 2009), hence a need to standardize analar ethanol and placed directly onto double-sided sticky stubs. for comparative purposes. Furthermore, guard hair is much more The fossil hairs were sputter- coated with gold and examined using regular in shape and easier to work with than fine underhair fi an FEI Quanta 400 E scanning electron microscope at magni ca- (Homan and Genoways, 1978). Prior to examination, the hair tions between 461 and 2538 times. Fossil hair cross sections were samples were ultrasonically cleaned with a mixture of absolute obtained only from naturally occurring breaks in the specimen. alcohol and sulphuric ether in equal proportions to remove the fi Hair identi cation was based on consultation of standard guides organic and inorganic dirt from the surface of the samples. The fi to hair identi cation by Hausman (1930), Lyne and McMahon hairs were washed in distilled water for three minutes and then air (1951), Appleyard (1978), Keogh (1979, 1983), Teerink (2003), and dried on a clean watch glass. To obtain cross sections, the hair comparison with our own collection of samples of guard hairs from samples were cut perpendicularly using a fine surgical blade. 15 taxa of indigenous southern African mammals (Table 2). The Finally, the hairs were mounted on stubs with double-sided sticky selection of these 15 animals was based on known Middle Pleis- tabs, sputter-coated with gold and examined using an FEI Quanta tocene fauna of the Florisian Land Mammal Age (Klein, 1980, 1984) 400 E scanning electron microscope following the same procedures reported for the Sterkfontein Valley (Lacruz et al., 2002, 2003), and as those used for the fossil specimens. mammals needed for comparison that are not represented in hair identification literature. Terminology used to describe hair features 3. Results follows that outlined by Keogh (1979), Teerink (2003) and Backwell et al. (2009). Guard hair samples were taken from the back of pelts 3.1. Modern comparative collection curated at the Johannesburg Zoo and Ditsong National Museum of Natural History in Tshwane (formerly Transvaal Museum, Pretoria). A selection of scanning electron micrographs of hairs from previously undocumented modern southern African mammal taxa is presented in Figs. 3e6. The images of hairs presented have scales

Table 2 Modern comparative mammals from which guard hair samples were collected.

Order Genus Species Common name

Perissodactyla Diceros bicornis Black rhinoceros Equus quagga (formerly Burchell’s zebra burchelli) Artiodactyla Antidorcas marsupialis Springbok Aepyceros melampus Impala Tragelaphus strepsiceros Greater kudu Connochaetes gnou Black wildebeest Connochaetes taurinus Blue wildebeest Syncerus caffer Cape buffalo Taurotragus oryx Eland Phacochoerus aethiopicus Warthog Raphicerus campestris Steenbok Lagomorpha Lepus saxatilis Scrub hare Pronolagus crassicaudatus Red rock hare Hyracoidea Procavia capensis Rock dassie fi Fig. 2. Block of calci ed cave sediment containing part of the hyaena latrine, showing Tubulidentata Orycteropus afer Aardvark the largest coprolite analysed (specimen C8). Scale ¼ 10 cm. P. Taru, L. Backwell / Journal of Archaeological Science 40 (2013) 3674e3685 3677

Fig. 3. Modern impala hair cross section (a), scale ¼ 30 mm and scale pattern (b), scale ¼ 40 mm; eland hair cross section (c) and scale pattern (d), scales ¼ 50 mm; steenbok hair cross section (e), scale ¼ 50 mm and scale pattern (f), scale ¼ 100 mm. ranging from 30 mmto100mm, to permit better observation of scale generally biconvex. The medulla is large but indistinct, with a thin features and independent assessment of hair samples. In cross cortex. The scale pattern of kudu hair is irregular waved mosaic and section, impala hair has a diagnostic triangular shape with blunted scale margins are generally smooth (Fig. 4b). The distance between corners. The cortex is thick and the medulla has numerous small scales is near to distant. In cross section, warthog hair is oval perforations (Fig. 3a). The scales are transverse relative to the (Fig. 4c). The scale pattern is highly irregular waved mosaic. The longitudinal axis of the hair. Scale pattern is irregular waved scale margins are rippled and the distance between scales is