Mossel Bay, Western Cape Province, South Africa)

Palaeogeography, Palaeoclimatology, Palaeoecology 302 (2011) 213–229

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Palaeogeography, Palaeoclimatology, Palaeoecology

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Environmental implications of micromammals accumulated close to the MIS 6 to MIS 5 transition at Pinnacle Point Cave 9 (Mossel Bay, Western Cape Province, South Africa)

Thalassa Matthews a,⁎, Amy Rector b, Zenobia Jacobs c, Andy I.R. Herries d, Curtis W. Marean b a Iziko South African Museum, 25 Queen Victoria Rd, Cape Town, 8000, South Africa b Institute of Human Origins, School of Human Evolution and Social Change, Arizona State University, Tempe, AZ 85287, United States c Centre for Archaeological Science, School of Earth and Environmental Sciences, University of Wollongong, NSW 2522, Australia d UNSW Archaeological Science Laboratory, integrative Palaeoecological and Anthropological Studies, School of Medical Sciences, University of New South Wales, 2052, NSW, Australia article info abstract

Article history: PP9C is a coastal cave situated on the south coast, Pinnacle Point, Mossel Bay, and contains both Received 9 September 2010 archeological deposits, as well as fossil micromammal accumulations. Micromammals were analyzed from Received in revised form 14 January 2011 facies accumulated close to the transition from MIS (Marine Isotope Stage) 6 to MIS 5, between 130±9 and Accepted 19 January 2011 120±7 ka. The taphonomy of the assemblages indicates that the majority of micromammals became Available online 25 January 2011 associated with the site when they were deposited in Spotted eagle owl or Barn owl pellets. Modern comparative studies suggest that these two predator species select a very similar suit of prey species from Keywords: Pinnacle Point the available micromammal population and the fossil micromammal assemblages they produce are Micromammal comparable when tracing palaeoenvironmental change. The fossil assemblages suggest that the period Palaeoenvironment during which the micromammals accumulated was warm and wet, and that the vegetation within the Taphonomy hunting range of the owls accumulating the micromammal assemblages was relatively dense. There is, Diversity however, some evidence that at the time of deposition of the older assemblages, conditions were colder, Middle Stone Age relatively grassier, and may reflect a rather more open environment. This difference may indicate the change from glacial MIS 6 to inter-glacial MIS 5e climatic conditions. These conclusions are supported both by the habitats and habits of the micromammal faunas,aswellasbymultivariateanalysis.E. edwardii and A. namaquensis appear only in the earliest fossil assemblages at PP9C, and their general scarcity suggests that the rocky and open habitat preferred by these two species was never available on a wide scale at Pinnacle Point. The Pinnacle Point fossil sites indicate that the soricid, C. flavescens,wasacommon component of the local micromammal fauna and occurred in the area during some of MIS 6, and into MIS 5, suggesting that this species was able to adapt to glacial/interglacial cycles, and may utilize drier habitats than commonly reported in the literature. The fossil evidence indicates that C. cyanea, S. infinitesimus and Saccostomys campestris are relative latecomers to the Mossel Bay region. S. campestris appears only in PP9C in surface sediments, and confirms previous suggestions that the arrival of this species to the southern Cape occurred sometime within the Holocene. © 2011 Elsevier B.V. All rights reserved.

1. Introduction rainwater, resulting in alkaline conditions in the sediments, and speleothem and tufa formation in the caves (Bar-Matthews et al., This paper presents a palaeoenvironmental analysis of micro- 2010). The result is that most of the sites at Pinnacle Point have mammal assemblages recovered from Cave 9C at Pinnacle Point, south excellent preservation of faunal material. PP9 currently presents coast, Western Cape Province, South Africa. PP9 is a large sea cave many excellent roosting spots, and many bird species live there today. complex formed in quartzite of the Table Mountain Sandstone Group PP9 consists of a number of smaller separate cavities named PP9A- (TMS) that outcrops along the southern Cape coast. The TMS in this E(Fig. 1). PP9A contains a deep, virtually sterile, dune in-ﬁll and was area is capped by calcretes and calcareous sands that buffer the partially excavated in 2000 (Marean et al., 2004). No access to PP9D and E is available due to their location up near the roof of the PP9A cavity. PP9B was excavated in 2006 and is located to the left (south) of the PP9A cavern. It is a small, tube-like deposit in-ﬁlled with eroded ⁎ Corresponding author. Tel.: +27 21 4813877; fax: +27 21 4813983. raised beach deposits, MSA occupation horizons, and a capping of E-mail address: [email protected] (T. Matthews). sterile dune sand. PP9B was found to contain only 2 murid mandibles

mals includes the murids (mice and rats), bathyergids (mole rats), soricids (shrews), macroscelids (elephant shrews) and Chiroptera (bats). The Pinnacle Point micromammal assemblages are being studied in order to trace palaeoenvironmental change over a time period which includes global glacial and inter-glacial cycles. This is of particular interest as little information is available on glacial South African micromammal populations, and the Pinnacle Point cave sites present a unique opportunity to study the reaction of micromammal taxa to extensive climatic and environmental change over time in a particular area. Another MSA cave site at Pinnacle Point, PP13B, lies some 300 m to the south of PP9 and an analysis of the micromammals from horizons dating to speciﬁc non-continuous time periods during MIS 6and5(Jacobs, 2010; Marean et al., 2010) has been published (Matthews et al., 2009). There is some overlap in the younger time periods represented by PP13B and PP9 which provides an opportunity to cross-check environmental information from the two cave sites.

2. Materials

In November 2005, prior to the controlled excavation of PP9C, a surface collection was taken of a dense, surface micromammal accumulation situated towards the rear of the cave. This collection is referred to as Surface Scraping 2005, and abbreviated as ‘SS:2005’, in this paper. Several species found in this horizon suggest that much of the micromammal material is modern, although there may have been some mixing with the underlying fossil horizons. Five excavation squares were made in the PP9C sediments in October 2006. Individual ‘Stratigraphic Units’ (StratUnits) were initially identified during excavation based on color and consistency of the deposit. StratUnits were later grouped into larger ‘Stratigraphic Fig. 1. The Pinnacle Point caves PP9A, PP9B and PP9C. Aggregates’ (StratAggs, similar to “layers”), which reflected a homogeneous set of formation processes of roughly the same time period. The micromammals were grouped into StratAggs for the and an incisor (see Appendix B) and will not be reported on any purpose of analysis. further in this paper. Directly above PP9B is PP9C which was also The stratigraphy and ages of the various StratAggs are shown in excavated in 2006 and contains a complex series of geological Table 1, and the abbreviations used in this paper for these are shown deposits and MSA occupations that will be described in detail in in brackets following the full name of each aggregate. The two Rear another paper. The entrance to PP9C involves accessing a 10 m rock Tunnel Entrance (RTE) StratAggs, OYCS and BYDS, as well as the two face from PP9A and fixed ropes were used for excavation and Rear of Rear Tunnel (RTR) StratAggs, BYSS and BYCS, are contiguous sampling of this cave. PP9C consists of a ‘front cavity’ that leads to the with one another. The age of the PP9C deposits relative to those of mouth of the larger PP9A cavern and a ‘rear tunnel’ that is accessed by the Pinnacle Point sites PP13B and PP5-6 (which is being excavated crawling through a small hole in the rear of the front cavity. This currently), and some other south coast sites, are summarized in constriction is due to the infill of the mouth of the tunnel with Fig. 2. sediment, which contains dense micromammal accumulations. The Excavation of the RTE area was made in the entrance to the rear constriction leads through to a ~5 m long tunnel that contains tunnel of PP9C, slightly beyond the sediment infill constriction. Its speleothem, ancient eroded calcified deposits, and a dune containing deposits have been dated to 126±9 and 120±7 ka, using optically micromammal deposits, which range in depth from 1.1 m at the stimulated luminescence (OSL) dating of individual sand-sized grains entrance, to 0.3 m at the rear of the tunnel. of quartz (Jacobs and Roberts, 2007). Both ages are statistically The excavations at PP9 form part of research being carried out by the South African Coast Paleoclimate, Paleoenvironment, Paleoecol- ogy, Paleoanthropology project (SACP4) which is currently underway in the Mossel Bay area. The project involves the study of a number of archeological sites in the Pinnacle Point area dating to the Middle Table 1 Stratigraphy and dating of PP9C. Stone Age (MSA) which are being excavated in order to study the response of early human behavior to climate change. The majority of StratAggs OSL ages these sites contain micromammal accumulations (Matthews et al., PP9C-SS:2005 Undated 2009), the greater part of which appear to have been accumulated by Barn or Spotted eagle owls via the deposition, and then disaggregation PP9C-RTE Olive Yellow Cave Sand (OYCS) 120±7 ka over time, of regurgitated pellets. Small mammals have small home Brown Yellow Dune Sand (BYDS) 126±9 ka ranges, do not undertake seasonal migrations, and many have precise habitat requirements. This makes them suitable for palaeoenviron- PP9C-RTR mental reconstruction (Avery, 1986, 1987, 1990, 1999a; Matthews et Brown Yellow Surface Sand (BYSS) Undated al., 2009). This palaeoenvironmental analysis of the PP9 micromam- Brown Yellow Cave Sand (BYCS) 130±9 ka T. Matthews et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 302 (2011) 213–229 215

PP9C Other South EPICA Delta D Rear Tunnel Rear of Rear Coast MSA Sites -460-440-420-400-380-360 0 Entrance Tunnel Entrance 0

10 10 PP5-6 PP13B 20 20 Blombos

30 30 Klasies River

40 40

50 50

60 60

70 70

80 80

90 90 (Ka) Age

100 100

Age (Ka) 110 110 Olive Yellow 120 Cave Sand * 120 * Brown Yellow 130 Brown Yellow * 130 Dune Sand Cave Sand 140 140

150 150

160 160

170 170

180 180

190 190

200 200

Fig. 2. Dating of the PP9C sediments relative to global climate as reﬂected by Epica δD record and other south coast sites.

