Palaeoecological Significance of Palms at Olduvai Gorge, Tanzania, Based on Phytolith Remains
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ARTICLE IN PRESS Quaternary International 193 (2009) 41–48 Palaeoecological significance of palms at Olduvai Gorge, Tanzania, based on phytolith remains Rosa Maria Alberta,Ã, Marion K. Bamfordb, Dan Cabanesc aCatalan Institution for Research and Advanced Studies (ICREA)/Department of Prehistory, Ancient History and Archaeology, University of Barcelona, c/ Montalegre, 6-8, E-08001 Barcelona, Spain bBernard Price Institute for Palaeontological Research, School of Geosciences, University of the Witwatersrand, P Bag 3, WITS 2050, South Africa. cInstitut Catala` de Paleoecologia Humana i Evolucio´ Social, A`rea de Prehisto`ria, Universitat Rovira i Virgili, Programa de Beques Fundacio´n Atapuerca, Plac-a Imperial Tarraco, 1, E-43005 Tarragona, Spain Available online 27 June 2007 Abstract Palms are among the most abundant, diverse and economically important families of plants in tropical and subtropical regions of the world; their number and diversity make them an important part of the ecosystem. Phytoliths are abundantly produced in palms, mainly the spheroid echinate type and, although their number decreases notably after their deposition in soils, they remain stable for long periods of time. Palm phytoliths were recovered from sediments at HWKEE, on the eastern palaeolake margin of the Olduvai basin during Lowermost Bed II (between Tuffs IF and IIA, 1.795–1.71 Ma). Their abundance in trench 107 (the only freshwater source recognised to date) occurs in three stratigraphically consecutive layers, the first one located immediately above the incision layer (40 cm above Tuff IF). Above these samples there are no palm phytoliths in the fossil record. The identification of the type of palms represented will be useful for determining the landscape distribution and nature of hominin activities along the lake margin and alluvial fan of the central Olduvai Basin. Here we use morphometric analysis of spheroid echinate phytoliths from fossil and modern palms (Phoenix reclinata and Hyphaene petersiana) and their corresponding soils to determine if the differences are significant and can be confidently applied to the fossil samples. Fossil and modern soil samples were found to be very similar to each other but the spheroid echinates from the fossil soils were bigger than the modern soil samples. Either the palms from the samples were different or there is a strong bias towards preserving phytoliths from the smaller end of the range. If this is the case then the implication would be that the fossil samples are the leaves and inflorescences of Phoenix reclinata. A wider variety of modern palms needs to be analysed. r 2007 Elsevier Ltd and INQUA. All rights reserved. 1. Introduction broad spatial scales, throughout exposed deposits of the Plio-Pleistocene Olduvai Basin with detailed palaeoenvi- Olduvai Gorge in northern Tanzania is a well-known site ronmental analysis focusing on terrain, hydrology and with a rich vertebrate fossil record including hominins and vegetation (Peters and Blumenschine, 1995; Blumenschine stone artifacts from Plio-Pleistocene lacustrine and tuffac- and Peters, 1998; Blumenschine et al., 2003). Models of eous deposits (Leakey, 1971; Hay, 1976; Potts, 1988; palaeolandscape ecology and hominin land use highlight Blumenschine et al., 2003). The site offers a good the central role of the landscape distribution and cover geographic and stratigraphic framework for the occupation abundance of herbaceous and woody vegetation in by hominins of the basin and palaeolake margins (Hay, determining the landscape distribution and nature of 1976, 1996) and this is being further refined by the hominin activities throughout the lake margin and alluvial multidisciplinary efforts of the OLAPP team members fan landscapes of the central Olduvai Basin. (Olduvai Landscape Palaeoanthropological Project). This A variety of methods have been employed to interpret work combines sampling of hominin activity traces over the palaeoenvironments of early hominin sites since it is generally accepted that the vegetation and climate ÃCorresponding author. Tel.: +34 934037525; fax: +34 934037541. affect the distribution and behaviour of the hominins and E-mail address: [email protected] (R.M. Albert). other faunas. The sedimentological record provides 1040-6182/$ - see front matter r 2007 Elsevier Ltd and INQUA. All rights reserved. doi:10.1016/j.quaint.2007.06.008 ARTICLE IN PRESS 42 R.M. Albert et al. / Quaternary International 193 (2009) 41–48 broad palaeoecological signatures such as fluvial (earthy phytoliths occurred at HWKEE, the freshwater source for claystone), lacustrine (waxy claystone), volcanic (tuffs) or the palaeolake (Hay, 1976) and only during a short time dune settings. Isotope analyses of carbonates can provide a interval. temperature and moisture regime. Faunal composition and Since palms are among the most abundant, diverse and comparison with modern faunas is another common economically important families of plants in the tropical method employed. Pollen analyses show