Anything more than a picnic? Re-considering arguments for ceremonial Macrozamia use in mid-Holocene Australia.
Article from:Archaeology in Oceania
Article date:October 1, 2008
Author: Asmussen, Brit
Abstract
Influential arguments have been advanced in Australian archaeology concerning the origins and development of social and economic change in the mid-late Holocene (Lourandos 1997). One example used to support this claim is the perceived existence of ceremonial feasting events held in the semi-arid and rugged sandstone gorge systems of central Queensland, attended by large groups of people for extended periods, and underwritten by large quantities of kernels from the cycad Macrozamia moorei (Beaton 1977, 1982; see also Lourandos 1997). However the reexamination of the macrobotanical evidence from archaeological sites in this region using taphonomic analysis, replicative processing experiments, recalculations of seed density and estimations of the minimum numbers of seeds, does not support this model. This re-examination questions the role of Macrozamia seeds in the context of socio-economic change and suggests new interpretations of Macrozamia resource use.
Keywords: ceremony, intensification, Macrozamia, central Queensland, mid-late Holocene
**********
Large-scale communal public rituals and ceremonies supported by feasts have been argued to have played an important role in the social, economic and political arenas of ancient cultures (Jennings 2005; Potter 1997). In Australia, continent-scale models concerning the development of mid-Holocene ceremonial events and resource intensification have been strongly influenced by Beaton's interpretations of the archaeobotanical records of Cathedral Cave, Wanderer's Cave and Rainbow Cave in the Central Queensland Highlands (CQH). These sites appeared to provide direct archaeological evidence for frequent, large-scale inter-group ceremonial feasting events supported by the processing and consumption of toxic Macrozamia seeds, dating back to the mid-Holocene (Beaton 1977, 1982, 1993). Beaton's arguments were highly influential and have been widely cited by researchers arguing for continental-scale transformative social processes in the mid-late Holocene (Jones 1978; Lourandos 1980a, 1980b, 1983a, 1983b, 1988, 1997; see also David and Denham 2006; Ross 2006).
There has been significant debate about ways of testing high-level models of intensification of socio-political complexity in Holocene Australia (Beaton 1995:798; Bird and Frankel 1991a, 1991b; Bird et al. 1997; Edwards and O'Connell 1995:776; Frankel 1988, 1991a, 1991b, 1993, 1995:654; Godfrey 1989; Hiscock 1981, 2002, 2008; Holdaway et al. 2002; Jones 1980; Lilley 2000; Pardoe 1995). However there have been few detailed reexaminations of archaeological data using the kinds of analytic approaches called for in these debates (although see Attenbrow 2004 and Hiscock 2008).
This paper presents the results of a detailed reexamination of the Macrozamia assemblages from these three sites from the Central Queensland Highlands, excavated by John Beaton in the mid 1970's, and held by the Queensland Museum. The first section of this paper presents an overview of the original model of ceremonial cycad use in the CQH. The second section of the paper presents the methods and results of experimental replicative processing techniques, taphonomic analyses to identify the non-human component of the assemblages, analysis of site formation, and techniques used to derive more accurate density and MNI estimates. The third section argues that the results of the reanalysis fail to support Beaton's arguments for either the use of large quantities of Macrozamia seeds over the last 4300 years, or the regular occurrence of Macrozamia supported ceremonial events in these sites, and argues for the subsistence use of these seeds by small groups of foragers.
Ceremonial cycad feasting in the Central Queensland Highlands
A particularly striking example of socio-economic intensification was argued to have developed 4300 years ago in the rugged sandstone gorges in the Central Queensland Highlands (Beaton 1977, 1991a, 1991b). Although the three rockshelter sites in this region, Cathedral Cave, Wanderer's Cave and Rainbow Cave, contained a range of excavated materials, the large, robust and highly visible seed shells from Macrozamia moorei plants became the focus of archaeological explanations (Beaton 1991a: 12, 24). Beaton estimated that the fractured and carbonised Macrozamia specimens had densities of 400-600 seeds per [m.sup.3] in each of these sites.
The density of Macrozamia shells led to the consideration of the circumstances in which such quantities of seeds would have been eaten and their shells discarded. Beaton's interpretation of the use of Macrozamia seeds in the highland sites was heavily based on the anthropological account made in the late 1970s by Meehan and Jones, who described the use of seeds from the related Cycas sp. by Aboriginal communities living along the Blythe River in Arnhem Land, in the Northern Territory (Meehan and Jones 1977 Appendix IV, in Beaton 1977:165-166; Morphy pets. com. to Beaton 1977). Here, large-scale multi-group ritual and initiation ceremonies and other large gatherings were underwritten by the collection and processing of large quantities of cycad seeds (Berndt 1951; Harvey 1945:191; Levitt 1981; Spencer 1914; Thomson 1938; for detailed recent accounts of cycad use in the Northern Territory see Bradley 2006). Beaton suggested that "the archaeological evidence is not unequivocal but it is highly suggestive" of the connection between Macrozamia seeds as a communion food attended by an "unusually large gathering" of people for ceremonial events (Beaton 1977:194-195). Beaton took support from Sullivan's arguments for Bunya Nut (Araucaria bidwillii) feasts in southeast Queensland (Sullivan 1977:60) and Flood's (1976:40-44) arguments for ceremonies supported by annual migrations of Bogong Moths (Agrotis infusa) in the southeastern Highlands (Beaton 1977).
But were the interpretations of the Macrozamia assemblages from the CQH correct? The original archaeological interpretations were based on a simple estimate of the density of Macrozamia specimens in the assemblage which was not rigorously quantified and without any taphonomic or site formation analysis to confirm the human origins of the assemblage.
Re-examining the Central Queensland Highland archaeological evidence
Since the time of Beaton's analysis it is now appreciated that macrobotanical remains in archaeological sites are often not a complete, exclusive or unambiguous record of human use, and that there are a diverse range of taphonomic processes that can affect the material deposited which must be analysed prior to forming behavioural interpretations (Schiffer 1976; see also Clarke 1988, 1999:83, Hansen 2001:401; Ladd 1988: 2; Miksicek 1987; Murphy 1992:9; Nelson 1992:240; Pennington and Weber 2004; Rossen et al. 1996:405: Spicer 1991:72). In addition, replicative plant processing experiments have allowed archaeologists to more adequately assess the economic importance of plant species as represented by fragmented archaeobotanical remains (for example see Dennell 1976; Diehl 1996; Margaritis and Jones 2006).
To test Beaton's model a detailed re-examination of the Macrozamia assemblage from Cathedral, Wanderer's and Rainbow Caves was undertaken. A better understanding of Macrozamia use was achieved by focusing on three analyses: 1) to identify and separate human and non-human depositors of the seeds via replicative processing experiments and taphonomic analyses, 2) the assessment of site formation processes affecting the assemblage, and 3) the re-calculation of the Number of Identified Specimens (NISP) and estimating the Minimum Number of Individuals (MNI) of the Macrozamia assemblages. A 100% sample of the Macrozamia specimens from each site was analysed.
