Sir Mark Mitchell Foundation, the Potter Foundation, the Sunshine Foundation, the. Myer Foundation and the Utah Foundation. Thanks are extended towards ...
SOME PROSPECTS FOR ARCHAEOLOGICAL PALAEOBOTANY IN AUSTRALIA: AN EXAMPLE FROM SOUTH AUSTRALIA W.E. Boyd and G.L. Pretty
SUMMARY The scope of archaeological palaeobotany is being continually expanded, and the techniques employed are constantly being refined. As much of this development is taking place abroad, primarily in Europe and North America, the substantive data have limited applications to Australian archaeological contexts. However, some fields look very promising, and the discussion presented here, based upon preliminary palaeobotanical investigations at the archaeological site at Roonka, South Australia, serves to illustrate the potential of the study of plant leaf surfaces and other fossil vegetative plant parts to archaeological investigation.
INTRODUCTION In recent years, several major advances have been made in the field of palaeobotany, especially in relation to its applications to archaeology. These focus on the development and refinement of techniques, and in the expansion of interest into the study of parts of the plants not previously examined. Many of these advances have been developed in Europe and North America, but the resulting expertise may equally be applied in a region such as Australia. Research outside Australia has, for example, shown a diagnostic value of cellular details and micro-sculpturing of seed and stem surfaces as an aid towards afuller understanding of cropped plants (e.g. Dickson 1987; Korber-Grohneand Piening 1980; Stemler and Falk 1981; and Yarnell 1976). Within the Australian context the focus of archaeobotanic studies has centred more upon the wide range of naturally occurring plants which are and were gathered and used for decorative, ritual and dietary purposes. In addition to this study of past resources utilisation of individual plants, archaeobotanical studies also contribute, along with other palaeoecological and palaeoenvironmentalstudies, to an understanding of the environment or environments within which prehistorical communities operated. The palaeoenvironmental approach may offer a framework for some form of cultural
Boyd, Pretty 41
developmental model in which apparent palaeocultural activity is related to palaeoenvironmental conditions and changes in these conditions (e.g. Pretty 1986). This paper discusses some of the newer techniques being developed by, and becoming available to environmental archaeologists. The discussion derives from the preliminary results of a pilot study (Harvey and Pretty 1984) undertaken on samples taken from Roonka. The results of this pilot study are, in themselves, of relatively little importance in that they tell little in the way of a palaeoenvironmental story; such information will be provided during the course of the full palaeobotanical investigation. The importance of the pilot study is that its results confirm that at an open, weathered site in the semi-arid zone, useful paleobotanical remains may be recovered. However, the results also, more importantly, indicate the direction for further productive investigation, emphasising the types of plant fossils that will contribute to an understanding of the paleoenvironment of this site. Since these fossil types, in particular, may be encountered at other archaeological sites, it is considered that an introductory discussion about them should be of interest to archaeologists. The purpose of this paper, therefore, is to introduce some aspects of paleobotanical research which may be usefully employed at archaeological sites. The paper is not a detailed treatise on archaeological techniques, nor is it a progress report on a particular site. However, it does use one archaeological site as a metaphorical coathanger upon which to display the cloth of palaeobotanical research. Perhapsthis display will entice onlookers to come in and feel the quality of the fabric!
SITE LOCATION AND MATERIALS The archaeological locality selected for exploratory study is at Roonka on the River Murray near Blanchetown (MO19's 13g036'E). Excavations by G.L.P at intervals between 1962 and 1977 explored five sites extending across a complete transect of the gorge through which the River Murray flows. Quantities of palaeobotanical material from contexts dated between 20,000 and 200 years BP (Pretty 1977, 1986; Pretty and Ward forthcoming) were recovered. All the sites principally comprise occupation residues, but three also contain tombs (216 in total), dating from throughout the full depth of the time sequence. The stratigraphic character of two sites, being dunes, is complex and difficult to interpret because of evidence for reworking in the past and of recent severe erosion. However, using thermoluminescent, palaeomagnetic, uranium uptake and radiocarbon analysis, synchronies have been established and the basis for a firm chronology created. The local sequence is presently divided into three periods, primarily based on shifts in the form and association of body postures within the tombs; the radiometric dating indicatesthree apparentlydiscrete periods, although occupation was probably more continuous, possibly with a hiatus during the last glacial maximum (c. 16,000 to 10,000 BP): Roonka I, c. 20,000 to 16,000 BP; Roonka II, c. 8000 to 7000 BP; Roonka Ill, c. 6000 to 200 BP A quarter of the tombs contain deliberate and sometimes elaborate settings of ornaments, domestic articles or offerings, a proportion of which are botanic in origin. These comprise fruits, bark and other plant tissue fragments. The palaeobotanical study material consists of 110 organic matter samples which contain plant tissue (leaf, bark and wood) with some seed and fruits, 370 samples of carbon, much of which consists of carbonised wood, and 350 soil samples collected in part to document contexts thought to contain pollen. Initial examination of specimens of tissue material show them to contain twigs and leaf tissue which, when subjected to suitable rehydration and mounting should be taxonomically identifiable.