consistent with each other at one standard error and it is not possible less standardized (e.g. Matthews, 1999, 2002, 2006; Manthi, 2002; to reliably place these ages in a particular marine isotope stage (MIS). Matthews et al., 2006; Dewar and Jerardino, 2007)andwasused These deposits may straddle the MIS 6 to MIS 5e transition at ~129 ka for recording the taphonomy of the PP9C assemblages. Only the (Cheng et al., 2009; Drysdale et al., 2009; Masson-Delmotte et al., taphonomic results relevant to identifying the predator/s respon- 2010), or may fall exclusively within MIS 5. Three small 50×50 cm sible for the fossil accumulations are presented in this paper as quadrants were also excavated in soft sediments at the rear of the these are pertinent to its focus, namely palaeoenvironmental PP9C rear tunnel, namely excavation area RTR. The fossils from these reconstruction. three small excavations were amalgamated for processing as the excavations were shallow (b30 cm) and contained similar sedimentary deposits with abundant micromammals. These deposits have 3.1. Taphonomy: recording predator-induced digestion patterns been dated to 130±9 ka. This age is statistically indistinguishable from the ages obtained for the RTE area and the error associated with Incisor digestion was studied in order to work out which the age prevents a clear association with a speciﬁc marine isotope predator/s had been responsible for the accumulation of the stage. Based on the age estimates alone, it appears that the micromammal assemblages. Ascertaining the predator/s responsi- micromammal assemblages date to a period close to the MIS 6 to ble for the accumulation of a fossil assemblage is essential if it is to MIS 5e transition. be used to extrapolate palaeoenvironmental change as changes in the identity of the predator accumulating the fossil assemblage may 3. Methodology lead to changes in the selection of prey species. Different predator species have different requirements when they select prey species, The use of the methodology initially proposed by Andrews and prey selection is affected by a number of variables which (1990) and Fernandez-Jalvo and Andrews (1992) for the tapho- include prey availability, prey size, hunting techniques, hunting nomic analysis of micromammal assemblages has become more or ranges and so on. 216 T. Matthews et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 302 (2011) 213–229

The following categories describe and record incisor digestion on level are noted as “Indeterminate bathyergid”, “Indeterminate both isolated and in situ incisors. Incisors that suffered damage and chrysochlorid”, etc. that could not be properly assessed were excluded. 3.3. Measurement of Crocidura ﬂavescens

Class 0: No visible digestion on the incisor. The early Various mandibular and tooth measurements of the soricid, stages of digestion may not be detectable with a Crocidura flavescens, were made in order to see if any size dif- light microscope and it is possible that some ferences could be observed between specimens from variously aged incisors falling into this category sustained very deposits as Avery (1990) found evidence that suggested that the light, but indiscernible digestion. tooth row length of C. flavescens fluctuated according to Bergmann's Class 1 (light): Light etching and removal of the upper layers of rule. The few specimens recovered from PP13B were included in small areas of enamel and/or dentine. the comparison. A description of the measurements made, and the Class 2 (moderate): The area of digestion may not be much greater results, may be found in Appendix C. The methodology used was than Class 1, but the digestion has penetrated based on, and closely follows, that used by Butler et al. (1989) and much deeper through enamel layers, down to, Butler and Greenwood (1979). The tooth nomenclature used for the or very close to dentine. The dentine shows a mandibles is I ,P,P,M ,M,andM (after Skinner and Smithers, deeper degree of penetration and loss. 2 1 4 1 2 3 1990). Measurements were made with a micrometer fitted into Class 3 (heavy): The area of digestion is more extensive than class the eyepiece of a binocular light microscope and are accurate to 2 digestion, with complete removal of enamel in 0.05 mm. areas, and digestion and removal of underlying dentine. 3.4. Assessing general diversity Class 4 (extreme): Extreme digestion of both enamel and dentine, with some teeth having all enamel, and much The Shannon Wiener index of general diversity was used to assess dentine, removed. The edges of the dentine or diversity of the fossil micromammal assemblages. This index accounts enamel may be collapsing in on themselves. for both the number of taxa present, and relative frequency (evenness of representation) of each taxon (Cruz-Uribe, 1988; Magurran, 1988). The Shannon Wiener index is calculated using the formula; 3.2. Identification of micromammalian taxa H = −∑Pi logePi; From this point onwards, the three molars of the murid mandible will be referred to as the M1–3, and the three maxillary molars, as the where Pi =n/N,that is, the proportion of the total sample represented 1–3 1 M respectively. Murid M and M1 teeth (in situ and isolated) were by each species. used to quantify the number of individuals of the various species present in the fossil assemblages. An exception to this rule was the 3.5. Univariate analyses Otomyinae (the vlei rats), where the M1 and the more diagnostic M3 were used to quantify this particular murid subfamily. Teeth of For communities of larger mammals, it has been shown that uncertain identification due to breakage, wear or digestion were community structure, as represented by substrate use and dietary excluded from analyses. adaptations of each included species, can be used to reconstruct Differentiating between Otomys saundersiae and Otomys irroratus habitats within a taxon-free framework that is comparable to other was complicated by the fact that the fossil material generally communities across time and space (Reed, 1997, 1998). When the consisted of isolated molars, or broken mandibles with incomplete large mammals in communities representing different modern tooth rows, as well as by the intra-specific variability in the size and ecological biotopes are considered in terms of these adaptations, shape of the laminate teeth of these two species. Specimens which habitats are differentiated based on the resulting community could not be allocated with certainty to either species were recorded structure. This method has been effectively used for reconstructing as O. saundersiae/O. irroratus. paleohabitats for fossil assemblages of larger mammals in eastern and BYCS (RTR excavation area) contained four isolated, and one in southern Africa (Andrews et al., 1979; Nesbit-Evans et al., 1981; situ,M3 molar which had the five laminate structure usually Andrews, 1989; Reed, 1997, 1998, 2008; Reed and Rector, 2007; associated with Otomys slogetti. The lack of any M1 molars which Rector and Reed, 2010), and while factor analysis has been applied to looked like O. slogetti eventually resulted in the M3s being identified as species abundances of micromammals in fossil assemblages to belonging to O. saundersiae. The identification of these specimens as O. reconstruct parameters such as temperature and rainfall (Avery, saundersiae is supported by the fact that Roberts (1951) and Ellerman 1982; Thackeray, 1987; Sénégas and Thackeray, 2008), studies of et al. (1953) noted that the M3 of O. irroratus and O. saundersiae may microfauna community structure comparing modern habitats with comprise 5–7 laminae. An O. saundersiae with five laminae was also fossil assemblages have not yet been applied. Here, we do so by recovered from Surface Sediments in PP13B. assigning micromammal species from 53 modern sites sampling Soricids were quantified differently to murids as the soricid bushland, desert, forest, grassland, scrubland and woodland habitats mandibles are always far better represented in fossil deposits than in Africa (see online supplementary information for all CA tables) to the relatively fragile soricid maxillary bones. The number of three adaptation categories: 1) locomotor, 2) dietary, and 3) substrate mandibles containing the M1 was used to quantify soricid species in (data primarily mined and modified from the National Center for the various facies. The lower first molars of soricids show the same Ecological Analysis and Synthesis Recent Mammal Database; http:// tendency as murid teeth to remain in situ. www.nceas.ucsb.edu/projects/2041). For this analysis, “locomotor” The bats (Order: Chiroptera), elephant shrews (Family: Macro- refers to where, in vertical space, the species spend the majority of scelididae) and mole rats (Family: Bathyergidae) were present in very their time, including arboreal, terrestrial, terrestrial/arboreal and low frequencies and were generally represented by mandibles, rather subterranean. “Substrate,” on the other hand, describes the type of than maxillae or isolated teeth, so the number of mandibles present landscape they prefer, including sandy, rocky, vlei (marshy or was used to quantify the MNI (minimum number of individuals) for swampy) and “forest.” Forest substrates are used to describe soils each species. Specimens which could not be identified to the Family that may or may not be rich in nutrients, but tend to be covered with T. Matthews et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 302 (2011) 213–229 217 a rich leaf-litter and organic layer (Comerford, 2006). Dietary categories included carnivore, frugivore, granivore, herbivore, invertivore, and omnivore. While some micromammals can be used as habitat indicator species, for the most part micromammals are generalists able to utilize a broad range of diets and substrates. Here, we attempted to create categories that were as specificas possible to maintain feasibility of usage, while also trying to recognize the variability inherent in studies of micromammals. Modern species used in this analysis are included, along with their category assign- ments, as are the databases, and results, of the univariate and multivariate analyses (see online supplementary data). The number of species in each dietary, locomotor and substrate category at each modern site was compared by habitats in a one-way Fig. 3. Incisor digestion patterns found in PP9C shown together with modern comparative Spotted eagle owl and Barn owl assemblages. ANOVA, with a null hypothesis stating that the number of species in each adaptation category would not vary in different habitats. Category membership was further analyzed with Tukey's post hoc “Honestly Significantly Different” (HSD) test for uneven sample sizes expected value with larger values reflecting more variance explained. between pairs of habitat for each adaptation. Only those categories When the row values were plotted using the first two – and thus most found to be significantly different at the pb0.05 level between three meaningful in terms of variance explained – dimensions, the sites that or more habitat types were retained for use in multivariate analyses to fell closest together were most similar in terms of species member- avoid using categories that do not reflect data relevant to the ship in the different categories of adaptations. The column values, or differences between habitat types, and thus muddying the subsequent adaptations, were also plotted, with the sites falling closest to them analyses. most affected by their variance. To compare the PP9C fossil assemblages (see online supplementary information, Table C) to the variance in modern sites described by the CA, the PP9C communities 3.6. Multivariate analyses were plotted onto the dimensions constructed in the CA using the functions created for the modern sites. Plotting the PP9C assemblages The proportions of adaptations found in mammalian communities using variation found in modern habitats allowed identification of the from the fossil sites were compared to data from extant habitats using most similar analogue based on charted proximity (Reed, 1996, 1998, correspondence analysis with STATISTICA 7.0 (StatSoft, 2005). 2008; Rector and Reed, 2010). Correspondence analysis (CA) is a multivariate exploratory technique that identifies similarities in datasets by maximizing the variation in a few dimensions, while resisting the effects of outliers (Shi, 19932), 4. Results and has been effectively applied to studies of paleohabitats (Reed, 2008; Rector and Reed, 2010). Functions were constructed by 4.1. Taphonomy: incisor digestion patterns maximizing variance in the number of species from each modern site (rows) in the adaptation categories (columns), based on their Sample size of murid incisors was inadequate for identification of deviations from an expected chi-square value of the total matrix the predator responsible for the fossil micromammal accumulations (Reed, 2008). A percentage of variance accounted for is reported for in all but the 3 largest units, namely the StratAggs BYSS and BYCS fi each dimension, which is the correlation coef cient between the row (RTR), and OYCS (RTE). Murid incisor digestion results are presented and column scores, and describes the degree of deviation from the in Table 2. In the other, smaller units, not shown in Table 2, incisors showed a similar pattern of digestion to the larger units, with the majority of incisors showing light and moderate digestion. Table 2 Murid incisor digestion patterns are illustrated in Fig. 3, together Murid incisor digestion patterns in PP9C. with that of typical Barn owl and Spotted eagle owl digestion patterns Digestion Isolated Isolated In situ mandibular Total % (modern samples of incisors from modern Barn owl and Spotted eagle category Mandibular Maxillary and maxillary owl pellets were examined to provide a comparative database incisors incisors incisors (Matthews, 2008)). No fossil or comparative incisors showed extreme StratAgg: RTE-Olive Yellow Cave Sand digestion and this digestion category is thus omitted. None 0 1 2 3 12.5 Light 4 7 5 16 66.7 Moderate 1 4 0 5 20.8 Heavy 0 0 0 0 0 Extreme 0 1 2 3 0 Table 3 Soricid percentage representation. StratAgg : RTR-Brown Yellow Surface Sand StratAggs Soricidae percentage Soricidae NISP None 1 4 0 5 14.7 representation (N) Light 8 9 9 26 76.5 (%) Moderate 0 2 0 2 5.9 Heavy 1 0 0 1 2.9 PP9C-SS:2005 28.2 44 156 Extreme 0 0 0 0 0 PP9C-RTE StratAgg: RTR-Brown Yellow Cave Sand Olive Yellow Cave Sand (OYCS) 24.4 21 86 None 1 6 2 9 11.54 Brown Yellow Dune Sand (BYDS) 4.8 1 21 Light 19 28 11 58 75.64 Moderate 0 5 0 5 6.41 PP9C-RTR Heavy 1 4 0 5 6.41 Brown Yellow Surface Sand (BYSS) 44.8 47 105 Extreme 0 0 0 0 0 Brown Yellow Cave Sand (BYCS) 38.4 139 362 218 T. Matthews et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 302 (2011) 213–229