Replicative processing experiments
Building on the work of Beck (1989) and Beck et al. (1988), a replicative experimental research programme was undertaken to identify seed specimens fractured as a result of human processing techniques. The methodology and test variables, including indentor type (hammerstone or wooden baton), presence of sarcotesta, and seed carbonisation extent (see Asmussen 2005) were based on traditional processing methods as described in ethnographic and historical documents (Gardner and Bennetts 1956; Goodale 1971; Levitt 1981; Maiden 1889, 1890; Meehan and Jones 1977; Roth 1901; Thomson 1938; Thozet 1878; Turner 1893).
As a result of this work, a set of diagnostic criteria indicative of human processing were derived using analysis of fracture patterns on individual seeds. The methods used to open the shells, end and side striking, required the very precise application of the right amount of force so as not to crush the kernel. This produced a very distinctive pattern of fracturing unlikely to be replicated by animals or other natural processes. Characteristic features of humanly processed seeds included ringcracks on the sides and ends of the seed where the indentor met the seed, irregularly fractured edges of seeds where the force propagated through the seed, and other small fractures on the seed shell (Asmussen 2005; see also Beck 1989 and Beck et al. 1988).
Identifying seed deposition and use by animals
A number of animals have been observed to actively collect, transport, modify and deposit Macrozamia seeds within and around archaeological sites. The outer fleshy layer of Macrozamia seeds (the sarcotesta) is brightly coloured and acts as a food attractor, and provides a nutritionally valuable starchy food reward (Jones 1993; Moore 1999; Norstog and Nicholls 1997; Renner 2003). If the sarcotesta has decayed or has already been eaten, the remaining inner seed shell (sclerotesta) can be opened to attain the edible internal kernel (Jones 1993: 60; Snow and Walter 2007). A literature review identified the likely animal agents as birds, fruit eating bats, bandicoots, possums, macropods, rodents, native cats and dogs (Ballardie and Whelan 1986: 65; Bauman and Yokohama 1978:73; Begg and Dunlop 1980: 65; Carter 1923; Hill 1984; Hill and Osborne 2001:5; Jones 1993:62; Kennedy 1993; Loafing 1952; Moore 1999; Parton 1952; Sanchez-Tinoco and Engleman 2004:36; Sargent 1928; Sedgwick 1952; Snow and Walter 2007; Stranger and Stranger 1970; Tang 1990; van der Pijl 1957; Vorster 1995:383: Watkinson and Powell 1997:347).
However taphonomic analysis of the archaeological specimens themselves indicated that the native rodent, Rattus fuscipes, and the native mouse, Mus sp., were the most important taphonomic agents involved in the formation and modification of the assemblage. R. fuscipes has been observed removing the outer flesh (sarcotesta) from seeds (Snow and Walter 2007: 595), and gnawing into the woody layer (sclerotesta) to extract the inner kernel (Ballardie and Whelan 1986: 101). R. fuscipes feeds on Macrozamia seeds where they rest on the ground near the parent plant, or remove seeds to other locations, including feeding refuges (Snow and Walter 2007: 595, 598), including within archaeological sites (Murphy 1992). More generally, rodent accumulated and gnawed seeds have been identified from several archaeological sites in Australia (for examples see Beck 1989: 46; Clarke 1988:123; Colvill 1995:16; Murphy 1992: 91).
The proportion of the Macrozamia assemblage gnawed, fractured and potentially deposited by rodents was identified using toothmarks, which were identified through direct 1:1 comparative analysis of rodent modified seeds obtained through field collection, actualistic feeding experiments (Asmussen 2005) and photographic evidence (Bang and Dahlstrom 1972; Begg and Dunlop 1980:68; Triggs 1996: 226).
Site formation
Macrozamia specimens were distributed fairly evenly in the cultural layers of each site, with no stratigraphic evidence of large scale depositional events of Macrozamia specimens. Site-specific analysis of site formation processes including geomorphological analysis, conjoin analysis of faunal, lithic and botanical components and analysis of thermal modification on conjoined faunal elements ruled out large-scale post-depositional disturbance at Rainbow and Wanderer's Caves. However, Cathedral Cave was subject to repeated flooding over the last 2910 years calBP, and the impact on the movement, winnowing and emplacement of Macrozamia specimens needed to be assessed. A sample of specimens created during the processing experiments were used to assess the potential for fluvial transport of complete and fractured, burnt and unburnt seed specimens (after Behrensmeyer 1975; Asmussen 2005).
Potential biases caused by differential spatial and temporal post-depositional preservation of plant remains were assessed using comparative statistical analysis of densities of carbonised and uncarbonised seeds and comparisons to pH variations (Asmussen 2005, see also Balme and Beck 2002:159, 164; Clarke 1999: 83; Karkanas et al. 1999; Zutter 1999). Charcoal was unable to be used as a comparative control as samples were not retained after excavation.
Results of experimental and taphonomic analysis
The Macrozamia assemblages from the three sites were analysed in comparison to the experimentally processed samples in order to determine the agents opening the seeds. The results of the experimental and taphonomic analyses supported Beaton's interpretation that the vast majority of shells in the highland archaeological sites were collected by humans and processed using traditional methods (Asmussen 2005).
Although ecological data indicated the potential for significant use of seeds by rodents, analysis of the seeds themselves indicated that rodents were not a major depositor of seeds. While there are significant rates of rodent toothmarking on the sclerotesta in some levels of the sites (discussed further below), the rate at which seeds were gnawed open is consistent with rates of gnawing observed on seeds found at the base of parent plants (Burbidge and Whelan 1982; Jones 1987; Snow and Walter 2007). Given the vast majority of seeds showed the direct evidence of human processing and very few seeds were gnawed open (less than 1% at each site), it is likely that gnawing of seeds had taken place prior to collection by human foragers.
At Cathedral Cave, the potential deposition and winnowing of portions of the Macrozamia assemblage through repeated fluvial events was assessed. A small scale experimental programme was used to identify the hydrodynamic behaviours of Macrozamia seed specimens of different sizes, shapes and taphonomic states. Transport potential was assessed by estimating a specimen's equivalence to quartz grains deposited under fluvial conditions in the site (see Behrensmeyer 1975). Sediment analysis was then conducted to determine whether floods at the site had enough velocity to winnow and or deposit noncultural Macrozamia seed remains. Comparisons between the results of the experimental trials and analysis of the archaeological specimens suggest that carbonised specimens were susceptible to water transport, and may have been removed from fluvially modified layers in the site. Cathedral Cave had the lowest rates of burnt specimens of all the three sites (Asmussen 2005). Taphonomic analysis also indicated that a small proportion (c. 5%) of Macrozamia specimens from Cathedral Cave may have been fluvially deposited; the presence of mechanical damage, abrasion and adhering silt indicating that they had spent part of their post-depositional history within a fluvial system. However, the overall impact of fluvial events was relatively small (Asmussen 2005).