42 Archaeological Palaeobotany in Australia
Figure 1.
Examples of features on the surfaces of plant leaves and other plant parts, showing some of the diversity occurring within such features. Note that the plants used as examples here are not necessarily Australian species. Also,note that the dimensions given are in microns (U), where 1p = 10* m. A-D (top left): the outer (A, B) and inner (C, D) epidermis of C.pedla pericarp in cross section and plan views. E: epidermis of a Sarnbucw stem, showing cuticle forming ridges inserted between the cells. F: epidermis of a Hdliborus leaf, showing cuticle with a wavy outer surface. G (top right): epidermis of grass (sugar cane, Saccharurn) showing a typical arrangement of guard and subsidiary cells around the stoma. H (top right): epidermis of a Saccharurn stem showing cork and silica cells. CL (bottom left): plan views of the epidermis of species in several genera, illustrating some
rubsidialry cell
gwrd
cork cell
WCJ, silica cell
of the patterns formed around the stoma by guard cells and other surrounding cells. M Y (bottom right): examples of trichomes. M: a simple hair from a Cktw leaf. N: a uniseriate hair from a Saintpaulia leaf. 0, P: tufted hairs from a leaf of cotton (Oouypium). Q: a stellate hair from a leaf of Sida. R: a dendroid hair from a leaf of L.v.ndul.. S: a short multicellular hair from a leaf of the potato (Solmum). T. U: a peltate Scale from an olive leaf (m). V: a bicellular hair from a Pdargonkrmstem. W-Y: epidermal hairs on a cotton ( m y p i u r n ) seed, showing young 0and mature M stage hairs. Z: a water vesicle of MeMlmbryanthemum. (Adapted from Esau 1960, Figs. 7.2, 7.3, 7.5 and 7.8).
44
Archaeological Palaeobotany in Australia
This resource of palaeobotanical data, particularly if supplemented by examination of terrestrial swamp sediments nearby, offers two lines of inquiry. First, the soil samples, carbonised wood and organic samples should provide the basis for an outline of the immediate region's palaeobotanic succession since the Late Pleistocene, and thus a palaeoenvironmental setting for the cultural development of this site. Secondly, the samples containing tissue, bark and fruits, particularly those originating from the tomb sediments, should also create opportunities to explore some of the resource utilisation at this site. A particular problem in this context is whether a narrow range of plants was deliberately selected for installation in the tombs, or whether a wider range of plants, perhaps collected at random, was deposited in the tombs. Foliage appears to have been widely used for the grave linings along the Murray and Darling Rivers (Meehan 1971) and in at least one instance from the Upper Darling (Parker 1905:85-87)there is reference to an unnamed species of tree being reserved for placement on the dead in graves and decoration of the mourners.
