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Proposal to export micromammal teeth, mandibles and maxillae for the purpose of stable oxygen and carbon isotope analyses





Case Type: 


EXECUTIVE SUMMARY The proposal is to expand on the conventional methods used for analyses of micromammal assemblages from archaeological sites by applying analyses of stable carbon and oxygen isotopes of the tooth enamel from selected micromammal species from the Howiesons Poort sequences at Klipdrift Shelter (KDS) (Henshilwood et al. 2014) and Klasies River main site (KRM) (Singer & Wymer 1982, Wurz 2002, Villa et al. 2010). The results will be compared to an extensive modern baseline derived from 10-20 owl pellet locations across the Western and Eastern Cape. Our study aims to provide a detailed local palaeoenvironmental record of C3 and C4 vegetation composition, humid/arid periods and potential rainfall seasonality shifts and establish a modern reference database to assess the dominant ecological drivers of isotopic change for the selected extant micromammal species. The micromammal species chosen for stable isotope analyses are based on several suitability measures: dietary and water consuming preferences, frequency of occurrence in the two archaeological assemblages, and comparability to similar studies from South Africa as per Leichliter et al. (2016, 2017). The sample for stable isotope analyses consist of in situ and isolated molars and incisors from eight species of micromammals that meet these criteria. Samples that are not subject to destructive analyses, will be returned to the KDS and KRM collections. INTRODUCTION & OBJECTIVES This proposal aims to maximize the information that micromammalian fauna can provide for reconstructing palaeoenvironments, using both modern and archaeological materials. Micromammals serve as suitable proxy data for fine-scale palaeoenvironmental reconstruction due to their limited territorial ranges, precise ecological requirements and short lifespans (Korpimäki et al. 2004; Heisler et al. 2016; Reed et al. 2019). Analyses of modern micromammal samples have demonstrated close correlation between relative abundance of species and composition of vegetation substrate near sample sites (Andrews, 1990; Avery et al., 2005, Reed et al. 2017). Local alterations in vegetation substrate and climatic conditions are reflected in presence/absence and/or variations in proportions of micromammal species in an archaeological assemblage (Nel 2013, Nel and Henshilwood 2016, Nel et al. 2018). Conventional methods of identification of micromammal species by dental morphology and subsequent palaeoenvironmental analyses based on extant species habitat requirements have the potential to be refined in order to increase the information output from archaeological assemblages. Our methodological approach focuses on refining micromammal palaeoclimatic datasets from the southern Cape region by analyses of the stable carbon and oxygen isotopes values of micromammal teeth from both modern and archaeological assemblages. These analyses will provide a detailed record of vegetation composition, evaporation/aridity, and potential rainfall shifts during the Howiesons Poort sequences at KDS and KRM (Fig. 1). At the same time our analyses will add essential information to improve knowledge of how the current environment is reflected (Leichliter et al. 2016, 2017) by inclusion of a modern baseline, represented by eight micromammal species from ten (potentially 20 – see explanation below) locations spanning the rainfall gradient (winter to summer) and variable vegetation environments in the Western and Eastern Cape (Avery et al. 2005, Avery et al. unpublished manuscript, Nel unpublished data) (Fig. 1). Our modern reference database will enable us to establish a sound methodology and baseline for interpretation of the archaeological samples, while also initiating a reference database for future stable isotopes analyses in the region. BACKGROUND & RATIONALE The MSA archaeological record in southern Africa displays variations in technological and cultural mechanisms and shifts in ecological niches, particularly between 100 and 50 ka (d’Errico et al. 2017). An essential question is the role of climatic and environmental changes as potential catalysts for human adaptability, changing mobility patterns and increased cultural complexity. Did global and regional climatic fluctuations affect local vegetation and precipitation patterns, and could potential local variations have influenced human settlement and subsistence strategies? To what extent were palaeoecological changes a factor in the occurrence and disappearance of techno-cultural complexes in the area? Reconstruction of environmental conditions is crucial for our understanding of resource procurement strategies, technological and cultural innovations, and site choices made by these early humans. The southern Cape boasts a rich record of archaeological sites containing material from prominent techno-cultural periods such as the Still Bay (SB) (76-72 ka) and Howiesons Poort (HP) (66-59 ka). There are conflicting hypotheses regarding the origin and disappearance of these periods. Some have suggested that they are indicative of human buffering mechanisms due to climatic and environmental instability associated with the Marine Isotope Stage (MIS) 5/4 and 4/3 transitions (see Bar-Matthews et al. 2010, Ziegler et al. 2013, d’Errico et al. 2017, Marean et al. 2020). Others have proposed that the southern Cape coast was hospitable refugia during the SB and HP and that the global and regional climatic and environmental record is not synced with human behavioural changes (Jacobs et al. 2008). The opposing hypotheses warrants the need for context-specific palaeoenvironmental reconstruction which can be directly associated with anthropogenic derived material and firmly