what plants were and subtropical regions of the world, their number and present in the area and show a general drying out trend diversity make them an important part of the ecosystem. from Uppermost Bed I through Bed II based on a They are found in a variety of habitats, from tropical combination of several sites. Wooded grassland was forests to mangrove swamps, from upland woodlands to present during Bed I times and by uppermost Bed I it oases in deserts (Henderson, 2002). The question posed is, was reduced to semidesert scrub. Above Tuff If (Lower- what climatic signature do the Olduvai palm phytoliths most Bed II) wooded grassland occurred and developed give? into Acacia woodland (Bonnefille, 1984). Palm pollen is Nine genera of palms occur naturally in East Africa and extremely rare with only one grain of Phoenix type pollen another three genera have become naturalised (Dransfield, from the site FLKN in Bed I deposits between Tuff ID and 1986). Two introduced taxa are restricted to the coast IF (Bonnefille, 1984). (Areca catechu and Cocos nucifera). Two indigenous taxa More recently detailed fossil macroplants and phytolith occur in gallery or swamp forest (Elaeis guineensis and studies have been performed in the area for Lowermost Raphia farinifera). The others are more widespread. Bed II (Albert et al., 2006; Bamford et al., 2006). Their Borassus aethiopium occurs in grasslands with high water studies were focused on interpreting the vegetation tables. Hyphaene compressa (doum palm) grows along the structure and composition of a portion of the eastern lake coast and along seasonal water courses. Hyphaene coriacea margin including localities FLK, VEK, HWKE, HWKEE, grows on sand dunes, creeks and rarely inland, whereas and MCK, over a distance of about 2 km and during Hyphaene petersiana is usually confined to alkaline soils lowermost Bed II times, about 80 kyr. They interpreted the where there is a relatively high water table. Phoenix vegetation during this time to be composed of a variety of reclinata (wild date palm) occurs along water courses, in grasses, sedges and dicotyledonous plants in different lowlands and also higher up rocky hills especially in proportions and in different areas. In one specific place protected steep valleys (Dransfield, 1986; Beentje, 1994). (HWKEE, T-107), phytoliths have shown palm trees to be Other naturally occurring taxa are the large single stemmed dominant (Fig. 1) and present at VEK. Borassus and climbing Eremospatha, Laccosperma, Calamus Macroplant remains of sedges, grasses and woody dicots and Chrysalidocarpus, which have a thin trunk and are are widespread (Hay, 1976; Bamford et al., 2006) but palm outliers of a Madagascan genus (Dransfield, 1986). macroplants have not been reported to date in any sites Phytoliths are produced abundantly in palm trees in Olduvai Gorge. It is interesting to note that palm (Bamford et al., 2006), mainly the spheroid echinate type Fig. 1. Map of the Olduvai Gorge showing the location of HWKEE, where T-107 has been studied. ARTICLE IN PRESS R.M. Albert et al. / Quaternary International 193 (2009) 41–48 43 (globular according to the ICPN, Madella et al., 2005) These palms characteristically have sharp spines on the and, although their number decreases notably after the basal portion of the fronds. deposition of plant parts in the soils, once deposited, Hyphaene petersiana, the ilala palm, grows in hot and they remain stable for long periods of time (Albert et al., dry environments, but will probably tolerate high rainfall if 2006). the soils are well drained. The large, round fruit, with a From previous studies on modern palm material hard white nut (hence the common name of vegetable (Bamford et al., 2006) we know that there are some ivory) is a favorite food of elephants. Based on years of differences between the phytoliths from different palm field work, Ellery and Ellery (1997) found that Phoenix species and also from different parts of the palm plant reclinata occurs close to freshwater along the edges of (leaves, flowers, dry rachis). Here we look more closely at islands and river banks in the Okavango Delta, Botswana. the phytoliths, using morphometric analyses, to determine It may form dense clumps and be associated with if the differences are significant and if they can be applied termitaria. Hyphaene petersiana frequently occurs towards confidently to the fossil samples from the site HWKEE on the middle of the islands (Ellery et al., 1997; personal the eastern palaeolake margin of the Olduvai basin during observation) and is associated with a high water table but the short time interval of Lowermost Bed II (between Tuffs its occurrence indicates the presence of saline groundwater IF and IIA, 1.795–1.71 Ma; (Hay, 1976; Blumenschine (Ellery and Ellery 1997). et al., 2003). The fresh palm material was collected from single trees and separated into leaves, inflorescence, rachis roots and 2. Materials and methods seeds and processed separately. Since palms produce huge amounts of palm phytoliths (Bamford et al., 2006), only Three types of samples are considered here, namely fossil one sample from each plant part was analysed.