DISCUSSION The excavation of many European and North American archaeological sites provides abundant and well-preserved macroscopic and microscopic plant fossils. The former group include seeds, fruit, wood fragments, leaf fragments and so on, for which specialist identification texts and keys are often available (e.g. Beijerinck 1947; Korber-Grohne 1964; Tomlinson 1985a, b). The latter group includes spores and pollen for which a diverse methodology of analysis exists in relation to arkhaeological problems (e.g. Bryant and Holloway 1983; Dimbleby 1985; Greig 1982) and based upon the wider study of Quaternary paleoecology (e.g. Berglund 1986; Moore and Webb 1978). In the Australian context, however, the options for palaeobotanical study are often limited for several reasons, such as availability of suitable fossil preservation conditions and the difficulty of applying analytical techniques which have been developed in the more temperate parts of the world. Although Martin (1973) and Clark (1985), for example, have demonstrated that the preservation of fossil pollen and spores (Martin) and macroscopic plant remains (Clark) may occur by desiccation rather than by waterlogging, preservation of plant remains in dry andfor biologically active soils is generally poor. Given the rarity of waterlogged archaeological sites in Australia, the prospects for archaeological pollen analysis, one of the most commonly employed paleobotanical techniques, are moderately poor. By way of parenthesis, it should be noted that Wyrie Swamp in the lower southeast of South Australia is an unusual Australian site in which archaeological artefacts, including wooden objects, and fossil pollen are preserved together (Dodson 1977; Luebbers 1975). However most of the fossil pollen data which is drawn upon for the archaeological study in Australia is usually derived from non-archaeological sites (e.g. Clark and Lampert 1981; Clark 1983; Hope et al. 1985; Kershaw 1986; Singh et al. 1981). The archaeological site at Roonka illustrates this. Although many potentially pollenbearing samples were collected, they are largely dry soil samples in which pollen preservation is poor. This problem was further increased by the samples having been allowed to dry out during storage. However, a preliminary examination of the samples did indicate that some pollen was present (J.R. Dodson pers. comm.), although it is debatable whether the pollen is entirely fossil or is partly modern contamination. Possibly more satisfactory evidence will be derived from swamp sediments from near the archaeologicalsites, but as dsewhere, this will sever the immediate contextual links betweenthe archaeologicaldata and the pollen analytical data. Giventhese difficulties, a further possibility may be applied with good effect at Rwnka. In situations where
pollen preservationis generally very poor, microscopic siliceous plant cell thickenings, known as phytoliths or plant opals, may be excellently preserved (Palmer 1976; Piperno l985a, b). Being siliceous, these plant fossils will survive moderately unaltered in most depositional environments. Piperno (1985a, b) has demonstrated their taxonomic value in a study of archaeological sediments in Panama, and indicates that for many plant families represented in Australia, phytoliths are taxonominally significant. Phytoloth analysis has already been applied in Australian archaeological contexts, with, for example, Fujiwara et al. (1985) identifying a limited range of species at archaeological sites in Kakadu (Northern Territory). Phytolith study in Australia may be worth pursuing, and its use in Australian archaeology may prove to be highly valuable, although several methodological problems, in particular the identification of the diverse phytolithtypes which occur even within one species, still require resolution Bowdery pers. comm.; Wilson 1985. In contrast to the inconclusive results of the preliminary study of plant microfossils in the palaeobotanical investigations at Roonka, Harvey's initial examination of the Roonka sediments indicates that the most promising paleobotanical work would be on the macroscopic fossils, in particular the leaf and stem, or axis, fragments occurring in many of the samples. Although analyses of such fossils, especially leaf fragments, are widely undertaken in pre-Quaternarypalaeobotanical studies, such fossils are less commonly examined from archaeological contexts, probably largely due to the presence of other suitable and often more readily analysable botanical fossils such as pollen, fruit and seeds. However, given the types of restrictions discussed above at Australian archaeological sites, it is pertinent to look at other types of plant fossils, and to discuss their applicability to archaeological paleobotanical study. Dealing with leaf fragments first, it is encouraging to note that plant leaf surfaces contain several diagnostic characters which may be used to identrfy the plant from which the fragment originally came (Fig. 1). On the outer surface of leaves there is a waxy layer, known as cuticle, which is more or less impervious to water, and the characteristicsof this leaf cuticle are commonly studied in the taxonomic examination of both fossil and modern groups (Kovack and Dilcher 1984; Wilkinson 1983). Early light microscopy studies of leaf cuticles concentrated upon gross features of the leaf epidermis (the outermost layer of cells in the leaf) as transcribed onto the cuticle (e.g. Stace 1965, 1966) and, understandably, considerable attention was given to the structure of the most conspicuous epidermal feature, a pore known as the stoma (plural: stomata, Fig. 1). The stomata1 structure and the arrangement of the surrounding subsidiary cells has been well documented for many plant families (e.g. Payne 1979). The form of other leaf structures, such as surface hairs and secretory devices (known collectively as trichomes, features which include a range of epidermis outgrowths variable in shape, size and function), may also have diagnostic value (Fig. 1). Surveys of plant hair types in plant families and genera containing Australian species have been presented, for example, for the family Goodeniaceae (Carolin 1971) and the genus Acacia (Pettigrew and Watson 1973), but secretory devices appear to have received less attention. The leaves of species of Myrtaceae, Rutaceae and related families contain numerous spherical oil-bearing bodies, the more external ones being linked with the leaf surface by several epidermal capping cells which may be distinguishedfrom other epidermal cells on the basis of their morphology and cell wall sculpturing. A recent survey of these families has shown that the epidermal and cuticular patterns are distinctive for groups of species and, in some cases, for individual species (Harvey and Pretty 1984).
