Ecosystem turnover in palaeoecological records: The ...

Ecosystem turnover in palaeoecological records: The sensitivity of pollen and phytolith proxies to detecting vegetation change in southwestern AmazoniaAuthors: Heather Plumptona*, Bronwen Whitneyb, Francis MayleaAuthor affiliations:aDepartment of Geography and Environmental Science, University of Reading, UKbDepartment of Geography and Environmental Sciences, Northumbria University, UK*corresponding author. Email address: hjplumpton@AbstractIdentification of ecosystem turnover in the palaeo-vegetation record is important for understanding the resilience of ecosystems to past environmental change. There is uncertainty over the ability of different types of palaeo-vegetation proxy to detect ecosystem turnover. The aim of this paper is to compare the sensitivity of two palaeo-vegetation proxies - pollen and phytoliths - to changes within and between three key tropical South American ecosystems: evergreen forest, dry forest and savannah. A quantitative approach is used to assess the sensitivity of these two proxies to vegetation changes, based on the variability of proxy assemblages from 1-hectare ecological plots in ecotonal south west Amazonia. This modern dataset of proxy variability within evergreen forest, dry forest and savannah plots is then used to define thresholds for proxy variability which differentiate floristic changes within an ecosystem from ecosystem turnover. These thresholds are applied to two palaeo-vegetation records from NE Bolivia. Our results show that pollen is more sensitive than phytoliths to changes within evergreen forest, but phytoliths are more sensitive than pollen to changes within dry forest. Both proxies were equally sensitive to changes within savannas. These are important considerations for palaeoecologists selecting proxies for the study of ecosystem turnover in the palaeo-record. Application of the thresholds to the palaeo-record demonstrated the utility of this quantitative approach for assessing the magnitude of vegetation change in the palaeo-record. This quantitative approach is therefore a useful tool to improve the identification of ecosystem turnover in the palaeo-record.KeywordsEcosystem turnover, variability, thresholds, pollen, phytoliths, south-west Amazonia, evergreen forest, dry forest, savannah, palaeoecology1. IntroductionEcosystem turnover is the process whereby one ecosystem is replaced by another, either in space or time ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1146/annurev.es.04.110173.000245","ISSN":"0066-4162","author":[{"dropping-particle":"","family":"Holling","given":"C S","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Annual Review of Ecology and Systematics","id":"ITEM-1","issue":"1","issued":{"date-parts":[["1973","11","28"]]},"page":"1-23","publisher":" Annual Reviews 4139 El Camino Way, P.O. Box 10139, Palo Alto, CA 94303-0139, USA ","title":"Resilience and Stability of Ecological Systems","type":"article-journal","volume":"4"},"uris":[""]}],"mendeley":{"formattedCitation":"(Holling, 1973)","plainTextFormattedCitation":"(Holling, 1973)","previouslyFormattedCitation":"(Holling, 1973)"},"properties":{"noteIndex":0},"schema":""}(Holling, 1973). For example, under drier climatic conditions a tropical rainforest may be replaced by savannah. The process of ecosystem turnover is a key area of ecological research for understanding the resilience of ecosystems to climatic change and for identifying tipping points between alternative states, such as humid forest and savannah ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1038/nature08227","ISSN":"0028-0836","abstract":"Early-warning signals for critical transitions","author":[{"dropping-particle":"","family":"Scheffer","given":"Marten","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Bascompte","given":"Jordi","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Brock","given":"William A.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Brovkin","given":"Victor","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Carpenter","given":"Stephen R.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Dakos","given":"Vasilis","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Held","given":"Hermann","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Nes","given":"Egbert H.","non-dropping-particle":"van","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Rietkerk","given":"Max","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Sugihara","given":"George","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Nature","id":"ITEM-1","issue":"7260","issued":{"date-parts":[["2009","9","3"]]},"page":"53-59","publisher":"Nature Publishing Group","title":"Early-warning signals for critical transitions","type":"article-journal","volume":"461"},"uris":[""]}],"mendeley":{"formattedCitation":"(Scheffer et al., 2009)","plainTextFormattedCitation":"(Scheffer et al., 2009)","previouslyFormattedCitation":"(Scheffer et al., 2009)"},"properties":{"noteIndex":0},"schema":""}(Scheffer et al., 2009). This area of research has gained particular attention due to the proposal of planetary boundaries beyond which the earth system will be outside the safe operating space for humanity ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1038/461472a","ISSN":"0028-0836","author":[{"dropping-particle":"","family":"Rockstr?m","given":"Johan","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Steffen","given":"Will","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Noone","given":"Kevin","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Persson","given":"?sa","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Chapin","given":"F. 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Palaeoecology has an important contribution to make to this area of research through the identification of ecosystem turnover and contributory environmental factors in the palaeo-record ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1111/1365-2745.12195","ISSN":"00220477","author":[{"dropping-particle":"","family":"Seddon","given":"Alistair W. R.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mackay","given":"Anson W.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Baker","given":"Ambroise G.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Birks","given":"Hilary John B. 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R.; Mackay, Anson W.; Baker, Ambroise G.; Birks, H. John B.; Breman, Elinor; Buck, Caitlin E.; Ellis, Erle C.; Froyd, Cynthia A.; Gill, Jacquelyn L.; Gillson, Lindsey; Johnson, Edward A.; Jones, Vivienne J.; Juggins, Stephen; Macias-Fauria, Marc; Mills, Keely; Morris, Jesse L.; Nogués-Bravo, David; Punyasena, Surangi W.; Roland, Thomas P.; Tanentzap, Andrew J.; Willis, Kathy J.; Aberhan, Martin; van Asperen, Eline N.; Austin, William E. N.; Battarbee, Rick W.; Bhagwat, Shonil; Belanger, Christina L.; Bennett, Keith D.; Birks, Hilary H.; Bronk Ramsey, Christopher; Brooks, Stephen J.; de Bruyn, Mark; Butler, Paul G.; Chambers, Frank M.; Clarke, Stewart J.; Davies, Althea L.; Dearing, John A.; Ezard, Thomas H. G.; Feurdean, Angelica; Flower, Roger J.; Gell, Peter; Hausmann, Sonja; Hogan, Erika J.; Hopkins, Melanie J.; Jeffers, Elizabeth S.; Korhola, Atte A.; Marchant, Robert; Kiefer, Thorsten; Lamentowicz, Mariusz; Larocque-Tobler, Isabelle; López-Merino, Lourdes; Liow, Lee H.; McGowan, Suzanne; Miller, Joshua H.; Montoya, Encarni; Morton, Oliver; Nogué, Sandra; Onoufriou, Chloe; Boush, Lisa P.; Rodriguez-Sanchez, Francisco; Rose, Neil L.; Sayer, Carl D.; Shaw, Helen E.; Payne, Richard; Simpson, Gavin; Sohar, Kadri; Whitehouse, Nicki J.; Williams, John W.; Witkowski, Andrzej)\n\nInteresting paper on the application of palaeoecology to ecological questions","page":"256-267","title":"Looking forward through the past: identification of 50 priority research questions in palaeoecology","type":"article-journal","volume":"102"},"uris":[""]}],"mendeley":{"formattedCitation":"(Seddon et al., 2014)","manualFormatting":"(see questions 16 and 17 in Seddon et al., 2014)","plainTextFormattedCitation":"(Seddon et al., 2014)","previouslyFormattedCitation":"(Seddon et al., 2014)"},"properties":{"noteIndex":0},"schema":""}(see questions 16 and 17 in Seddon et al., 2014). For example, the latest IPCC report on 1.5°C of warming uses three palaeo time periods (Holocene Thermal Maximum, Last Interglacial, Mid Pliocene Warm Period) as analogues of warmer periods when feedbacks in the earth system led to ecosystem turnover including retreat of tropical forests and expansion of savannah ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.5194/cp-12-1519-2016","ISSN":"1814-9332","abstract":"Abstract. The mid-Piacenzian is known as a period of relative warmth when compared to the present day. A comprehensive understanding of conditions during the Piacenzian serves as both a conceptual model and a source for boundary conditions as well as means of verification of global climate model experiments. In this paper we present the PRISM4 reconstruction, a paleoenvironmental reconstruction of the mid-Piacenzian (?～?3Ma) containing data for paleogeography, land and sea ice, sea-surface temperature, vegetation, soils, and lakes. Our retrodicted paleogeography takes into account glacial isostatic adjustments and changes in dynamic topography. Soils and lakes, both significant as land surface features, are introduced to the PRISM reconstruction for the first time. Sea-surface temperature and vegetation reconstructions are unchanged but now have confidence assessments. The PRISM4 reconstruction is being used as boundary condition data for the Pliocene Model Intercomparison Project Phase?2 (PlioMIP2) experiments.","author":[{"dropping-particle":"","family":"Dowsett","given":"Harry","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Dolan","given":"Aisling","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Rowley","given":"David","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Moucha","given":"Robert","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Forte","given":"Alessandro M.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mitrovica","given":"Jerry X.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Pound","given":"Matthew","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Salzmann","given":"Ulrich","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Robinson","given":"Marci","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Chandler","given":"Mark","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Foley","given":"Kevin","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Haywood","given":"Alan","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Climate of the Past","id":"ITEM-1","issue":"7","issued":{"date-parts":[["2016","7","13"]]},"page":"1519-1538","title":"The PRISM4 (mid-Piacenzian) paleoenvironmental reconstruction","type":"article-journal","volume":"12"},"uris":[""]}],"mendeley":{"formattedCitation":"(Dowsett et al., 2016)","plainTextFormattedCitation":"(Dowsett et al., 2016)","previouslyFormattedCitation":"(Dowsett et al., 2016)"},"properties":{"noteIndex":0},"schema":""}(Dowsett et al., 2016). The conventional approach to identifying ecosystem turnover in the palaeo-record is through qualitative interpretation of pollen and other palaeo-vegetation and -environmental proxies. A more quantitative approach is to use the variability in pollen records as an indicator of ecosystem turnover; the more variable the record, the more likely vegetation change is taking place ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/0033-5894(85)90074-2","ISSN":"0033-5894","abstract":"Dissimilarity coefficients measure the difference between multivariate samples and provide a quantitative aid to the identification of modern analogs for fossil pollen samples. How eight coefficients responded to differences among modern pollen samples from eastern North America was tested. These coefficients represent three different classes: (1) unweighted coefficients that are most strongly influenced by large-valued pollen types, (2) equal-weight coefficients that weight all pollen types equally but can be too sensitive to variations among rare types, and (3) signal-to-noise coefficients that are intermediate in their weighting of pollen types. The studies with modern pollen allowed definition of critical values for each coefficient, which, when not exceeded, indicate that two pollen samples originate from the same vegetation region. Dissimilarity coefficients were used to compare modern and fossil pollen samples, and modern samples so similar to fossil samples were found that most of three late Quaternary pollen diagrams could be “reconstructed” by substituting modern samples for fossil samples. When the coefficients indicated that the fossil spectra had no modern analogs, then the reconstructed diagrams did not match all aspects of the originals. No modern analogs existed for samples from before 9300 yr B.P. at Kirchner Marsh, Minnesota, and from before 11,000 yr B.P. at Wintergreen Lake, Michigan, but modern analogs existed for almost all Holocene samples from these two sites and Brandreth Bog, New York.","author":[{"dropping-particle":"","family":"Overpeck","given":"J. T.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Webb","given":"T.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Prentice","given":"I. C.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Quaternary Research","id":"ITEM-1","issue":"01","issued":{"date-parts":[["1985","1","20"]]},"page":"87-108","publisher":"Cambridge University Press","title":"Quantitative Interpretation of Fossil Pollen Spectra: Dissimilarity Coefficients and the Method of Modern Analogs","type":"article-journal","volume":"23"},"uris":[""]}],"mendeley":{"formattedCitation":"(Overpeck et al., 1985)","plainTextFormattedCitation":"(Overpeck et al., 1985)","previouslyFormattedCitation":"(Overpeck et al., 1985)"},"properties":{"noteIndex":0},"schema":""}(Overpeck et al., 1985). This approach has been used in temperate and tropical regions to identify periods of high vegetation turnover in the pollen record and their relation to climatic changes ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/J.QUASCIREV.2009.04.005","ISSN":"0277-3791","abstract":"We use a series of tests to evaluate two competing hypotheses about the association of climate and vegetation trends in the northeastern United States over the past 15kyrs. First, that abrupt climate changes on the scale of centuries had little influence on long-term vegetation trends, and second, that abrupt climate changes interacted with slower climate trends to determine the regional sequence of vegetation phases. Our results support the second. Large dissimilarity between temporally close fossil pollen samples indicates large vegetation changes within 500 years across >4° of latitude at ca 13.25–12.75, 12.0–11.5, 10.5, 8.25, and 5.25ka. The evidence of vegetation change coincides with independent isotopic and sedimentary indicators of rapid shifts in temperature and moisture balance. In several cases, abrupt changes reversed long-term vegetation trends, such as when spruce (Picea) and pine (Pinus) pollen percentages rapidly declined to the north and increased to the south at ca 13.25–12.75 and 8.25ka respectively. Abrupt events accelerated other long-term trends, such as a regional increase in beech (Fagus) pollen percentages at 8.5–8.0ka. The regional hemlock (Tsuga) decline at ca 5.25ka is unique among the abrupt events, and may have been induced by high climatic variability (i.e., repeated severe droughts from 5.7 to 2.0ka); autoregressive ecological and evolutionary processes could have maintained low hemlock abundance until ca 2.0ka. Delayed increases in chestnut (Castanea) pollen abundance after 5.8 and 2.5ka also illustrate the potential for multi-century climate variability to influence species' recruitment as well as mortality. Future climate changes will probably also rapidly initiate persistent vegetation change, particularly by acting as broad, regional-scale disturbances.","author":[{"dropping-particle":"","family":"Shuman","given":"Bryan N.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Newby","given":"Paige","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Quaternary Science Reviews","id":"ITEM-1","issue":"17-18","issued":{"date-parts":[["2009","8","1"]]},"page":"1693-1709","publisher":"Pergamon","title":"Abrupt climate change as an important agent of ecological change in the Northeast U.S. throughout the past 15,000 years","type":"article-journal","volume":"28"},"uris":[""]},{"id":"ITEM-2","itemData":{"DOI":"10.1177/0959683614556383","ISSN":"0959-6836","abstract":"A long-standing question in palaeoecology has been to determine the importance of climate driving vegetation change since the last deglaciation. Here, we investigate the local-to-regional dynamics of vegetation change during the Lateglacial and the Holocene in Northern Europe. We extracted sites from the European Pollen Database and used the squared-chord distance (SCD) dissimilarity metric to identify time periods of high pollen assemblage turnover representing periods of abrupt vegetation change. In addition, a set of generalized additive mixed models were applied to investigate the underlying dynamics of two periods of higher rates of turnover: the Younger Dryas–early Holocene transition (YD-EH; 11.6–9.0?kyr) and early–middle Holocene (EMH; 9.0–6.0?kyr). Results revealed a high frequency of turnover events between 12.75–11.5, 10.75–11, 10.25–10, 7.75–7.25, 3.25–3.0 and 1.75–.25?kyr. Furthermore, there was a strong linear relationship between pollen assemblage turnover and large directional temperature ...","author":[{"dropping-particle":"","family":"Seddon","given":"Alistair WR","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Macias-Fauria","given":"Marc","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Willis","given":"Kathy J","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"The Holocene","id":"ITEM-2","issue":"1","issued":{"date-parts":[["2015","1","9"]]},"page":"25-36","publisher":"SAGE PublicationsSage UK: London, England","title":"Climate and abrupt vegetation change in Northern Europe since the last deglaciation","type":"article-journal","volume":"25"},"uris":[""]},{"id":"ITEM-3","itemData":{"DOI":"10.1007/978-90-481-2672-9_12","abstract":"An analysis of rates of ecological change (RoC) from thirteen pollen records from tropical South America is presented here. The analysis aims to identify the periods of fastest change since the last glacial maximum (LGM) and possible driving mechanisms. Despite rapid cooling periods, region-wide profound droughts, fire and human disturbances, RoC analysis showed that the speed of these climate changes never exceed the species response capabilities. Our results legitimize concerns regarding the resilience of species to accommodate future change and emphasize the urgency for integrative environmental measures.","author":[{"dropping-particle":"","family":"Urrego","given":"Dunia H","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Bush","given":"Mark B","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Silman","given":"Miles R","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Correa-Metrio","given":"Alexander","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Ledru","given":"Marie-Pierre","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Paduano","given":"Gina","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Valencia","given":"Bryan G","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Past climate variability in South America and surrounding regions","id":"ITEM-3","issued":{"date-parts":[["2009"]]},"page":"283-300","title":"Millennial-Scale Ecological Changes in Tropical South America Since the Last Glacial Maximum","type":"chapter"},"uris":[""]},{"id":"ITEM-4","itemData":{"author":[{"dropping-particle":"","family":"Bush","given":"Mark B.