mosaic. The structure of scale margins is generally smooth and the extremely close (Fig. 4d). The cross section of Burchell’s zebra is distance between scale margins is near to distant (Fig. 3b). The oblong (Fig. 4e). The scale pattern is irregular waved mosaic and the cross section of eland hair (Fig. 3c) is generally circular tending to scale margins are rippled. The distance between scale margins is oval. The cortex is very thick and the medulla is small to medium in near to close (Fig. 4f). Scrub-hare hair cross section (Fig. 5a) is size. The scale pattern (Fig. 3d) is irregular waved mosaic and the generally biconvex to oval. The medulla usually contains a few large scale margins are moderately rippled. The distance between scales cavities. The scales form a lanceolate pectinate (comb-like) pattern. is near. In cross section, steenbok hair (Fig. 3e) is generally reniform The distance between scales is distant and scale margins are tending to concavo-convex. The medulla is very large and spongy. moderately rippled (Fig. 5b). Red rock hare hair cross section is The scales are transverse and the scale pattern is irregular waved dumb-bell shaped (Fig. 5c). The medulla contains large cavities. mosaic. Scale margins are smooth and the distance between scales There is a broad, deep groove along the length of the hair. Both is near to distant (Fig. 3f). The cross section of kudu hair (Fig. 4a) is sides of the groove show a coronal scale pattern. The distance 3678 P. Taru, L. Backwell / Journal of Archaeological Science 40 (2013) 3674e3685

Fig. 4. Modern kudu hair cross section (a) and scale pattern (b), scales ¼ 50 mm; warthog hair cross section (c) and scale pattern (d), scales ¼ 100 mm; Burchell’s zebra cross section(e) and scale pattern (f), scales ¼ 50 mm. between scales is near and the scale margins are smooth (Fig. 5d). identification impossible, whilst eight were identifiable to five Black rhino hair cross section is approximately oval and the me- species, and seven could not be identified. Some fossil hairs showed dulla is ill-defined (Fig. 5e). Scales are very coarse and the pattern is reasonably clear scale morphologies but could not be conclusively irregular waved. The scale margins are moderately smooth to matched in the comparative collections. These hairs may represent rippled and the distance between scales is near (Fig. 5f). Blue taxa for which we do not have a modern comparative sample, or wildebeest hair cross section (Fig. 6a) is concavo-convex. The cortex the hair of a mammal now extinct. Considering that few fossil hairs is thick and the medulla is small to medium in size. Scale position is are preserved in the African fossil record to enable comparison with transversal and scale pattern is irregular waved mosaic. The scale those from Gladysvale, we are obliged to use modern taxa as a margins are generally smooth to moderately rippled and the dis- proxy for fossil forms. A selection of scanning electron micrographs tance between scales is near (Fig. 6b). Rock dassie hair cross section and descriptions of fossil hairs is presented in Figs. 7e9. In fossil is approximately oval although it tends to oblong (Fig. 6c). The scale hair specimen 1, the scale pattern is imbricate, i.e. the cuticular pattern is ill-defined and there are cracks on the cuticular surface scales overlap. The scales lie transverse to the longitudinal direction (Fig. 6d). of the hair and the scale margins are generally smooth to moder- ately rippled (Fig. 7b). In cross section, the hair is generally oval and 3.2. Fossil hairs shows no lumen (Fig. 7a). The medulla is amorphous, lacking a definite shape, a feature of human hair. This combination of char- Of the 48 fossil hairs extracted from the 12 coprolites, 33 were acters closely resembles those observed on modern African and extremely degraded with ill-defined scale patterns, rendering hair European hair samples, and fossil human hairs reported by P. Taru, L. Backwell / Journal of Archaeological Science 40 (2013) 3674e3685 3679

Fig. 5. Modern scrub-hare hair cross section (a), scale ¼ 30 mm, and scale pattern (b), scale ¼ 50 mm; red rock hare hair cross section (c) and scale pattern (d); Black rhino hair crosssection (e) and scale pattern (f), scales ¼ 50 mm.