46 Archaeological Palaeobotany in Australia
varcular
Figure 2
bundles
Examples of the internal anatomical structure of plant stems, showing some of the range of variation exhibited by these features. Note that the examples used here are not necessarily Australian plants. A-C (left): cross sections of herbaceous dicotyledonous (Le. flowering plant) stems. A: Pebrgonium (original diameter, 1 cm). B: Ruruncuiw (original diameter, c. 0.5 cm). C: Cucurbita (originaldiameter c. 0.9 cm). D, E (top right): cross sections of the stem of the vine Aristolochia, in early (D) and later (E) stages of
Boyd, Pretty 47
periderm
1
secondary xylem
crushed pith
I
xylem
-
secondary (i.e. woody) growth; the original diameter are 0.5 cm (D) and 1 cm (E). F-M (bottom right): cross section sketches of petioles (G, I, K, M) and leaf midveins (F, H, J, K) of various plants, showing the different arrangements of vascular tissues. F, G: Songuiunba. H, I: Vitis. J, K: Pelargonium. L, M: Juglam. (Adapted from Esau 1960, Figs. 17.5 and 18.4).
48
Archaeological Palaeobotany in Australia
Stomata, hairs and secretory devices are thus readily observable on isolatedfragments of leaf cuticle. However, the isolated cuticle is translucent and even with staining, light microscope examination of cuticles show a range of outlines and shapes of microstructureswhich are frequently superimposed upon each other because of their position on either side of the cuticle. The application of interference phase contrast microscopy has allowed better discrimination of cuticle features of different thickness and optical density, although scanning electron microscopy (S.E.M.) offers the best technique presently available, even though separate preparations are required for the examination of outer and inner surfaces of the cuticle (Barker 1970; Lange 1969). Indeed, the use of the S.E.M., for example, has long been identified as a powerful technique in environmental archaeology (Brothwell 1969; Ford 1979; Korber-Grohne 1981; Yarnell 1976). This type of S.E.M. work relies on the presence on the leaf surface of many plants of a surface coating of wax or other organic deposits secreted by the leaf, as well as a range of adcrustations such as fungal hyphae and spores which are largely dependant upon the environment. These external features persist on the isolated leaf cuticle and can be obsenred by S.E.M. examination of the outer surface. S.E.M. examination of the inner surface will provide information on the arrangement and micro-relief of the epidermal periclinal walls (i.e. cell walls parallel to the leaf surface) and the anticlinal walls (i.e. cell walls perpendicular to the leaf surface); the anticlinal walls of the epidermis are represented on the cuticle, for example, by flanges of cuticular material of differing morphologies, and whereas the periclinal walls may be smooth, in some cases, there may be pitting, striations or other textural features which will assist in the identification. These characters are regarded as constant across the leaf and thus are useful in the diagnosis and description of the leaf. The characters are also accepted as taxonomicallv reliable within a narrow suite of taxa and thus useful in distinguis'hing subspecies within species or species within genera (e.g. Wilkinson 1983).