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Silman","given":"Miles R.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Urrego","given":"Dunia H.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Science","id":"ITEM-4","issued":{"date-parts":[["2004"]]},"note":"Bush et al 2004 paper - rates of change during glacial-Holocene transition","page":"827-829","title":"48,000 years of Climate and Forest Change in a Biodiversity Hot Spot","type":"webpage"},"uris":[""]}],"mendeley":{"formattedCitation":"(Bush et al., 2004; Seddon et al., 2015; Shuman and Newby, 2009; Urrego et al., 2009)","plainTextFormattedCitation":"(Bush et al., 2004; Seddon et al., 2015; Shuman and Newby, 2009; Urrego et al., 2009)","previouslyFormattedCitation":"(Bush et al., 2004; Seddon et al., 2015; Shuman and Newby, 2009; Urrego et al., 2009)"},"properties":{"noteIndex":0},"schema":""}(Bush et al., 2004; Seddon et al., 2015; Shuman and Newby, 2009; Urrego et al., 2009). Variation in the pollen record is measured through calculation of multivariate distance between pollen assemblages through time. Multivariate distance takes both the number and abundance of pollen taxa into account and gives a measure of the “difference” between two pollen assemblages ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1007/978-94-007-2745-8","ISBN":"9789400727441","ISSN":"00134651","abstract":"First book covering full range of modern data-analytical and statistical techniques used in paleolimnology and paleoecology Features exploratory data analysis, error estimation, clustering, ordination and modern statistical learning techniques Includes case studies on human impact, lake development and climate change ? Numerical and statistical methods have rapidly become part of a palaeolimnologist’s tool-kit. They are used to explore and summarise complex data, reconstruct past environmental variables from fossil assemblages, and test competing hypotheses about the causes of observed changes in lake biota through history. This book brings together a wide array of numerical and statistical techniques currently available for use in palaeolimnology and other branches of palaeoecology.","author":[{"dropping-particle":"","family":"Legendre","given":"Pierre","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Birks","given":"H John B","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Tracking Environmental Change using Lakes Sediment: Volume 5 Data Handling and Numerical Techniques","editor":[{"dropping-particle":"","family":"Birks","given":"H John B","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Lotter","given":"Andre F","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Juggins","given":"Steve","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Smol","given":"John P","non-dropping-particle":"","parse-names":false,"suffix":""}],"id":"ITEM-1","issued":{"date-parts":[["2012"]]},"note":"Section on distances methods","page":"201-248","publisher":"Springer","title":"Chapter 8: From Classical to Canonical Ordination","type":"chapter"},"uris":[""]},{"id":"ITEM-2","itemData":{"DOI":"10.1007/s004420100716","ISSN":"0029-8549","author":[{"dropping-particle":"","family":"Legendre","given":"Pierre","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Gallagher","given":"Eugene D.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Oecologia","id":"ITEM-2","issue":"2","issued":{"date-parts":[["2001","10","1"]]},"page":"271-280","publisher":"Springer Berlin Heidelberg","title":"Ecologically meaningful transformations for ordination of species data","type":"article-journal","volume":"129"},"uris":[""]}],"mendeley":{"formattedCitation":"(Legendre and Birks, 2012; Legendre and Gallagher, 2001)","plainTextFormattedCitation":"(Legendre and Birks, 2012; Legendre and Gallagher, 2001)","previouslyFormattedCitation":"(Legendre and Birks, 2012; Legendre and Gallagher, 2001)"},"properties":{"noteIndex":0},"schema":""}(Legendre and Birks, 2012; Legendre and Gallagher, 2001). A large distance indicates a large difference between two pollen assemblages and hence a difference in the parent vegetation ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/0033-5894(85)90074-2","ISSN":"0033-5894","abstract":"Dissimilarity coefficients measure the difference between multivariate samples and provide a quantitative aid to the identification of modern analogs for fossil pollen samples. How eight coefficients responded to differences among modern pollen samples from eastern North America was tested. These coefficients represent three different classes: (1) unweighted coefficients that are most strongly influenced by large-valued pollen types, (2) equal-weight coefficients that weight all pollen types equally but can be too sensitive to variations among rare types, and (3) signal-to-noise coefficients that are intermediate in their weighting of pollen types. The studies with modern pollen allowed definition of critical values for each coefficient, which, when not exceeded, indicate that two pollen samples originate from the same vegetation region. Dissimilarity coefficients were used to compare modern and fossil pollen samples, and modern samples so similar to fossil samples were found that most of three late Quaternary pollen diagrams could be “reconstructed” by substituting modern samples for fossil samples. When the coefficients indicated that the fossil spectra had no modern analogs, then the reconstructed diagrams did not match all aspects of the originals. No modern analogs existed for samples from before 9300 yr B.P. at Kirchner Marsh, Minnesota, and from before 11,000 yr B.P. at Wintergreen Lake, Michigan, but modern analogs existed for almost all Holocene samples from these two sites and Brandreth Bog, New York.","author":[{"dropping-particle":"","family":"Overpeck","given":"J. T.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Webb","given":"T.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Prentice","given":"I. C.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Quaternary Research","id":"ITEM-1","issue":"01","issued":{"date-parts":[["1985","1","20"]]},"page":"87-108","publisher":"Cambridge University Press","title":"Quantitative Interpretation of Fossil Pollen Spectra: Dissimilarity Coefficients and the Method of Modern Analogs","type":"article-journal","volume":"23"},"uris":[""]},{"id":"ITEM-2","itemData":{"DOI":"10.1007/BF00193530","ISSN":"0930-7575","author":[{"dropping-particle":"","family":"Grimm","given":"Eric C","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Jacobson","given":"George L","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Climate Dynamics","id":"ITEM-2","issue":"3-4","issued":{"date-parts":[["1992","1"]]},"page":"179-184","publisher":"Springer-Verlag","title":"Fossil-pollen evidence for abrupt climate changes during the past 18 000 years in eastern North America","type":"article-journal","volume":"6"},"uris":[""]},{"id":"ITEM-3","itemData":{"DOI":"10.1177/0959683614556383","ISSN":"0959-6836","abstract":"A long-standing question in palaeoecology has been to determine the importance of climate driving vegetation change since the last deglaciation. Here, we investigate the local-to-regional dynamics of vegetation change during the Lateglacial and the Holocene in Northern Europe. We extracted sites from the European Pollen Database and used the squared-chord distance (SCD) dissimilarity metric to identify time periods of high pollen assemblage turnover representing periods of abrupt vegetation change. In addition, a set of generalized additive mixed models were applied to investigate the underlying dynamics of two periods of higher rates of turnover: the Younger Dryas–early Holocene transition (YD-EH; 11.6–9.0?kyr) and early–middle Holocene (EMH; 9.0–6.0?kyr). Results revealed a high frequency of turnover events between 12.75–11.5, 10.75–11, 10.25–10, 7.75–7.25, 3.25–3.0 and 1.75–.25?kyr. Furthermore, there was a strong linear relationship between pollen assemblage turnover and large directional temperature ...","author":[{"dropping-particle":"","family":"Seddon","given":"Alistair WR","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Macias-Fauria","given":"Marc","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Willis","given":"Kathy J","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"The Holocene","id":"ITEM-3","issue":"1","issued":{"date-parts":[["2015","1","9"]]},"page":"25-36","publisher":"SAGE PublicationsSage UK: London, England","title":"Climate and abrupt vegetation change in Northern Europe since the last deglaciation","type":"article-journal","volume":"25"},"uris":[""]},{"id":"ITEM-4","itemData":{"DOI":"10.1016/J.QUASCIREV.2005.03.018","ISSN":"0277-3791","abstract":"Temporal differences among fossil pollen samples from eastern North America provide a measure of the amount of climatic change that occurred on a broad spatial scale since the last glacial maximum (21,000calyrBP). Square-chord distances (SCDs) quantify the difference between pollen samples, and, thus, represent the potential magnitude of climatic change underlying the pollen record. The magnitude varied significantly among time scales with small changes common at centennial to millennial scales and large changes common at multi-millennial (i.e. orbital) scales. SCDs measured across 3000-yr intervals averaged 0.20, and often exceeded the maximum difference expected from samples collected within the same biome (0.15). SCDs across individual millennia were smaller and averaged 0.08. SCDs across the millennia at the beginning (13,000–12,000calyrBP) and end (12,000–11,000calyrBP) of the Younger Dryas chronozone (12,900–11,600calyrBP), however, averaged 0.20 and 0.18, respectively. These rapid step changes, large at sub-millennial scales, equal about 5–25% of the total glacial–interglacial transition. Large magnitude progressive changes in insolation, ice sheet extent, and atmospheric composition parallel the dominant trends in the SCD data, which show that progressive change comprises the first-order climatic pattern of the Holocene rather than stable or oscillatory patterns.","author":[{"dropping-particle":"","family":"Shuman","given":"Bryan","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Bartlein","given":"Patrick J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Webb","given":"Thompson","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Quaternary Science Reviews","id":"ITEM-4","issue":"20-21","issued":{"date-parts":[["2005","11","1"]]},"page":"2194-2206","publisher":"Pergamon","title":"The magnitudes of millennial- and orbital-scale climatic change in eastern North America during the Late Quaternary","type":"article-journal","volume":"24"},"uris":[""]}],"mendeley":{"formattedCitation":"(Grimm and Jacobson, 1992; Overpeck et al., 1985; Seddon et al., 2015; Shuman et al., 2005)","plainTextFormattedCitation":"(Grimm and Jacobson, 1992; Overpeck et al., 1985; Seddon et al., 2015; Shuman et al., 2005)","previouslyFormattedCitation":"(Grimm and Jacobson, 1992; Overpeck et al., 1985; Seddon et al., 2015; Shuman et al., 2005)"},"properties":{"noteIndex":0},"schema":""}(Grimm and Jacobson, 1992; Overpeck et al., 1985; Seddon et al., 2015; Shuman et al., 2005). If this variation in the vegetation is significant, it could represent replacement of one ecosystem by another; i.e. ecosystem turnover. Therefore, the magnitude of variation in the pollen record gives an indication that ecosystem turnover may be taking place. Additionally, the choice of palaeo-vegetation proxy may be important, as different proxies may differ in their sensitivity to ecosystem turnover. Alongside pollen, phytoliths are becoming an important palaeoecological proxy in the Neotropics as they have been shown to differentiate tropical ecosystems such as evergreen forest, dry forest and savannah ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.revpalbo.2013.01.004","ISSN":"00346667","abstract":"The interpretation of Neotropical fossil phytolith assemblages for palaeoenvironmental and archaeological reconstructions relies on the development of appropriate modern analogues. We analyzed modern phytolith assemblages from the soils of ten distinctive tropical vegetation communities in eastern lowland Bolivia, ranging from terra firme humid evergreen forest to seasonally-inundated savannah. Results show that broad ecosystems – evergreen tropical forest, semi-deciduous dry tropical forest, and savannah – can be clearly differentiated by examination of their phytolith spectra and the application of Principal Component Analysis (PCA). Differences in phytolith assemblages between particular vegetation communities within each of these ecosystems are more subtle, but can still be identified. Comparison of phytolith assemblages with pollen rain data and stable carbon isotope analyses from the same vegetation plots show that these proxies are not only complementary, but significantly improve taxonomic and ecosystem resolution, and therefore our ability to interpret palaeoenvironmental and archaeological records. Our data underline the utility of phytolith analyses for reconstructing Amazon Holocene vegetation histories and pre-Columbian land use, particularly the high spatial resolution possible with terrestrial soil-based phytolith studies.","author":[{"dropping-particle":"","family":"Dickau","given":"Ruth","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Whitney","given":"Bronwen S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"José","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Soto","given":"J. Daniel","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Metcalfe","given":"Phil","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Street-Perrott","given":"F. Alayne","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Loader","given":"Neil J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Ficken","given":"Katherine J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Review of Palaeobotany and Palynology","id":"ITEM-1","issued":{"date-parts":[["2013","6"]]},"page":"15-37","publisher":"Elsevier B.V.","title":"Differentiation of neotropical ecosystems by modern soil phytolith assemblages and its implications for palaeoenvironmental and archaeological reconstructions","type":"article-journal","volume":"193"},"uris":[""]},{"id":"ITEM-2","itemData":{"abstract":"Accurate archaeological and palaeoenvironmental reconstructions using phytoliths relies on the study ofmodern reference material. In eastern Acre, Brazil, weexamined whether the five most common forest types present todaywere able to be differentiated by their soil phytolith assemblages, and thus provide analogues with which to compare palaeoecological assemblages from pre-Columbian earthwork sites in the region. Surface soils and vegetation from dense humid evergreen forest, dense humid evergreen forestwith high palm abundance, palm forest, bamboo forest and fluvial forest were sampled and their phytoliths analysed. Relative phytolith frequencies were statistically compared using Principal Components Analyses (PCAs). We found the major differences in species composition to be well-represented by the phytolith assemblages as all forest types, apart from the two sub-types of dense humid evergreen forest, could be differentiated. Larger phytoliths from the sand fraction were found to be more ecologically diagnostic than those from the silt fraction. The surface soil phytolith assemblages we analysed can therefore be used as analogues to improve the accuracy of archaeological and palaeoecological reconstructions in the region. ? 2015 Elsevier B.V. All rights reserved","author":[{"dropping-particle":"","family":"Watling","given":"Jennifer","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"José","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Whitney","given":"Bronwen S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Consuelo","given":"E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Castro","given":"W.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Schaan","given":"Denise","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Feldpausch","given":"Ted R.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Review of Palaeobotany and Palynology","id":"ITEM-2","issued":{"date-parts":[["2016"]]},"note":"Could differentiate evergreen forest, palm forest, bamboo forest, fluvial forest - using PCA. Sand fraction phytoliths more useful than silt fraction - MUST separate for soil samples.\nVegetation composition of evergreen forest very specific to region of Acre (page 4) - need other studies from other regions to extend this work!