Backwell et al. (2009) from a single coprolite from the same de- rippled. The cross section of these specimens is not available due to posit. The scale morphology of fossil hair specimens 2 (Fig. 7c), 3 the absence of naturally occurring breaks in the samples. The dis- (Fig. 7d and 4 (Fig. 7e) is slightly obscured, but a closer examination tance between fossil hair scale margins is near to close, a scale shows an irregular waved mosaic pattern and smooth scale mar- pattern comparable to that of modern Burchell’s zebra (Fig. 4f). The gins that are near to distant. This combination of features resembles hair scale morphology of fossil specimen 7 is not preserved (Fig. 8c). those found in modern impala (Fig. 3a and b). Even though the In cross section, it is generally biconvex to oval (Fig. 8d), a feature cross section of fossil hair specimen 2 (Fig. 7c) could not be ob- shared with modern kudu (Fig. 4a). Although the cross section of tained because of the absence of naturally occurring breaks in the fossil hair specimen 8 (Fig. 8e) is distorted, it is reniform or sample, the fossil hair specimen can be tentatively attributed to concavo-convex, a shape similar to that of modern steenbok hair modern impala based on scale morphology. In cross section, fossil (Fig. 3e). The scale pattern of fossil hair specimen 8 is poorly pre- hair specimen 4 (Fig. 7f) is a triangle with blunted corners, a feature served, but nonetheless shows an irregular waved pattern (Fig. 8f). shared with modern impala (Fig. 3a). There are no clear small Unlike modern steenbok scale margins, which are smooth (Fig. 3f), perforations in the fossil medulla, which is to be expected from a the fossil hair scale margins are moderately rippled. The scale cast. Although ill-defined, the scale pattern of fossil hair specimen 4 morphology of fossil hair specimen 8 (Fig. 8f) could not be (Fig. 7e) is irregular waved mosaic, and the scale margins are conclusively matched in the comparative collections, and hence an smooth, as in modern impala (Fig. 3b). identification could not be made. Fossil hair specimens 9 (Fig. 9a) Scale morphology of specimens 5 (Fig. 8a and 6 (Fig. 8b) is and 10 (Fig. 9b) show an irregular waved mosaic scale morphology, irregular waved mosaic and the scale margins are moderately and the distance between scale margins is close. Scale margins are 3680 P. Taru, L. Backwell / Journal of Archaeological Science 40 (2013) 3674e3685

Fig. 6. Blue wildebeest hair cross section (a) and scale pattern (b); Rock dassie hair cross section (c) and scale pattern (d), scales ¼ 50 mm. rippled. Although the cross section of these fossil hair specimens is as guard hairs, under hairs, and whiskers from a variety of species not available due to the absence of naturally occurring breaks in the (Bonnichsen et al., 2001). samples, their scale morphology closely resembles that of modern Hair identification reference keys are limited to guard hairs warthog (Fig. 4d), being highly irregular waved. The scale because of their characteristic features, but this makes hair iden- morphology of fossil hair specimen 11 shows a regular waved tification of less distinct or developed features challenging, espe- pattern (Fig. 9c). Scale margins are smooth and the distance be- cially when previous researchers have shown that there are tween scales is distant. Even though the scale morphology is well significant differences in hair morphology between hairs taken defined, it cannot be matched in the comparative collection. There from different sites on the animal (Riggot and Wyatt, 1981). Sea- is no cross section for this specimen. sonality, age and sex differences (Brothwell, 1993) can also complicate the taxonomic identification of hair, as demonstrated by 4. Discussion Keogh (1975), who investigated the effect of age on individual scale patterns of rodent hair, and found significant variation in the hairs 