Overall, Macrozamia seeds were extremely well preserved due to the reasonably thick (1-3 ram) and highly lignified sclerotesta (seed coat), and the majority of seeds were carbonised, increasing preservation potential (as is generally the case with macrobotanical specimens see Hansen 2001: 405; Jones 1987; Ladd 1988:10; Miksicek 1987; Pearsall 1989:229, 440). There is some evidence for the possible loss of uncarbonised seeds as a result of sediment acidity in levels lower than 30 cm depth in Wanderer's Cave (Asmussen 2005). Extensive carbonisation at Wanderer's Cave was likely to have reduced the MNI calculations and increased NISP. pH data was not collected at Cathedral Cave, and differential preservation could not be assessed.
Recalculating Maerozamia MNI and NISP
Having established that the majority of the specimens were brought into the caves and humanly opened, various measures of seed density were calculated to assess the argument that there were large numbers of specimens in these sites. While Beaton had argued that the amount of Macrozamia in the sites was far more than could be explained by any possible everyday subsistence or "mundane use", the data suggest otherwise.
Calculations were made for the number of identifiable specimens (NISP) and minimum number of individuals (MNI) for each of the sites. MNI calculations were possible as each seed has two ends, the micropylar and the attachment end, each of which have distinct external and internal features. MNI was based on estimating the completeness of the diagnostic end represented on each specimen (Lyman 1994). These percentages were tallied for each end of the seed to generate an MNE, and the largest MNE for either end was taken as the MNI. Identification of either end (on which MNI is based) was possible using both internal and external features, and identification was reduced only when specimens were smaller than 2.0 mm in maximum dimension rather than as a result of taphonomic alteration (for example weathering, acid dissolution or carbonisation). Estimates of the total volume of excavated sediment and total volume of sediment in each site were used to extrapolate a total NISP and MNI for each site. The figures are summarised in Table 1.
Reconsidering models for the ceremonial use of Macrozamia seeds
It is clear from Table 1 that the Macrozamia assemblages from these sites do not support the idea that Macrozamia were used to underwrite large-scale ceremonial activities conducted in or around the sites. Analysis of the NISP and MNI of Macrozamia seeds in the highland sites indicates that there are not enough specimens when compared with the quantity that would be expected if ceremonies had occurred.
Female M. moorei plants produce seeds on a large strobilus (megasporophyll) which typically contain 300-360 seeds (Jones 1993:44; Low 1991:138; Rolf Kyburz pers.com; pets. obs). Extrapolating over the entire site at Cathedral Cave, which has by far the largest NISE the results indicate an average rate of deposition of 13 MNI per year and 83.5 NISP per year. Experimental studies (Asmussen 2005) indicate that an average of three fragments are generated per processed Macrozamia seed, implying the observed density of Macrozamia could be generated by the deposition of approximately 28 seeds per year, or the seeds from one strobilus every twelve years. Even taking the most generous approach and allowing a count of one seed per NISR this still only represents one strobilus every four years. This is hardly prima facie evidence of intense use--ceremonial or otherwise.
As a further point of comparison it is worth considering the expected quantities of Macrozamia specimens generated by the kinds of ceremonies being considered by Beaton. Beaton based his account of the ceremonial feasting on the ethnographic accounts of Meehan and Jones (1977). The number of people and the duration are not specified but it is reported that 45 kg of seeds were collected and processed per day for a seven day period, giving 315 kg in total, for a total weight of kernels of around 180 kg. To generate 180 kg of Macrozamia kernels would require the processing of 11,000-12,000 seeds, generating more than 33,000 fragments (NISP), assuming a mean of three fragments generated per seed. It would take almost 400 years to accumulate this number of specimens at Cathedral Cave, which has the highest rate of deposition of the three sites. The densities of 400-600 [m.sup.3] estimated by Beaton is much more consistent with mundane use.
Examining the economic role of Macrozamia resources in the CQH
A more detailed picture of the use of this resource can be gained by analysis of the ecological processes occurring in plant populations prior to human collection. Some of these ecological processes leave taphonomic traces on the seeds themselves, which can be used to infer the age of seeds at the time of collection and the collection of seeds from masting events. "Mast seeding" occurs in the genus, indicated by the synchronous production of seeds in one year by a high proportion of the plants in a population (c. 70%), but is usually followed by a long interval where few seeds are set (Ballardie and Whelan 1986: 101).
Age of seed at time of collection
In Macrozamia populations, seeds of three main developmental stages can be found, which can be used in subsistence: 1) fresh seeds which have just been produced by plants, 2) old seeds produced in a prior seed production event, and 3) and seeds which are in the initial stages of germination.
Seeds which have just been produced by plants have adhering fleshy sarcotesta (flesh). The flesh usually decays or is eaten by animals in the first six months following seed release (Baird 1939:155; Ballardie and Whelan 1986; Ornduff 1990, 1991; Tang 1990). Fresh seeds are toxic, and need to be processed either by roasting in the shells, or leaching of the kernel. Those archaeological specimens with remnants of sarcotesta still attached were almost certainly collected while fresh.
Older seeds produced in prior seed production events can often become trapped in the leaves of parent plants, or can remain around the bases of parent plants for several years, even in the presence of predators as, once the flesh has been removed, seeds are less attractive to predators (Burbidge and Whelan 1982; Snow and Walter 2007). Well-aged seeds can be eaten without processing (Beck 1985; Meehan and Jones 1977). While it has been suggested that aged seeds can be identified by a colour change in the seed from mid-brown to grey-white (Levitt 1981:48), actualistic experiments indicate that seeds collected and processed while fresh can assume a weathered appearance post-discard and may not indicate collection of aged seeds (Asmussen pers. obs.).
Most Macrozamia seeds cannot germinate immediately following seed release, as the embryo has an "after ripening period", the length of which varies between species in the genus from a few weeks to several months. In M. moorei this period is between three and 12 months (Jones 1993). Hunter-gatherers can utilize early germinating seeds by removing the embryo from the kernel, however once the root becomes too advanced it cannot be used in subsistence. Aged seeds which have just started to germinate can be identified by a characteristic hole on the micropylar end which remains identifiable when the seed has been processed using side-striking techniques.
Collection of seed from masting events
Differences in the overall rate of rodent toothmarking on the outside of seeds may reflect differences in the timing of forager seed collection strategies in relation to masting events. A number of studies have indicated that rodents intensify their activities during masting events in seeding plants. For example, Li and Zhang (2007) found greater caching of seeds and gnawing into seeds of Prunus armeniaca (Apricot) during masting events.