\nTook 10 soil samples feom each plot - I don't have that kind of replication except for Alta Vista... particularly important for palm overepresentation (see page 8). Phytoliths are SO LOCAL in deposition. \n\nPossiblitiy that palm phytoliths might be distinguishable to genus level - I could test this using my phytolith reference collection as I build it??","page":"30-43","title":"Differentiation of neotropical ecosystems by modern soil phytolith assemblages and its implications for palaeoenvironmental and archaeological reconstructions II: Southwestern Amazonian forests","type":"article-journal","volume":"226"},"uris":[""]}],"mendeley":{"formattedCitation":"(Dickau et al., 2013; Watling et al., 2016)","plainTextFormattedCitation":"(Dickau et al., 2013; Watling et al., 2016)","previouslyFormattedCitation":"(Dickau et al., 2013; Watling et al., 2016)"},"properties":{"noteIndex":0},"schema":""}(Dickau et al., 2013; Watling et al., 2016). This ability suggests they would be sensitive to turnover between these ecosystems, but their relative sensitivity compared to pollen is poorly understood. As most plant taxa cannot be identified to the same taxonomic resolution by both pollen and phytoliths, it is not possible to compare the sensitivity of the two proxies to floristic changes within ecosystems based on individual indicator taxa. Therefore, to directly compare the two proxies, it is necessary to use a quantitative approach which standardises the full proxy assemblage. The variation in proxy assemblages indicates the sensitivity of the proxy to vegetation changes in that ecosystem; high variability indicates high sensitivity. This information can be used to inform proxy choice by palaeoecologists studying these ecosystems. Furthermore, analysis of variability in modern pollen and phytolith assemblages provides context and a modern training dataset for identification of periods of high variability representing ecosystem turnover in the palaeo-record.Noel Kempff Mercado National Park (NKMNP), NE Bolivia, was chosen as the study site to develop this modern training dataset and compare the sensitivity of pollen and phytoliths to vegetation changes because both modern pollen and phytolith datasets are available from ecological plots from a range of ecosystem types within this park, covering all the major plant communities in south-west Amazonia (evergreen forest, dry forest and savannah) ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.yqres.2005.08.008","ISSN":"00335894","abstract":"The paucity of modern pollen-rain data from Amazonia constitutes a significant barrier to understanding the Late Quaternary vegetation history of this globally important tropical forest region. Here, we present the first modern pollen-rain data for tall terra firme moist evergreen Amazon forest, collected between 1999 and 2001 from artificial pollen traps within a 500?×?20 m permanent study plot (14°34′50″S, 60°49′48″W) in Noel Kempff Mercado National Park (NE Bolivia). Spearman's rank correlations were performed to assess the extent of spatial and inter-annual variability in the pollen rain, whilst statistically distinctive taxa were identified using Principal Components Analysis (PCA). Comparisons with the floristic and basal area data of the plot (stems ≥10 cm d.b.h.) enabled the degree to which taxa are over/under-represented in the pollen rain to be assessed (using R-rel values). Moraceae/Urticaceae dominates the pollen rain (64% median abundance) and is also an important constituent of the vegetation, accounting for 16% of stems ≥10 cm d.b.h. and ca. 11% of the total basal area. Other important pollen taxa are Arecaceae (cf. Euterpe), Melastomataceae/Combretaceae, Cecropia, Didymopanax, Celtis, and Alchornea. However, 75% of stems and 67% of the total basal area of the plot ≥10 cm d.b.h. belong to species which are unidentified in the pollen rain, the most important of which are Phenakospermum guianensis (a banana-like herb) and the key canopy-emergent trees, Erisma uncinatum and Qualea paraensis.","author":[{"dropping-particle":"","family":"Gosling","given":"William D.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Tate","given":"Nicholas J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"collection-title":"Late Quaternary Tropical Ecosystem Dynamics","container-title":"Quaternary Research","id":"ITEM-1","issue":"3","issued":{"date-parts":[["2005","11"]]},"page":"284-297","title":"Modern pollen-rain characteristics of tall terra firme moist evergreen forest, southern Amazonia","type":"article-journal","volume":"64"},"uris":[""]},{"id":"ITEM-2","itemData":{"DOI":"10.1016/j.revpalbo.2008.06.007","ISSN":"0034-6667","abstract":"Accurate differentiation between tropical forest and savannah ecosystems in the fossil pollen record is hampered by the combination of: i) poor taxonomic resolution in pollen identification, and ii) the high species diversity of many lowland tropical families, i.e. with many different growth forms living in numerous environmental settings. These barriers to interpreting the fossil record hinder our understanding of the past distributions of different Neotropical ecosystems and consequently cloud our knowledge of past climatic, biodiversity and carbon storage patterns. Modern pollen studies facilitate an improved understanding of how ecosystems are represented by the pollen their plants produce and therefore aid interpretation of fossil pollen records. To understand how to differentiate ecosystems palynologically, it is essential that a consistent sampling method is used across ecosystems. However, to date, modern pollen studies from tropical South America have employed a variety of methodologies (e.g. pollen traps, moss polsters, soil samples). In this paper, we present the first modern pollen study from the Neotropics to examine the modern pollen rain from moist evergreen tropical forest (METF), semi-deciduous dry tropical forest (SDTF) and wooded savannah (cerrad?o) using a consistent sampling methodology (pollen traps). Pollen rain was sampled annually in September for the years 1999–2001 from within permanent vegetation study plots in, or near, the Noel Kempff Mercado National Park (NKMNP), Bolivia. Comparison of the modern pollen rain within these plots with detailed floristic inventories allowed estimates of the relative pollen productivity and dispersal for individual taxa to be made (% pollen/% vegetation or ‘p/v’). The applicability of these data to interpreting fossil records from lake sediments was then explored by comparison with pollen assemblages obtained from five lake surface samples. Pollen productivity is demonstrated to vary inter-annually and conforms to a consistent hierarchy for any given year: METF > SDTF > cerrad?o. This suggests an association between pollen productivity and basic structural characteristics of the ecosystem, i.e. closed canopy vs. open canopy vs. savannah. Comparison of modern pollen and vegetation revealed that some important floristic elements were completely absent from the pollen: Qualea and Erisma (METF), Bauhinia, Simira and Guazuma (SDTF), and Pouteria and Caryocar (cerrad?o). Anadenanthera was foun…","author":[{"dropping-particle":"","family":"Gosling","given":"William D.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Tate","given":"Nicholas J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Review of Palaeobotany and Palynology","id":"ITEM-2","issue":"1–2","issued":{"date-parts":[["2009","1"]]},"note":"Uses a surface sample from Mandiore = 51.6% Poaceae pollen at 0-1cm depth.","page":"70-85","title":"Differentiation between Neotropical rainforest, dry forest, and savannah ecosystems by their modern pollen spectra and implications for the fossil pollen record","type":"article-journal","volume":"153"},"uris":[""]},{"id":"ITEM-3","itemData":{"DOI":"10.1016/j.palaeo.2010.05.009","ISSN":"0031-0182","abstract":"An ongoing controversy in Amazonian palaeoecology is the manner in which Amazonian rainforest communities have responded to environmental change over the last glacial–interglacial cycle. Much of this controversy results from an inability to identify the floristic heterogeneity exhibited by rainforest communities within fossil pollen records. We apply multivariate (Principal Components Analysis) and classification (Unweighted Pair Group with Arithmetic Mean Agglomerative Classification) techniques to floral-biometric, modern pollen trap and lake sediment pollen data situated within different rainforest communities in the tropical lowlands of Amazonian Bolivia. Modern pollen rain analyses from artificial pollen traps show that evergreen terra firme (well-drained), evergreen terra firme liana, evergreen seasonally inundated, and evergreen riparian rainforests may be readily differentiated, floristically and palynologically. Analogue matching techniques, based on Euclidean distance measures, are employed to compare these pollen signatures with surface sediment pollen assemblages from five lakes: Laguna Bella Vista, Laguna Chaplin, and Laguna Huachi situated within the Madeira-Tapajós moist forest ecoregion, and Laguna Isirere and Laguna Loma Suarez, which are situated within forest patches in the Beni savanna ecoregion. The same numerical techniques are used to compare rainforest pollen trap signatures with the fossil pollen record of Laguna Chaplin.\n\nPollen assemblages of pollen traps situated within riparian forest communities are most similar to surface sediment samples from Lagunas Bella Vista and Chaplin. Pollen derived from terra firme forests also comprises a significant proportion of these assemblages. Together, these pollen spectra successfully identify riparian and terra firme rainforest communities surrounding the two lakes today. Close similarity between terra firme liana pollen trap assemblages and surface samples obtained from Laguna Huachi, a lake surrounded by relatively undisturbed liana forest, suggests liana forest pollen rain may also be identified within lake sediment records. Pollen spectra obtained from surface sediment samples from lakes situated within gallery forests of the Beni savanna ecoregion are significantly different to those obtained from the Madeira-Tapajós ecoregion, reflecting their different floristic compositions. By applying our findings to the previously published Laguna Chaplin Quaternary pollen record, we show that…","author":[{"dropping-particle":"","family":"Burn","given":"Michael J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Palaeogeography, Palaeoclimatology, Palaeoecology","id":"ITEM-3","issue":"1–2","issued":{"date-parts":[["2010","9"]]},"page":"1-18","title":"Pollen-based differentiation of Amazonian rainforest communities and implications for lowland palaeoecology in tropical South America","type":"article-journal","volume":"295"},"uris":[""]},{"id":"ITEM-4","itemData":{"DOI":"10.1016/j.revpalbo.2011.01.001","ISSN":"00346667","abstract":"The majority of vegetation reconstructions from the Neotropics are derived from fossil pollen records extracted from lake sediments. However, the interpretation of these records is restricted by limited knowledge of the contemporary relationships between the vegetation and pollen rain of Neotropical ecosystems, especially for more open vegetation such as savannas. This research aims to improve the interpretation of these records by investigating the vegetation and modern pollen rain of different savanna ecosystems in Bolivia using vegetation inventories, artificial pollen traps and surface lake sediments. Two types of savanna were studied, upland savannas (cerrado), occurring on well drained soils, and seasonally-inundated savannas occurring on seasonally water-logged soils. Quantitative vegetation data are used to identify taxa that are floristically important in the different savanna types and to allow modern pollen/vegetation ratios to be calculated. Artificial pollen traps from the upland savanna site are dominated by Moraceae (35%), Poaceae (30%), Alchornea (6%) and Cecropia (4%). The two seasonally-inundated savanna sites are dominated by Moraceae (37%), Poaceae (20%), Alchornea (8%) and Cecropia (7%), and Moraceae (25%), Cyperaceae (22%), Poaceae (19%) and Cecropia (9%), respectively. The modern pollen rain of seasonally-inundated savannas from surface lake sediments is dominated by Cyperaceae (35%), Poaceae (33%), Moraceae (9%) and Asteraceae (5%). Upland and seasonally-flooded savannas were found to be only subtly distinct from each other palynologically. All sites have a high proportion of Moraceae pollen due to effective wind dispersal of this pollen type from areas of evergreen forest close to the study sites. Modern pollen/vegetation ratios show that many key woody plant taxa are absent/under-represented in the modern pollen rain (e.g., Caryocar and Tabebuia). The lower-than-expected percentages of Poaceae pollen, and the scarcity of savanna indicators, in the modern pollen rain of these ecosystems mean that savannas could potentially be overlooked in fossil pollen records without consideration of the full pollen spectrum available.","author":[{"dropping-particle":"","family":"Jones","given":"Huw T.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Pennington","given":"R. Toby","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Review of Palaeobotany and Palynology","id":"ITEM-4","issue":"3-4","issued":{"date-parts":[["2011","4"]]},"page":"223-237","title":"Characterisation of Bolivian savanna ecosystems by their modern pollen rain and implications for fossil pollen records","type":"article-journal","volume":"164"},"uris":[""]},{"id":"ITEM-5","itemData":{"DOI":"10.1016/j.revpalbo.2013.01.004","ISSN":"00346667","abstract":"The interpretation of Neotropical fossil phytolith assemblages for palaeoenvironmental and archaeological reconstructions relies on the development of appropriate modern analogues. We analyzed modern phytolith assemblages from the soils of ten distinctive tropical vegetation communities in eastern lowland Bolivia, ranging from terra firme humid evergreen forest to seasonally-inundated savannah. Results show that broad ecosystems – evergreen tropical forest, semi-deciduous dry tropical forest, and savannah – can be clearly differentiated by examination of their phytolith spectra and the application of Principal Component Analysis (PCA). Differences in phytolith assemblages between particular vegetation communities within each of these ecosystems are more subtle, but can still be identified. Comparison of phytolith assemblages with pollen rain data and stable carbon isotope analyses from the same vegetation plots show that these proxies are not only complementary, but significantly improve taxonomic and ecosystem resolution, and therefore our ability to interpret palaeoenvironmental and archaeological records. Our data underline the utility of phytolith analyses for reconstructing Amazon Holocene vegetation histories and pre-Columbian land use, particularly the high spatial resolution possible with terrestrial soil-based phytolith studies.","author":[{"dropping-particle":"","family":"Dickau","given":"Ruth","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Whitney","given":"Bronwen S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"José","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Soto","given":"J. Daniel","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Metcalfe","given":"Phil","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Street-Perrott","given":"F. Alayne","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Loader","given":"Neil J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Ficken","given":"Katherine J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Review of Palaeobotany and Palynology","id":"ITEM-5","issued":{"date-parts":[["2013","6"]]},"page":"15-37","publisher":"Elsevier B.V.","title":"Differentiation of neotropical ecosystems by modern soil phytolith assemblages and its implications for palaeoenvironmental and archaeological reconstructions","type":"article-journal","volume":"193"},"uris":[""]}],"mendeley":{"formattedCitation":"(Burn et al., 2010; Dickau et al., 2013; Gosling et al., 2005, 2009; Jones et al., 2011)","plainTextFormattedCitation":"(Burn et al., 2010; Dickau et al., 2013; Gosling et al., 2005, 2009; Jones et al., 2011)","previouslyFormattedCitation":"(Burn et al., 2010; Dickau et al., 2013; Gosling et al., 2005, 2009; Jones et al., 2011)"},"properties":{"noteIndex":0},"schema":""}(Burn et al., 2010; Dickau et al., 2013; Gosling et al., 2005, 2009; Jones et al., 2011). These modern pollen and phytolith samples are from within permanent, 1-hectare, vegetation study plots within the RAINFOR network ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"URL":"","accessed":{"date-parts":[["2018","10","25"]]},"author":[{"dropping-particle":"","family":"SALVIAS","given":"","non-dropping-particle":"","parse-names":false,"suffix":""}],"id":"ITEM-1","issued":{"date-parts":[["2004"]]},"title":"SALVIAS: Synthesis and Analysis of Local Vegetation Inventories Across Scales","type":"webpage"},"uris":[""]},{"id":"ITEM-2","itemData":{"author":[{"dropping-particle":"","family":"Malhi","given":"Y.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Phillips","given":"O.L.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Llloyd","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Baker","given":"T.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Wright","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Almedia","given":"S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Arroyo","given":"L.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Journal of Vegetation Science","id":"ITEM-2","issue":"3","issued":{"date-parts":[["2002"]]},"page":"439-450","title":"An international network to monitor the structure, composition and dynamics of Amazonian forests (RAINFOR)","type":"article-journal","volume":"13"},"uris":[""]}],"mendeley":{"formattedCitation":"(Malhi et al., 2002; SALVIAS, 2004)","plainTextFormattedCitation":"(Malhi et al., 2002; SALVIAS, 2004)","previouslyFormattedCitation":"(Malhi et al., 2002; SALVIAS, 2004)"},"properties":{"noteIndex":0},"schema":""}(Malhi et al., 2002; SALVIAS, 2004), which has inventories of all woody stems > 10 cm d.b.h. This provides a unique opportunity to compare pollen and phytolith assemblages from the same vegetation using a common, quantitative methodology. This is the first direct quantitative comparison of these two proxies. The aim of this paper is to compare the sensitivity of pollen and phytolith assemblages to changes within and between three key tropical South American ecosystems: humid evergreen rainforest, semi-deciduous dry forest, and terra firme (cerrado) and seasonally-flooded savannah. The specific questions to be addressed are: Which proxy – pollen or phytoliths – is more sensitive to changes within and between evergreen forest, dry forest and savannah? Does this relationship vary by ecosystem type?What are the implications of proxy variability for reconstructing ecosystem turnover in the palaeo-record? What can this quantitative method of identifying ecosystem turnover add to the traditional qualitative approach? Here we use a numerical approach to differentiating ecosystem turnover from floristic turnover in palaeo vegetation records between evergreen forest, dry forest and savannah in south west Amazonia. For the purposes of this study, we define floristic turnover as the process of floristic change within an ecosystem, for example an increase in abundance of Anadenanthera trees within a dry forest, and ecosystem turnover as the process of change from one ecosystem to another i.e. dry forest to savannah. A modern training set of phytolith and pollen assemblages from these three key ecosystems is used to quantify the variability in modern assemblages within an ecosystem and between ecosystems. This modern dataset is used to set thresholds for floristic turnover and ecosystem turnover, based on the magnitude of variability in the dataset. These thresholds are then applied to palaeo-vegetation records from south-west Amazonia, where change from one ecosystem to another has been identified through interpretation of the fossil record. The thresholds assess whether the magnitude of variability in the palaeo-record enables identification of ecosystem turnover at the same horizons as the qualitative interpretation of the record. The pairwise distances between assemblages downcore in the palaeo-record and the full matrix of distances between all assemblages at all depths are compared with the thresholds defined from the modern dataset. 