4.1. Fossil hair preservation of young and old animals. Many factors complicate morphological analysis, including protein degradation, abrasion, fungal and bac- Fossil hairs were extremely difficult to identify compared to terial attack, all of which can obscure or alter scale and medullary modern hairs, which record clear scale patterns and margins for the features (Bonnichsen et al., 2001). This is quite evident in the fossil entire length of the specimen. Most of the fossil hairs were very hairs from this research, many of which had abraded scales and small shaft fragments (1 mm), so only a few features were present obscure medullary features (e.g. specimens 7, Fig. 8c and d, e). DNA and available for use in identification. Scanning electron micro- analysis could have resolved the taxonomic identity of the fossil scope analysis of fossil hairs revealed that the extent of degradation hairs, but their age and mode of preservation do not permit it. varied significantly between specimens. This is most likely due to Despite this, some fossil hairs (e.g. Fig. 7a and b) record remarkably microbial activity and diagenetic processes (Martill et al., 2000), but well preserved casts of the original hair scales, providing sufficient could in part be attributed to the position from which the fossil detail to make tentative identifications of the mammal taxa hairs originated on the animal. This is because those from the an- represented. imal’s back and tail (guard hairs) are generally more robust than those from the neck and stomach, enabling them to resist degra- 4.2. Fossil hair identification dation. Alternatively, the hair was degraded before consumption, having been exposed to the elements for some days or weeks. Some fossil hairs had well preserved cuticular surface and cross Whatever the case, variable degradation between guard and under sectional morphology, which is rarely observed in ancient hair (e.g. hair warrants further investigation, especially because ancient hair specimen 1, Fig. 7a and b), and permitted visual comparison with assemblages are known to contain a wide range of hair types such modern hair samples. Scanning electron microscope analysis was P. Taru, L. Backwell / Journal of Archaeological Science 40 (2013) 3674e3685 3681

Fig. 7. Fossil hair specimen 1 cross section (a), scale ¼ 30 mm, and scale pattern (b), scale ¼ 50 mm; specimen 2 scale pattern (c); specimen 3 scale pattern (d); specimen 4 sca- lepattern (e), scales ¼ 100 mm, and cross section (f), scale ¼ 50 mm. used for all the hair samples and resulted in relatively clear mi- zebra hairs. Skinner and van Aarde (1981) analysed hairs from crographs. Visual comparison of hair can be subjective and is open scats from brown hyaena latrines from Wolfsbaai, Namibia, and to interpretation (Steck-Flynn, 2009). In a study conducted by the found that seal and jackal hair predominated. Kuhn et al. (2008) Federal Bureau of Investigation, 11% of hairs deemed to be matches analysed brown hyaena scats and examined faunal remains at upon visual inspection were found to be non-matches after DNA nine dens, and concluded that brown hyaenas along the Nami- testing (Saferstein, 2004). As with most forensic evidence, the in- bian Coast predominantly feed on seals. These are only a few formation obtained from hair is expressed in terms of probabilities examples of evidence of brown hyaenas consuming locally of a match rather than an absolute match (Crocker, 1999). available animals (see Faith, 2007; Kuhn et al., 2010), making The cuticular scale surface detail of fossil hairs in this research them good environmental indicators. The Burchell’s zebra, impala is sufficiently intact to be of use in species identification, and on and warthog represented in the coprolites are commonly found this evidence tentative identifications have been made, with in savannas. Plains zebra live in open savannas and