established chronologies, in particular concerning the impact of observed global and regional climatic fluctuations on a local scale (Roberts et al. 2016, d’Errico et al. 2017). The spatial and chronological dissonance between regional climatic records (Ziegler et al. 2013, d’Errico et al. 2017) and archaeological sites in the southern Cape is best addressed by creating local palaeoenvironmental datasets. Stable carbon and oxygen isotope analyses of micromammal teeth from archaeological samples have the potential to form the basis of such ‘on-site’ palaeoclimatic and palaeoenvironmental datasets. The foundation for studies that utilize faunal isotopes to infer paleoenvironment is based on the notion that the isotopic character of an animal’s consumed food is reflected in that animal’s isotopic profile. Thus, the environment from which the predator (e.g. a predatory bird or human population) selects prey items is reflected in the isotopic composition of the fossil assemblages (Lee-Thorp and Talma, 2000; Lee-Thorp, 2002; Ecker et al., 2018). Isotopic analysis of tooth enamel from large mammals recovered in archaeological contexts has been widely employed to reconstruct past vegetation composition and climate in Africa (e.g. Koch et al. 1994; Cerling et al. 1997; Sponheimer & Lee-Thorp, 2003; Lee-Thorp et al., 2007). A similar methodological approach is applicable to micromammal species (Hynek et al. 2012; Jeffrey et al., 2016; Roberts et al., 2017), yet this remains in the early stages of exploration in a South African context (Leichliter et al. 2016; 2017, Williams et al. 2020). Stable carbon isotope analyses of small mammals are likely to reflect fluctuations in C3 and C4 vegetation which, in the southern Cape, are primarily indicative of rainfall seasonality shifts and humid/arid periods. Stable oxygen isotopes could provide information relating to precipitation source, temperature, or evaporation/aridity (Leichliter et al. 2016, 2017). However, there is a need for control studies based on modern samples in order to better understand the number of factors that influence oxygen isotopic composition in particular. These factors are leaf water and plant carbohydrates (influenced by evaporative enrichment under arid conditions), small-scale variability in the oxygen isotopic composition of different plant parts, and animal reliance on either environmental water sources or plant water (Kohn et al., 1996; Sponheimer & Lee-Thorp, 1999; Williams et al. 2020). Application of the methodology on the southern Cape has been limited due to a lack of knowledge of micromammal isotopic ecology and difficulties with sampling of the small teeth. Thus, interpretation of the archaeological sample firstly require a modern reference database in order to establish the dominant ecological drivers of isotopic change for extant micromammals. The latter is particularly missing for the region but holds interpretative promise due to the regions climatic gradients. Furthermore, it is essential to establish which species that are likely to provide solid isotopic signals, and thus control analyses of modern samples from known environments are required. We also aim to establish an improved understanding of stable carbon and oxygen isotopes for various feeder categories such as insectivores, herbivores and omnivores (as per Leichliter et al. 2016, 2017; Williams et al. 2020). Facilities for analyses Stable carbon and oxygen isotope analysis of micromammal tooth enamel will be undertaken within the Stable Isotope Laboratory, Department of Archaeology, Max Planck Institute for the Science of Human History, Jena, Germany by Dr. Nel under the guidance of Dr. Patrick Roberts. This is one of the few state-of-the-art dedicated isotope laboratories for archaeological research globally, and Dr. Roberts is a world-leading expert in isotopic and archaeological research in South Africa, with a particular focus on the development of ‘on-site’ palaeoenvironmental records. Dr. Roberts has always followed through on his SAHRA reports in a timely and effective manner. Samples will be analysed using a Thermo V Isotope Ratio Mass Spectrometer connected to a Gasbench II device. The strong basis of theoretical knowledge in relation to South African archaeological science and palaeoecology makes the Stable Isotope Laboratory in Jena a logical location for sample analysis as part of this project. MATERIAL The materials for the analyses are micromammal molars and incisors, in situ or isolated, extracted from coarse (3 mm) and fine fraction (1.5 mm) sieved material, from KDS and KRM (see Appendix 1 for list of samples). The modern micromammal molars, incisors, mandibles and maxillae are from barn owl pellet assemblages from 10 barn owl locations in the Western and Eastern Cape (Fig. 1 and Appendix 2). A further 10 locations (Fig. 1) will be sampled once access to South Africa and Iziko South African Museum is possible after the corona lockdown and travel ban. These references will create a baseline for our archaeological samples and be vital in developing an interpretive framework for carbon and oxygen isotope analyses of small mammals in South Africa. The molars and incisors have been identified to species by morphological traits following standard methodology (Andrews 1990, Nel 2013, Nel and Henshilwood 2016). Prior to the analyses, all samples will be photo-documented. The analyses will commence in January 2021 and the material will be returned to South Africa by January 2022 (dates to be adjusted accordingly, with the permit application process), as per agreement with Iziko South African Museum, Cape Town (see attachment). In terms of the National Heritage Resources Act (Act 25 of 1999), section 35(4), a temporary export permit must be obtained prior to analysis. METHOD Stable carbon isotope (δ¹³C) of small mammal tooth enamel will reflect that of the food