2. Study area and site selectionThe study area is Noel Kempff Mercado National Park (NKMNP), a 15,230km2 protected area in north-eastern Bolivia near the southern margin of Amazonia containing a diverse mixture of ecosystems including evergreen forest, semi-deciduous dry forest and savannah (Fig. 1). The east of the park is dominated by upland (cerrado) savannas on the Huanchaca plateau (600-900m a.s.l.), while the west of the park beyond the plateau is dominated by evergreen forests, with areas of seasonally inundated savannah near to river courses and small patches of terra firme (cerrado) savannah in raised areas. Towards the south of the park is the ecotone with semi-deciduous dry forest (Fig. 1). The climate is highly seasonal, with a 6-month dry season, annual precipitation between 1400 and 1500mm, mean annual temperature of 25-26°C and occasional cold fronts from Patagonia during austral winter which bring temperatures down to 10°C for several days ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"author":[{"dropping-particle":"","family":"Killeen","given":"T.J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Schulenberg","given":"T.S.","non-dropping-particle":"","parse-names":false,"suffix":""}],"id":"ITEM-1","issued":{"date-parts":[["1998"]]},"publisher":"Conservation International","publisher-place":"Washington, DC","title":"A biological assessment of Parc Nacional Noel Kempff Mercado, Bolivia, RAP Working Papers 10","type":"book"},"uris":[""]}],"mendeley":{"formattedCitation":"(Killeen and Schulenberg, 1998)","plainTextFormattedCitation":"(Killeen and Schulenberg, 1998)","previouslyFormattedCitation":"(Killeen and Schulenberg, 1998)"},"properties":{"noteIndex":0},"schema":""}(Killeen and Schulenberg, 1998). The only recent history of human disturbance in the park over the last century is rubber tapping in the early 1900s and low-intensity, selective logging of mahogany prior to the establishment of the national park in 1988 ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"author":[{"dropping-particle":"","family":"Killeen","given":"T.J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Schulenberg","given":"T.S.","non-dropping-particle":"","parse-names":false,"suffix":""}],"id":"ITEM-1","issued":{"date-parts":[["1998"]]},"publisher":"Conservation International","publisher-place":"Washington, DC","title":"A biological assessment of Parc Nacional Noel Kempff Mercado, Bolivia, RAP Working Papers 10","type":"book"},"uris":[""]}],"mendeley":{"formattedCitation":"(Killeen and Schulenberg, 1998)","plainTextFormattedCitation":"(Killeen and Schulenberg, 1998)","previouslyFormattedCitation":"(Killeen and Schulenberg, 1998)"},"properties":{"noteIndex":0},"schema":""}(Killeen and Schulenberg, 1998). This lack of disturbance makes it an ideal site for modern pollen and phytolith studies. Furthermore, a network of permanent vegetation study plots (part of the RAINFOR network) has been set up in and around the park with detailed botanical inventories of each vegetation type ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"URL":"","accessed":{"date-parts":[["2018","10","25"]]},"author":[{"dropping-particle":"","family":"SALVIAS","given":"","non-dropping-particle":"","parse-names":false,"suffix":""}],"id":"ITEM-1","issued":{"date-parts":[["2004"]]},"title":"SALVIAS: Synthesis and Analysis of Local Vegetation Inventories Across Scales","type":"webpage"},"uris":[""]},{"id":"ITEM-2","itemData":{"author":[{"dropping-particle":"","family":"Malhi","given":"Y.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Phillips","given":"O.L.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Llloyd","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Baker","given":"T.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Wright","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Almedia","given":"S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Arroyo","given":"L.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Journal of Vegetation Science","id":"ITEM-2","issue":"3","issued":{"date-parts":[["2002"]]},"page":"439-450","title":"An international network to monitor the structure, composition and dynamics of Amazonian forests (RAINFOR)","type":"article-journal","volume":"13"},"uris":[""]}],"mendeley":{"formattedCitation":"(Malhi et al., 2002; SALVIAS, 2004)","plainTextFormattedCitation":"(Malhi et al., 2002; SALVIAS, 2004)","previouslyFormattedCitation":"(Malhi et al., 2002; SALVIAS, 2004)"},"properties":{"noteIndex":0},"schema":""}(Malhi et al., 2002; SALVIAS, 2004).[Figure 1 - Map of Noel Kempff Mercado National Park (NKMNP) showing major vegetation communities and location of plots MV1, AC2, CP1, LF1, CH1, CH2, FC1, TO1, FT1. Inset map of South American countries with red-shaded area showing location of NKMNP.]The evergreen forest ecosystem class contains terra firme evergreen forest at plot LF1 and evergreen liana forest at plots CH1 and CH2. The evergreen forest at LF1 is tall, mature and dominated by tall trees such as Pseudolmedia laevis and the arboreal herbaceous plant Phenakospermum guianensis ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.yqres.2005.08.008","ISSN":"00335894","abstract":"The paucity of modern pollen-rain data from Amazonia constitutes a significant barrier to understanding the Late Quaternary vegetation history of this globally important tropical forest region. Here, we present the first modern pollen-rain data for tall terra firme moist evergreen Amazon forest, collected between 1999 and 2001 from artificial pollen traps within a 500?×?20 m permanent study plot (14°34′50″S, 60°49′48″W) in Noel Kempff Mercado National Park (NE Bolivia). Spearman's rank correlations were performed to assess the extent of spatial and inter-annual variability in the pollen rain, whilst statistically distinctive taxa were identified using Principal Components Analysis (PCA). Comparisons with the floristic and basal area data of the plot (stems ≥10 cm d.b.h.) enabled the degree to which taxa are over/under-represented in the pollen rain to be assessed (using R-rel values). Moraceae/Urticaceae dominates the pollen rain (64% median abundance) and is also an important constituent of the vegetation, accounting for 16% of stems ≥10 cm d.b.h. and ca. 11% of the total basal area. Other important pollen taxa are Arecaceae (cf. Euterpe), Melastomataceae/Combretaceae, Cecropia, Didymopanax, Celtis, and Alchornea. However, 75% of stems and 67% of the total basal area of the plot ≥10 cm d.b.h. belong to species which are unidentified in the pollen rain, the most important of which are Phenakospermum guianensis (a banana-like herb) and the key canopy-emergent trees, Erisma uncinatum and Qualea paraensis.","author":[{"dropping-particle":"","family":"Gosling","given":"William D.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Tate","given":"Nicholas J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"collection-title":"Late Quaternary Tropical Ecosystem Dynamics","container-title":"Quaternary Research","id":"ITEM-1","issue":"3","issued":{"date-parts":[["2005","11"]]},"page":"284-297","title":"Modern pollen-rain characteristics of tall terra firme moist evergreen forest, southern Amazonia","type":"article-journal","volume":"64"},"uris":[""]}],"mendeley":{"formattedCitation":"(Gosling et al., 2005)","plainTextFormattedCitation":"(Gosling et al., 2005)","previouslyFormattedCitation":"(Gosling et al., 2005)"},"properties":{"noteIndex":0},"schema":""}(Gosling et al., 2005). The evergreen liana forest at CH1 and CH2 has a low, dense canopy dominated by liana growth ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"author":[{"dropping-particle":"","family":"Killeen","given":"T.J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Schulenberg","given":"T.S.","non-dropping-particle":"","parse-names":false,"suffix":""}],"id":"ITEM-1","issued":{"date-parts":[["1998"]]},"publisher":"Conservation International","publisher-place":"Washington, DC","title":"A biological assessment of Parc Nacional Noel Kempff Mercado, Bolivia, RAP Working Papers 10","type":"book"},"uris":[""]}],"mendeley":{"formattedCitation":"(Killeen and Schulenberg, 1998)","plainTextFormattedCitation":"(Killeen and Schulenberg, 1998)","previouslyFormattedCitation":"(Killeen and Schulenberg, 1998)"},"properties":{"noteIndex":0},"schema":""}(Killeen and Schulenberg, 1998). The semi-deciduous dry forest ecosystem class contains seasonally inundated (plot MV1) as well as terra firme (plots AC2, CP1) forest. The three most abundant species in the terra firme dry forest are Caesalpinia floribunda, Tabebuia roseo-alba, and Anadenanthera colubrina ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.revpalbo.2008.06.007","ISSN":"0034-6667","abstract":"Accurate differentiation between tropical forest and savannah ecosystems in the fossil pollen record is hampered by the combination of: i) poor taxonomic resolution in pollen identification, and ii) the high species diversity of many lowland tropical families, i.e. with many different growth forms living in numerous environmental settings. These barriers to interpreting the fossil record hinder our understanding of the past distributions of different Neotropical ecosystems and consequently cloud our knowledge of past climatic, biodiversity and carbon storage patterns. Modern pollen studies facilitate an improved understanding of how ecosystems are represented by the pollen their plants produce and therefore aid interpretation of fossil pollen records. To understand how to differentiate ecosystems palynologically, it is essential that a consistent sampling method is used across ecosystems. However, to date, modern pollen studies from tropical South America have employed a variety of methodologies (e.g. pollen traps, moss polsters, soil samples). In this paper, we present the first modern pollen study from the Neotropics to examine the modern pollen rain from moist evergreen tropical forest (METF), semi-deciduous dry tropical forest (SDTF) and wooded savannah (cerrad?o) using a consistent sampling methodology (pollen traps). Pollen rain was sampled annually in September for the years 1999–2001 from within permanent vegetation study plots in, or near, the Noel Kempff Mercado National Park (NKMNP), Bolivia. Comparison of the modern pollen rain within these plots with detailed floristic inventories allowed estimates of the relative pollen productivity and dispersal for individual taxa to be made (% pollen/% vegetation or ‘p/v’). The applicability of these data to interpreting fossil records from lake sediments was then explored by comparison with pollen assemblages obtained from five lake surface samples. Pollen productivity is demonstrated to vary inter-annually and conforms to a consistent hierarchy for any given year: METF > SDTF > cerrad?o. This suggests an association between pollen productivity and basic structural characteristics of the ecosystem, i.e. closed canopy vs. open canopy vs. savannah. Comparison of modern pollen and vegetation revealed that some important floristic elements were completely absent from the pollen: Qualea and Erisma (METF), Bauhinia, Simira and Guazuma (SDTF), and Pouteria and Caryocar (cerrad?o). Anadenanthera was foun…","author":[{"dropping-particle":"","family":"Gosling","given":"William D.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Tate","given":"Nicholas J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Review of Palaeobotany and Palynology","id":"ITEM-1","issue":"1–2","issued":{"date-parts":[["2009","1"]]},"note":"Uses a surface sample from Mandiore = 51.6% Poaceae pollen at 0-1cm depth.","page":"70-85","title":"Differentiation between Neotropical rainforest, dry forest, and savannah ecosystems by their modern pollen spectra and implications for the fossil pollen record","type":"article-journal","volume":"153"},"uris":[""]}],"mendeley":{"formattedCitation":"(Gosling et al., 2009)","plainTextFormattedCitation":"(Gosling et al., 2009)","previouslyFormattedCitation":"(Gosling et al., 2009)"},"properties":{"noteIndex":0},"schema":""}(Gosling et al., 2009). In the seasonally inundated dry forest common taxa include Inga ingoides, Physocalymma scaberimum and Hirtella gracilipes ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.revpalbo.2013.01.004","ISSN":"00346667","abstract":"The interpretation of Neotropical fossil phytolith assemblages for palaeoenvironmental and archaeological reconstructions relies on the development of appropriate modern analogues. We analyzed modern phytolith assemblages from the soils of ten distinctive tropical vegetation communities in eastern lowland Bolivia, ranging from terra firme humid evergreen forest to seasonally-inundated savannah. Results show that broad ecosystems – evergreen tropical forest, semi-deciduous dry tropical forest, and savannah – can be clearly differentiated by examination of their phytolith spectra and the application of Principal Component Analysis (PCA). Differences in phytolith assemblages between particular vegetation communities within each of these ecosystems are more subtle, but can still be identified. Comparison of phytolith assemblages with pollen rain data and stable carbon isotope analyses from the same vegetation plots show that these proxies are not only complementary, but significantly improve taxonomic and ecosystem resolution, and therefore our ability to interpret palaeoenvironmental and archaeological records. Our data underline the utility of phytolith analyses for reconstructing Amazon Holocene vegetation histories and pre-Columbian land use, particularly the high spatial resolution possible with terrestrial soil-based phytolith studies.","author":[{"dropping-particle":"","family":"Dickau","given":"Ruth","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Whitney","given":"Bronwen S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"José","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Soto","given":"J. 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The savannah ecosystem class also contains both terra firme (plot FC1) and seasonally inundated (plots FT1, TO1) communities. Both communities are dominated by Panicoideae grasses, but the terra firme (cerrado) savannah contains Myrtaceae, Borreria sp. and Dilleniaceae, whereas the seasonally inundated savannah contains Cyperaceae, Eriocaulaceae and Xyridaceae ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.revpalbo.2013.01.004","ISSN":"00346667","abstract":"The interpretation of Neotropical fossil phytolith assemblages for palaeoenvironmental and archaeological reconstructions relies on the development of appropriate modern analogues. We analyzed modern phytolith assemblages from the soils of ten distinctive tropical vegetation communities in eastern lowland Bolivia, ranging from terra firme humid evergreen forest to seasonally-inundated savannah. Results show that broad ecosystems – evergreen tropical forest, semi-deciduous dry tropical forest, and savannah – can be clearly differentiated by examination of their phytolith spectra and the application of Principal Component Analysis (PCA). Differences in phytolith assemblages between particular vegetation communities within each of these ecosystems are more subtle, but can still be identified. Comparison of phytolith assemblages with pollen rain data and stable carbon isotope analyses from the same vegetation plots show that these proxies are not only complementary, but significantly improve taxonomic and ecosystem resolution, and therefore our ability to interpret palaeoenvironmental and archaeological records. 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Table 1 – plant community and broad ecosystems for each 1-hectare study plot, number of samples per plot and reference to original publication of raw data. SI = seasonally-inundated, TF = terra firme. EcosystemPlant communityPlot codeNo. phytolith samplesNo. pollen samplesReferencesDry forestSI Dry forestMV164ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.revpalbo.2011.01.001","ISSN":"00346667","abstract":"The majority of vegetation reconstructions from the Neotropics are derived from fossil pollen records extracted from lake sediments. However, the interpretation of these records is restricted by limited knowledge of the contemporary relationships between the vegetation and pollen rain of Neotropical ecosystems, especially for more open vegetation such as savannas. This research aims to improve the interpretation of these records by investigating the vegetation and modern pollen rain of different savanna ecosystems in Bolivia using vegetation inventories, artificial pollen traps and surface lake sediments. Two types of savanna were studied, upland savannas (cerrado), occurring on well drained soils, and seasonally-inundated savannas occurring on seasonally water-logged soils. Quantitative vegetation data are used to identify taxa that are floristically important in the different savanna types and to allow modern pollen/vegetation ratios to be calculated. Artificial pollen traps from the upland savanna site are dominated by Moraceae (35%), Poaceae (30%), Alchornea (6%) and Cecropia (4%). The two seasonally-inundated savanna sites are dominated by Moraceae (37%), Poaceae (20%), Alchornea (8%) and Cecropia (7%), and Moraceae (25%), Cyperaceae (22%), Poaceae (19%) and Cecropia (9%), respectively. The modern pollen rain of seasonally-inundated savannas from surface lake sediments is dominated by Cyperaceae (35%), Poaceae (33%), Moraceae (9%) and Asteraceae (5%). Upland and seasonally-flooded savannas were found to be only subtly distinct from each other palynologically. All sites have a high proportion of Moraceae pollen due to effective wind dispersal of this pollen type from areas of evergreen forest close to the study sites. Modern pollen/vegetation ratios show that many key woody plant taxa are absent/under-represented in the modern pollen rain (e.g., Caryocar and Tabebuia). 