Similarly, ecological studies indicate that rodent predation on Macrozamia seeds is likely to be "density responsive", with strong relationships between the frequency of rodent predation and the size of the seed bank (Ballardie and Whelan 1986; Jones 1993; Ornduff 1990; Schnurr et al. 2002; Zhang et al. 1997). Rodent seed predation on M. communis seeds in NSW over one year has been studied by Ballardie and Whelan (1986:103-4). They found that rodents gnawed seeds open more frequently (18.9%) in a masting population, where the majority of the population (in this case 70%) was synchronously producing large quantities of seed, in comparison to an non-masting population (1.8% gnawed open, 8% of the population producing seed) (see also Farrera 2004:307; Sanchez-Tinoco and Engleman 2004:35; Vovides 1990:1539).
Interestingly, rodent opened specimens are extremely uncommon in the archaeological assemblages. This is most likely because people deliberately avoided collecting rodent opened shells. However, it appears rodents will often eat the sarcotesta without gnawing through the shell. Snow and Walter's (2007) study of predation in M. lucida populations found that rodents removed the flesh of seeds but largely ignored seeds without the fleshy sarcotesta. Given this, it is highly likely that any intensified rodent activity will be reflected in the rate of toothmarking overall (see Table 2).
Results of ecological analyses
The ecological attributes are presented in Table 2. It should be noted that the assemblage at 10-15 cm depth from Wanderer's Cave was unable to be analysed as it was not in the Queensland Museums' collection. Seven specimens from the final two cultural strata at Cathedral Cave were not included in this analysis due to small sample size. In addition, specimens from Level 2 (N=43) and from Level 4 (N=79) are not discussed here as they were emplaced by fluvial events and do not reflect human use. Trends from Rainbow Cave were unable to be analysed due to small sample size (N= 106).
Age of seed at time of collection
The evidence indicates that people were using both fresh and aged seeds through time at both sites. Most levels have seeds with sarcotesta still attached, indicating that they were collected while fresh, and seeds which had started to germinate, indicating the use of well-aged seeds remaining in the environment from prior coning events. (See also Beck et al. (1988:143-145), who concluded that both fresh and aged seeds were processed c.1000-2000 BP at Wanderer's Cave).
The rate of burning of seeds was examined as a possible indicator of deliberate roasting of fresh seeds. However taphonomic analysis on bone from the sites indicates that the seeds were almost certainly exposed to the heat from campfires after deposition. Thus burning cannot be taken as evidence of deliberate roasting (Asmussen 2005).
Germinating seeds will generally be found as part of a much larger cohort of aged seeds from a prior coning event either at the base of the plant or trapped within its leaves. Only seeds in the early stages of germination will be useful as a food resource by hunter-gatherers. Recent ecological study of M. lucida indicated that only 26% of the seeds germinated even though 93% of the seed were viable (Snow and Walter 2007:597, Figure 4). This indicates that it is likely that at least 3-4 ungerminated aged seeds were collected for each germinated seed in the assemblage, suggesting the use of 30% or more aged seeds from most layers of the sites.
Similarly, not all fresh seeds will retain sarcotesta at the time of collection. Sarcotesta does not survive well, as it is eaten by mammals, attacked by insects and decomposes shortly after seed dispersal. However, sarcotesta may survive better archaeologically if it is burnt, as it is no longer attractive to animal consumption. Sarcotesta only survived as small patches on archaeological specimens. The rate of sarcotesta in most levels of the sites is below 10%. This represents a substantial underestimation of the actual number of fresh seeds collected by foragers. The high rate of sarcotesta at Wanderer's Cave between 15 and 20 cm depth is probably the result of an individual event where a whole strobilus of fresh seeds with adhering sarcotesta was cooked in a fire.
Collection of seed from masting events
Rates of toothmarking vary between the two sites, with an average rate of toothmarking of 6.8% at Cathedral Cave and 13.5% at Wanderer's Cave. Both sites display an increased rate of toothmarking in the upper layers.
The increases in the frequency of rodent toothmarking in the upper layers may indicate increased frequency of masting events in the last 2000 years due to late Holocene climatic amelioration in the CQH. Current data suggests masting events may be correlated with increased rainfall. Global patterns of masting in 570 different species of perennial plants indicate that variability in seed production in the Southern Hemisphere is related to rainfall variability (Kelly and Sork 2002). During the last 1000 to 2000 years, climatic conditions along the eastern coast of Australia appear to have ameliorated, with warm temperatures and increased rainfall when compared with the preceding period (Harrison and Dodson 1993:276, 279; McGlone et al. 1992:435). Palaeoenvironmental data from the CQH indicates slightly wetter conditions may have been more common in the last 2000 years. Recent increased rainfall events are indicated by phytolith data from Kenniff Cave, also within the CQH (Bowdery 2006), and extensive palaeoflood records from the Nogoa River (Bell et al. 1989) and Cathedral Cave (Asmussen 2005; Beaton 1991b).
While some of the general increase in rate of toothmarking between the two sites may reflect general environmental change, the difference in timing of the increase suggests that the changes also reflect a difference in human behaviour related to the specific use of the sites. At Wanderer's Cave, the high overall rate of toothmarking may reflect a systematic targeting of seeds after masting events, or the deliberate use of the site following masting events. However the low rate of discard suggests small-scale local collecting, with the pattern of using aged and fresh seed consistent with a strategy of opportunistic collection of seeds which are available in the local environment around the sites.
Discussion
Both sites show similarities in the general patterns of use of the resources. The presence of both fresh and aged seeds in both sites is a pattern that is consistent with modern ecological data for seed availability in Macrozamia populations. This suggests that fresh cones seem to have been collected off plants, and isolated seeds from prior coning events were also collected. Both patterns of use are consistent with small-scale local collection (Lepovsky and Lyons 2003:1361) by small groups, indicating the use of the seeds as everyday subsistence use within a broad based hunter-gatherer subsistence strategy rather than associated with ceremonial use.
The data suggest that Macrozamia was not a substantial food resource in the region in the mid-late Holocene. Additionally, NISP and MNI of faunal remains support the view that occupation of these sites was by small forager groups. The thermal modification of the faunal component, and evidence for significant occupation of these shelters by dingoes, suggests that occupation of these sites was relatively infrequent and of short duration (Asmussen 2005).
The use at Cathedral Cave may reflect the opportunistic exploitation of Macrozamia as part of the intermittent transitory use of the shelter. Cathedral Cave is situated in the middle of a rugged sandstone gorge system. The likely small numbers of Macrozamia plants along the gorge in the walk to the site, and the relatively small quantities of seeds in the local environment suggests opportunistic seed collection as people were moving through the landscape, with Macrozamia processing one of number of activities undertaken within the site. Occupation of the site was not necessarily timed to take advantage of local seed availability.
However the higher rate of toothmarking at Wanderer's Cave suggests occupation of the site may have been more regularly timed, or occurred closely in time to masting events. At the time of excavation, Beaton (1991a:19) noted a large grove of 60 plants within 100 m of the site. Ecological studies indicate that only a limited number of Macrozamia seeds disperse and then only to a limited distance, resulting in clumps of plants which are slow to spread (Snow and Walter 2007:597, 598). Given that these are long-lived plants, it is probable that this local population is of some significant antiquity.