Fig. 4. Percentage representation of Steatomys krebsii and Rhabdomys pumilio. Fig. 6. Representation of the Soricidae versus the Otomyinae.

Interpretation of the incisor digestion patterns of the SS:2005 dominant taxa varies from region to region. This patterning was sediments is hampered by poor sample size (N=15 murid incisors), observed in PP9C, and the relative proportions of the more common but they may have been accumulated by a Spotted eagle owl as 13 taxa are compared in Figs. 4–7. The percentage representation in the incisors showed ‘light’ digestion, while one showed ‘moderate’, and five largest StratAggs of two of the more common species, namely another ‘heavy’ digestion. Generally speaking, the older BYCS and Rhabdomys pumilio (Striped Fieldmouse) and Steatomys krebsii BYSS StratAggs from RTR show a similar trend of digestion to OYCS (Kreb's Fat Mouse) are compared in Fig. 4. (RTE). As mentioned previously, the StratAggs OYCS and BYDS (RTE), and also BYSS and BYCS (RTR), are contiguous, and in both areas of the 4.2. Species representation cave these horizons contain similar percentages of Steatomys and Rhabdomys. Interestingly, Rhabdomys and Steatomys appear to The micromammalian taxa found in PP9C and PP9B are listed in experience a converse relationship in the two excavation areas of Appendix B, together with the number of different species, and the the cave, but in SS:2005 appear in close to equal proportions. NISP (Number of Individual Specimens), in each StratAgg. The percentage representation of O. saundersiae versus O. irroratus Mus minutoides, Elephantulus edwardii, Rhinolophus clivosus, and in each StratAgg is illustrated in Fig. 5. The converse relationship Cryptomys hottentotus occur only in the RTR deposits. Aethomys between the RTE and RTR areas seen in Fig. 4 is repeated (Fig. 5), with namaquensis is found only in the latter, and in the SS:2005 O. irroratus (Vlei Rat) dominating the otomyine assemblages in RTE, assemblage. Crocidura cyanea, Suncus infinitesimus and Saccostomys while O. saundersiae (Saunder's Vlei Rat) dominates in RTR. Vlei rats campestris are found only in SS:2005. are relatively less common in SS:2005 where O. irroratus dominates. In order to illustrate the marked dominance of soricids in certain The percentage representation of the Otomyinae versus the facies, Table 3 shows the percentage representation of all soricid Soricidae is also illustrated (Fig. 6). In the RTR, the soricids make up species, relative to that of the combined total of murids, macroscelids 38–44% of the assemblage, and the Otomyinae make up approxi- and chrysochlorids. A total of the actual number of soricids (N) mately 25%. In the OYCS (RTE) the situation is reversed, with the represented by the percentage, as well as the total micromammal Otomyinae making up approximately 40% and the Soricidae 24%. NISP, including murids, macroscelids, soricids and chrysochlorids, are The species representation in the BYDS StratAgg shows a similar given. Only the Stratigraphic Aggregates with a NISP of ≥15 are pattern to the similarly aged OYCS, but with a more marked discussed in this paper, unless otherwise stated. As Table 3 clearly dominance of Otomyinae. The difference between BYDS and OYCS illustrates, the soricids dominate the deposits in the contiguous BYSS maybeexplainedbytheunsatisfactorysamplesizeforBYDS. and BYCS (RTR) StratAggs. Both of these assemblages are of a good SS:2005 has a similar percentage of soricids to OYCS, but fewer sample size, with NISPS of 105 and 362 for BYSS and BYCS, Otomyinae. respectively. The percentage representation of the various soricid species, Certain micromammal taxa are frequently found in relatively high relative to each other, is illustrated (Fig. 7). An unidentified soricid frequencies in Spotted eagle owl and Barn owl pellets, although the

Fig. 5. Percentage representation of Otomys saundersiae versus Otomys irroratus. Fig. 7. Percentage representation of the Soricidae in PP9C. T. Matthews et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 302 (2011) 213–229 219 species was found in BYCS (RTE) and was represented by only 1 mandible. This specimen looked most similar to C. cyanea (Reddish Grey Musk Shrew), but differed in several aspects and could not be identified to species. Some 2 mandibles representing another unidentified, Myosorex-like species (Forest shrew) were found in RTR in a small StratUnit named Light Yellow Decalcified Sand (LYDS). The unidentified Myosorex species from PP9 were compared to Myosorex longicaudatus, a species found only in the Knysna area and discovered only in the late 1970s (Meester and Dippenaar, 1978), as well as the Myosorex species currently found in the western and eastern Cape, namely, M. varius, M. cafer and M. sclateri. The fossil species were clearly differentiated from these taxa, and may represent new subspecies/species. Myosorex varius and Suncus varilla appear to occur in relatively similar abundances in the BYSS, BYCS and OYCS StratAggs, while Crocidura flavescens appears to be much less represented in the RTE deposits. In terms of soricid species representation, the percentage Fig. 8. The relationship between NISP and (H) in PP9C and PP13B. representation of S. varilla is similar throughout the four largest assemblages shown in Table 3. M. varius appears well represented in all the fossil horizons, but is poorly represented in SS:2005. C. flavescens shows a notable increase in representation in both SS:2005, significantly different between the habitat types (online supplemen- and in BYSS (RTR). As noted above, Appendix C contains the results of tary information, Table D). Additional Tukey's Post-Hoc analyses various tooth and mandibular measurements made on C. flavescens indicated that of the remaining categories, terrestrial/arboreal mandibles and teeth from PP9C and PP13B. No significant differences locomotion, granivore diets and rocky substrates were only signifi- were noted between the ranges and averages of the various tooth and cantly different between one or two habitat types, and thus were also mandibular measurements. not included in the CA. The remaining categories, including arboreal, subterranean, and terrestrial locomotion, carnivore, frugivore, herbi- 4.3. General diversity vore, and omnivore diets, and forest substrate use, were used in the CA to describe the variation in these categories between and among The Shannon Wiener index for all units in PP9C and PP13B with a the 53 modern sites. fi NISP of ≥15 is shown in Table 4. In PP9C diversity is highest in The rst two dimensions constructed in the CA explain a SS:2005 and in OYCS, and lowest in the RTR facies. The Shannon combined 77.1% of the inertia (variation) in the sample (online diversity index is plotted against NISP for PP9 and PP13B (Fig. 8). supplementary information, Table E). When the scores calculated for the sites (rows) and adaptation categories (columns) for the first two constructed dimensions (online supplementary information, 4.4. Results: univariate and multivariate analyses Tables F and G), were plotted together, a gradient of habitat types is illustrated (Fig. 9). Though there is a significant amount of overlap Results of the one-way ANOVA revealed that the numbers of between the intermediate biotopes, primarily because the sites species in the aquatic locomotion adaptation, the invertivore dietary categorized as woodland habitats overlap with bushlands, the forest categories, and the sandy and vlei substrate categories were not and desert habitat groups are clearly separate, and there is some differentiation between the remaining habitats apart from the woodlands group. The most congested area of the Figure, where