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In this paper, we present the first modern pollen study from the Neotropics to examine the modern pollen rain from moist evergreen tropical forest (METF), semi-deciduous dry tropical forest (SDTF) and wooded savannah (cerrad?o) using a consistent sampling methodology (pollen traps). Pollen rain was sampled annually in September for the years 1999–2001 from within permanent vegetation study plots in, or near, the Noel Kempff Mercado National Park (NKMNP), Bolivia. Comparison of the modern pollen rain within these plots with detailed floristic inventories allowed estimates of the relative pollen productivity and dispersal for individual taxa to be made (% pollen/% vegetation or ‘p/v’). The applicability of these data to interpreting fossil records from lake sediments was then explored by comparison with pollen assemblages obtained from five lake surface samples. Pollen productivity is demonstrated to vary inter-annually and conforms to a consistent hierarchy for any given year: METF > SDTF > cerrad?o. This suggests an association between pollen productivity and basic structural characteristics of the ecosystem, i.e. closed canopy vs. open canopy vs. savannah. Comparison of modern pollen and vegetation revealed that some important floristic elements were completely absent from the pollen: Qualea and Erisma (METF), Bauhinia, Simira and Guazuma (SDTF), and Pouteria and Caryocar (cerrad?o). Anadenanthera was foun…","author":[{"dropping-particle":"","family":"Gosling","given":"William D.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Tate","given":"Nicholas J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Review of Palaeobotany and Palynology","id":"ITEM-1","issue":"1–2","issued":{"date-parts":[["2009","1"]]},"note":"Uses a surface sample from Mandiore = 51.6% Poaceae pollen at 0-1cm depth.","page":"70-85","title":"Differentiation between Neotropical rainforest, dry forest, and savannah ecosystems by their modern pollen spectra and implications for the fossil pollen record","type":"article-journal","volume":"153"},"uris":[""]},{"id":"ITEM-2","itemData":{"DOI":"10.1016/j.revpalbo.2013.01.004","ISSN":"00346667","abstract":"The interpretation of Neotropical fossil phytolith assemblages for palaeoenvironmental and archaeological reconstructions relies on the development of appropriate modern analogues. We analyzed modern phytolith assemblages from the soils of ten distinctive tropical vegetation communities in eastern lowland Bolivia, ranging from terra firme humid evergreen forest to seasonally-inundated savannah. Results show that broad ecosystems – evergreen tropical forest, semi-deciduous dry tropical forest, and savannah – can be clearly differentiated by examination of their phytolith spectra and the application of Principal Component Analysis (PCA). Differences in phytolith assemblages between particular vegetation communities within each of these ecosystems are more subtle, but can still be identified. Comparison of phytolith assemblages with pollen rain data and stable carbon isotope analyses from the same vegetation plots show that these proxies are not only complementary, but significantly improve taxonomic and ecosystem resolution, and therefore our ability to interpret palaeoenvironmental and archaeological records. Our data underline the utility of phytolith analyses for reconstructing Amazon Holocene vegetation histories and pre-Columbian land use, particularly the high spatial resolution possible with terrestrial soil-based phytolith studies.","author":[{"dropping-particle":"","family":"Dickau","given":"Ruth","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Whitney","given":"Bronwen S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"José","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Soto","given":"J. Daniel","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Metcalfe","given":"Phil","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Street-Perrott","given":"F. Alayne","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Loader","given":"Neil J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Ficken","given":"Katherine J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Review of Palaeobotany and Palynology","id":"ITEM-2","issued":{"date-parts":[["2013","6"]]},"page":"15-37","publisher":"Elsevier B.V.","title":"Differentiation of neotropical ecosystems by modern soil phytolith assemblages and its implications for palaeoenvironmental and archaeological reconstructions","type":"article-journal","volume":"193"},"uris":[""]}],"mendeley":{"formattedCitation":"(Dickau et al., 2013; Gosling et al., 2009)","manualFormatting":"Dickau et al., 2013; Gosling et al., 2009","plainTextFormattedCitation":"(Dickau et al., 2013; Gosling et al., 2009)","previouslyFormattedCitation":"(Dickau et al., 2013; Gosling et al., 2009)"},"properties":{"noteIndex":0},"schema":""}Dickau et al., 2013; Gosling et al., 2009TF Liana forestCH1, 265ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.palaeo.2010.05.009","ISSN":"0031-0182","abstract":"An ongoing controversy in Amazonian palaeoecology is the manner in which Amazonian rainforest communities have responded to environmental change over the last glacial–interglacial cycle. Much of this controversy results from an inability to identify the floristic heterogeneity exhibited by rainforest communities within fossil pollen records. We apply multivariate (Principal Components Analysis) and classification (Unweighted Pair Group with Arithmetic Mean Agglomerative Classification) techniques to floral-biometric, modern pollen trap and lake sediment pollen data situated within different rainforest communities in the tropical lowlands of Amazonian Bolivia. Modern pollen rain analyses from artificial pollen traps show that evergreen terra firme (well-drained), evergreen terra firme liana, evergreen seasonally inundated, and evergreen riparian rainforests may be readily differentiated, floristically and palynologically. Analogue matching techniques, based on Euclidean distance measures, are employed to compare these pollen signatures with surface sediment pollen assemblages from five lakes: Laguna Bella Vista, Laguna Chaplin, and Laguna Huachi situated within the Madeira-Tapajós moist forest ecoregion, and Laguna Isirere and Laguna Loma Suarez, which are situated within forest patches in the Beni savanna ecoregion. The same numerical techniques are used to compare rainforest pollen trap signatures with the fossil pollen record of Laguna Chaplin.\n\nPollen assemblages of pollen traps situated within riparian forest communities are most similar to surface sediment samples from Lagunas Bella Vista and Chaplin. Pollen derived from terra firme forests also comprises a significant proportion of these assemblages. Together, these pollen spectra successfully identify riparian and terra firme rainforest communities surrounding the two lakes today. Close similarity between terra firme liana pollen trap assemblages and surface samples obtained from Laguna Huachi, a lake surrounded by relatively undisturbed liana forest, suggests liana forest pollen rain may also be identified within lake sediment records. Pollen spectra obtained from surface sediment samples from lakes situated within gallery forests of the Beni savanna ecoregion are significantly different to those obtained from the Madeira-Tapajós ecoregion, reflecting their different floristic compositions. By applying our findings to the previously published Laguna Chaplin Quaternary pollen record, we show that…","author":[{"dropping-particle":"","family":"Burn","given":"Michael J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Palaeogeography, Palaeoclimatology, Palaeoecology","id":"ITEM-1","issue":"1–2","issued":{"date-parts":[["2010","9"]]},"page":"1-18","title":"Pollen-based differentiation of Amazonian rainforest communities and implications for lowland palaeoecology in tropical South America","type":"article-journal","volume":"295"},"uris":[""]},{"id":"ITEM-2","itemData":{"DOI":"10.1016/j.revpalbo.2013.01.004","ISSN":"00346667","abstract":"The interpretation of Neotropical fossil phytolith assemblages for palaeoenvironmental and archaeological reconstructions relies on the development of appropriate modern analogues. We analyzed modern phytolith assemblages from the soils of ten distinctive tropical vegetation communities in eastern lowland Bolivia, ranging from terra firme humid evergreen forest to seasonally-inundated savannah. Results show that broad ecosystems – evergreen tropical forest, semi-deciduous dry tropical forest, and savannah – can be clearly differentiated by examination of their phytolith spectra and the application of Principal Component Analysis (PCA). Differences in phytolith assemblages between particular vegetation communities within each of these ecosystems are more subtle, but can still be identified. Comparison of phytolith assemblages with pollen rain data and stable carbon isotope analyses from the same vegetation plots show that these proxies are not only complementary, but significantly improve taxonomic and ecosystem resolution, and therefore our ability to interpret palaeoenvironmental and archaeological records. Our data underline the utility of phytolith analyses for reconstructing Amazon Holocene vegetation histories and pre-Columbian land use, particularly the high spatial resolution possible with terrestrial soil-based phytolith studies.","author":[{"dropping-particle":"","family":"Dickau","given":"Ruth","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Whitney","given":"Bronwen S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"José","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Soto","given":"J. Daniel","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Metcalfe","given":"Phil","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Street-Perrott","given":"F. Alayne","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Loader","given":"Neil J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Ficken","given":"Katherine J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Review of Palaeobotany and Palynology","id":"ITEM-2","issued":{"date-parts":[["2013","6"]]},"page":"15-37","publisher":"Elsevier B.V.","title":"Differentiation of neotropical ecosystems by modern soil phytolith assemblages and its implications for palaeoenvironmental and archaeological reconstructions","type":"article-journal","volume":"193"},"uris":[""]}],"mendeley":{"formattedCitation":"(Burn et al., 2010; Dickau et al., 2013)","manualFormatting":"Burn et al., 2010; Dickau et al., 2013","plainTextFormattedCitation":"(Burn et al., 2010; Dickau et al., 2013)","previouslyFormattedCitation":"(Burn et al., 2010; Dickau et al., 2013)"},"properties":{"noteIndex":0},"schema":""}Burn et al., 2010; Dickau et al., 2013SavannahTF SavannahFC164ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.revpalbo.2011.01.001","ISSN":"00346667","abstract":"The majority of vegetation reconstructions from the Neotropics are derived from fossil pollen records extracted from lake sediments. However, the interpretation of these records is restricted by limited knowledge of the contemporary relationships between the vegetation and pollen rain of Neotropical ecosystems, especially for more open vegetation such as savannas. This research aims to improve the interpretation of these records by investigating the vegetation and modern pollen rain of different savanna ecosystems in Bolivia using vegetation inventories, artificial pollen traps and surface lake sediments. Two types of savanna were studied, upland savannas (cerrado), occurring on well drained soils, and seasonally-inundated savannas occurring on seasonally water-logged soils. Quantitative vegetation data are used to identify taxa that are floristically important in the different savanna types and to allow modern pollen/vegetation ratios to be calculated. Artificial pollen traps from the upland savanna site are dominated by Moraceae (35%), Poaceae (30%), Alchornea (6%) and Cecropia (4%). The two seasonally-inundated savanna sites are dominated by Moraceae (37%), Poaceae (20%), Alchornea (8%) and Cecropia (7%), and Moraceae (25%), Cyperaceae (22%), Poaceae (19%) and Cecropia (9%), respectively. The modern pollen rain of seasonally-inundated savannas from surface lake sediments is dominated by Cyperaceae (35%), Poaceae (33%), Moraceae (9%) and Asteraceae (5%). Upland and seasonally-flooded savannas were found to be only subtly distinct from each other palynologically. All sites have a high proportion of Moraceae pollen due to effective wind dispersal of this pollen type from areas of evergreen forest close to the study sites. Modern pollen/vegetation ratios show that many key woody plant taxa are absent/under-represented in the modern pollen rain (e.g., Caryocar and Tabebuia). The lower-than-expected percentages of Poaceae pollen, and the scarcity of savanna indicators, in the modern pollen rain of these ecosystems mean that savannas could potentially be overlooked in fossil pollen records without consideration of the full pollen spectrum available.","author":[{"dropping-particle":"","family":"Jones","given":"Huw T.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Pennington","given":"R. Toby","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Review of Palaeobotany and Palynology","id":"ITEM-1","issue":"3-4","issued":{"date-parts":[["2011","4"]]},"page":"223-237","title":"Characterisation of Bolivian savanna ecosystems by their modern pollen rain and implications for fossil pollen records","type":"article-journal","volume":"164"},"uris":[""]},{"id":"ITEM-2","itemData":{"DOI":"10.1016/j.revpalbo.2013.01.004","ISSN":"00346667","abstract":"The interpretation of Neotropical fossil phytolith assemblages for palaeoenvironmental and archaeological reconstructions relies on the development of appropriate modern analogues. We analyzed modern phytolith assemblages from the soils of ten distinctive tropical vegetation communities in eastern lowland Bolivia, ranging from terra firme humid evergreen forest to seasonally-inundated savannah. Results show that broad ecosystems – evergreen tropical forest, semi-deciduous dry tropical forest, and savannah – can be clearly differentiated by examination of their phytolith spectra and the application of Principal Component Analysis (PCA). Differences in phytolith assemblages between particular vegetation communities within each of these ecosystems are more subtle, but can still be identified. Comparison of phytolith assemblages with pollen rain data and stable carbon isotope analyses from the same vegetation plots show that these proxies are not only complementary, but significantly improve taxonomic and ecosystem resolution, and therefore our ability to interpret palaeoenvironmental and archaeological records. Our data underline the utility of phytolith analyses for reconstructing Amazon Holocene vegetation histories and pre-Columbian land use, particularly the high spatial resolution possible with terrestrial soil-based phytolith studies.","author":[{"dropping-particle":"","family":"Dickau","given":"Ruth","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Whitney","given":"Bronwen S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"José","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Soto","given":"J. Daniel","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Metcalfe","given":"Phil","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Street-Perrott","given":"F. Alayne","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Loader","given":"Neil J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Ficken","given":"Katherine J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Review of Palaeobotany and Palynology","id":"ITEM-2","issued":{"date-parts":[["2013","6"]]},"page":"15-37","publisher":"Elsevier B.V.","title":"Differentiation of neotropical ecosystems by modern soil phytolith assemblages and its implications for palaeoenvironmental and archaeological reconstructions","type":"article-journal","volume":"193"},"uris":[""]}],"mendeley":{"formattedCitation":"(Dickau et al., 2013; Jones et al., 2011)","manualFormatting":"Dickau et al., 2013; Jones et al., 2009","plainTextFormattedCitation":"(Dickau et al., 2013; Jones et al., 2011)","previouslyFormattedCitation":"(Dickau et al., 2013; Jones et al., 2011)"},"properties":{"noteIndex":0},"schema":""}Dickau et al., 2013; Jones et al., 2009SI SavannahFT134ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.revpalbo.2011.01.001","ISSN":"00346667","abstract":"The majority of vegetation reconstructions from the Neotropics are derived from fossil pollen records extracted from lake sediments. However, the interpretation of these records is restricted by limited knowledge of the contemporary relationships between the vegetation and pollen rain of Neotropical ecosystems, especially for more open vegetation such as savannas. This research aims to improve the interpretation of these records by investigating the vegetation and modern pollen rain of different savanna ecosystems in Bolivia using vegetation inventories, artificial pollen traps and surface lake sediments. Two types of savanna were studied, upland savannas (cerrado), occurring on well drained soils, and seasonally-inundated savannas occurring on seasonally water-logged soils. Quantitative vegetation data are used to identify taxa that are floristically important in the different savanna types and to allow modern pollen/vegetation ratios to be calculated. Artificial pollen traps from the upland savanna site are dominated by Moraceae (35%), Poaceae (30%), Alchornea (6%) and Cecropia (4%). The two seasonally-inundated savanna sites are dominated by Moraceae (37%), Poaceae (20%), Alchornea (8%) and Cecropia (7%), and Moraceae (25%), Cyperaceae (22%), Poaceae (19%) and Cecropia (9%), respectively. The modern pollen rain of seasonally-inundated savannas from surface lake sediments is dominated by Cyperaceae (35%), Poaceae (33%), Moraceae (9%) and Asteraceae (5%). Upland and seasonally-flooded savannas were found to be only subtly distinct from each other palynologically. All sites have a high proportion of Moraceae pollen due to effective wind dispersal of this pollen type from areas of evergreen forest close to the study sites. Modern pollen/vegetation ratios show that many key woody plant taxa are absent/under-represented in the modern pollen rain (e.g., Caryocar and Tabebuia). The lower-than-expected percentages of Poaceae pollen, and the scarcity of savanna indicators, in the modern pollen rain of these ecosystems mean that savannas could potentially be overlooked in fossil pollen records without consideration of the full pollen spectrum available.","author":[{"dropping-particle":"","family":"Jones","given":"Huw T.