Even though there seems to be large quantities of seed available for collection around Wanderer's Cave, the relatively low numbers present in the site suggest exploitation by a small group. Low numbers of people occupying the site is also supported by the low MNI of key mammalian faunal resources, while its intermittent use is supported by the high-level of use of the site by dingoes (Asmussen 2005).
Of course, this evidence can not rule out intense use of the resource away from the rockshelters (that is, seeds collected, processed and discarded in other contexts). Meehan and Jones (1977) indicate that women often set up specific-purpose field processing sites and from these locations journeyed several kilometres into different areas of the forest to collect seeds. Archaeological data often suggest the relatively infrequent or particularistic use of rockshelters. It is possible the majority of human activity occurred outside such sediment and behavioural traps (Bird and Frankel 1991b:189; Walthall 1998).
However, the interpretation of small-scale opportunistic use is supported by more recent ecological data which suggests that large-scale ceremonies were unlikely to have been underwritten by Macrozamia seeds. Although seed production has been modelled as large-scale and easily manipulated, analyses of average yearly seed production indicates consistently low levels of natural seed production. In addition, Macrozamia are not as amenable or receptive to human manipulation as originally argued, and ecological data fails to support the notion that plants reliably respond to fire by generating large numbers of seeds (Asmussen 2005; Jones 1993). Important ecological and environmental differences have been identified between cycad plants in the Northern Territory, and Macrozamia plants which may explain why it was possible that large-scale cycad seed use for ceremonial events was indeed possible in the Northern Territory but not possible with Macrozamia seeds in the CQH (Asmussen 2005). Ecological data also suggests significant constraints on the human use of Macrozamia in arid and semi-arid environments across Australia.
Conclusion
A substantial leap was taken when going from the relatively small amount of plant remains in the Central Queensland Highland sites to the existence of large-scale ceremonies supported by toxic Macrozamia seeds. The macrobotanical evidence does not fit with a model of intensive late-Holocene ceremonial use of this resource. Although the reanalysis confirmed that the majority of seeds were humanly processed in these sites, the number of seeds, either calculated by MNI or NISE is not consistent with the quantities required to feed several hundred people for several weeks at a time. There is no positive archaeological evidence of large-scale intensive collection, economic production nor the sustained, pyrogenically facilitated strategic Macrozamia resource exploitation as previously described (Beaton 1977, 1991a, 1991b; Lourandos 1997:143).
Overall, archaeological, taphonomic and ecological analyses suggest the intermittent low-intensity, subsistence use of seeds and plants by small groups within a broad based hunter-gatherer subsistence strategy. This is strongly supported by the analysis of the faunal remains from these sites (Asmussen 2005).
The reappraisal of arguments about the role of Macrozamia in the CQH has implications for our understanding of the prehistory of the CQH, but also has implications for high-level models which have used macrobotanical arguments from these sites to support the idea of wider continental-scale changes in Aboriginal societies in the mid-Holocene. Large-scale inter-group ceremonial activities quite possibly occurred in this, and other regions throughout Australian prehistory. However the Macrozamia remains from these sites can no longer be used as direct evidence of continent-wide mid-late Holocene economic intensification, or evidence of increasingly complex and competitive inter-group relations (including feasting, ceremonies and exchange) (Lourandos 1997).
Acknowledgements
This paper was written with the support of a Wenner-Gren Richard Carley Hunt Postdoctoral Fellowship, and while I was a Visiting Fellow in the School of Archaeology at the Australian National University. This research was part of a larger Doctoral research programme conducted at the Australian National University and made possible by an Australian Postgraduate Award. I sincerely thank Prof. John Beaton for his support of this research and access to unpublished site archives. Many thanks to Paul McInnes, Val Attenbrow, Peter Hiscock, Pat Faulkner and referees for their constructive comments on a prior version of this paper.
References
Asmussen, B., 2005, Dangerous Harvest revisited: taphonomy, methodology and intensification in the Central Queensland Highlands, Australia. Unpublished PhD thesis, Australian National University, Canberra.
Attenbrow, V., 2004, What's changing: population size or land-use patterns? The archaeology of Upper Mangrove Creek, Sydney Basin. Terra Australis 21.
Baird, A., 1939, A contribution to the life history of Macrozamia reidlei. Journal of the Royal Society of Western Australia 25:153-174.
Ballardie, R.T. and R.J. Whelan, 1986, Masting, seed dispersal and seed predation in the cycad Macrozamia communis. Oecologia 70:100-105.
Balme, J. and W. Beck, 2002, Starch and charcoal: useful measures of activity areas in archaeological shelters. Journal of Archaeological Research 29:157-166.
Bang, P. and P. Dahlstrom, 1972, Collins guide to animal tracks and signs of British and European mammals and birds. London: Collins Publishing.
Bauman, A.J. and H. Yokoyama, 1978, Seed coat carotenoids of the cycad genera Dioon, Encephalartos, Macrozamia and Zamia: evolutionary significance. Biological Systematics and Ecology 4:73-74.
Beaton, J., 1977, Dangerous Harvest: investigations in the late prehistoric occupation of upland south-east central Queensland. Unpublished PhD Thesis, Australian National University, Canberra.
Beaton, J., 1982, Fire and water: aspects of Aboriginal management of cycads. Archaeology in Oceania 17(1): 5158.
Beaton, J., 1991a, Wanderer's Cave and Rainbow Cave: two rockshelters in the Carnarvon Range of Central Queensland. Queensland Archaeological Research 8:3-33.
Beaton, J., 1991b, Cathedral Cave: a rockshelter in Carnarvon Gorge, Queensland. Queensland Archaeological Research 8:33-84.
Beaton, J., 1993, The improbable story of cycads as human food: archaeological and ethnographic evidence of more than 4,000 years. In D. Stevenson and K. Norstog, eds, Proceedings of CYCAD 90, the Second International Conference on Cycad Biology, pp: 371 (abstract). Brisbane: Palm and Cycad Societies of Australia.
Beaton, J., 1995, The transition on the coastal fringe of greater Australia. Antiquity 69:798-806.
Beck, W., 1985. Technology, Toxicity and Subsistence. A Study of Australian Aboriginal Plant Food Processing. Unpublished PhD Thesis, LaTrobe University, Melbourne.
Beck, W., 1989, The taphonomy of plants. In W. Beck, A. Clarke and L. Head, eds, Plants in Australian Archaeology. Tempus 1:31-49.
Beck, W., R. Fullagar and N. White, 1988, Archaeology from ethnography: the Aboriginal use of cycad as an example. In B. Meehan and R. Jones eds, Archaeology with Ethnography an Australian Perspective, pp. 137-147. Canberra: Australian National University Department of Prehistory, Research School of Pacific and Asian Studies.
Begg, R.J. and C.R. Dunlop, 1980, Security eating, and diet in the large rock-rat, Zygomys woodwardi (Rodentia:Muridae). Australian Wildlife Research 7:63-70.