partial to eight fossil hairs identified to species level in five cases. Based on open woodlands. They are a daily and seasonal migratory our collection of modern mammals from southern Africa, this mammal that moves in search of better grazing areas and water study has established that impala, Burchell’s zebra and warthog supplies, but are highly dependent upon water and are never hair predominated in the coprolites from Gladysvale cave. In more than 10e12 km from a source (Skinner and Smithers, 1990). accordance with Kruuk (1972), who found one taxon represented This species is predominantly a grazer preferring short grasses, in most spotted hyaena faeces, this research showed the same but will also browse and feed on herbs. Warthogs are found in pattern for brown hyaena coprolites. The only exception is in open grasslands, floodplains, open woodlands and open scrub. coprolite specimen 4, which preserved impala and Burchells’s They are selective feeders, preferring short grasses growing in 3682 P. Taru, L. Backwell / Journal of Archaeological Science 40 (2013) 3674e3685

Fig. 8. Fossil hair specimen 5 scale pattern (a), scale ¼ 50 mm; specimen 6 scale pattern (b); specimen 7 scale pattern (c), scales ¼ 100 mm and cross section (d), scale ¼ 50 mm; specimen 8 cross section (e) and scale pattern which could not be matched (f), scales ¼ 50 mm. freshly burnt and damp areas. They also root for underground environment of today. Amid a scarce fossil and archaeological rhizomes and will consume sedges, herbs, shrubs and wild fruit. record for this time period, these results contribute data to the Warthogs are not dependent upon water, but are usually found local Middle Pleistocene fossil faunal record, and insight into the close to it. Impala are associated with woodland, preferring light environment in which archaic and emerging modern humans in open associations. In southern Africa, they are associated partic- the interior of the African subcontinent lived. ularly with Acacia and Mopane woodland, occurring on the On the evidence of scale morphology preserved on the fossil ecotone of open grassland and woodland. Cover is an essential hairs studied here, rodents and other carnivores in general, and cats habitat requirement, but impala occasionally graze on open in particular, can be excluded as potentially represented. Rodent grassland when it is fresh and green (Skinner and Smithers, hair is typically grooved along the entire length of the hair and is 1990). A single fossil hair exhibits features characteristic of hu- usually coronal in scale morphology (Keogh, 1985). Carnivore hair man hair, supporting the previous tentative identification of hu- scale pattern consist of closely packed scales forming an irregular man hair in a single coprolite from the same deposit (Backwell pattern (Keogh, 1979), unlike that on the fossil hairs analysed in this et al., 2009). The cuticular scale pattern on the fossil human research. In this regard, the possibility that the hyaena ingested its hair (Fig. 7b) is imbricated, with a ‘regular wave’ morphology and own hair during grooming can be eliminated. Cats, like other car- the margins on the human hair scales (African and European) are nivores, have irregular waved patterns, but the scale margins are smooth and moderately rippled, and relatively evenly spaced typically frilly, and none of the fossil hairs record this feature. Our (Backwell et al., 2009). Apart from humans, the identified animals results indicate that the previously undocumented scub-hare (Lepus are associated with savanna grasslands, much like the Highveld saxatilis) scale pattern (Fig. 5b) closely resembles that of vlei rat hair P. Taru, L. Backwell / Journal of Archaeological Science 40 (2013) 3674e3685 3683

Fig. 9. Fossil hair specimen 9 scale pattern (a), scale ¼ 100 mm; specimen 10 scale pattern (b) and specimen 11 scale pattern (c), scales ¼ 50 mm.