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Pennington","given":"R. Toby","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Review of Palaeobotany and Palynology","id":"ITEM-1","issue":"3-4","issued":{"date-parts":[["2011","4"]]},"page":"223-237","title":"Characterisation of Bolivian savanna ecosystems by their modern pollen rain and implications for fossil pollen records","type":"article-journal","volume":"164"},"uris":[""]},{"id":"ITEM-2","itemData":{"DOI":"10.1016/j.revpalbo.2013.01.004","ISSN":"00346667","abstract":"The interpretation of Neotropical fossil phytolith assemblages for palaeoenvironmental and archaeological reconstructions relies on the development of appropriate modern analogues. We analyzed modern phytolith assemblages from the soils of ten distinctive tropical vegetation communities in eastern lowland Bolivia, ranging from terra firme humid evergreen forest to seasonally-inundated savannah. Results show that broad ecosystems – evergreen tropical forest, semi-deciduous dry tropical forest, and savannah – can be clearly differentiated by examination of their phytolith spectra and the application of Principal Component Analysis (PCA). Differences in phytolith assemblages between particular vegetation communities within each of these ecosystems are more subtle, but can still be identified. Comparison of phytolith assemblages with pollen rain data and stable carbon isotope analyses from the same vegetation plots show that these proxies are not only complementary, but significantly improve taxonomic and ecosystem resolution, and therefore our ability to interpret palaeoenvironmental and archaeological records. Our data underline the utility of phytolith analyses for reconstructing Amazon Holocene vegetation histories and pre-Columbian land use, particularly the high spatial resolution possible with terrestrial soil-based phytolith studies.","author":[{"dropping-particle":"","family":"Dickau","given":"Ruth","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Whitney","given":"Bronwen S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"José","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Soto","given":"J. Daniel","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Metcalfe","given":"Phil","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Street-Perrott","given":"F. Alayne","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Loader","given":"Neil J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Ficken","given":"Katherine J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Review of Palaeobotany and Palynology","id":"ITEM-2","issued":{"date-parts":[["2013","6"]]},"page":"15-37","publisher":"Elsevier B.V.","title":"Differentiation of neotropical ecosystems by modern soil phytolith assemblages and its implications for palaeoenvironmental and archaeological reconstructions","type":"article-journal","volume":"193"},"uris":[""]}],"mendeley":{"formattedCitation":"(Dickau et al., 2013; Jones et al., 2011)","manualFormatting":"Dickau et al., 2013; Jones et al., 2009","plainTextFormattedCitation":"(Dickau et al., 2013; Jones et al., 2011)","previouslyFormattedCitation":"(Dickau et al., 2013; Jones et al., 2011)"},"properties":{"noteIndex":0},"schema":""}Dickau et al., 2013; Jones et al., 2009TO134ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.revpalbo.2013.01.004","ISSN":"00346667","abstract":"The interpretation of Neotropical fossil phytolith assemblages for palaeoenvironmental and archaeological reconstructions relies on the development of appropriate modern analogues. We analyzed modern phytolith assemblages from the soils of ten distinctive tropical vegetation communities in eastern lowland Bolivia, ranging from terra firme humid evergreen forest to seasonally-inundated savannah. Results show that broad ecosystems – evergreen tropical forest, semi-deciduous dry tropical forest, and savannah – can be clearly differentiated by examination of their phytolith spectra and the application of Principal Component Analysis (PCA). Differences in phytolith assemblages between particular vegetation communities within each of these ecosystems are more subtle, but can still be identified. Comparison of phytolith assemblages with pollen rain data and stable carbon isotope analyses from the same vegetation plots show that these proxies are not only complementary, but significantly improve taxonomic and ecosystem resolution, and therefore our ability to interpret palaeoenvironmental and archaeological records. Our data underline the utility of phytolith analyses for reconstructing Amazon Holocene vegetation histories and pre-Columbian land use, particularly the high spatial resolution possible with terrestrial soil-based phytolith studies.","author":[{"dropping-particle":"","family":"Dickau","given":"Ruth","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Whitney","given":"Bronwen S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"José","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Soto","given":"J. 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However, the interpretation of these records is restricted by limited knowledge of the contemporary relationships between the vegetation and pollen rain of Neotropical ecosystems, especially for more open vegetation such as savannas. This research aims to improve the interpretation of these records by investigating the vegetation and modern pollen rain of different savanna ecosystems in Bolivia using vegetation inventories, artificial pollen traps and surface lake sediments. Two types of savanna were studied, upland savannas (cerrado), occurring on well drained soils, and seasonally-inundated savannas occurring on seasonally water-logged soils. Quantitative vegetation data are used to identify taxa that are floristically important in the different savanna types and to allow modern pollen/vegetation ratios to be calculated. Artificial pollen traps from the upland savanna site are dominated by Moraceae (35%), Poaceae (30%), Alchornea (6%) and Cecropia (4%). The two seasonally-inundated savanna sites are dominated by Moraceae (37%), Poaceae (20%), Alchornea (8%) and Cecropia (7%), and Moraceae (25%), Cyperaceae (22%), Poaceae (19%) and Cecropia (9%), respectively. The modern pollen rain of seasonally-inundated savannas from surface lake sediments is dominated by Cyperaceae (35%), Poaceae (33%), Moraceae (9%) and Asteraceae (5%). Upland and seasonally-flooded savannas were found to be only subtly distinct from each other palynologically. All sites have a high proportion of Moraceae pollen due to effective wind dispersal of this pollen type from areas of evergreen forest close to the study sites. Modern pollen/vegetation ratios show that many key woody plant taxa are absent/under-represented in the modern pollen rain (e.g., Caryocar and Tabebuia). 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Toby","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Review of Palaeobotany and Palynology","id":"ITEM-2","issue":"3-4","issued":{"date-parts":[["2011","4"]]},"page":"223-237","title":"Characterisation of Bolivian savanna ecosystems by their modern pollen rain and implications for fossil pollen records","type":"article-journal","volume":"164"},"uris":[""]}],"mendeley":{"formattedCitation":"(Dickau et al., 2013; Jones et al., 2011)","manualFormatting":"Dickau et al., 2013; Jones et al., 2009","plainTextFormattedCitation":"(Dickau et al., 2013; Jones et al., 2011)","previouslyFormattedCitation":"(Dickau et al., 2013; Jones et al., 2011)"},"properties":{"noteIndex":0},"schema":""}Dickau et al., 2013; Jones et al., 2009The first site chosen for application of the ecosystem turnover thresholds to the palaeo-record is Laguna Granja, as it has both pollen and phytolith records for the past 6,000 years ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1177/0959683615581204","ISBN":"0959-6836","ISSN":"0959-6836","abstract":"The nature and extent of pre-Columbian (pre-AD 1492) human impact in Amazonia is a contentious issue. The Bolivian Amazon has yielded some of the most impressive evidence for large and complex pre-Columbian societies in the Amazon basin, yet there remains relatively little data concerning the land use of these societies over time. Palaeoecology, when integrated with archaeological data, has the potential to fill these gaps in our knowledge. We present a 6000-year record of anthropogenic burning, agriculture and vegetation change, from an oxbow lake located adjacent to a pre-Columbian ring ditch in north-east Bolivia (13°15′44″S, 63°42′37″W). Human occupation around the lake site is inferred from pollen and phytoliths of maize (Zea mays L.) and macroscopic charcoal evidence of anthropogenic burning. First occupation around the lake was radiocarbon dated to ~2500 calibrated years before present (BP). The persistence of maize in the record from ~1850 BP suggests that it was an important crop grown in the ring-ditch region in pre-Columbian times, and abundant macroscopic charcoal suggests that pre-Columbian land management entailed more extensive burning of the landscape than the slash-and-burn agriculture practised around the site today. The site was occupied continuously until near-modern times, although there is evidence for a decline in agricultural intensity or change in land-use strategy, and possible population decline, from ~600–500 BP. The long and continuous occupation, which predates the establishment of rainforest in the region, suggests that pre-Columbian land use may have had a significant influence on ecosystem development at this site over the last ~2000 years.","author":[{"dropping-particle":"","family":"Carson","given":"J. F.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Watling","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"F. E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Whitney","given":"B. S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Prumers","given":"H.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Soto","given":"J. D.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"The Holocene","id":"ITEM-1","issue":"8","issued":{"date-parts":[["2015","5","1"]]},"note":"From Duplicate 2 (Pre-Columbian land use in the ring-ditch region of the Bolivian Amazon - Carson, J. F.; Watling, J.; Mayle, F. E.; Whitney, B. S.; Iriarte, J.; Prumers, H.; Soto, J. D.)\n\nUsed phytoliths and pollen from oxbow lake in NE Bolivia - oxbow 6000 yrs old.\n\nNice passage about phytoliths from lakes and their source area compared to pollen - assumes local representation due to modes of transport available.","page":"1285-1300","title":"Pre-Columbian land use in the ring-ditch region of the Bolivian Amazon","type":"article-journal","volume":"25"},"uris":[""]}],"mendeley":{"formattedCitation":"(Carson et al., 2015)","plainTextFormattedCitation":"(Carson et al., 2015)","previouslyFormattedCitation":"(Carson et al., 2015)"},"properties":{"noteIndex":0},"schema":""}(Carson et al., 2015). L. Granja is located 250km north-west of NKMNP, at the geological boundary between the Pre-Cambrian Shield which supports terra firme evergreen forest and the sedimentary basin of the Beni region which supports a mosaic of seasonally inundated savannah and forest islands. L. Granja is a small (0.2km2) oxbow lake surrounded by riparian forest at the lake margins, transitioning to terra firme evergreen forest further away from the lake. A small area (0.3km2) to the east of the lake has been cleared for agriculture ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1177/0959683615581204","ISBN":"0959-6836","ISSN":"0959-6836","abstract":"The nature and extent of pre-Columbian (pre-AD 1492) human impact in Amazonia is a contentious issue. The Bolivian Amazon has yielded some of the most impressive evidence for large and complex pre-Columbian societies in the Amazon basin, yet there remains relatively little data concerning the land use of these societies over time. Palaeoecology, when integrated with archaeological data, has the potential to fill these gaps in our knowledge. We present a 6000-year record of anthropogenic burning, agriculture and vegetation change, from an oxbow lake located adjacent to a pre-Columbian ring ditch in north-east Bolivia (13°15′44″S, 63°42′37″W). Human occupation around the lake site is inferred from pollen and phytoliths of maize (Zea mays L.) and macroscopic charcoal evidence of anthropogenic burning. First occupation around the lake was radiocarbon dated to ~2500 calibrated years before present (BP). The persistence of maize in the record from ~1850 BP suggests that it was an important crop grown in the ring-ditch region in pre-Columbian times, and abundant macroscopic charcoal suggests that pre-Columbian land management entailed more extensive burning of the landscape than the slash-and-burn agriculture practised around the site today. The site was occupied continuously until near-modern times, although there is evidence for a decline in agricultural intensity or change in land-use strategy, and possible population decline, from ~600–500 BP. The long and continuous occupation, which predates the establishment of rainforest in the region, suggests that pre-Columbian land use may have had a significant influence on ecosystem development at this site over the last ~2000 years.","author":[{"dropping-particle":"","family":"Carson","given":"J. F.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Watling","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"F. E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Whitney","given":"B. S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Prumers","given":"H.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Soto","given":"J. D.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"The Holocene","id":"ITEM-1","issue":"8","issued":{"date-parts":[["2015","5","1"]]},"note":"From Duplicate 2 (Pre-Columbian land use in the ring-ditch region of the Bolivian Amazon - Carson, J. F.; Watling, J.; Mayle, F. E.; Whitney, B. S.; Iriarte, J.; Prumers, H.; Soto, J. D.)\n\nUsed phytoliths and pollen from oxbow lake in NE Bolivia - oxbow 6000 yrs old.\n\nNice passage about phytoliths from lakes and their source area compared to pollen - assumes local representation due to modes of transport available.","page":"1285-1300","title":"Pre-Columbian land use in the ring-ditch region of the Bolivian Amazon","type":"article-journal","volume":"25"},"uris":[""]}],"mendeley":{"formattedCitation":"(Carson et al., 2015)","plainTextFormattedCitation":"(Carson et al., 2015)","previouslyFormattedCitation":"(Carson et al., 2015)"},"properties":{"noteIndex":0},"schema":""}(Carson et al., 2015). The second site chosen for application of the pollen ecosystem turnover threshold to the palaeo-record is Laguna Chaplin, as it is a well-studied example of ecosystem turnover from savannah to evergreen forest ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.yqres.2003.12.004","ISSN":"00335894","abstract":"Pollen and charcoal records from two large, shallow lakes reveal that throughout most of the past 50,000 yr Noel Kempff Mercado National Park, in northeastern lowland Bolivia (southwestern Amazon Basin), was predominantly covered by savannas and seasonally dry semideciduous forests. Lowered atmospheric CO2 concentrations, in combination with a longer dry season, caused expansion of dry forests and savannas during the last glacial period, especially at the last glacial maximum. These ecosystems persisted until the mid-Holocene, although they underwent significant species reassortment. Forest communities containing a mixture of evergreen and semideciduous species began to expand between 6000 and 3000 14C yr B.P. Humid evergreen rain forests expanded to cover most of the area within the past 2000 14C yr B.P., coincident with a reduction in fire frequencies. Comparisons between modern pollen spectra and vegetation reveal that the Moraceae-dominated rain forest pollen spectra likely have a regional source area at least 2–3 km beyond the lake shore, whereas the grass- and sedge-dominated savanna pollen spectra likely have a predominantly local source area. The Holocene vegetation changes are consistent with independent paleoprecipitation records from the Bolivian Altiplano and paleovegetation records from other parts of southwestern Amazonia. The progressive expansion in rain forests through the Holocene can be largely attributed to enhanced convective activity over Amazonia, due to greater seasonality of insolation in the Southern Hemisphere tropics driven by the precession cycle according to the Milankovitch Astronomical Theory.","author":[{"dropping-particle":"","family":"Burbridge","given":"Rachel E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Quaternary Research","id":"ITEM-1","issue":"2","issued":{"date-parts":[["2004","3"]]},"note":"chapling and Bella Vista records - dry forest/savanna until 2000 bp when precip increased - humid evergreen","page":"215-230","title":"Fifty-thousand-year vegetation and climate history of Noel Kempff Mercado National Park, Bolivian Amazon","type":"article-journal","volume":"61"},"uris":[""]},{"id":"ITEM-2","itemData":{"DOI":"10.1126/science.290.5500.2291","ISSN":"0036-8075","PMID":"11125139","abstract":"Amazonian rain forest-savanna boundaries are highly sensitive to climatic change and may also play an important role in rain forest speciation. However, their dynamics over millennial time scales are poorly understood. Here, we present late Quaternary pollen records from the southern margin of Amazonia, which show that the humid evergreen rain forests of eastern Bolivia have been expanding southward over the past 3000 years and that their present-day limit represents the southernmost extent of Amazonian rain forest over at least the past 50,000 years. This rain forest expansion is attributed to increased