Table 4 multiple habitat groups converge, is characterized by sites with high General micromammalian diversity in PP9C and PP13B. rainfall for each category (online supplementary information, Table A). Given that micromammals tend not to be as constrained by No. of species (H) NISP fi present (All taxa) ecological factors as larger mammals, it is interesting to nd that the CA results suggest that their community structures can effectively PP9C StratAggs differentiate most of the habitat types, and that those sites with PP9C-SS:2005 19 2.36 158 PP9C-RTE higher rainfall fall within close proximity on the charted Olive Yellow Cave Sand (OYCS) 13 2.03 87 dimensions. Brown Yellow Dune Sand (BYDS) 10 1.98 22 When the fossil assemblages from PP9C are plotted on the PP9C-RTR dimensions created using the variation in modern sites and habitat Brown Yellow Surface Sand (BYSS) 13 0.96 106 Brown Yellow Cave Sand (BYCS) 20 1.04 366 types, they fall closest to the area of overlapping habitats (Fig. 9). The majority of the StratAgg assemblages are at the edge of, or PP13B: StratAggs Western area outside of the habitat groups formed by the modern sites. However, Surface sediments 11 2.15 20 they lie between forests and woodland/bushlands in the wetter LBG Sand 1 17 1.78 207 portion of the Figure, and can be presumed to be indicative of LB Silt 9 1.57 57 LB Sand 12 2.13 39 somewhat closed and wetter habitats. There is some suggestion that DB Sand 3 1.46 9 28 9 1.46 28 the RTR deposits (BYSS and BYCS) may reflect a more open DB Sand 2 1.54 9 20 9 1.54 20 woodland/bushland environment, compared to the RTE deposits (OYCS and BYDS), which lie in the direction of the modern forested PP13B: StratAggs Eastern area Upper Roof Spall 1.76 7 17 7 1.76 17 sites. Though it may be possible to interpret a slight change in Lower Roof Spall 2.58 13 27 13 2.58 27 habitat through time with some periods slightly more closed than others, results of this analysis suggest that, in general, the PP9C PP13B: StratAggs LC-MSA area deposits reported here were characterized by communities reflect- LC-MSA Lower 1.98 12 63 12 1.98 63 ing relatively closed, moist habitats. 220 T. Matthews et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 302 (2011) 213–229

Fig. 9. Plot of ﬁrst two dimensions constructed by the CA using those variables found to be signiﬁcantly different between multiple modern habitat types, with adaptation categories, modern African sites, and PP9C assemblages plotted.

5. Discussion study made of these two species living in the Serengeti region of northern Tanzania. Reed (2005) concluded that Barn Owls and 5.1. Taphonomy of the fossil assemblages Spotted Eagle Owls may be considered isotaphonomic with regards to prey composition unless very sensitive measures of relative As far as can be assessed, the facies which contained only small abundance are required. This has also been reported by other authors samples of incisors, and which have been excluded from Fig. 3, such as Brain (1981) and Mendelsohn (1989). The PP9C micromam- showed a similar pattern of digestion to the larger units, with the mal assemblages are thus thought to be taxonomically comparable to majority of incisors showing light and moderate digestion. those from PP13B, where the main predator responsible for the The incisor digestion patterns of all three fossil assemblages are micromammal accumulations appears to be the Barn owl (Matthews more similar to the Spotted eagle owl than the Barn owl in that they et al., 2009). Both Barn and Spotted eagle owls are likely to produce contain lower numbers of incisors showing no sign of digestion, and prey assemblages which provide a good representation of the slightly higher percentages of incisors with moderate and heavy available micromammal population living in the area. The degree of digestion. The evidence is not clear-cut, however, as for all three breakage of prey bones and teeth in Spotted eagle owl assemblages comparative Spotted eagle owl assemblages studied, 25–26% of the from the south coast was found not to be as advanced as reported by incisors showed moderate digestion. The percentage of incisors Andrews (1990) (Matthews, 2008), and it is considered very unlikely showing moderate plus heavy digestion for OYCS (RTE) and for the that breakage would have affected species representation to any two RTR facies (BYSS and BYCS) are lower at 20%, 13% and 8%, significant degree. respectively. Given the degree of variation in the incisor digestion patterns produced by different predator species, the agents of 5.2. The soricids from PP9C accumulation appear more likely to be Spotted eagle owl, but the Barn owl cannot be entirely ruled out. Avery (1990) noted a decrease in tooth length of Crocidura The effect that a Spotted eagle owl, as opposed to a Barn owl, may flavescens during the Holocene at Boomplaas as indicating a have had on the selection of prey species is not entirely clear, but progressive rise in mean temperature from 10 ka to the present. should not have had too great an influence on taxonomic represen- Crocidura flavescens is present in PP9C in SS:2005, OYCS (RTE), BYCS tation for the following reasons. The Spotted eagle owl is reported as and BYSS (RTR). The fact that no marked changes occur over time in nocturnal in most of the literature (e.g. Andrews, 1990), however, the various mandibular and tooth measurements of C. flavescens at modern comparative Spotted eagle owl assemblages from various PP9C indicates that the population did not respond to climatic change sites around the south coast were found to contain species such as by size change and indicates stability in the local population in terms Rhabdomys, which is a crepuscular species, and species from the of the variables measured. Otomyinae, which are diurnal. Generally, the species and proportions New genetic evidence indicates that modern Myosorex varius is of micromammals found in Barn and Spotted eagle owl pellets were genetically differentiated into a Cape Town group, and a second very similar in the modern south coast pellet assemblages collected, population which stretches from Port Elizabeth up to KwaZulu–Natal and both species appeared to take a very similar suite of animals, with (Willows-Munro, pers. comm.). This was not picked up by previous a numerically similar emphasis on certain prey species (Matthews, morphological and morphometric studies of the crania and teeth of 2008). Reed (2005) found identical taxonomic representation in a this species, and highlights the problems faced when identifying fossil study of the prey taken by Tyto alba affinis and Bubo africanus in a soricid species. New research based on parsimony and Bayesian T. Matthews et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 302 (2011) 213–229 221 analyses of sequence data derived from the mtDNA control region and Diversity in PP9C-RTE assemblages is similar relative to micro- the nuclear intron STAT suggests that M. longicaudatus became mammal assemblages from other MIS 5 archeological sites along the established around 6.7 Ma, when it diverged from the east African south and east coast, such as Klasies River (H ranged from 1.9 to 2.3 M. geata (Willows-Munro and Matthee, 2009). A subsequent single (Avery, 1987)) and Die Kelders 1 (H=2.27–1.75 for MSA deposits) colonization event at ~5.7 Ma then gave rise to a second monophyletic (Avery, 1986) as well as the Holocene, Nelson Bay Cave (H=1.62– clade comprising the forest adapted M. sclateri and M. cafer, and the 1.42) (Avery, 1986). The diversity in the PP9C-RTR is lower than all of more generalist M. varius (Willows-Munro and Matthee, 2009). The the above. taxonomic affinities of the three unidentified soricid taxa from PP9 are uncertain at this point. 5.4. General comments on species representation