(Otomys irroratus) studied by Keogh (1975). Similarities in scale used as a den for rearing pups. Spotted hyaenas (Crocuta crocuta)do pattern were also observed between red rock hare (Fig. 5d) and four- not defacate in one place, and striped hyaenas (Hyaena hyaena) striped mouse (Rhabdomys pumilio) documented by Keogh (1975). mark a large territory around the entrance (Watts and Holekamp, The only difference lies in the fact that the scale pattern on either 2007). In accordance with the most common foraging behaviour side of the groove is coronal in red rock hare (Fig. 5d) and petal reported for modern brown hyaena (Skinner and Ilani, 1979; mosaic in four-striped mouse. This is not surprising because of Skinner et al., 1980; Mills, 1990; Maude and Mills, 2005; Kuhn, possible overlaps between mammal hair features, as discussed by 2001, 2005, 2006; Kuhn et al., 2009, 2010; Kuhn, 2011), the ani- Hess et al. (1985). These similarities are noteworthy because of the mal(s) responsible for the coprolites, scavenged the identified problem of misidentification, and thus misinterpretation of the mammals. Brown hyaenas are reportedly inefficient predators, and fossil record and palaeoenvironment. Perrin and Campbell (1980) their food is rarely obtained by hunting (Maude, 2005). Researchers report that rock dassies have a hair cuticular scale pattern unlike have observed that if hunting occurs, it is targeted towards smaller most small mammals, in that it is flattened mosaic in the proximal mammals only (Skinner, 1976; Mills, 1978), as evidenced by brown region and changes to a waved pattern distally. The rock dassie hair hyaenas killing seal pups on the Namibian Coast (Wiesel, 2006; studied here preserves no scales, and shows cracking of the shaft Kuhn et al., 2008). Some brown hyaenas show specialization of (Fig. 6d). A lack of hair scales has been documented in human hair hunting techniques towards certain prey species (Wiesel, 2006), subject to pathology (Brown and Crounse, 1980), a condition such as southern Kalahari brown hyaenas hunting springbok lambs observed when studying our diabetic colleague’s hair as part of the (Antidorcas marsupialis) and korhaans (Eupodotis spp) as reported human comparative sample. In the case of the rock dassie, a lack of by Mills (1990), and brown hyaenas killing small livestock (Kuhn, scales could be due to pathology, abrasion of the hair resulting from 2011). They are said to be poorly equipped for running and hunt- inhabiting rock crevices, or perhaps preparation of the pelt by a ing, especially of large mammals, such as the zebra and kudu rep- taxidermist. To our knowledge, the effect of taxidermy on hair resented in the coprolites. If this behaviour is true of extant brown preservation is unknown, and warrants further investigation. hyaena, it should hold that Middle Pleistocene hyaena scavenged on the large mammals represented in the coprolites, which were 4.3. Parahyaena brunnea ecology and palaeoenvironment killed by other carnivores, most likely large cats. However, ac- cording to Brain (1981), brown hyaenas are keen hunters of large The existence of fossil hairs in Middle Pleistocene hyaena cop- mammals only when rearing pups, hence the possibility that the rolites from Gladysvale cave is attributed by Berger et al. (2009) to Middle Pleistocene hyaena(s) responsible for the coprolites actively the activities of Parahyaena brunnea. This is based on the contents hunted the mammals represented by the fossil hairs as part of their and position of the latrine in the cave, which suggests that it was pup-rearing behaviour. The fact that Middle Pleistocene hyaena fed 3684 P. Taru, L. Backwell / Journal of Archaeological Science 40 (2013) 3674e3685 on the identified mammals is consistent with previous researchers Backwell, L.R., Pickering, R., Brothwell, D., Berger, L.R., Witcomb, M., Martill, D., who found that hyaenas accumulate a wide range of faunal remains Penkman, K., Wilson, A., 2009. Probable human hair found in a fossil hyaena coprolite from Gladysvale cave. South Africa. J. Arch. Sci. 36 (6), to varying degrees and that their foraging behaviour is variable 1269e1276. (Scott and Klein, 1981; Faith, 2007; Kuhn et al., 2010). Apart from Benfer, R.A., Typpo, J.T., Graf, V.B., Picket, E.E., 1978. Mineral analysis of ancient e the five mammals identified in this research, our findings show that Peruvian hair. Am. J. Phys. Anthropol. 