Plant remains are often preserved in dry sediments in a dehydrated state, and if they are present, may be recovered from such sediments by sieving (wet or dry) or by floatation methods (the sediment may need some assistance to disaggregate, and washing with water or a mild alkali should not do too much damage to the plant remains). In such cases, these organic remains may be readily identifiable are plant material by their non-mineral appearance, translucent and usually brownish colour and their apparent cellular structure. The dehydrated remains have often not been mineralised, and may be restored by rehydration, using a wetting agent, and then stored in a liquid preservative. The resultant plant remains will then be fragile, but should be capable of handling, and can be treated for examination as for modern plant material. Turning to the second group of plant macrofossils indicated in the Roonka pilot study as being potentially important botanical fossils in such a situation, it is possible to illustratethe significant advances that have been made to the knowledge of plant stem anatomy and their application to taxonomy by referring to Metcalfe and Chalk's two editions (1950, 1979) of an encyclopaedic work which emphasised stem and petiole (the stalk of the leaf) characters of each angiosperm (the non-coniferous flowering plants) family. In the second edition of that work (Metcalfeand Chalk 1979) contributors provide reviews of the many post-1950 publicationswhich include, of importance here, the application of plant stem anatomy to taxonomy. Stem anatomy continues to be a significant part of taxonomic reviews of contemporary plant groups, but compression and distortion of plant stems in fossil deposits may restrict the use of anatomical detail in palaeosystematic studies. In archaeological deposits, however, material often undergoes only minimal compression, and in practice dehydrated fragments can be
Boyd, Pretty 49
rehydrated (if necessary), and stored and handled as for leaf fragments, embedded in a suitable medium and sectioned for microscopy. Unlike the cuticular studies outlined above, observation of the anatomical details of stem tissue types can be achieved with light microscopy alone, although loss or minor distortion of some detail through imperfect preservation should be expected. Useful characters in primary plant axes (i.e. stems and petioles without wood) include the extent of the cortical parenchyma tissue (the unspecialised tissue between the conducting and the mechanical tissues), the disposition of the xylem and phloem (the tissues which conduct fluids through the plant), and the location and structure of other special inclusions such as oil bodies, sclerified cells or mineralisedcells, many of which have a distinctive shape (Esau 1960; Fig. 2). In secondary stems, the presence of wood tissues provides additional characters such as the presence or absence of growth rings and the distribution, size and arrangement of the different vessel cells, fibre cells and parenchyma cells in the wood (Jane 1970). Other characters relate to the cell wall structure of these tissues, particularly the pits which occur along the long axis of the cells. Woody stems have, for a long time, been analysed from samples collected at archaeological sites, largely since wood and, especially carbonised wood (i.e. charcoal) may have a high survival rate in many preserving conditions. It has also been possible to handle wood and charcoal fragments with instruments of equally limited resolution, often with little specialist equipment (for well-preserved specimens a simple low-power binocular microscope may be all that is required) although greater resolution may be derived using higher-power (e.g. to X 250 magnification) microscope. One of us (W.E.B.), for example, has made good use of such a microscope in the study of carbonised cereal caryopses (grains) and wood fragments from many European archaeological sites. The important point here, however, is that identification of species present as fossils at an archaeological site need not necessarily be restricted to those species which have woody stems, as has often been the case in the past, and that a wider range stem material may be usefully studied. There is clearly a wide range of diagnostic characters which may be used for identification. Whatever to fossil type, however, to make the fullest use of these in all instances of taxonomic identification, the determinations are dependent upon comparisons with known, modern materials. In a field such as this, the necessity for access to as complete as possible a collection of reference material as can be obtained is imperative. The establishment and eventual use of such a reference collection may be a moderately time-consuming exercise, but one which will inevitably enhance the value of the research undertaken.
CONCLUSIONS With the growth of environmental archaeology, especially in Europe, as an increasingly important field of research within archaeology, the scope and techniques of palaeobiological research are widening. In such palaeobotanic work, many developments show great archaeological promise, and although some, such as pollen analysis, may be less directly applicable within archaeological research than in, for example, Europe, others such as the study of vegetative plant parts offer a valuable potential input. In particular, the preliminary investigations at Roonka offer promise that studies of plant leaf and stem fragments form a potentially sensitive alternative avenue of taxonomic identification in sites such as those in arid Australia where conditions impede the employment of the classic Northern Hemisphere palaeobotanical techniques.
