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Turtles and molluscs appear at almost every site in relative high occurrence and abundance.
Future archaeozoological research should focus on standardizing quantification techniques, and providing fine-grained identification of turtle specimens to more accurately investigate shifting subsistence strategies, paleoenvironmental change, and seasonal occupation of sites throughout mainland Southeast Asia during the Pleistocene and Holocene.
Meta-analyses of archaeozoological data provide explanations of fine-grained questions using large databases Conolly et al. A classic example of a meta-analysis is the recognition that we only have evidence for Clovis-era hunter-gatherers in North America exploiting two of the 35 large mammal genera that became extinct during the late Pleistocene Grayson and Meltzer Aside from qualitative syntheses of Southeast Asian faunas Gorman b , little archaeozoological meta-analysis has occurred in this region. Identifying subsistence change at the Pleistocene-Holocene transition is a popular topic in archaeozoological endeavors Conolly et al.
This transition is significant during human prehistory because of worldwide climatic change that transpired, away from a glacial period into the climatic and environmental regime that we see today Straus et al. In mainland Southeast Asia, this transition is significant but few meta-analyses attempt to understand the inherent subsistence changes that may have emerged cf. Rabett In this paper I conduct a compositional overview of the late Pleistocene and Holocene archaeozoological record in Thailand and Peninsular Malaysia to establish the methodological and quantitative patterns found within this large faunal dataset.
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I then use nestedness and squared chord distance metrics to explore shifting faunal exploitation in Thai-Malay assemblages spanning the Pleistocene-Holocene transition. The primary goal of this analysis is to examine trends in forager subsistence activity in Thailand and Peninsular Malaysia, thus this sample is limited to a total of 28 sites within both regions that span the late Pleistocene to Holocene and have archaeozoological deposits that are primarily pre-agricultural in nature Figure 1 ; Table 1. I use the presence of ceramics as a coarse proxy for agriculture.
There are two important caveats to this.
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Location of sites discussed in text. Archaeological sites, locations, methods of faunal recording, available date ranges and key references for Thai-Malay sites. This table uses published radiocarbon dates, including mollusk dates Bulbeck , as such; the dates reported here should not be regarded as an absolute chronological range. Instead these dates act as a coarse date range. For each site, see cited literature for exact chronological details. First, many of the reported radiocarbon dates for these sites come from uncontrolled deposits not in stratigraphic association , or from material molluscs that have uncorrected dates which do not account for reservoir effects or diagenetic processes Bulbeck Second, while I am primarily interested in forager subsistence strategies, several of the sites span the Neolithic transition into the Protohistoric period.
Commonly included in these later Holocene assemblages are increased ceramic accumulations and adzes, both suggesting a sedentary agricultural adaptation. At Khao Toh Chong Rockshelter, these artifact classes increase rapidly in abundance after 7, years ago Van Vlack Thus, it is difficult to tell when faunal remains were deposited due to mobile hunter-gatherer groups or agriculturalists in nearby settlements.
I do not attempt to dichotomize faunal data to differentiate these groups although this would be a worthwhile and interesting endeavor in the future. Instead, I simply use all available data during the span of these sites and assume that hunter-gatherer groups are the primary accumulating agents. For this reason, I specifically exclude sites and faunal data from known settlement agricultural sites generally open sites in flood plain valleys.
Assemblages chosen for this analysis span the late Pleistocene through the Holocene, approximately 45, years ago to the present, in Thailand and Peninsular Malaysia Table 1 ; Figure 1. This provides the ability to understand stratigraphic and chronological sequences of faunal exploitation and subsistence activities over long temporal spans Anderson ; Anderson ; Anderson ; Straus ; Straus , but also may indicate that site type bias is occurring in this region Higham Aside from similar chronologies and site types, these sites also share characteristics in their lithic techno-complex assemblages.
Cave and rockshelter sites in Thailand and Peninsular Malaysia dating to the late Pleistocene and Holocene often contain deposits described as Hoabinhian Bronson and White ; Dunn ; Forestier et al. Recent research on the lithic assemblages from Tham Lod TL and Ban Rai BR in northwest Thailand show that paleoenvironmental processes drove considerable variability in the procurement strategies and technological adaptations of so-called Hoabinhian foragers Marwick Some archaeologists thus avoid the term Hoabinhian to describe general archaeological periods, ethnicities or economies and instead explicitly use the term for conveying information on a distinct artifact typology i.
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All 28 sites reviewed for this work date to the late Pleistocene or Holocene and have some Hoabinhian type material within their deposits. I also chose the 28 sites in this review based on availability of published or gray literature. Excluded sites include Buang Baeb and Pak Om Rockshelters, which both have large faunal assemblages, but are recorded in Thai language reports only Reynolds ; Shoocongdej a ; Pers. Cholawit Thongcharoenchaikit The 28 sites included here represent some of the best evidence for prehistoric hunter-gatherer occupations in mainland Southeast Asia see previous syntheses by Adi ; Shoocongdej a , and are historically used in other reviews of archaeological material from this region Marwick ; Reynolds ; Shoocongdej a ; Shoocongdej There is a considerable literature on the benefits and potential bias derived from using NISP versus MNI quantification in vertebrate faunal studies see Lyman for overview.