5.3. General diversity In the RTR excavation area, BYSS has the potential to contain modern micromammal material. However, the similarity between The relationship between sample size and diversity in a fossil site BYSS and the immediately underlying StratAgg, BYCS, in terms of is well-known (Grayson, 1979). The measurements of diversity (H)in species representation, in addition to the lack of any ‘modern’ species, PP9C did not show this trend and were surprisingly informative suggests mixing with modern material has not occurred to any (Table 4). None of the PP9C fossil assemblages are as diverse as those noticeable extent. This is also suggested by the similarity in found in the SS: 2005 sediments, although OYCS comes close. This percentage representation of many of the species held in common relatively high diversity may be attributable to the mixing of modern between both aggregates. The greater number of species found in and fossil assemblages but, as mixing does not appear to be at all BYCS (N=20, as opposed to N=13 in BYSS) is clearly related to extensive (see Section 5.5.3 for further discussion of this issue), is relative sample size. likely to reflect a genuinely higher diversity. In excavation area RTE, OYCS likewise shows a similar percentage The RTR sediments, which were of good sample size (NISP=105 representation of species to the underlying StratAgg BYDS, with the for BYSS and NISP=362 for BYCS), showed a lower diversity than any exception that soricids are much better represented in OYCS. These of the other fossil assemblages, even those with small sample size, and results indicate similarities between the micromammals in different including those from PP13B. The relatively low diversity in these areas horizons within the same excavation area, a pattern also observed in thus appears to be real. Diversity in BYDS (RTE) is surprisingly high PP13B. given the small sample size and supports the trend shown by the The Dendromurinae (Climbing mouse) are never common, and are superimposed OYCS, which is that the RTE StratAggs contain found in frequencies of 3–5% in all the larger PP9C assemblages, assemblages with a relatively high diversity. including SS:2005. Dendromus mystacalis appears to be more common The diversity indices from Cave PP13B may be compared to the than D. mesomelas as the former is found in six of the StratAggs, as results from PP9C. The LC-MSA Lower sediments, which are well as SS:2005. D. mesomelas is found only in SS:2005, and in the RTR approximately 40 ka older and date to MIS 6 (162±6 ka), had a is represented by only one specimen in each StratAgg (LYDS and diversity of H=1.98 (Matthews et al., 2009), the same as BYDS (RTE). BYCS). D. mesomelas was absent from all the PP13B StratUnits, In the Western area of PP13B, the facies Light Brown Grey Sand 1 although D. mystacalis appeared in a number of StratUnits dating to (LBG Sand 1) revealed ages consistent with two pulses of deposition MIS 5, confirming that D. mystacalis appears to have been present in at 124±5 ka and 99±4 ka. The diversity index for the faunal the area for a long period (Matthews et al., 2009). assemblages contained within these sediments is H=1.78, which is Gerbilliscus afra (Cape Gerbil) is represented by 1–3 individuals in greater than PP9C-RTR, and less than PP9C-RTE. The highest diversity the RTR area in BYSS and BYCS, and in RTE, in OYCS and BYDS. This in PP13B (H=2.58) was found in the Lower Roof Spall Facies in the species was found in low frequencies in several PP13B StratUnits Eastern area, which is slightly younger than the RTR and RTE facies dating to both MIS 5 and MIS 6 (Matthews et al., 2009). at 114±4 ka. This diversity is greater than that found in any of the The southwestern Cape endemic, Myomyscus verreauxi, (Ver- PP9C assemblages, despite the fact it was of relatively poor sample reaux's Mouse) is present in relatively low frequencies in both size. Of all the PP9C and PP13B assemblages, SS:2005 comes closest excavation areas of PP9C, and was present in some of the MIS 5 PP13B to the diversity shown by the Lower Roof Spall Facies. The PP13B facies, albeit in low frequencies. This species is one of several small StratUnit, Light Brown Sand 1 (LB Sand 1), contained a relatively small rodents that have been associated with the pollination of Protea sample size (NISP=39) but is worth mentioning. This StratUnit has species (David, 1978; David and Jarvis, 1983), along with A. been dated to 90±4 ka, and has an H value of 2.13, which is slightly namaquensis, Acomys subspinosus and E. edwardii (Fleming and higher than the diversity in OYCS and BYDS (RTE), and quite a bit Nicolson, 2002), and may well be indicative of fynbos vegetation. higher than the RTR facies. The presence of numerous micromammal taxa from PP9C to The evidence from both PP9 and PP13B indicates there is some PP13B, which date to MIS 5e–5a and MIS 6, and which are currently congruence in the diversity indices from sedimentary deposits that associated with fynbos, indicates a fynbos component in the area date to periods within both MIS 6 and MIS 5, although this may be in during these periods. Bar-Matthews et al. (2010) hypothesize that the part due to the fact that the MIS 6 deposits represented by LC-MSA now submerged coastal platform provided a likely refuge zone for Lower may represent a stadial, and a period of climatic amelioration. fynbos vegetation, which may have followed the coastline during sea There is, however, some evidence from PP13B and PP9C that there level regressions, while current coastal locations were enveloped in a was a relative increase in diversity of the resident micromammalian more mixed C3–C4 vegetation. The situation is, however, complex fauna at Pinnacle Point as MIS 5 progressed. with some fynbos endemics scarce or lacking in the fossil assem- Traditionally, low general diversity in micromammal assemblages blages. An example of such a species is A. subspinosus, a true fynbos was interpreted as representing harsher environmental conditions endemic, which is noted as currently having a distribution confined to and high diversity with climatic equitability (e.g. Avery, 1986; Brown, the southwestern parts of the Cape Province, stretching from Citrusdal 1973), but this link has not been clearly established. Avery (1999b) in the west, to Knysna in the east (Skinner and Smithers, 1990). This found no correlation between diversity and climatic variables in a species did not make an appearance in either PP9 or PP13B. study of micromammal assemblages from Barn owl pellets. The PP9C Elephantulus edwardii was found only in RTR in PP9C, and in PP13B data likewise suggests that the link between diversity and climatic in only one facies (LBG Sand 1) dated 124±5 and 99±4 ka (Jacobs, change is complex. 2010). 222 T. Matthews et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 302 (2011) 213–229

Zelotomys cf. woosnami and Thamnomys dolichurus were found in grassy hillsides (Stuart and Stuart, 2001). Interpreting the PP13B, but did not appear at all in PP9C. The reasons for their non- environmental implications of the two species in fossil assem- appearance are uncertain at this stage as similarly aged deposits are blages is also complicated by the fact that the distribution of O. represented in PP9C. saundersiae is under-reported in the literature (Avery et al., 2005; Based on age estimates alone, it appears that the micromammal Matthews, 2008), indicating that this species is currently, and was assemblages date to a period close to the MIS 6 to MIS 5e transition. in the past (Avery, 1977, 1992a), far more widespread than Any significant changes in the microfauna assemblages may, generally reported. Also, O. irroratus and O. saundersiae are therefore, provide better estimates of whether or not the lowermost frequently found together in owl pellet assemblages, indicating assemblages represent the MIS 6 terminus, or whether it is more likely they can occupy the same areas simultaneously, although they may that all micromammals were accumulated during the same climatic inhabit different microhabitats (Matthews et al., 2009). O. irroratus period. An inverse relationship between relative species representa- is the dominant Otomyine in several PP13B StratUnits dating to tion in the RTE and RTR deposits (Figs. 4–6) may indicate that the two MIS5, whereas O. saundersiae is dominant in the PP13B LC-MSA excavation areas represent different sides of the MIS 6 to MIS 5e Lower deposits dating to MIS 6. However, the dominance of transition. This is discussed further below. The discussion focuses on O. irroratus in interglacials and O. saundersiae during glacial periods the larger fossil samples, namely OYCS (RTE), BYSS and BYCS (RTR), is not a clearly defined pattern, as the latter dominates in the and SS:2005. eastern area of PP13B in the Upper Roof Spall Facies, dating from later MIS 5 (Matthews et al., 2009). The assemblage is of poor 5.5. Environmental implications of the micromammals sample size (N=17), however, and the data may be problematic. ThepredominanceofO. saundersiae during cooler periods would, Results of the taxon-free CA suggest relatively dense habitats, however, concur with Thackeray's (1987) results whereby tem- with good rainfall for both the RTE and RTR assemblages, with a perature indices were generated from multivariate analyses relatively slightly more open environment in the latter. Palaeoen- (Factor Analysis) of the relative abundances of micromammals vironmental reconstruction based on the micromammalian faunas is from South African cave deposits dating the Late Pleistocene and discussed in the following sections. Information pertinent to the Holocene. This methodology was used subsequently to investigate following sub-sections, relating to the breeding habits and habitats Plio-Pleistocene fossil sites from the Cradle of Humankind utilized by some of the micromammals mentioned, may be found in (Sénégas and Thackeray, 2008) and at Klasies River and Border Appendix D. Cave (Thackeray and Avery, 1990). Loadings on the first factor (F1), The micromammals represent a small and discreet environment associated with certain rodent species analyzed by Thackeray relative to the large fauna from the site. For example, the murid (1987), fell within the range of 1 and −1, and taxa with extreme Rhabdomys pumilio reportedly has an average home range size of only loadings were compared. Taxa with relatively high loadings were 0.001 km2 to 0.015 km2 (Schraden and Pillay, 2005). The owls represented by species found today in relatively warm environ- accumulating the fossil micromammal population are likely to have ments, whereas those found in cooler environments, or at high had a hunting range of 400–500 m, though ranges may be as large as altitudes, had lower F1 loadings (see Thackeray (1987) for further 3km(Andrews 1990, Avery 1992b). According to a bathymetric 3D details). At Boomplaas and Nelson Bay Cave, O. saundersiae,which coastline model for the south coast (Marean et al., 2007) the coastal Thackeray's (1987) results suggest is a colder tolerant species, plain uncovered by lower sea levels predicts that through most of dominated assemblages which dated to periods of global cooling. If MIS5 there was a coastal plain of several kilometers in front of the these results are applied to the relative abundances of the two cave. However, during MIS 5e the sea was higher, and the cave sat Otomys species found in the PP9C assemblages, the fact that O. above a rough wave-dominated zone. The owl accumulating the fossil irroratus is relatively more abundant than O. saundersiae in the RTE assemblages is presumably more likely to have hunted on the cliff assemblages (dated to 120±7 ka and 126±9 ka) provides further tops, than below the cave. evidence that the RTR assemblages represent different, and probably warmer conditions, to the slightly older (circa 130± 5.5.1. PP9C-RTE 9ka)O. saundersiae-dominated RTR assemblages. In OYCS and BYDS the percentage representation is very similar for M. varius and S. varilla are two other relatively common species in most species, with the majority of species held in common appearing OYCS. M. varius is likely to be associated with densely vegetated, moist in frequencies of under 10%. Rhabdomys is one of the relatively more environments on the south coast (Stuart and Stuart, 2001), but is common species at ~14% in both StratAggs. An exception to this adaptable and found in drier conditions on the arid west coast today general similitude of the two StratAggs is the Soricidae, which are where its presence appears to depend on dense succulent vegetation poorly represented in BYDS and show none of the diversity observed (Bigalke, 1979). in OYCS. Three, out of the four, most common species in OYCS (O. Rhabdomys is a very versatile species (see Appendix D) and lives in irroratus, M. varius and S. varilla) are frequently associated with habitats which range from desert fringe to high rainfall areas (Stuart relatively moist, dense habitats, while the abundance of R. pumilio and Stuart, 2001). Clearly conditions were optimal for this species at indicates good ground cover and rainfall. Another indication for a the time of accumulation in PP9C, which suggests that there was a relatively moist environment is the fact that the Soricidae make up reasonable amount of cover available, and sufficient rainfall to 24.4% of the micromammal assemblage. The StratAgg BYDS produce food to stimulate recruitment (Bond et al. 1980; Perrin appears to be representing a period when soricids were less 1980, 1981). R. pumilio and S. krebsii are both seasonal breeders, have commonly preyed upon, but whether this was because they were a similar diet of seeds and insects, and could easily inhabit the same scarce in the landscape is uncertain as it may also reflect cyclical habitat. In PP13B, R. pumilio is found in very low frequencies, with the changes in local populations, changes in the area the owl was exception of LBG Sand 1 in PP13B, which has been dated to 124±5 hunting, or occupation of the roost in a different season which has and 99±4 ka and thus (in terms of the earlier date) contains deposits been noted as affecting sample structure (Avery, 2002c), rather of a similar age to OYCS. than any environmental change. The general picture produced by In the RTE deposits the Otomyinae are the most common murid an analysis of the RTE micromammals is that conditions were taxa, with O. irroratus dominating these assemblages. Otomys relatively warm and moist with relatively dense vegetation. This is irroratus is associated with moist, marshy habitats, but may also be consistent with ages of 120±7 and 126±9 ka, indicating accu- found in drier habitats (Rowe-Rowe and Meester, 1982), such as mulation during MIS 5e. T. Matthews et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 302 (2011) 213–229 223