Behrensmeyer, A.K., 1975, The taphonomy and palaeoecology of Plio-Pleistocene vertebrate assemblages East of Lake Rudolf, Kenya. Bulletin of the Museum of Comparative Zoology 145(10):473-578.
Bell, C.J.E., B.L. Finlayson and A.P. Kershaw, 1989, Pollen analysis and dynamics of a peat deposit in Carnarvon National Park, central Queensland. Australian Journal of Ecology 14:449-456.
Berndt, R.M., 1951, Kunapipi. Melbourne: Cheshire.
Bird, C.F.M. and D. Frankel, 1991a, Chronology and explanation in western Victoria and south-east South Australia. Archaeology in Oceania 26: 1-16.
Bird, C.F.M and D. Frankel, 1991b, Problems in constructing a prehistoric regional sequence: Holocene south-east Australia. World Archaeology 23(2):179-192.
Bird, C.F.M., D. Frankel and N.V. Van Waarden, 1997, New radiocarbon determinations from the Grampians-Gariewerd region, western Victoria. Archaeology in Oceania 33:31-36.
Bowdery, D., 2006, Phytolith analysis report on seven sediments from Kenniff Cave, Queensland. Unpublished report to Dr. Peter Hiscock.
Bradley, J.J., 2006, The social, economic and historical construction of cycad palms among the Yanyuwa. In B. David, B. Barker, and I. McNiven, eds, The Social Archaeology of Australian Indigenous Societies, pp. 161-181. Canberra: Aboriginal Studies Press.
Burbidge, A.H. and R.J. Whelan, 1982, Seed dispersal in a cycad, Macrozamia reidlei. Australian Journal of Ecology 7: 63-67.
Carter, T., 1923, Birds of the Broom Hill district. Emu 23:125-142.
Clarke, A., 1988, Archaeological and Ethnographic Interpretations of Plant Remains from Kakadu National Park, Northern Territory. In B. Meehan and R. Jones, eds., Archaeology with Ethnography: an Australian perspective, pp. 123-136. Canberra: Australian National University Research School of Pacific and Asian Studies.
Clarke, A., 1999, Preservation profiles: a case study from Anbangbang 1, Kakadu National Park, Northern Territory. In M. Mountain and D. Bowdery, eds, Taphonomy: The Analysis of Processes from Phytoliths to Megafauna, pp. 83-91. Canberra: Department of Archaeology and Natural History, Research School of Pacific and Asian Studies, Australian National University.
Colvill, M., 1995, Off the shelf- out of the museum. The retrieval of plant material from the Australian archaeological record. Unpublished Bachelor of Arts Honours Thesis, School of Archaeology, Australian National University, Canberra.
David, B. and T.P. Denham. 2006, Unpacking Australian prehistory. In B. David, B. Barker and I. McNiven, eds., The social archaeology of indigenous societies: essays on Aboriginal and Tortes Strait Islander history in honour of Harry Lourandos, pp. 52-71. Canberra: Aboriginal Studies Press.
Dennell, R.W., 1976, The economic importance of plant remains represented on archaeological sites. Journal of Archaeological Science 3:229-247.
Diehl, M.W., 1996, The intensity of Maize processing and production in upland Mogollon pit house Villages A.D. 200-1000. American Antiquity 61(1):102-115.
Edwards, D. and J. O'Connell, 1995, Broad-spectrum diets in arid Australia. Antiquity 69:769-783.
Farrera, M.A., 2004, Spatial distribution, population structure, and fecundity of Ceratozamia matusai Lundell (Zamiaceae) in El Triunfo Biosphere Reserve, Chipas, Mexico. The Botanical Review 70(2):299-311.
Flood, J. 1976. Man and ecology in the highlands of southeastern Australia: a case study. In: N. Peterson, ed., Tribes and Boundaries in Australia. Australian Institute of Aboriginal Studies, Canberra, pp. 30-49.
Frankel, D., 1988, Characterising change in prehistoric sequences: a view from Australia. Archaeology in Oceania 23:41-48.
Frankel, D., 1991a, Problems in constructing a prehistoric regional sequence: Holocene south-east Australia. World Archaeology 23(2): 179-192.
Frankel, D., 1991b, Remains to be seen: archaeological insights into Australian prehistory. South Melbourne: Longman Cheshire.
Frankel, D., 1993, Pleistocene chronological structures and explanations: a challenge. In M.A. Smith, M. Spriggs and B. Fankhauser, eds, Sahul in Review: Pleistocene Archaeology in Australia, New Guinea and Island Melanesia, pp. 24-33. Canberra: Australian National University, Research School of Pacific Studies, Department of Prehistory.
Frankel, D., 1995, The Australian transition: real and perceived boundaries. Antiquity 69:649-55.
Gardner, C.A. and H.W. Bennetts, 1956, The toxic plants of Western Australia. West Australia Newspapers, Ltd., Perth, Australia.
Godfrey, M. 1989, Shell midden chronology in southwestern Victoria: reflections of change in prehistoric population and subsistence. Archaeology in Oceania 24:65-69.
Goodale J.C., 1971, Tiwi Wives: a study of the women of Melville Island, north Australia. Seattle: University of Washington Press.
Hansen, J., 2001, Macroscopic plant remains from Mediterranean caves and rockshelters: avenues of interpretation. Geoarchaeology 16(4):401-432.
Harrison, S.P. and J. Dodson, 1993, Climates of Australia and New Guinea since 18.000 yr BP. In H.E. Wright, Jr., J.E. Kutzbach, T. Webb III, W.E Ruddiman, F. Alayne Street-Perrott and P.J. Bartlein, eds, Global Climates since the Last Glacial Maximum, pp. 265-92. Minneapolis: University of Minnesota Press.
Harvey, A. 1945, Food preservation in Australian tribes. Mankind 3(7): 191-192.
Hill, K., 1984, Cycads of Australia. Australian Plants 13(4): 3-23.
Hill, K. and R. Osborne, 2001, Cycads of Australia. East Roseville: Kangaroo Press.
Hiscock, P. 1981, Comments on the use of chipped stone artefacts as a measure of intensity of site usage. Australian Archaeology 13:30-34.
Hiscock, P. 2002, Pattern and context in the Holocene proliferation of backed artefacts in Australia. In R.G. Elston and S.L. Kuhn, eds, Thinking Small: Global Perspectives on Microlithicization, pp. 163-177. Archaeological Papers of the American Anthropological Association (AP3A) No. 12.
Hiscock, P., 2008, Archaeology of ancient Australia. London: Routledge.
Holdaway, S.J., P.C. Fanning, M. Jones, J. Shiner, D. Witter, and G. Nicholls 2002, Variability in the chronology of late Holcene Australian Aboriginal occupation on the arid margin of southeastern Australia. Journal of Archaeological Science 29:351-363.