Since this is a meta-analysis, I am limited by counts of taxonomic abundance provided in the published or gray literature. Separate quantification issues occur when counting the taxonomic abundance of invertebrate faunas Claassen I do not quantify invertebrate faunas from these 28 sites, due to dissimilar recording techniques, and thus my investigation of these issues is limited. Additionally, there is inconsistency on screen size use or reporting sieving methods in the archaeozoological literature from this region. While some excavations used small-sized screens e.
It is possible that several animal types, including fish, birds, small mammals and small reptiles, are absent from this faunal dataset due to screen size biases see Nagaoka ; Nagaoka ; Quitmyer for examples. Non-human accumulation of faunal material represents another potential bias in this dataset. I am unaware of any robust taphonomic analyses in mainland Southeast Asian archaeozoological contexts see Forestier et al. Rodents, raptors, mammalian carnivores and some reptilian carnivores could potentially accumulate faunal remains in cave and rockshelter sites, independent of human activity Blumenschine ; Faith ; Lyman ; Schmitt and Lupo , but analyses focused on this topic in Southeast Asia are lacking.
Furthermore, some fauna from these 28 sites likely accumulated through natural death events, specifically bat species Higham ; Rabett et al. There is a tendency in Southeast Asian archaeozoology to assume that all fauna in cave and rockshelter sites accumulated through human activity.
Without re-investigation of faunal assemblages focused on taphonomic processes, it is difficult to determine which taxa from these sites truly reflect human subsistence strategies. I expect that the signal of human subsistence activity should be clearly represented in this analysis based on large abundances of specific prey types that were consumed throughout the late Pleistocene and Holocene in this region.
As such, I do not discard any faunal data in this research, even though it was potentially accumulated through non-anthropogenic means. Seasonal bias in human occupation of cave and rockshelter sites is an additional potential limitation in this analysis. Interestingly, this question remains relatively unanswered during this temporal period.
Based upon the association of wet and dry season resources freshwater gastropods and bivalves in the same stratigraphic contexts at Spirit Cave SC in northwest Thailand, Gorman a ; b ; argued that this represented year-round site occupation by hunter-gatherer groups. Yet, more recent research at Lang Kamnan Cave LK in central Thailand suggests that only a wet season occupation is represented in the assemblage based upon the presence of botanical and land snail remains Shoocongdej b ; Shoocongdej At Lang Rongrien Rockshelter LR in southern Thailand, faunal data suggests that the site was intermittently occupied during the late Pleistocene and Holocene, in both the wet and dry seasons Anderson ; Anderson ; Mudar and Anderson Clearly, additional research is required before identifying site-specific seasonal occupations in mainland Southeast Asia.
Typically, only the qualitative association of wet and dry season resources in cave and rockshelter sites is used to determine seasonal occupation events. These resources are useful as paleoecological proxies for seasonality, but depositional resolution of cultural activity remains low. Archaeozoologists in mainland Southeast Asia must develop new techniques to identify what is likely low intensity, short occupation, seasonal events in sites. Micromorphology of sediments to examine micro-stratigraphic layers and geochemical analyses of fauna may be useful techniques in this regard.
Finally, some prehistoric sites in mainland Southeast Asia may represent specialist activities. Tham Phaa Can does not have an identified and published faunal assemblage, only a descriptive account remains Higham Still, this account suggests that large artiodactyls and turtles or tortoises dominate the assemblage. Similarly, Nong Nor is dominated by approximately 5. There is little evidence for specialist subsistence strategies at other sites in Thailand and Peninsular Malaysia.
Since the faunal assemblage is not quantified from Tham Phaa Can, any potential bias is limited from this site and it is included here. Nong Nor is excluded due to the considerable specialist activity represented in the assemblage. Although some bias is present in Thailand and Peninsular Malaysian faunas, specifically non-human accumulation of remains and seasonal occupation events, the nuances of these processes remain unknown.
Thus, I include all sites and faunal data except for Nong Nor to investigate shifts in methodological techniques, faunal composition, and changing subsistence strategies during the late Pleistocene and Holocene. First, I aggregate all faunal publications by decade to explore how archaeozoological research emerged during the twentieth and twenty-first centuries. Decadal divisions of quantitative units employed by Southeast Asian archaeozoologists provide insight into the shifting methodological techniques used in this same period. Thai-Malay faunal research has its own discrete temporal and methodological qualities that are expressed by these analyses.