5.5.2. PP9C-RTR in savanna, but encroach into the southwest Cape, Southwest arid, BYSS and BYCS differ from OYCS (RTE) in several aspects, but and lowland forest biomes. also show a high soricid abundance, although C. flavescens appears Interpreting the environmental implications of the dominant in unprecedentedly high frequencies. The Soricidae are more taxa, namely O. irroratus, O. saundersiae, S. krebsii, S. varilla, S. varius common than the Otomyinae, and these two groups of taxa, and C. flavescens is complicated due to factors mentioned previously, together with S. krebsii dominate the faunal assemblages. Unlike and the fact that all of these taxa are found in a variety of the RTE excavation area, Rhabdomys occurs in relatively low environments (Appendix D). The dominance and species diversity proportions. of the soricids as a group, however, indicates a relatively dense, and The elephant shrew, E. edwardii, and the bat, R. clivosus,were moist environment. This is supported by the presence of D. recovered only from BYCS (RTR) and were represented by only 1–2 mystacalis which favors tall grasses and thick vegetation and is individuals. E. edwardii is associated with rocky slopes and hard, relatively abundant in BYCS. In addition, species which might be sandy ground with sparse vegetation (Skinner and Smithers 1990). expected to reflect drier, more sparsely vegetated or sandier This species was also scarce in PP13B and was found only in the environments, such as E. rupestris, A. namaquensis and G. afra,are Laminated Facies which was dated to 385±17 ka (Jacobs, 2010), not found in greater abundance in the StratAggs rich in S. krebsii, and in LBG Sand 1 dated to two discrete sediment pulses at 124±5 which, together with the abundance of soricids, suggests this and 99±4 ka. A. namaquensis (Namaqua Rock Rat) is also a species was occupying a habitat which may have differed from the relatively scarce species and is found only in SS:2005, and in BYSS sandy environment it is frequently associated with. A grassy and BYCS. The relative scarcity of this species seems to be confirmed component to the vegetation may be indicated from the presence by the fact that only one specimen was found at PP13B, in the of both S. krebsii and O. saundersiae, and the latter may indicate that StratAgg ‘Truncation Fill’, which may date to approximately 35– somewhat colder conditions existed relative to the period of 39 ka. It is possible that the presence of both E. edwardii and A. deposition of the RTE deposits (as discussed in the previous namaquensis in BYCS indicates a rocky component in the immediate section). This, together with the presence of A. namaquensis and E. environment. The general scarcity at Pinnacle Point of both species edwardii, may indicate somewhat more open, and possibly grassier, in both PP13B and PP9C may be attributable to predator choice, conditions than when the RTE sediments accumulated, but the however the consistency in the evidence between PP13B and PP9C evidence for this is not unequivocal and there are no indications of a suggests that the rocky and open habitat which these two species substantial change in environment. A similar pattern emerges from prefer, was never available on a wide scale immediately around PP13B. Not only is there no indication of marked environmental Pinnacle Point. change over the time periods dating to MIS 5 represented by the O. saundersiae is more abundant than O. irroratus, the opposite Western area, but there is a marked consistency in the pattern of pattern to that seen in the RTE facies. The possible relevance of this species abundance over time (Matthews, et al., 2009). has been discussed in Section 5.5.1. The low diversity in the RTR facies, commented on previously, has In PP9C soricids make up 38–44% of micromammal taxa in BYSS resulted from an emphasis on a number of abundant species which and BYCS, respectively. There is thus a marked surge in the soricid appear to have been experiencing a seasonal boost, probably due to population in these StratAggs. In these StratAggs, as well as in conditions favorable to breeding. The high percentages of seasonal SS:2005, Myosorex, Suncus,andCrocidura are all well represented. breeders in the assemblage suggest that these animals had responded The very high percentage of C. flavescens (36%) found in BYSS to the onset of the rainy season (this instigates breeding in the case of occurs only in SS:2005. The versatility of the habitats occupied by many rodent species). The soricids are generally seasonal breeders in Myosorex and Suncus (Appendix D) complicates elucidation of their the wet summer months, as is S. krebsii (Stuart and Stuart 2001), and environmental significance. Interpreting the abundance of C. it is possible that the dominance of these taxa is reflecting their flavescens is also problematic as reports regarding its current seasonal abundance in the landscape. RTR appears to be presenting a distribution (see Appendix D) note that this species is confined to brief snapshot in time. areas with a mean annual rainfall of between 500 and 750 mm. This is erroneous as C. flavescens was recovered from modern owl roosts 5.5.3. PP9C-SS:2005 on the west coast (Avery, 1999a; Avery et al., 2005)wherethe Similarities between some features of the faunal assemblages from annual rainfall falls between 275 and 150 mm (Weather Bureau, SS:2005 and the underlying BYSS in terms of the relative represen- 1981). Fossil evidence also indicates that this species enjoyed a tation and abundance of soricid species (Fig. 7), and the proportion of wider distribution in the past (Avery, 1992a, 1997). C. flavescens Otomyinae to Soricidae (Fig. 6), may indicate some mixing of fossil was found mainly in the older StratUnits of the three excavated and younger material. The evidence is rather contradictory though as areas of PP13B, namely the LC-MSA Lower which date to MIS 6 differences also exist between SS:2005 and BYSS in the relative (162±6 ka) and LB silt (157±10ka), but also occurred in the proportions of S. krebsii and R. pumilio (Fig. 4), in the relative Lower Roof Spall sediments (111±4 ka). The fossil evidence to proportions of the two Otomyinae species (Fig. 5), and in the date indicates that this species was present at Pinnacle Point during substantially higher diversity (H) of the SS:2005 assemblage. In MIS 6, MIS 5e and MIS 5c. addition, species representation in SS:2005 differs to that of the fossil The relatively high frequencies in which S. krebsii (Kreb's Fat assemblages in the following features; Rhabdomys and Steatomys Mouse) is present in BYCS and BYSS are quite different to many appear to experience a converse relationship in the fossil horizons, but areas of PP9C where this species is absent, or present in low in SS:2005 they appear in close to equal proportions, and the frequencies (ie. OYCS and BYDS). S. krebsii is thought to prefer a otomyines and soricids dominate to a lesser extent than in the fossil sandy substrate, and to occur in dry, sandy grassland conditions, as assemblages. This suggests that, if there is mixing, it is not extensive. well as in sandy alluvium (Skinner and Smithers, 1990). It is Two soricid (C. cyanea and S. infinitesimus) and one murid species therefore somewhat unexpected that this species is found in highest (S. campestris), are found only in SS:2005. Their presence confirms frequencies in the units in which soricids (which are generally that this assemblage represents a relatively ‘modern’ collection. This is accepted as indicators of “moisture” and dense vegetation) are inferred from the fact that another modern surface collection made dominating the micromammal assemblage. This abundance was from a small cave slightly to the south of PP9C, dubbed Caterpillar also unexpected in that S. krebsii was represented by only one Cave, was also found to contain C. cyanea and S. infinitesimus, as well specimen in PP13B in a StratUnit dating to 90 ±4 ka. Nel (1969) as M. varius, S. varilla and C. flavescens (Matthews, pers. ob.). In terms notes that species comprising the genus Steatomys occur primarily of Soricidae, the fossil horizons from PP13B contained only M. varius, 224 T. Matthews et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 302 (2011) 213–229