Jennings, J., 2005, La Chichera y El Patron: Chicha and the energetics of feasting in the Prehistoric Andes. Archaeological Papers of the American Anthropological Association 14:241-259.
Jones, R., 1978, Calories and Bytes: towards a history of the Australian Islands, The 1978 Wentworth Lecture
http://www.aiatsis.gov.au/lbry/dig_prgm ... rth/a33458 3_a.pdf
Jones, R., 1980, Different strokes for different folks: sites, scale and strategy. In I. Johnston, ed., Holier than Thou, pp. 151-171. Canberra: Department of Prehistory, Research School of Pacific Studies, Australian National University.
Jones, M. 1987, Plant macrofossils. In P. Mellars, ed., Research priorities in archaeological science, pp. 11-13. London: Council for British Archaeology
Jones, D., 1993, Cycads of the World. Sydney: Reed Books. Karkanas, P. N., Kyparissi-Apostolika, O. Bar Yosef, and S. Weiner, 1999, Mineral assemblages in Theopetra, Greece: A framework for understanding diagenesis in a prehistoric cave. Journal of Archaeological Science 26:1171-1180.
Kelly, D. and V.L. Sork, 2002, Mast seeding in perennial plants: why, how, where? Annual Review of Ecological Systematics 33:427-447.
Kennedy, P., 1993, Macrozamia communis. Palms and Cycads 40:1-6.
Ladd, P.G., 1988, Floral analysis and archaeology: is Australia any different? Australian Aboriginal Studies 2: 2-18.
Lepovsky, D. and N. Lyons, 2003, Modelling ancient plant use on the northwest coast: towards an understanding mobility and sedentism. Journal of Archaeological Science 30:1357-1371.
Levitt, J., 1981, Plants and people: Aboriginal use of plants on Groote Eylandt. Canberra: Australian Institute of Aboriginal Studies.
Li, H., and Zhang, Z. 2007. Effects of mast seeding and rodent abundance on seed predation and dispersal by rodents in Prunus armeniaca (Rosaceae). Forest Ecology and Management 242:511-517.
Lilley, I., 2000, So near and yet so far: reflections on archaeology in Australia and Papua New Guinea, intensification and culture contact. Australian Archaeology 50:36-44.
Loaring, W.A., 1952, Birds eating the fleshy outer coat of zamia [sic] seeds. West Australian Naturalist 3:94.
Lourandos, H., 1980a, Change or stability? Hydraulics, hunter-gatherers and population in temperate Australia. World Archaeology 11:245-266.
Lourandos, H., 1980b, Forces of change: Aboriginal technology and population in south-western Victoria. Unpublished PhD thesis, University of Sydney.
Lourandos, H., 1983a, Intensification: a late Pleistocene-Holocene archaeological sequence from southwestern Victoria. Archaeology in Oceania 18:81-94.
Lourandos, H., 1983b, 10,000 years in the Tasmanian highlands. Australian Archaeology 16:39-47.
Lourandos, H., 1988, Palaeopolitics: resource intensification in Aboriginal Australia and Papua New Guinea. In T. Ingold, D. Riches and J. Woodburn, eds, Hunters and gatherers: history, evolution and social change, pp. 148-60. Oxford: Berg.
Lourandos, H., 1997, Continent of hunter-gatherers. New perspectives in Australian prehistory. Cambridge: Cambridge University Press.
Low, T. 1991, Wild Food Plants of Australia. Sydney: Angus and Robertson.
Lyman, R.L., 1997, Vertebrate Taphonomy. Cambridge: Cambridge University Press.
Margaritis, E. and M. Jones, 2006, Beyond cereals: crop processing and Vitis vinifera L. Ethnography, experiment and charred grape remains from Hellenistic Greece. Journal of Archaeological Science 33:784805.
Maiden, J.H., 1889, The useful native plants of Australia. Sydney: Technology Museum.
Maiden, J.H., 1890, Native food plants. New South Wales: Department of Agriculture.
McGlone, M.S., A.P. Kershaw, V. Markgraf, 1992, El Nino/Southern Oscillation climatic variability in Australasian and South American palaeoenvironmental records. In H.F. Diaz and V. Markgraf, eds, El Nino: historical and palaeoclimatic aspects of the Southern Oscillation, pp. 435-62. Cambridge: Cambridge University Press.
Meehan, B. and R. Jones, 1977, Preliminary comments on the preparation of Cycas Media by the Gidjingali of coastal Arnhem Land. Appendix 4 in J. Beaton, 1977, Dangerous Harvest: Investigations in the late prehistoric occupation of upland south-east central Queensland. Unpublished PhD Thesis, Australian National University, Canberra.
Miksicek, C.H., 1987, Formation processes of the archaeobotanical record. Advances in Archaeological Method and Theory 10:211-247.
Moore, RD., 1999, Ecology: a shrike for mobility. Nature 397:21-23.
Murphy, D.A., 1992, Plant taphonomy in rockshelters: a study of plant material in sandstone rockshelters near Coonabarabran, N.S.W. Unpublished MA Thesis,
Department of Archaeology and Palaeoanthropology, the University of New England.
Nelson, M., 1992, Shell midden deposits and the archaeobotanical record: a case study from the northwest coast. In J. Stein, ed., Deciphering a Shell Midden, pp. 239-259. California: Academic Press.
Norstog, K.J. and T.J. Nicholls, 1997, The biology of the cycads. New York: Cornell University Press.
Ornduff, R., 1990, Geographic variation in the reproductive behavior and size structure of the Australian cycad Macrozamia communis (Zamiaceae). American Journal of Botany 77(1):92-99.
Ornduff, R., 1991, Coning phenology of the cycad Macrozamia reidlei (Zamiaceae) over a five-year interval. Bulletin of the Torrey Botanical Club 118:6-11.
Pardoe, C., 1995, Riverine, biological and cultural evolution in southeastern Australia. Antiquity 69(265):696-713.
Parton, G.F., 1952, Encounter with a native cat. West Australian Naturalist 3:93.
Pearsall, D. 1989, Palaeoethnobotany: a handbook of procedures. New York: Academic Press.
Pennington, H.L. and S.A. Weber, 2004, Paleoethnobotany: modern research connecting ancient plants and ancient peoples. Critical Reviews in Plant Sciences 23(1): 13-20.
Potter, J.M., 1997, Communal ritual and faunal remains: an example from the Dolores Anasazi. Journal of Field Archaeology 24(3):353-364.
Renner, S.S., 2003, Plant animal interactions: a somewhat evolutionary approach. American Journal of Botany 90(2):330-332.
Ross, A. 2006 Comment on Ian Keen's Constraints on the Development of Enduring Inequalities in Late Holocene Australia. Current Anthropology 47(1):24-25.
Rossen, J., T. Dillehay and D. Ugent, 1996, Ancient cultigens or modern intrusions? Evaluating plant remains in an Andean Case Study. Journal of Archaeological Science 23:391-407.