consumed and, indirectly, vegetation patterning and habitat (Jeffrey et al., 2015). In the context of the southern Cape, vegetation δ¹³C is dictated by precipitation-controlled ecological variability. Plants in the winter rainfall zone are primarily C3 plants, with relatively low ¹³C values (globally -24 to -32‰), while vegetation in the year-round rainfall zone includes some C4 plants with higher (-10 and -14‰) ¹³C values (Smith and Epstein, 1971; Vogel et al., 1978). Although CAM plants, with values that span these groups, can complicate interpretation, while CAM plants in the C3-dominated winter rainfall zone of the southern Cape coast of South Africa today have ‘C3’ ¹³C, those in the year-round rainfall region have ¹³C spanning C3 and C4 values (Rundel et al., 1999). These differences are tracked into the tooth enamel 13C of consumers, with herbivorous small mammals being considered as often tracking the local availability of certain vegetation types and omnivorous small mammals capturing wider regional vegetation availability (Leichliter et al., 2017). Stable oxygen isotope (δ¹⁸O) analysis of small mammal tooth enamel reflects the blood bicarbonate which is primarily influenced by body water (Passey et al., 2005). This, in turn, is derived from imbibed water and water from food (Luz et al., 1984). Environmental water can be influenced by precipitation source, temperature, geographical location, and evaporation potential (Buchmann et al., 1997). Different mammals will reflect environmental variation to different extents. Those animals that obtain the majority of their water from plants will strongly reflect those environmental factors that influence plant transpiration and plant δ¹⁸O, namely aridity and evaporative potential (Kohn et al., 1996; Levin et al., 2006; Carter and Bradbury, 2016). Meanwhile, those that obtain most of their water from groundwater will primarily be influenced by the δ¹⁸O of precipitation. Depending on the small mammals sampled, they will therefore be expected to document variations in precipitation δ¹⁸O linked to the seasonality and origins of rainfall across the southern Cape or seasonal aridity (Roberts et al., 2016). In the context of this study, molars and incisors will be used for stable isotopic analysis. In general, for rodents, molars form in vitro, while incisors grow continuously during the rodent’s lifespan. We will thus use molars for the analyses, while incisors are included for the purpose of understanding intra-specimen variations of stable carbon and oxygen isotopes between incisors and molars on both the modern and archaeological sample. We will create a baseline of these variations as the modern material have incisors and molars in situ, while, due to the fragmentary nature of the archaeological assemblage, this is only possible for some specimens from selected species from the KDS and KRM samples (see list of samples). The external edge of the tooth will be cleaned using air-abrasion to remove any adhering external material. The roots and dentine of the incisor and molar teeth will be removed when possible. Due to their small sizes, the remaining enamel will then be crushed to a fine powder using an agate mortar and pestle following Gehler et al. (2012) and Jeffrey et al. (2015). All enamel powder will be pre-treated using a published technique consisting of washes in 1.5% sodium hypochlorite for 60 minutes, followed by three rinses in purified H2O and centrifuging, before 0.1M acetic acid will be added for 10 minutes, followed by another three rinses in purified H2O (Lee-Thorp et al., 2012; Sponheimer et al., 2005). Following reaction with 100% Phosphoric Acid, gases evolved from the samples will be measured for their δ¹³C and δ¹⁸O values using a Thermo Gas Bench II connected to a Thermo Delta V Advantage Mass Spectrometer at the Department of Archaeology, Max Planck Institute for the Science of Human History. Values will be compared and corrected using linear regressions based on the measured and expected values of international and in-house carbonate standards. An in-house enamel standard will be used as a test for overall variation caused by enamel pre-treatment and measurement (see Roberts et al., 2018; Ventresca Miller et al., 2018). (POTENTIAL) Significance Our study will provide a detailed local vegetation and evaporation record for the prominent Howiesons Poort period at KDS and KRM. The results will add vital information regarding the environmental factors which influenced human adaptability, changing mobility patterns and increased cultural complexity during this period. We will contribute with important information that sheds light on the observed global and regional climatic fluctuations on a local scale, enabling a greater understanding of environmental responses to climatic fluctuations in the southern Cape region. We are contributing to the future development of science in South Africa by creating a modern baseline of stable carbon and oxygen values from eight micromammal species. This contribution will not only create an interpretative framework for the archaeological samples in our study but will also be beneficial for future analyses of stable isotopes from micromammal archaeological assemblages in the region undertaken by local scholars. At the same time, we will improve our understanding of the reflection of the environment in small mammal herbivores, omnivores and insectivores in a South African context. The results of our analyses will be presented in at least two papers in peer-reviewed journals and at international conferences such as SAFA (Society of Africanist Archaeologists) 2021, PANAF (Pan African Archaeological Association of Prehistory and Related Studies) 2022 and INQUA (the International Union for Quaternary Research) 2023.


Monday, November 2, 2020 - 10:56





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