S. varilla, and C. flavescens (Matthews et al., 2009). C. cyanea, S. micromammals to be generalists and capable of adapting to a variety of infinitesimus and one S. campestris were not recovered from PP13B, ecological situations. Therefore, it is possible to categorize one species not even in the modern, surface sediments (Matthews et al., 2009). in multiple ways, and for a CA only one categorization can be used. The appearance of S. infinitesimus at PP9C in surface sediments Further explorations of ways in which these adaptations can be confirms that this species has a more widespread distribution than defined and assigned may result in better habitat description. reported. This is supported by the fact that S. infinitesimus is currently Additionally, this analysis did not include any habitats that were found to have a greater distribution along the south coast, including defined as fynbos, and some of the reconstructions of paleohabitats the Mossel Bay area, than reported in the literature (Matthews, 2008). from PP9C that did not show clear affiliations with any modern habitat The presence of S. campestris is also indicative of a modern component type may, in fact, be reflecting fynbos ecology. Because fynbos in the assemblage. It is not found in any other fossil horizon from vegetation is unique to South Africa, it is rarely included in broad Pinnacle Point, and Avery (1977) noted that archeological evidence scale studies of African paleoecology, and thus modern sampling of the suggests that S. campestris arrived in the southern Cape at the biotope for that purpose is rare. While fynbos communities have been beginning of the Holocene, which places the ‘modern’ component of included in paleoecological reconstructions for larger mammals with SS:2005 sometime in the Holocene. some success (Rector and Reed, 2010), future work with community It is acknowledged that the mixed nature of SS:2005 renders a structure analyses of specifically modern fynbos communities will direct comparison with other sites unsatisfactory. It is worth noting, help elucidate how these habitats can be reconstructed in the fossil however, that differences observed between the SS:2005 and fossil record. assemblages suggest that recent conditions may have differed quite The RTR sediments appear to be representing a brief period in time considerably to the periods documented by the fossil deposits, as this when the dominance of soricids and otomyine taxa, and S. krebsii, scenario is supported by the micromammal assemblages from PP13B reflect their seasonal abundance in the landscape. The brief time and Caterpillar Cave. Large mammal faunas from the Western Cape period represented may have affected general diversity, as diversity is also suggest that conditions during MIS 1 differed from those markedly low in the RTR facies, as compared with the PP9C RTE observed in MIS 5 and MIS 6 (Klein, 1983; Rector and Verrelli, StratAggs, the PP13B fossil assemblages, and with other sites such as 2010). Anthropogenic alteration of the landscape around Pinnacle Klasies River, Die Kelders 1 and Nelson's Bay Cave. Point has been considerable and has undoubtedly had a large effect on The RTR and RTE excavation areas show an interesting converse the resident micromammal population. relationship between the proportion of soricids to otomyines, S. krebsii to R. pumilio, and the domination of O. saundersiae versus O. 6. Conclusions irroratus. This may be attributable to the time periods in which they accumulated, with the RTR deposits accumulating at the end of MIS 6/ It appears that C. cyanea,andS. infinitesimus are relative beginning of MIS 5e, and the RTE deposits accumulating during 5e. latecomers to the Mossel Bay region as these species were found During the accumulation of the RTR deposits, there are some only in surface sediments in SS:2005, and in modern sediments from indications that conditions were relatively grassier, and may reflect Caterpillar Cave. S. campestris has been recovered only from the a rather more open, and somewhat colder, environment relative to surface sediments of PP9C and suggests a Holocene age for the when the RTE facies were deposited. However, given the adaptability modern component of this assemblage (Avery, 1977). and versatility of the taxa involved, the evidence is not unequivocal. The Pinnacle Point fossil sites indicate that C. flavescens was a Variation in the exposure of the coastal platform off Pinnacle Point common component of the local micromammal fauna in the older would have affected the local environment and the available hunting fossil sediments, and was present at Pinnacle Point during MIS 6 and area below the cave available to the owls accumulating the fossil MIS 5e and 5c. It also appears in surface sediments dating to sometime assemblages. The general scarcity of E. edwardii and A. namaquensis at within the Holocene. This species appears to be very adaptable to PP suggests that the rocky and open habitat which these two species climatic change and can utilize much drier habitats than reported in prefer, was never available on a wide scale at PP. the literature, which complicates elucidation of the habitat of fossil Generally speaking, the micromammal assemblages of the RTE and representatives. The diversity of species of Soricidae observed in the RTR areas indicate that the periods during which the micromammals Holocene PP sediments may have been matched in the past as some 3 accumulated were warm and wet, and that the vegetation within the unidentified soricid species from the PP sediments have been recovered. hunting range of the owls accumulating the micromammal assem- To date these have only been tentatively allocated to genera. blages was relatively dense. This conclusion is supported both by the The presence of numerous micromammal taxa from PP9C and habitats and habits of the micromammal faunas in the various PP13B, which date to MIS 5e–5a and MIS 6, and which are currently StratAggs, as well as by multivariate analysis. associated with fynbos, indicates a fynbos component in the area during these periods. Acknowledgements Isotopic (18O and 13C) speleothem data from the Pinnacle Point caves indicates that during periods when global conditions were Thank you to the National Science Foundation (USA) (grants # warmer there was an increase in winter rainfall, and in C3 grasses BCS-9912465, BCS-0130713, and BCS-0524087 to C.W. Marean), the (Bar-Matthews et al., 2010). In cooler periods, there was an increase Hyde Family Foundation, IHO, and ASU for funding the excavations, in summer rainfall and in C4 grasses, with possibly grassy fynbos or analysis, and write-up. We are grateful to the National Center for thicket vegetation, or a mosaic of the two, existing at Pinnacle Point Ecological Analysis and Synthesis for use of the Recent Mammal (Bar-Matthews et al., 2010). D. mystacalis is found in frequencies of 3– Database, as well as Dr. Robert Williams, Dr. Kaye Reed, and Amy 5% in all the larger PP9C assemblages. Today it is found on the eastern Shapiro for assistance with manipulation of the data. Our thanks too, coast in Albany thicket, and its presence in the PP13B and PP9C fossil to the many people who have worked on PP9C and contributed to this sites in horizons dating to MIS 5 provide some supporting evidence research. AIRH would like to acknowledge additional support from the that this biome, or certain elements of this biome, extended further UNSW Faculty of Medicine and ARC Discovery grant DP0877603. down the south coast in the past. A first attempt at using micromammal community structure of Appendix A. Supplementary data modern African habitats in a correspondence analysis to reconstruct palaeohabitats was relatively successful in discriminating between Supplementary data to this article can be found online at habitat types, but was in all likelihood affected by the tendency of doi:10.1016/j.palaeo.2011.01.014. T. Matthews et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 302 (2011) 213–229 225

Appendix

Appendix B Micromammal species (NISP) present in the PP9 Stratigraphic Aggregates.

StratAgg NISP (all Number of 1 2 34567 8 9 1011121314151617181920212223242526272829 taxa excl. identiﬁed bathyergids) species

PP9B Yellow brown 11 1 sand Olive Yellow 11 1 Sand Mixed contact Only 1 murid incisor recovered horizon PP9C: SS:2005 156 18 3 20 9 1 30 7 2 2 6 31 1 16 2 8 16 2 * *

PP9C: FC Yellow Brown 11 1 Sandy Silt Clay Yellow brown 4411 1 1 clay Light Yellow 8 6 131 111 * Brown Sandy Clay 1 Light yellow 11 1 brown sand Dark yellow 95 4 1 12 1 brown sand Mixed Surface 15 6–7 7112 1 1 1 1 Sand Brown clay 1 15 5 1 1 1 2 9 1 Rubiﬁed clay 111 units

PP9C:RTE Olive Yellow Cave 86 13 8 25 1 7 12 2 4 1 3 1 8 10 2 * 2 Sand Brown yellow 21 10 1 8 1 1 2 3 1 1 1 1 1 * dune sand Raised beach2 3 2 2 1 Yellow cemented 7215 1 sand

PP9C:RTR Light yellow 10 8 1 1 1 1 1 2 1 2 decalciﬁed sand Brown Yellow 105 13 18 9 1 3 2 3 3 3 16 14 16 17 * Surface Sand Brown Yellow 362 20 78 14 4 3 20 14 19 1 15 2 46 56 54 4 24 1 1 1 1 * 3 4 Cave Sand Note: * = taxa represented only by maxillae and/maxillary teeth. Bathyergids are excluded from the NISP given above as this taxa was generally represented by single teeth or maxillary fragments.

Key to species

Muridae Common name Common name Macroscelididae Common name (Macroscelidea)

Otomyinae Gerbillinae 23 Elephantulus edwardii Cape rock elephant shrew 1 Otomys saundersiae Saunders vlei rat 12 Gerbilliscus afra Cape gerbil Chiroptera (Rhinolophidae) 2 Otomys irroratus Vlei rat Cricetomyinae 24 Rhinolophus sp. 3 Indeterminate Otomys sp. 13 Steatomys krebsii Kreb's fat mouse 25 Rhinolophus clivosus Geoffrey's horseshoe bat 4 O.irroratus/O. saundersiae 14 Saccostomys campestris Pouched mouse Bathyergidae Murinae Insectivora (Soricidae) Bathyergus 5 Aethomys namaquensis Namaqua rock mouse 15 Suncus sp. 26 Indet. mole rat 6 Myomyscus verreauxi Verreaux's mouse 16 Suncus varilla Lesser dwarf shrew 27 Bathyergus suillus Cape dune molerat 7 Rhabdomys pumilio Striped mouse 17 Suncus inﬁnitesimus Least dwarf shrew Cryptomys 8 Mus minutoides Pygmy mouse 18 Myosorex varius Forest shrew 28 Cryptomys hottentotus Common molerat 9 Mystromys albicaudatus White-tailed mouse 19 Myosorex sp. Chrysochloridae Dendromurinae 20 Crocidura ﬂavescens Greater musk shrew 29 Indet. chrysochlorid 10 Dendromus mesomelas Brant's climbing mouse 21 Crocidura cyanea Reddish-grey musk shrew 11 Dendromus mystacalis Chestnut climbing mouse 22 Indeterminate soricid 226 T. Matthews et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 302 (2011) 213–229

Appendix C Measurements made on Pinnacle Point Crocidura ﬂavescens (Soricidae) mandibles.

Abbreviation Description of measurement

I2P Combined length of I2 and P4, in lingual view

ML1–3 Total length of tooth row of the M1,M2 and M3 measured in lingual view

ML1–2 Total length of tooth row of the M1 and M2, measured in lingual view

ML1length Maximum lingual length of M1

ML1breath Maximum lingual width of M1

ML2length Maximum lingual length of M2

ML2breadth Maximum lingual width of M2

R Depth of the horizontal ramus below M3. Measurement was taken from the middle of the last cusp of M3 F Greatest anteroposterior diameter of the temporalis fossa SCL Subcondylar length, from the subcondylar embayment to the anterior edge of the ascending process

ARL Length of the ascending ramus, from the alveolus of M3 to the posterior edge of the condyle COH Coronoid height, from the top of the process to the ventral embayment anterior to the angular process CH Maximum height of the condyle, on the medial side

Results of measurements made on Pinnacle Point Crocidura ﬂavescens (Soricidae) mandibles.