48, 277 282. Berger, L.R., 1992. Early hominin fossils discovered at Gladysvale Cave, South Africa. brown hyaenas fed on at least one mammal that could not be S. Afr. J. Sci. 88, 362. matched in the comparative collection (e.g. specimen 11, Fig. 9c and Berger, L.R., 1993. A preliminary estimate of the age of the Gladysvale australo- specimen 8, Fig. 8f), suggesting that some as yet unidentified spe- pithecine site. Palaeontol. Afr. 30, 51e55. Berger, L.R., Keyser, A.W., Tobias, P.V., 1993. Gladysvale: first early hominid site cies are represented in the fossil hair samples. discovered in South Africa since 1948. Am. J. Phys. Anthropol. 92, 107e111. Berger, L.R., Pickering, R., Kuhn, B., Backwell, L.R., Hancox, P.J., Kramers, J.D., Boshoff, P.A., 2009. A Mid-Pleistocene in situ fossil brown hyaena (Parahyaena 5. Conclusion brunnea) latrine from Gladysvale Cave, South Africa. Palaeogeogr. Palae- oclimatol. Palaeoecol. 279, 131e136. Scanning electron microscope analysis of modern hair samples Bonnichsen, R., Hodges, L., Ream, W., Field, K.G., Kirner, D.L., Selsor, K., Taylor, R.E., fi 2001. Methods for the study of ancient hair: radiocarbon dates and gene se- of known taxa revealed ne details of scale and cross sectional quences from individual hairs. J. Arch. Sci. 28 (7), 775e785. morphology of hair, and showed that cuticular scale pattern and Brain, C.K., 1981. The Hunters or the Hunted? An Introduction to African Cave cross section shape can be used as definitive criteria in species Taphonomy. University of Chicago Press. fi Brothwell, D.R., Spearman, R., 1963. The hair of earlier peoples. In: Brothwell, D., identi cation, and the pertinence of this technique to palae- Higgs, E. (Eds.), Science in Archaeology. A Comprehensive Survey of Progress ontological research. The identified fossil hairs show that between and Research. Thames and Hudson, pp. 427e436. 257 and 195 ka, Parahyaena brunnea shared the Sterkfontein Valley Brothwell, D.R., 1993. Ancient hair analysis and the question of diet and disease. e with warthog, impala, zebra, kudu and humans. Whether the hy- Pact 38, 317 326. Brothwell, D.R., Grime, G., 2003. The analysis of the hair of the Neolithic Iceman. In: aena(s) responsible for the coprolites hunted or scavenged from Lynnerup, N., Andreasen, C., Berglund, J. (Eds.), Mummies in a New Millenium. these animals is unclear. The identification of five species of Danish Polar Center, Copenhagen, pp. 66e69. mammal represented in the coprolites in the form of fossil hairs Brown, A.C., Crounse, R.G., 1980. Hair, Trace Elements and Human Illness. Praeger fi Publishers, New York. support the previous tentative identi cation of fossil human hair in Brunner, H., Coman, B.J., 1974. The Identification of Mammalian Hair. Inkata Press, a single coprolite, provides a new source of information on the local Victoria, Australia. Middle Pleistocene fossil mammal community, and rare insight into Budworth, G., McCord, M., Priston, A., Stead, I., 1986. The artefacts. In: Stead, I.M., Bourke, J.B., Brothwell, D. (Eds.), Lindow Man, the Body in the Bog. British the environment in which archaic and modern humans in the Museum Publications, pp. 38e40. interior of the African subcontinent lived. Hair provides evidence of Chang, B.S., Hong, W.S., Lee, E., Yeo, S.M., Bang, I.S., Chung, Y.H., Lim, D.S., Mun, G.H., inland occupation by archaic Homo sapiens or modern humans. Kim, J., Park, S.O., Shin, D.H., 2005. Ultramicroscopic observations on morpho- logical changes in hair during 25 years of weathering. Forensic. Sci. Int. 151 (2), Amid a scarce fossil and archaeological record for this time period, 193e200. these results contribute data to the ongoing debate about the role of Clark, J., Plug, I., 2008. Animal exploitation strategies during