50 Archaeological Palaeobotanyin Australia
ACKNOWLEDGMENTS The archaeological excavations, under the direction of G.L.I?, were funded by the Australian Institute of Aboriginal Studies, the Australian Research Grants Scheme, the Sir Mark Mitchell Foundation, the Potter Foundation, the Sunshine Foundation, the Myer Foundation and the Utah Foundation. Thanks are extended towards those scientistswho have contributed advice at various stages in this project: Drs W.K. Harris and J.R. Dodson provided exploratory palynological assessments, and Drs D.C. Christophel, W.J. Harvey, J.F? Jessop and G. Singh have all advised on the design of the Roonka palaeobotany project.
REFERENCES Baker, E.A. 1970 The morphology and composition of isolated plant cuticles. New Phytologist 69: 1053-1OS8 Beijerinck, W. 1947 Zadenatlas der Nederlandsche Flora. Veenman and Zonen: Wageningen Berglund, B.E. (ed.) 1986 Handbook of Holocene Palaeoecology and Palaeohydrology. John Wiley: Chichester Brothwell, D. 1969 The study of archaeological materials by means of the scanning electron microscope; a new important field. In Brothwell, D. and E. Higgs (eds) Science in Archaeology: A Survey of Progress and Research pp.564-66. Thames and Hudson: London Bryant, V.M. and R.G. Holloway 1983 The role of palynology in archaeology. Advances in Archaeological Method and Theo/y 6: 191-224 Carolin, R.C. 1971 The trichomes of the Goodeniaceae. Proceedings of the Linnean Society of New South Wales 96:8-22 Clark, A. 1985 A preliminary archaeobotanical analysis of the Anbangbang 1 site. In R. Jones (ed.) Archaeological Research in Kakadu National Park pp.77-96. Australian National Parks and Wildlife Service, Special Publication 13: Canberra Clark, R.L. 1983 Pollen and charcoal evidence for the effects of Aboriginal burning on the vegetation of Australia. Archaeology in Oceania 18:32-7 Clark, R.L. and R.J. Lampert 1981 Past changes in burning regime as markers of man's activity on Kangaroo Island, South Australia. Terra Australis 5: 186-89 Dickson, C.A. 1987 The identification of cereal from ancient bran fragments. Circaea 4:95-102 Dimbleby, G.W. l985 The Palynology of Archaeological Sites. Academic Press: London Dodson, J.R. 1977 Late Quaternary palaeoecology in Wyrie Swamp, southeastern South Australia. Quaternary Research 8:97-114 Esua, K. 1960 Anatomy of Seed Plants. John Wiley: New York Ford, R.I. 1979 Palaeoethnobotany in American archaeology. Advances in Archaeological Method and Theory 2:285-336 Fujiwara, H., R. Jones and S. Brockwell 1985 Plant opals (phytoliths) in Kakadu archaeological sites: a preliminary report. In R. Jones (ed.) Archaeological Research in Kakadu National Park pp. 155-64. Australian National Parks and Wildlife Service, Special Publication 13: Canberra Harvey, W.J., and G.L. Pretty. 1984 Archaeobotany in Australia. Unpublished manuscript, Archaeology Laboratory, South Australian Museum.
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Greig, J.R.H. 1982 The interpretation of pollen spectra from urban archaeological deposits. In A.R. Hall and H.K. Kenward (eds) Environmental Archaeology in the Urban Context pp.47-65. Council for British Archaeology, Research Report 43: London Hope, G., PJ. Hughes and J. Russell-Smith 1985 Geomorphological fieldwork and the evolution Archaeological Research of the landscape in Kakadu National Park. In R. Jones (d.) in Kakadu National Park pp.229-40. Australian National Parks and Wildlife Service, Special Publication 13: Canberra Jane, F.W. 1970 The Structure of Wood. Adarn and Charles Black: London Kershaw, A.F? 1986 Climatic change and Aboriginal burning in north-east Australia during the last two glacial/interglacial cycles. Nature 322:47-9 Korber-Grohne, U. l 9 6 4 BestimmungsschliisseI fijr subfossile Juncus-samen und Gramineen-fruchte. Problerne der Kiistenforschung im sudlichen Nordseegebiet 7. Hildeheim Korber-Grohne, U. 1981 Distinguishing prehistoric cereal grains of Triticum and Secale on the basis of their