By examining Thai sites and Peninsular Malay sites alone, differences and similarities between geographic regions, dominant faunas and total abundances appear. All numerical and taxonomic data employed in this research is available as supplemental information in the University of New Mexico digital electronic repository, including source code for analyses conducted in R 3. I use nestedness analysis to explore change in composition of these faunas through time. Nestedness uses a presence-absence matrix to establish whether faunas within a set are subsets of each other Atmar and Patterson ; Jones ; Jones ; Lyman ; Patterson ; Staniczenko, Kopp and Allesina As an ecological metric, nestedness is important because it allows inference into how random or non-random a faunal assemblage is, and is thus useful in studies of culturally accumulated remains Kougioumoutzis, Simaiakis and Tiniakou ; Simaiakis and Strona Originally created to deal with species level extinctions, nestedness provides a quantitative means to test if the diversity of species is broadening or narrowing through time, or across space, when coupled with analysis of changing species richness NTAXA Atmar and Patterson ; Patterson By quantifying the degree to which species enter and leave a population e.
A benefit of nestedness analyses in archaeozoological research is that quantitative units in the original literature can easily be transformed into presence-absence matrices, even to the level of basic description. While methods of quantifying nestedness have changed over time Atmar and Patterson ; Patterson , the basic principles remain.
A perfectly nested assemblage occurs when successive populations have a perfect subset number of species in relation to the population prior Atmar and Patterson ; Jones ; Jones ; Jones Using the open-source software NeD Strona et al. Here, I use squared chord distance analysis to understand similarity in faunas through time. Squared chord distance analyses otherwise known as dissimilarity coefficients originated in the ecological literature describing the similarity between biological assemblages Gavin et al.
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By quantifying faunal classes as a weighted percentage, an emphasis on the relative proportion of each taxonomic class or species relative to all others but not to their absolute values gives squared chord distance its uniqueness as a broadly applicable measurement Ludwig and Reynolds Regardless of how fauna is reported, given that the same reporting technique is used for the complete assemblage or compared contexts , squared chord distance is applicable.
Here, shifts in taxonomic composition are not correlated with taphonomic processes when quantified using chord distance Faith In Thai-Malay sites, fauna is recorded differentially as NISP or MNI counts, and squared chord distance provides a proxy measure to understand similarity in taxonomic composition through time between sites with contrastingly recorded quantitative units. The Thai-Malay archaeofaunal record is abundant and diverse. At least distinct taxonomic classifications are represented in Thailand and Peninsular Malaysian archaeological sites with a total NISP of 29, Tables 1 and 2 ; Appendix 1 Table A , but this number is only a minimum because of the number of publications in Thai or Malay that I am unable to access.
As a whole, molluscs bivalves and gastropods appear most frequently in sites followed closely by sambar deer C. Due to scattered recording techniques applied to molluscan assemblages, their analytical value in this study is low and they are not explored in greater detail Table 2. Thailand and Peninsular Malaysia have a comparable division of faunal publications using different recording techniques Table 3 , but their use through time has shifted. Methodological recording of molluscan fauna from Thai-Malay sites.
Unknown indicates that mollusc data could not be identified in the site literature. Present indicates that a passing note of molluscan presence is provided for the site, but no absolute counts. Raw count is molluscan data that is aggregated into broad categories, for example shellfish or mollusc, and recorded in the literature. Total count of Thailand and Peninsular Malaysia sites with differential faunal recording methodologies. Decadal division of faunal publications using differential methodological recording techniques during the s—s.
A full range of size classifications are also represented in this collection, from large sized elephants to small sized mice, rats, and bats. Clearly, select species and habitats were exploited more intensely than others throughout this region. Large abundances of turtles, tortoises, deer and pigs indicate that both aquatic and terrestrial species were consumed.
Testudines clearly dominate the fauna with over 14, identified specimens, with the next closest classes being monitor lizards Varanidae and deer Cervidae , but this regional trend is primarily driven by the dominance of Testudines from Moh Khiew Cave II MKII. Furthermore, when Testudines are removed primates, deer and monitor lizards are more evenly distributed and all share relatively high NISP values Figure 5. Peninsular Malaysian sites show a slightly different trend. Here, NISP values are substantially lower than Thai sites and thus the spread between sites and classes is considerably less.
A surprising transition in the Peninsular Malaysian data is the lower abundance of deer compared to other classes. Deer appear in less abundance than primates, birds and fish in this region. The significance of this change is discussed below. Nestedness values suggest that during the late Pleistocene and Holocene, hunter-gatherer subsistence strategies were variable, but clearly influenced by archaeozoological investigator bias Table 4.
Only sites with NISP recording have significantly nested faunas. At SC a nested fauna appears without being identified as NISP values, but this is potentially due to the high degree of documentation provided by both Gorman b and Higham MKI is also nested, but less strongly, only the T metric is significant.