seasonal latitudinal migration of the Intertropical Convergence Zone, which can in turn be explained by Milankovitch astronomic forcing.","author":[{"dropping-particle":"","family":"Mayle","given":"F E","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Burbridge","given":"R","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"T J","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Science (New York, N.Y.)","id":"ITEM-2","issue":"2000","issued":{"date-parts":[["2000"]]},"page":"2291-2294","title":"Millennial-scale dynamics of southern Amazonian rain forests.","type":"article-journal","volume":"290"},"uris":[""]}],"mendeley":{"formattedCitation":"(Burbridge et al., 2004; Mayle et al., 2000)","plainTextFormattedCitation":"(Burbridge et al., 2004; Mayle et al., 2000)","previouslyFormattedCitation":"(Burbridge et al., 2004; Mayle et al., 2000)"},"properties":{"noteIndex":0},"schema":""}(Burbridge et al., 2004; Mayle et al., 2000). However, only pollen data are available from this site. L. Chaplin is a large (4-6km diameter), flat-bottomed lake situated within NKMNP on Pre-Cambrian Shield bedrock, surrounded by terra firme evergreen forest with areas of riparian forest lining nearby rivers and streams ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.yqres.2003.12.004","ISSN":"00335894","abstract":"Pollen and charcoal records from two large, shallow lakes reveal that throughout most of the past 50,000 yr Noel Kempff Mercado National Park, in northeastern lowland Bolivia (southwestern Amazon Basin), was predominantly covered by savannas and seasonally dry semideciduous forests. Lowered atmospheric CO2 concentrations, in combination with a longer dry season, caused expansion of dry forests and savannas during the last glacial period, especially at the last glacial maximum. These ecosystems persisted until the mid-Holocene, although they underwent significant species reassortment. Forest communities containing a mixture of evergreen and semideciduous species began to expand between 6000 and 3000 14C yr B.P. Humid evergreen rain forests expanded to cover most of the area within the past 2000 14C yr B.P., coincident with a reduction in fire frequencies. Comparisons between modern pollen spectra and vegetation reveal that the Moraceae-dominated rain forest pollen spectra likely have a regional source area at least 2–3 km beyond the lake shore, whereas the grass- and sedge-dominated savanna pollen spectra likely have a predominantly local source area. The Holocene vegetation changes are consistent with independent paleoprecipitation records from the Bolivian Altiplano and paleovegetation records from other parts of southwestern Amazonia. The progressive expansion in rain forests through the Holocene can be largely attributed to enhanced convective activity over Amazonia, due to greater seasonality of insolation in the Southern Hemisphere tropics driven by the precession cycle according to the Milankovitch Astronomical Theory.","author":[{"dropping-particle":"","family":"Burbridge","given":"Rachel E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Quaternary Research","id":"ITEM-1","issue":"2","issued":{"date-parts":[["2004","3"]]},"note":"chapling and Bella Vista records - dry forest/savanna until 2000 bp when precip increased - humid evergreen","page":"215-230","title":"Fifty-thousand-year vegetation and climate history of Noel Kempff Mercado National Park, Bolivian Amazon","type":"article-journal","volume":"61"},"uris":[""]}],"mendeley":{"formattedCitation":"(Burbridge et al., 2004)","plainTextFormattedCitation":"(Burbridge et al., 2004)","previouslyFormattedCitation":"(Burbridge et al., 2004)"},"properties":{"noteIndex":0},"schema":""}(Burbridge et al., 2004). 3. MethodsThis paper contains new analyses of previously published datasets ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.revpalbo.2008.06.007","ISSN":"0034-6667","abstract":"Accurate differentiation between tropical forest and savannah ecosystems in the fossil pollen record is hampered by the combination of: i) poor taxonomic resolution in pollen identification, and ii) the high species diversity of many lowland tropical families, i.e. with many different growth forms living in numerous environmental settings. These barriers to interpreting the fossil record hinder our understanding of the past distributions of different Neotropical ecosystems and consequently cloud our knowledge of past climatic, biodiversity and carbon storage patterns. Modern pollen studies facilitate an improved understanding of how ecosystems are represented by the pollen their plants produce and therefore aid interpretation of fossil pollen records. To understand how to differentiate ecosystems palynologically, it is essential that a consistent sampling method is used across ecosystems. However, to date, modern pollen studies from tropical South America have employed a variety of methodologies (e.g. pollen traps, moss polsters, soil samples). In this paper, we present the first modern pollen study from the Neotropics to examine the modern pollen rain from moist evergreen tropical forest (METF), semi-deciduous dry tropical forest (SDTF) and wooded savannah (cerrad?o) using a consistent sampling methodology (pollen traps). Pollen rain was sampled annually in September for the years 1999–2001 from within permanent vegetation study plots in, or near, the Noel Kempff Mercado National Park (NKMNP), Bolivia. Comparison of the modern pollen rain within these plots with detailed floristic inventories allowed estimates of the relative pollen productivity and dispersal for individual taxa to be made (% pollen/% vegetation or ‘p/v’). The applicability of these data to interpreting fossil records from lake sediments was then explored by comparison with pollen assemblages obtained from five lake surface samples. Pollen productivity is demonstrated to vary inter-annually and conforms to a consistent hierarchy for any given year: METF > SDTF > cerrad?o. This suggests an association between pollen productivity and basic structural characteristics of the ecosystem, i.e. closed canopy vs. open canopy vs. savannah. Comparison of modern pollen and vegetation revealed that some important floristic elements were completely absent from the pollen: Qualea and Erisma (METF), Bauhinia, Simira and Guazuma (SDTF), and Pouteria and Caryocar (cerrad?o). Anadenanthera was foun…","author":[{"dropping-particle":"","family":"Gosling","given":"William D.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Tate","given":"Nicholas J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Review of Palaeobotany and Palynology","id":"ITEM-1","issue":"1–2","issued":{"date-parts":[["2009","1"]]},"note":"Uses a surface sample from Mandiore = 51.6% Poaceae pollen at 0-1cm depth.","page":"70-85","title":"Differentiation between Neotropical rainforest, dry forest, and savannah ecosystems by their modern pollen spectra and implications for the fossil pollen record","type":"article-journal","volume":"153"},"uris":[""]},{"id":"ITEM-2","itemData":{"DOI":"10.1016/j.palaeo.2010.05.009","ISSN":"0031-0182","abstract":"An ongoing controversy in Amazonian palaeoecology is the manner in which Amazonian rainforest communities have responded to environmental change over the last glacial–interglacial cycle. Much of this controversy results from an inability to identify the floristic heterogeneity exhibited by rainforest communities within fossil pollen records. We apply multivariate (Principal Components Analysis) and classification (Unweighted Pair Group with Arithmetic Mean Agglomerative Classification) techniques to floral-biometric, modern pollen trap and lake sediment pollen data situated within different rainforest communities in the tropical lowlands of Amazonian Bolivia. Modern pollen rain analyses from artificial pollen traps show that evergreen terra firme (well-drained), evergreen terra firme liana, evergreen seasonally inundated, and evergreen riparian rainforests may be readily differentiated, floristically and palynologically. Analogue matching techniques, based on Euclidean distance measures, are employed to compare these pollen signatures with surface sediment pollen assemblages from five lakes: Laguna Bella Vista, Laguna Chaplin, and Laguna Huachi situated within the Madeira-Tapajós moist forest ecoregion, and Laguna Isirere and Laguna Loma Suarez, which are situated within forest patches in the Beni savanna ecoregion. The same numerical techniques are used to compare rainforest pollen trap signatures with the fossil pollen record of Laguna Chaplin.\n\nPollen assemblages of pollen traps situated within riparian forest communities are most similar to surface sediment samples from Lagunas Bella Vista and Chaplin. Pollen derived from terra firme forests also comprises a significant proportion of these assemblages. Together, these pollen spectra successfully identify riparian and terra firme rainforest communities surrounding the two lakes today. Close similarity between terra firme liana pollen trap assemblages and surface samples obtained from Laguna Huachi, a lake surrounded by relatively undisturbed liana forest, suggests liana forest pollen rain may also be identified within lake sediment records. Pollen spectra obtained from surface sediment samples from lakes situated within gallery forests of the Beni savanna ecoregion are significantly different to those obtained from the Madeira-Tapajós ecoregion, reflecting their different floristic compositions. By applying our findings to the previously published Laguna Chaplin Quaternary pollen record, we show that…","author":[{"dropping-particle":"","family":"Burn","given":"Michael J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Palaeogeography, Palaeoclimatology, Palaeoecology","id":"ITEM-2","issue":"1–2","issued":{"date-parts":[["2010","9"]]},"page":"1-18","title":"Pollen-based differentiation of Amazonian rainforest communities and implications for lowland palaeoecology in tropical South America","type":"article-journal","volume":"295"},"uris":[""]},{"id":"ITEM-3","itemData":{"DOI":"10.1016/j.revpalbo.2011.01.001","ISSN":"00346667","abstract":"The majority of vegetation reconstructions from the Neotropics are derived from fossil pollen records extracted from lake sediments. However, the interpretation of these records is restricted by limited knowledge of the contemporary relationships between the vegetation and pollen rain of Neotropical ecosystems, especially for more open vegetation such as savannas. This research aims to improve the interpretation of these records by investigating the vegetation and modern pollen rain of different savanna ecosystems in Bolivia using vegetation inventories, artificial pollen traps and surface lake sediments. Two types of savanna were studied, upland savannas (cerrado), occurring on well drained soils, and seasonally-inundated savannas occurring on seasonally water-logged soils. Quantitative vegetation data are used to identify taxa that are floristically important in the different savanna types and to allow modern pollen/vegetation ratios to be calculated. Artificial pollen traps from the upland savanna site are dominated by Moraceae (35%), Poaceae (30%), Alchornea (6%) and Cecropia (4%). The two seasonally-inundated savanna sites are dominated by Moraceae (37%), Poaceae (20%), Alchornea (8%) and Cecropia (7%), and Moraceae (25%), Cyperaceae (22%), Poaceae (19%) and Cecropia (9%), respectively. The modern pollen rain of seasonally-inundated savannas from surface lake sediments is dominated by Cyperaceae (35%), Poaceae (33%), Moraceae (9%) and Asteraceae (5%). Upland and seasonally-flooded savannas were found to be only subtly distinct from each other palynologically. All sites have a high proportion of Moraceae pollen due to effective wind dispersal of this pollen type from areas of evergreen forest close to the study sites. Modern pollen/vegetation ratios show that many key woody plant taxa are absent/under-represented in the modern pollen rain (e.g., Caryocar and Tabebuia). The lower-than-expected percentages of Poaceae pollen, and the scarcity of savanna indicators, in the modern pollen rain of these ecosystems mean that savannas could potentially be overlooked in fossil pollen records without consideration of the full pollen spectrum available.","author":[{"dropping-particle":"","family":"Jones","given":"Huw T.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Pennington","given":"R. Toby","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Review of Palaeobotany and Palynology","id":"ITEM-3","issue":"3-4","issued":{"date-parts":[["2011","4"]]},"page":"223-237","title":"Characterisation of Bolivian savanna ecosystems by their modern pollen rain and implications for fossil pollen records","type":"article-journal","volume":"164"},"uris":[""]},{"id":"ITEM-4","itemData":{"DOI":"10.1016/j.revpalbo.2013.01.004","ISSN":"00346667","abstract":"The interpretation of Neotropical fossil phytolith assemblages for palaeoenvironmental and archaeological reconstructions relies on the development of appropriate modern analogues. We analyzed modern phytolith assemblages from the soils of ten distinctive tropical vegetation communities in eastern lowland Bolivia, ranging from terra firme humid evergreen forest to seasonally-inundated savannah. Results show that broad ecosystems – evergreen tropical forest, semi-deciduous dry tropical forest, and savannah – can be clearly differentiated by examination of their phytolith spectra and the application of Principal Component Analysis (PCA). Differences in phytolith assemblages between particular vegetation communities within each of these ecosystems are more subtle, but can still be identified. Comparison of phytolith assemblages with pollen rain data and stable carbon isotope analyses from the same vegetation plots show that these proxies are not only complementary, but significantly improve taxonomic and ecosystem resolution, and therefore our ability to interpret palaeoenvironmental and archaeological records. Our data underline the utility of phytolith analyses for reconstructing Amazon Holocene vegetation histories and pre-Columbian land use, particularly the high spatial resolution possible with terrestrial soil-based phytolith studies.","author":[{"dropping-particle":"","family":"Dickau","given":"Ruth","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Whitney","given":"Bronwen S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"José","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Soto","given":"J. Daniel","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Metcalfe","given":"Phil","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Street-Perrott","given":"F. Alayne","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Loader","given":"Neil J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Ficken","given":"Katherine J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Review of Palaeobotany and Palynology","id":"ITEM-4","issued":{"date-parts":[["2013","6"]]},"page":"15-37","publisher":"Elsevier B.V.","title":"Differentiation of neotropical ecosystems by modern soil phytolith assemblages and its implications for palaeoenvironmental and archaeological reconstructions","type":"article-journal","volume":"193"},"uris":[""]},{"id":"ITEM-5","itemData":{"abstract":"At present there is uncertainty over the response of neotropical ecosystems to the climatic changes of the Quaternary. The majority of vegetation reconstructions from the region are derived from fossil pollen records extracted from lake sediments. However, the interpretation of these records is restricted by limited knowledge of the contemporary relationships between the vegetation and pollen rain of neotropical ecosystems, especially for more open vegetation such as savanna and dry forest. This research aims to improve the interpretation of these records by investigating the relationship between the vegetation and modern pollen rain of different savanna and seasonally dry tropical forest (SDTF) ecosystems in Bolivia using artificial pollen traps and surface lake sediments to analyse the modern pollen rain. Vegetation data is used to identify taxa that are floristically important within the different ecosystems and to allow modern pollen/vegetation ratios to be calculated. The modern pollen rain from the upland savanna is dominated by Moraceae/Urticaceae (35.1%), Poaceae (29.6%), Alchornea (6.1%) and Cecropia (4.1%), whilst the seasonally-inundated savanna sites are dominated by Moraceae/Urticaceae (30.7%), Poaceae (19.5%), Cyperaceae (14.0%) and Cecropia (7.9%). These two different savanna ecosystems are only slightly differentiated by their modern pollen rain. The main taxa in the modern pollen rain of the upland SDTF are Moraceae/Urticaceae (25.8%), Cecropia (10.5%), Acalypha (7.6%) and Combretaceae/Melastomataceae (6.7%). Seasonally-inundated SDTF is dominated by Cecropia pollen to the extent that it was removed from the pollen sum and the main non-Cecropia pollen types are Moraceae/Urticaceae (39.0%), unknown type df 61 (6.4%), Asteraceae (6.3%), Celtis (6.0%) and Physocalymma scaberrimum (4.9%). These two SDTF ecosystems are well differentiated by their modern pollen rain, implying that they may be defined in fossil pollen records. The modern pollen rain obtained from the surface lake samples is generally complementary to that obtained from the artificial pollen traps for a given ecosystem. All sites have a high Moraceae/Urticaceae pollen signal due to effective dispersal of this pollen type from areas of evergreen forest in close proximity to the study sites. The savanna sites show lower Poaceae percentages than have been previously reported in the literature by some authors and this raises the possibility than the extent of this ecosystem in the…","author":[{"dropping-particle":"","family":"Jones","given":"Huw T.","non-dropping-particle":"","parse-names":false,"suffix":""}],"id":"ITEM-5","issued":{"date-parts":[["2009"]]},"publisher":"The University of Edinburgh","title":"Characterisation of neotropical savanna and seasonally dry forest ecosystems by their modern pollen rain","type":"thesis"},"uris":[""]}],"mendeley":{"formattedCitation":"(Burn et al., 2010; Dickau et al., 2013; Gosling et al., 2009; Jones, 2009; Jones et al., 2011)","plainTextFormattedCitation":"(Burn et al., 2010; Dickau et al., 2013; Gosling et al., 2009; Jones, 2009; Jones et al., 2011)","previouslyFormattedCitation":"(Burn et al., 2010; Dickau et al., 2013; Gosling et al., 2009; Jones, 2009; Jones et al., 2011)"},"properties":{"noteIndex":0},"schema":""}(Burn et al., 2010; Dickau et al., 2013; Gosling et al., 2009; Jones, 2009; Jones et al., 2011) that have been utilised with authors’ permission (raw data plotted in Figures SI and S2). Field and laboratory methods used by each of these authors are summarised below; see original papers for full details. 