surface patterns using scanning electron microscope. Journal of Archaeological Science 8: 197-204 Korber-Grohne, U. and U. Peining 1980 Microstructure of the surfaces of carbonised and non-carbonised grains of cereals as observed in scanning electron and light microscopes as an additional aid in determining prehistoric findings. Flora 170:189-228 Kovack, W.L. and D.L. Dilcher 1984 Dispersed cuticles from the Eocene of North America. Botanical Journal of the Linnean Society 88:63-104 Lange, R.T. 1969 Concerning the morphology of isolated plant cuticles. New Phytologist 68163-104 Luebbers, R.A. 1973 Ancient boomerangs discovered in South Australia. Nature 253:39 Martin, H.H. 1973 Palynology and historical ecology of some cave excavations in the Australian Nullabor. Australian Journal of Botany 21:283-316 Meehan, B. 1971 The Form, Distribution and Antiquity of Australian Mortuary Practices. Unpublished M.A. thesis, University of Sydney Metcalfe, C.R. and L. Chalk 1950 Anatomy of the Dicotyledons. Clarendon Press: Oxford Metcalfe, C.R. and L. Chalk (eds) 1979 Anatomyof theDicotyledons (2nd ed.). Clarendon Press: Oxford Moore, PD. and J.A. Webb 1978 An Illustrated Guide to PollenAnalysis. Hodder and Stoughton: London Palmer, PG. 1976 Grass cuticles: a new palaeoecological tool for East African lake sediments. Canadian Journal of Botany 54: 1725-34 Parker, K.L. 1905 The Euahlayi Tribe. Constable: London Payne, W.W. 1979 Stomata patterns in Embryophytes: their evolution, ontogeny and interpretation. Taxon 28:117-32 Pettigrew, C.J. and L. Watson 1973 On the identification of sterile Acacias and the feasibility of establishing an automatic key-generating system. Australian Journal of Botany 21:l4l-5O Piperno, D.R. 1985a Phytolith taphonomy and distributions in archaeological sediments from Panama. Journal of Archaeological Science 12:247-67
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Piperno, D.R. 1985b Phytolith analysis and tropical paleo-ecology: production and taxonomic significance of siliceous forms in New World plant domesticates and wild species. Review of Palaeobotany and Palynology 45: 185-228
Pretty, G.L. 1977 The cultural chronology of the Roonka Flat: a preliminary consideration. In R.V.S. Wright (d.) Stone Tools as Cultural Markers: Change, Evolution and Complexity pp.288-331. Australian Institute of Aboriginal Studies: Canberra Pretty, G.L. l986 The prehistory of South Australia. In E. Richards (ed.) The Flinders Hisfory of South Australia pp.36-62. Wakefield Press: Adelaide Singh, G., A I ! Kershaw and R. Clark 1981 Quaternary vegetation and fire history in Australia. In A.M. Gill, R A . Groves and I.R. Noble (eds) Fire and the Australian Biota pp.23-54. Australian Academy of Science Stace, C.A. 1965 Cuticular studies as an aid to plant taxonomy. Bulletin of the British Museum (Natural History) 4:3-79 Stace, C.A. 1966 The use of epidermal characters in phytogenetic considerations. New Phytologist 65:3O4- 18 Stemler, A.M. and R.H. Falk 1981 S.E.M. of archaeological plant specimens. Scanning Electron 1981 :19 1-96 M~C~OSCOPY Tomlinson, F! 1985a Use of vegetative remains in the identification of dyeplants from waterlogged 9th-l 0th century A. D. deposits from York. Journal of Archaeological Science 12:269-83 Tomlinson, F! 1985b An aid to the identification of fossil buds, bud-scales and catkin-bracts of Briiish trees and shrubs. Circaea 3:45-130 Wilkinson, H.P 1983 Leaf anatomy of Gluta (L.) Ding Hon (Anacardiaceae). Botanical Journal of the Linnean Society 86:375-403 Wilson, S.M. 1985 Phytolith analysis at Kuk, an early agrictutural site in Papua New Guinea. Archaeology in Oceania, 2090-97 Yarnell, A. 1976 Early plant husbandry in eastern North America. In C.E. Cleland (ed.) Cultural Change and Continuity pp.265-75. Academic Press: New York
1.
School of Resource Science and Management, University of New England, Northern Rivers, Lismore NSW 2480
Archaeology Department, South Australian Museum, Adelaide SA 5000