Site and StratAgg Accession number Measurement (mm)

I2P ML1–3 ML1–2 ML1 ML1 breath ML2 length ML2 breadth R F SCL ARL COH CH

PP13B 13B 99516 5.8 4.3 2.2 1.6 1.9 1.5 2.9 13B 99516 5.9 3.7 2.2 1.7 2.0 1.5 2.7 13B 99389 2.8 5.6 3.1 2.0 2.0 1.7 1.6 2.7 13B 99676 2.1 1.6 2.7 2.8 3.5 7.2 2.1 13B 99511 5.6 4.0 2.1 1.6 2.0 1.5 2.8 2.4 3.5 7.7

PP9C SS 100344 4.2 2.0 1.5 2.0 1.2 SS 100345 2.1 1.6 0.0 0.0 0.0 0.0 7.3 0.0 SS 100270 5.8 4.4 2.1 1.8 2.1 1.5 2.5 2.0 3.5 6.3 7.3 2.2 SS 100271 4.1 2.0 1.7 2.0 1.4 2.1 SS 100272 3.0 4.4 2.1 1.7 2.0 1.5 2.6 2.2 3.5 7.7 SS 100273 3.1 5.8 4.2 2.2 1.8 2.0 1.4 2.8 2.3 3.9 6.4 7.8 2.4 SS 100274 3.1 4.0 2.0 1.7 1.9 1.4 2.6 2.3 3.5 7.9 7.5 2.2 SS 100275 5.7 4.2 2.1 1.7 2.0 1.5 2.5 2.4 3.5 6.3 7.2 2.1 SS 100276 4.3 2.1 1.8 2.0 1.5 2.6 SS 100277 2.9 5.5 3.6 2.0 1.6 2.0 1.5 2.6 SS 100278 2.9 4.1 2.1 1.8 1.9 1.5 SS 100279 4.1 2.2 1.7 2.0 1.5 SS 100280 1.7 2.0 0.0 SS 100281 4.4 2.2 1.6 2.1 1.4 2.3 SS 100283 2.2 SS 100282 5.6 3.6 2.1 1.9 2.8 2.1 4.0 6.5 7.7 2.4 BYCS 100435 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 BYCS 100435 2.7 5.6 3.6 2.0 1.6 2.0 1.4 2.5 1.8 3.6 5.8 6.9 2.2 BYCS 100435 4.5 2.2 1.8 2.0 1.5 2.1 2.4 3.7 7.6 2.1 BYCS 100435 2.6 5.2 3.8 2.0 1.5 1.7 1.3 2.4 2.1 3.2 5.6 6.8 2.0 BYCS 100435 5.2 3.8 2.0 1.5 1.9 1.4 2.3 2.2 3.2 5.7 6.8 2.0 BYCS 100435 5.5 4.0 2.0 1.8 2.0 1.5 2.5 2.1 3.5 6.1 2.1 BYCS 100435 2.4 BYCS 100435 4.0 2.1 1.6 2.0 1.5 BYCS 100439 1.5 1.2 1.7 BYCS 100438 4.2 2.1 1.6 2.0 1.4 1.8 1.9 3.2 6.3 1.8 BYCS 100438 4.4 2.1 1.6 2.2 1.4 2.0 1.9 3.3 6.6 2.0 BYCS 100432 5.7 4.2 2.1 1.7 2.0 1.5 2.6 2.2 3.7 6.1 7.5 2.0 BYCS 100428 5.6 4.5 2.0 1.8 2.0 1.6 2.8 2.7 3.5 6.3 7.5 2.1 BYCS 100428 5.7 4.3 2.2 1.7 2.0 1.5 2.7 1.9 3.4 6.2 7.6 2.3 BYCS 100428 6.0 4.5 2.3 1.9 2.2 1.5 2.8 2.2 3.6 6.4 7.6 2.5 BYCS 100428 3.2 5.7 4.1 2.0 1.6 1.9 1.3 2.9 2.0 3.9 6.2 7.6 2.3 BYCS 100428 5.9 4.3 2.3 1.7 2.0 1.5 2.5 2.2 3.5 6.0 7.7 2.1 BYCS 100428 3.0 5.6 4.2 2.2 1.7 1.9 1.5 2.4 2.0 3.5 6.2 7.3 2.2 BYCS 100428 3.0 5.5 4.0 2.1 1.6 1.9 1.4 2.5 2.3 3.5 5.8 6.9 2.0 BYCS 100428 4.3 2.2 1.6 2.0 1.4 2.0 1.9 2.1 BYCS 100428 2.1 1.6 2.2 1.9 1.9 BYCS 100425 3.8 2.1 1.8 2.2 1.4 2.4 2.3 3.6 2.2 BYCS 100367 3.2 5.7 4.1 2.1 1.8 1.9 1.5 2.8 2.5 3.5 6.4 7.4 2.6 BYCS 100366 4.1 2.1 2.0 BYSS 100430 2.5 2.1 3.6 7.2 2.4 BYSS 100422 5.8 4.1 2.1 1.7 2.1 1.5 2.5 3.4 2.0 BYSS 100422 2.8 5.4 3.8 2.0 1.6 1.9 1.4 2.6 2.2 3.5 5.5 7.1 1.9 BYSS 100422 3.2 0.0 4.4 2.1 1.8 2.0 1.5 2.6 2.1 4.0 8.3 7.6 2.4 BYSS 100422 2.9 5.6 4.0 2.0 1.7 2.0 1.5 2.9 2.1 3.7 6.2 7.3 2.3 BYSS 100422 4.0 2.1 1.7 1.9 1.4 3.0 2.1 3.9 7.5 2.4 T. Matthews et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 302 (2011) 213–229 227

Appendix(continued B ()continued) Site and StratAgg Accession number Measurement (mm)

I2P ML1–3 ML1–2 ML1 ML1 breath ML2 length ML2 breadth R F SCL ARL COH CH

PP9C BYSS 100422 4.2 2.1 1.6 2.0 1.4 2.4 BYSS 100422 4.1 2.0 1.6 2.0 1.3 2.0 2.3 3.7 7.3 2.3 BYSS 100422 5.9 1.9 BYSS 100412 5.7 3.8 2.2 1.7 2.0 1.4 2.7 2.2 3.6 6.4 7.5 2.2 BYSS 100412 5.8 4.3 2.1 1.8 1.9 1.5 2.8 2.5 3.5 6.5 7.2 2.3 BYSS 100408 1.8 2.3 3.5 7.2 2.0 BYSS 100408 5.5 3.9 2.0 1.5 2.0 1.4 2.3 2.1 3.3 5.6 6.7 1.9 BYSS 100399 6.0 4.3 2.2 1.8 2.1 1.5 2.7 2.4 3.5 6.5 8.0 2.5 BYSS 100399 2.7 2.4 3.6 6.3 7.9 2.3 OYCS 100398 2.1 1.7 3.0 2.4 LYBS 100401 5.8 4.0 2.0 1.6 2.0 1.5 2.5 2.4 4.0 6.4 7.9 2.3

Caterpillar Cave SS 100237 2.1 1.4 2.5

Key to PP9C aggregate names: SS = Surface Sand, BYCS = Brown Yellow Cave Sand, BYSS = Brown Yellow Surface Sand, OYCS = Olive Yellow Cave Sand, and LYBS = Light Yellow Brown Sandy Clay 1.

Appendix D Additional notes on the habits and habitats of some murid and soricid species from PP9C.

Muridae

Rhabdomys pumilio Breeding: This species is a cyclical, r-selected breeder and breeding took place from September to March in a study of a Cape Flats population (in a winter rainfall area) (Henschel et al., 1982) and in summer, in a summer rain-fall area (the Fish River Valley) (Perrin, 1981). Perrin (1980) demonstrated that dietary changes in R. pumilio were significantly correlated with rainfall, and this in turn significantly affected pregnancy and recruitment. Perrin (1981) notes that it appears that R. pumilio rapidly increases the number of progeny it has when conditions are optimal, but declining recruitment parallels resource depletion. Habitat: The great versatility of this species is demonstrated by the fact that in the arid succulent Karoo region R. pumilio lives in social groups, but in the moist grasslands of South Africa is solitary (Schraden and Pillay, 2005). This species is reported as needing grass cover in the literature (eg. Bond et al. 1980), but David and Jarvis (1983) note that in a study at Ladysmith in the southern Cape it was found on fairly exposed slopes with sparse cover, although it showed a strong preference for more densely covered, south-facing slopes, which had more grass, as well as closely packed Protea bushes. Aethomys Habitat: Bond et al. (1980) noted that this species was never found in areas with more than 75% total shrub cover in the Swartberg, and that it namaquensis favors rocky and/or more open habitats. Steatomys krebsii Breeding: Breeds in wet summer months, as is S. krebsii (Stuart and Stuart 2001). Habitat: Steatomys species are adapted to unpredictable environments as they are heterothermic, that is they have the ability to maintain a constant body temperature, but can also allow body temperature to fluctuate, and can regularly enter energy-saving periods of torpor (Perrin and Richardson, 2005). Diet appears to be eclectic as Stuart and Stuart (2001) noted that this species primarily eats seeds, but may eat bulbs and insects. Rautenbach and Nel (1980) noted that in the Cedarberg stomach contents consisted solely of insects. Otomys irroratus Breeding: Otomys irroratus is a k-breeder, produces several small litters throughout the year, and utilizes poor quality, but abundant foods to maintain a stable population (Perrin, 1980). Habitat: Otomys irroratus is associated with moist, marshy habitats, but may also be found in drier habitats (Rowe-Rowe and Meester, 1982), such as grassy hillsides (Stuart and Stuart, 2001). Soricidae Suncus varilla Breeding: Reproduction of Suncus is uncertain (Stuart and Stuart, 2001). Habitat: S. varilla occupies a range of habitats, and is associated with the termite mounds this species inhabits (Stuart and Stuart, 2001) Suncus infinitesimus Breeding: Reproduction of Suncus is uncertain (Stuart and Stuart, 2001). Habitat: S. infinitesimus occupies a range of habitats, and is associated with the termite mounds this species inhabits (Stuart and Stuart, 2001) Myosorex varius Breeding: M. varius is a seasonal breeder in the warm, wet, summer months (Stuart and Stuart, 2001). Habitat: Rowe-Rowe and Meester (1982) noted that M. varius is associated with a number of environments, including grasslands and fynbos. Crocidura flavescens Breeding: C. flavescens is a seasonal breeder in the warm, wet, summer months (Stuart and Stuart, 2001). Habitat: Skinner and Smithers, (1990) cite Meester (1963) who noted that this species is confined to areas with a mean annual rainfall of between 500 and 750 mm This species is reportedly found in association with moist habitats and dense vegetation.

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Glossary C: Carnivore FG: Frugivore Habitat types H: Herbivore O: Omnivore BL: Bushland FS: Forest substrate D: Desert F: Forest GL: Grassland Strategraphic aggregates SL: Shrubland WL: Woodland SS: Surface Scrapings:2005 MSS: Mixed Surface Sand Adaptations BC1: Brown Clay 1 OYCS: Olive Yellow Cave Sand BYDS: Brown Yellow Dune Sand A: Arboreal BYSS: Brown Yellow Surface Sand SU: Subterranean BYCS: Brown Yellow Cave Sand T: Terrestrial