the South African environment in the evolution of modern humans (Mitchell, 2008; Middle Stone Age: Howiesons Poort and post-Howiesons Poort fauna from . J. Hum. Evol. 54, 886e898. Schiegl and Conard, 2006; Deacon and Deacon,1999; Wadley, 2006; Cole, H.I., 1924. Taxonomic value of hair in Chiroptera. Philipp. J. Sci. 14, 117e121. Clark and Plug, 2008; d’Errico and Banks, 2012). This study high- Crocker, J.E., 1999. Trace evidence. In: Chayko, G.M., Gulliver, E.D. (Eds.), Forensic lights the importance of researching all aspects of the fossil record, Evidence in Canada, second ed. Canadian Law Books Inc., Aurora. and the contribution of microscopy to palaeontology. In addition to Daniel, C.R., Piraccini, B.M., Tosti, A., 2004. The nail and hair in forensic science. J. Am. Acad. Dermatol. 50, 258e261. expanding the taxonomic representation of modern southern Afri- Davis, S.J.M., Robert, I., Zilhao, J., 2007. Caldeirao cave (Central Portugal)-whose can mammal hair samples, we propose that researchers involved in home? Hyaena man bearded vulture. Cour. Forschungsinsti. Senckenb. 259, e the study of hair document when possible, in addition to guard 213 226. Deacon, H.J., Deacon, J., 1999. Human Beginnings in South Africa. Uncovering the hairs, examples from different sites on the body, and from young Secrets of the Stone Age. David Philip Publishers, Cape Town. and old individuals, factors that affect hair morphology, and moti- d’Errico, F., Banks, W.E., 2012. Identifying mechanisms behind Middle Paleolithic e vate for the expansion of the range of features considered diagnostic and Middle Stone Age cultural trajectories. Curr. Anthropol. 39, S1 S44. Faith, J.T., 2007. Sources of variation in carnivore tooth - mark frequencies in a of a species. We hope that the presentation of our data will assist in modern spotted hyaena (Crocuta crocuta) den assemblage, Amboseli Park, other fields of research and stimulate further inquiry. Kenya. J. Arch. Sci. 34 (10), 1601e1609. Fraser, R.B., MacRae, T.P., Rogers, G.E., 1972. Keratins: their Composition, Structure and Biosynthesis. Charles E. Thomas, Springfield. Acknowledgements Gilbert, M.T.P., Tomsho, L.P., Rendulic, S., Packard, M., Drautz, D.I., Sher, A., Tikhonov, A., Dalen, L., Kuznetsova, T., Kosintsev, P., Campos, P.F., Higham, T., Collins, M.J., Wilson, A.S., Shidlovskiy, F., Buigues, B., Ericson, P.G.P., LB acknowledges the National Research Foundation (NRF) of Germonpre, M., Gotherstrom, A., Iacumin, P., Nikolaev, N., Nowak-Kemp, M., South Africa for a Thuthuka Women in Research grant. PT is grateful Willerslev, E., Knight, J.R., Irzyk, G.P., Perbost, C.S., Fredrikson, K.M., Harkins, T.T., to the University of the Witwatersrand for an FRC bursary and a Sheridan, S., Miller, W., Schuster, S.C., 2007. Whole-genome shotgun sequencing e fi of mitochondria from ancient hair shafts. Science 317, 1927 1930. Postgraduate Merit Award, and to the Palaeontological Scienti c Green, C.S., Patterson, M., Biek, L., 1981. A Roman coffin burial from the crown Trust (PAST) for a research grant. We are thankful to Professor buildings site. Dorchester: with particular reference to the head of well pre- Michael Witcomb for his invaluable technical assistance during served hair. In: Proc. Dorset Nat. Hist. Archaeol. Soc., 103, pp. 67e100. Hausman, L.A., 1920. Structural characteristics of the hair of mammals. Am. Nat. 54, scanning electron microscope analysis of all the hair samples. Dr. 496e523. Theresa Kearney and the Ditsong National Museum of Natural His- Hausman, L.A., 1930. Recent studies of hair structure relationships. Sci. Month 30, tory and Mrs. Louise Matschke and the Johannesburg Zoo are also 258e277. thanked for providing us with hair samples for our investigation. We Hess, W.M., Flinders, J.T., Pritchett, C.L., Allen, J.V., 1985. Characterization of hair morphology in Families Tayassuidae and Suidae with scanning electron micro- are grateful to Precious Somerai for proof reading this work. scopy. J. Mammal. 66 (1), 75e84. Homan, J.A., Genoways, H.H., 1978. 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