Roth, W.E. 1901, Food: its search, capture and preparation. North Queensland Ethnography Bulletin 3:114-115. Brisbane: G. A Vaughne.
Sanchez-Tinoco, M. Y. and E.M. Engleman, 2004, Seed Coat Anatomy of Ceratozamia mexicana (Cycadales). The Botanical Review 70(1):24-38.
Sargent, O.H., 1928, Reaction between the birds and plants. Emu 27:185-92.
Schiffer, M., 1976, Behavioral archaeology. New York: Academic Press.
Schnurr, J.L., R.S. Ostfeld, and C.D. Canham, 2002, Direct and indirect effects of masting on rodent populations and tree seed survival. Oikos 96(3):402.
Sedgwick, E.H., 1952, Birds and zamia seeds. West Australian Naturalist 3:117.
Snow, E.L. and G.H. Walter, 2007, Large seeds, extinct vectors and contemporary ecology: testing dispersal in a locally distributed cycad, Macrozamia lucida (Cycadales). Australian Journal of Botany 55:592-600.
Spencer, W.B., 1914, Native Tribes of the Northern Territory. Macmillan and Co, London.
Spicer, R.A., 1991, Plant taphonomic processes. In P.A. Allison and D.E. Briggs, eds, Taphonomy: releasing the data locked in the fossil record, pp. 72-108. Topics in Geobiology 9: 72-108
Stranger, M.J. and R.H. Stranger, 1970, Grey butcher bird feeding on Macrozamia fruit. West Australian Naturalist 11:145.
Sullivan, M. 1977, Aboriginal Gatherings in South-East Queensland. Unpublished BA thesis, Australian National University, Canberra.
Tang, W., 1990, Reproduction in the cycad Zamia pumila in a fire-climax habitat; an eight-year study. Bulletin of the Torrey Botanical Club 117:368-374.
Thomson, D., 1938, The Australian native woman as food producer: catering in Arnhem Land--the preparation of cycad nuts. Illustrated London News (Oct 22) 1938: 730-731.
Thozet, A., 1878, Notes on some of the roots, tubers, bulbs and fruits used as vegetable food by the Aboriginals of North Queensland, Australia. In R. Smyth-Brough, ed., The Aborigines of Victoria with notes relating to the habits of the natives of other parts of Australia and Tasmania, pp. 227-233. Melbourne: Government Printer.
Triggs, B., 1996, Tracks, scats and other traces. A field guide to Australian mammals. South Melbourne: Oxford University Press.
Turner, F., 1893, The zamia palm and its relation to the disease known as rickets in cattle. Agricultural Gazette of New South Wales 4:158-161.
van der Pijl, L., 1957, The dispersal of plants by bats. Acta Botanica 6: 291-315.
Vorster, P., 1995, Aspects of the reproduction of cycads II. An annotated review of known information. In P. Vorster, ed., Proceedings of the third international conference on Cycad Biology, pp. 379-388. Stellenbosch: Cycad Society of South Africa.
Vovides, A.P., 1990, Spatial distribution, survival, and fecundity of Dioon edule (Zamiaceae) in tropical deciduous forest in Veracruz, Mexico, with notes on its habitat. American Journal of Botany 77(12):1532-1543.
Walthall, J.A., 1998, Rockshelters and hunter-gatherer adaptation to the Pleistocene/Holocene Transition. American Antiquity 63(2):223-238.
Watkinson, A.R. and J.C. Powell, 1997, The life history and population structure of Cycas armstrongii in monsoonal northern Australia. Oecologica 111(3):341-349.
Zhang, J., F.A. Drummond, M. Liebman, A. Hartke, 1997, Insect predation of seeds and plant population dynamics. Maine Agricultural and Forest Experiment Station Technical Bulletin 163:1-25.1
Zutter, C., 1999, Congruence or concordance in archaeobotany: Assessing micro and macro botanical data sets from Icelandic middens. Journal of Archaeological Science 26:833-844.
Visiting Fellow, School of Archaeology, The Australian National University, Canberra ACT 0200. email:
brit.asmussen@exemail.com.au" onclick="window.open(this.href);return false
Table 1. Quantification of Macrozamia moorei in the three
sites in the Central Queensland Highlands.
CathedralRainbow Wanderer's
Attributes Cave+Cave Cave
NISP 2748 113 1514
MNI *426 23 135
Approximate excavated volume 4.7 1.0 1.0
Approximate density NISP per 587.2113.01514.0
cubic meter
Approximate density MNI per 91.1 23 135
cubic meter
Estimated occupational 317.8300 85.50
sediment volume
Extrapolated NISP in entire 186593 33900129447
site
Extrapolated MNI in entire 289506900 11543
site
Earliest date of Macrozamia 2234 1650 4823
use, calBP
Average NISP per year over 83.5 20.5 26.8
whole site
Average MNI per year over13.0 4.2 2.4
whole site
+ Figures for Cathedral Cave exclude flood-deposited layers;
* MNI is based on highest sum of end completeness.
Table 2. NISP of Macrozamia specimens and ecological data at
Wanderer's and Cathedral Cave, CQH.All samples were charcoal
processed by conventional methods.
Site and Radiocarbon Laboratory NISP
excavation layer age (BP)Number
Wanderer 00-05 NA NA 151
Wanderer 05-10 820 [+ or -] 70 ANU 1539 543
Wanderer 10-15 ^ 1090 [+ or -] 60ANU 12013NA
Wanderer 15-20 2080 [+ or -] 80ANU 12016205
Wanderer 20-25 3340 [+ or -] 70ANU 12014248
Wanderer 25-30 4150 [+ or -] 70ANU 12015256
Wanderer 30-35 NA NA 106
Wanderer 35-40 4320 [+ or -] 80ANU 1522 5
Wanderer 40-45 NA NA 0
Wanderer 45-50 NA NA 0
Cathedral Level 1 1040 [+ or -] 190 ANU 113161434
Cathedral Level 2 *NA NA 43
Cathedral Level 3 2300 [+ or -] 70ANU 1761 1314
Cathedral Level 4 *NA NA 79
Site and % %%
excavation layer sarcotesta gnaw germinated
Wanderer 00-05 2.9 27.2 11
Wanderer 05-10 6.4 15.5 6.62
Wanderer 10-15 ^ NA NA NA
Wanderer 15-20 42.914.6 10.2
Wanderer 20-25 4.3 7.3 3.22
Wanderer 25-30 9.6 7.8 8.2
Wanderer 30-35 15.010.4 0
Wanderer 35-40 0.0 20.0 0
Wanderer 40-45 0 00
Wanderer 45-50 0 00
Cathedral Level 1 4.5 10.1 8.4
Cathedral Level 2 *6.2 4.7 0.0
Cathedral Level 3 2.6 3.7 15.8
Cathedral Level 4 *0.0 0.0 0.0
^ Assemblage unavailable to be analysed. * Layer containing
Macrozamia specimens that were deposited by flooding.