3.1 Field methodsPermanent 1-hectare (500 x 20m) vegetation study plots from Noel Kempff Mercado National Park (NKMNP) were sampled for pollen rain and soil-surface phytoliths. Pollen rain was collected from artificial pollen traps ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.yqres.2005.08.008","ISSN":"00335894","abstract":"The paucity of modern pollen-rain data from Amazonia constitutes a significant barrier to understanding the Late Quaternary vegetation history of this globally important tropical forest region. Here, we present the first modern pollen-rain data for tall terra firme moist evergreen Amazon forest, collected between 1999 and 2001 from artificial pollen traps within a 500?×?20 m permanent study plot (14°34′50″S, 60°49′48″W) in Noel Kempff Mercado National Park (NE Bolivia). Spearman's rank correlations were performed to assess the extent of spatial and inter-annual variability in the pollen rain, whilst statistically distinctive taxa were identified using Principal Components Analysis (PCA). Comparisons with the floristic and basal area data of the plot (stems ≥10 cm d.b.h.) enabled the degree to which taxa are over/under-represented in the pollen rain to be assessed (using R-rel values). Moraceae/Urticaceae dominates the pollen rain (64% median abundance) and is also an important constituent of the vegetation, accounting for 16% of stems ≥10 cm d.b.h. and ca. 11% of the total basal area. Other important pollen taxa are Arecaceae (cf. Euterpe), Melastomataceae/Combretaceae, Cecropia, Didymopanax, Celtis, and Alchornea. However, 75% of stems and 67% of the total basal area of the plot ≥10 cm d.b.h. belong to species which are unidentified in the pollen rain, the most important of which are Phenakospermum guianensis (a banana-like herb) and the key canopy-emergent trees, Erisma uncinatum and Qualea paraensis.","author":[{"dropping-particle":"","family":"Gosling","given":"William D.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Tate","given":"Nicholas J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"collection-title":"Late Quaternary Tropical Ecosystem Dynamics","container-title":"Quaternary Research","id":"ITEM-1","issue":"3","issued":{"date-parts":[["2005","11"]]},"page":"284-297","title":"Modern pollen-rain characteristics of tall terra firme moist evergreen forest, southern Amazonia","type":"article-journal","volume":"64"},"uris":[""]},{"id":"ITEM-2","itemData":{"DOI":"10.1016/j.revpalbo.2008.06.007","ISSN":"0034-6667","abstract":"Accurate differentiation between tropical forest and savannah ecosystems in the fossil pollen record is hampered by the combination of: i) poor taxonomic resolution in pollen identification, and ii) the high species diversity of many lowland tropical families, i.e. with many different growth forms living in numerous environmental settings. These barriers to interpreting the fossil record hinder our understanding of the past distributions of different Neotropical ecosystems and consequently cloud our knowledge of past climatic, biodiversity and carbon storage patterns. Modern pollen studies facilitate an improved understanding of how ecosystems are represented by the pollen their plants produce and therefore aid interpretation of fossil pollen records. To understand how to differentiate ecosystems palynologically, it is essential that a consistent sampling method is used across ecosystems. However, to date, modern pollen studies from tropical South America have employed a variety of methodologies (e.g. pollen traps, moss polsters, soil samples). In this paper, we present the first modern pollen study from the Neotropics to examine the modern pollen rain from moist evergreen tropical forest (METF), semi-deciduous dry tropical forest (SDTF) and wooded savannah (cerrad?o) using a consistent sampling methodology (pollen traps). Pollen rain was sampled annually in September for the years 1999–2001 from within permanent vegetation study plots in, or near, the Noel Kempff Mercado National Park (NKMNP), Bolivia. Comparison of the modern pollen rain within these plots with detailed floristic inventories allowed estimates of the relative pollen productivity and dispersal for individual taxa to be made (% pollen/% vegetation or ‘p/v’). The applicability of these data to interpreting fossil records from lake sediments was then explored by comparison with pollen assemblages obtained from five lake surface samples. Pollen productivity is demonstrated to vary inter-annually and conforms to a consistent hierarchy for any given year: METF > SDTF > cerrad?o. This suggests an association between pollen productivity and basic structural characteristics of the ecosystem, i.e. closed canopy vs. open canopy vs. savannah. Comparison of modern pollen and vegetation revealed that some important floristic elements were completely absent from the pollen: Qualea and Erisma (METF), Bauhinia, Simira and Guazuma (SDTF), and Pouteria and Caryocar (cerrad?o). Anadenanthera was foun…","author":[{"dropping-particle":"","family":"Gosling","given":"William D.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Tate","given":"Nicholas J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Review of Palaeobotany and Palynology","id":"ITEM-2","issue":"1–2","issued":{"date-parts":[["2009","1"]]},"note":"Uses a surface sample from Mandiore = 51.6% Poaceae pollen at 0-1cm depth.","page":"70-85","title":"Differentiation between Neotropical rainforest, dry forest, and savannah ecosystems by their modern pollen spectra and implications for the fossil pollen record","type":"article-journal","volume":"153"},"uris":[""]},{"id":"ITEM-3","itemData":{"DOI":"10.1016/j.palaeo.2010.05.009","ISSN":"0031-0182","abstract":"An ongoing controversy in Amazonian palaeoecology is the manner in which Amazonian rainforest communities have responded to environmental change over the last glacial–interglacial cycle. Much of this controversy results from an inability to identify the floristic heterogeneity exhibited by rainforest communities within fossil pollen records. We apply multivariate (Principal Components Analysis) and classification (Unweighted Pair Group with Arithmetic Mean Agglomerative Classification) techniques to floral-biometric, modern pollen trap and lake sediment pollen data situated within different rainforest communities in the tropical lowlands of Amazonian Bolivia. Modern pollen rain analyses from artificial pollen traps show that evergreen terra firme (well-drained), evergreen terra firme liana, evergreen seasonally inundated, and evergreen riparian rainforests may be readily differentiated, floristically and palynologically. Analogue matching techniques, based on Euclidean distance measures, are employed to compare these pollen signatures with surface sediment pollen assemblages from five lakes: Laguna Bella Vista, Laguna Chaplin, and Laguna Huachi situated within the Madeira-Tapajós moist forest ecoregion, and Laguna Isirere and Laguna Loma Suarez, which are situated within forest patches in the Beni savanna ecoregion. The same numerical techniques are used to compare rainforest pollen trap signatures with the fossil pollen record of Laguna Chaplin.\n\nPollen assemblages of pollen traps situated within riparian forest communities are most similar to surface sediment samples from Lagunas Bella Vista and Chaplin. Pollen derived from terra firme forests also comprises a significant proportion of these assemblages. Together, these pollen spectra successfully identify riparian and terra firme rainforest communities surrounding the two lakes today. Close similarity between terra firme liana pollen trap assemblages and surface samples obtained from Laguna Huachi, a lake surrounded by relatively undisturbed liana forest, suggests liana forest pollen rain may also be identified within lake sediment records. Pollen spectra obtained from surface sediment samples from lakes situated within gallery forests of the Beni savanna ecoregion are significantly different to those obtained from the Madeira-Tapajós ecoregion, reflecting their different floristic compositions. By applying our findings to the previously published Laguna Chaplin Quaternary pollen record, we show that…","author":[{"dropping-particle":"","family":"Burn","given":"Michael J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Palaeogeography, Palaeoclimatology, Palaeoecology","id":"ITEM-3","issue":"1–2","issued":{"date-parts":[["2010","9"]]},"page":"1-18","title":"Pollen-based differentiation of Amazonian rainforest communities and implications for lowland palaeoecology in tropical South America","type":"article-journal","volume":"295"},"uris":[""]},{"id":"ITEM-4","itemData":{"DOI":"10.1016/j.revpalbo.2011.01.001","ISSN":"00346667","abstract":"The majority of vegetation reconstructions from the Neotropics are derived from fossil pollen records extracted from lake sediments. However, the interpretation of these records is restricted by limited knowledge of the contemporary relationships between the vegetation and pollen rain of Neotropical ecosystems, especially for more open vegetation such as savannas. This research aims to improve the interpretation of these records by investigating the vegetation and modern pollen rain of different savanna ecosystems in Bolivia using vegetation inventories, artificial pollen traps and surface lake sediments. Two types of savanna were studied, upland savannas (cerrado), occurring on well drained soils, and seasonally-inundated savannas occurring on seasonally water-logged soils. Quantitative vegetation data are used to identify taxa that are floristically important in the different savanna types and to allow modern pollen/vegetation ratios to be calculated. Artificial pollen traps from the upland savanna site are dominated by Moraceae (35%), Poaceae (30%), Alchornea (6%) and Cecropia (4%). The two seasonally-inundated savanna sites are dominated by Moraceae (37%), Poaceae (20%), Alchornea (8%) and Cecropia (7%), and Moraceae (25%), Cyperaceae (22%), Poaceae (19%) and Cecropia (9%), respectively. The modern pollen rain of seasonally-inundated savannas from surface lake sediments is dominated by Cyperaceae (35%), Poaceae (33%), Moraceae (9%) and Asteraceae (5%). Upland and seasonally-flooded savannas were found to be only subtly distinct from each other palynologically. All sites have a high proportion of Moraceae pollen due to effective wind dispersal of this pollen type from areas of evergreen forest close to the study sites. Modern pollen/vegetation ratios show that many key woody plant taxa are absent/under-represented in the modern pollen rain (e.g., Caryocar and Tabebuia). The lower-than-expected percentages of Poaceae pollen, and the scarcity of savanna indicators, in the modern pollen rain of these ecosystems mean that savannas could potentially be overlooked in fossil pollen records without consideration of the full pollen spectrum available.","author":[{"dropping-particle":"","family":"Jones","given":"Huw T.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Pennington","given":"R. Toby","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Review of Palaeobotany and Palynology","id":"ITEM-4","issue":"3-4","issued":{"date-parts":[["2011","4"]]},"page":"223-237","title":"Characterisation of Bolivian savanna ecosystems by their modern pollen rain and implications for fossil pollen records","type":"article-journal","volume":"164"},"uris":[""]}],"mendeley":{"formattedCitation":"(Burn et al., 2010; Gosling et al., 2005, 2009; Jones et al., 2011)","plainTextFormattedCitation":"(Burn et al., 2010; Gosling et al., 2005, 2009; Jones et al., 2011)","previouslyFormattedCitation":"(Burn et al., 2010; Gosling et al., 2005, 2009; Jones et al., 2011)"},"properties":{"noteIndex":0},"schema":""}(Burn et al., 2010; Gosling et al., 2005, 2009; Jones et al., 2011) and phytoliths were sampled from the soil layer underlying the soil leaf litter layer ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.revpalbo.2013.01.004","ISSN":"00346667","abstract":"The interpretation of Neotropical fossil phytolith assemblages for palaeoenvironmental and archaeological reconstructions relies on the development of appropriate modern analogues. We analyzed modern phytolith assemblages from the soils of ten distinctive tropical vegetation communities in eastern lowland Bolivia, ranging from terra firme humid evergreen forest to seasonally-inundated savannah. Results show that broad ecosystems – evergreen tropical forest, semi-deciduous dry tropical forest, and savannah – can be clearly differentiated by examination of their phytolith spectra and the application of Principal Component Analysis (PCA). Differences in phytolith assemblages between particular vegetation communities within each of these ecosystems are more subtle, but can still be identified. Comparison of phytolith assemblages with pollen rain data and stable carbon isotope analyses from the same vegetation plots show that these proxies are not only complementary, but significantly improve taxonomic and ecosystem resolution, and therefore our ability to interpret palaeoenvironmental and archaeological records. Our data underline the utility of phytolith analyses for reconstructing Amazon Holocene vegetation histories and pre-Columbian land use, particularly the high spatial resolution possible with terrestrial soil-based phytolith studies.","author":[{"dropping-particle":"","family":"Dickau","given":"Ruth","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Whitney","given":"Bronwen S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"José","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Soto","given":"J. Daniel","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Metcalfe","given":"Phil","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Street-Perrott","given":"F. Alayne","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Loader","given":"Neil J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Ficken","given":"Katherine J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Review of Palaeobotany and Palynology","id":"ITEM-1","issued":{"date-parts":[["2013","6"]]},"page":"15-37","publisher":"Elsevier B.V.","title":"Differentiation of neotropical ecosystems by modern soil phytolith assemblages and its implications for palaeoenvironmental and archaeological reconstructions","type":"article-journal","volume":"193"},"uris":[""]}],"mendeley":{"formattedCitation":"(Dickau et al., 2013)","plainTextFormattedCitation":"(Dickau et al., 2013)","previouslyFormattedCitation":"(Dickau et al., 2013)"},"properties":{"noteIndex":0},"schema":""}(Dickau et al., 2013). Sample collection was evenly spaced along a central transect through each plot. The ten pollen samples per plot were spaced 50m apart (Gosling et al., 2005) and the three phytolith samples per plot were 250m apart (Dickau et al., 2013). The spatial variability of pollen rain within terra firme evergreen forest plots was found to be high for some taxa (Hyeronmia, Trema, Cecropia, Didymopanax) and for Pollen Accumulation Rates (PARs). However, all samples were still dominated by Moraceae/Urticaceae, and there was no discernible bias in the pollen variability towards the taxon of the nearest tree. The majority of the spatial variability within the plots was captured by analysis of five of the ten pollen trap samples (Gosling et al., 2005). The samples used in this analysis are a subset of the full published datasets. The plots chosen for this analysis were selected based on the availability of both pollen and phytolith datasets; only plots with both datasets were included in the analysis. The nine plots that met these criteria span six plant communities and three ecosystem classes (Table 1). Twelve samples from within each ecosystem class were selected from the full published datasets. Samples were chosen to reflect an even distribution between different plant communities within each ecosystem class e.g. terra firme and seasonally-inundated savannas. Within plots, samples were chosen to represent an even spatial distribution within each plot (Table 1). Detailed vegetation inventories of forest plots record all taxa representing >1% of the total number of stems >10cm d.b.h. ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.yqres.2005.08.008","ISSN":"00335894","abstract":"The paucity of modern pollen-rain data from Amazonia constitutes a significant barrier to understanding the Late Quaternary vegetation history of this globally important tropical forest region. Here, we present the first modern pollen-rain data for tall terra firme moist evergreen Amazon forest, collected between 1999 and 2001 from artificial pollen traps within a 500?×?20 m permanent study plot (14°34′50″S, 60°49′48″W) in Noel Kempff Mercado National Park (NE Bolivia). Spearman's rank correlations were performed to assess the extent of spatial and inter-annual variability in the pollen rain, whilst statistically distinctive taxa were identified using Principal Components Analysis (PCA). Comparisons with the floristic and basal area data of the plot (stems ≥10 cm d.b.h.) enabled the degree to which taxa are over/under-represented in the pollen rain to be assessed (using R-rel values). Moraceae/Urticaceae dominates the pollen rain (64% median abundance) and is also an important constituent of the vegetation, accounting for 16% of stems ≥10 cm d.b.h. and ca. 11% of the total basal area. Other important pollen taxa are Arecaceae (cf. Euterpe), Melastomataceae/Combretaceae, Cecropia, Didymopanax, Celtis, and Alchornea. However, 75% of stems and 67% of the total basal area of the plot ≥10 cm d.b.h. belong to species which are unidentified in the pollen rain, the most important of which are Phenakospermum guianensis (a banana-like herb) and the key canopy-emergent trees, Erisma uncinatum and Qualea paraensis.","author":[{"dropping-particle":"","family":"Gosling","given":"William D.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Tate","given":"Nicholas J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"collection-title":"Late Quaternary Tropical Ecosystem Dynamics","container-title":"Quaternary Research","id":"ITEM-1","issue":"3","issued":{"date-parts":[["2005","11"]]},"page":"284-297","title":"Modern pollen-rain characteristics of tall terra firme moist evergreen forest, southern Amazonia","type":"article-journal","volume":"64"},"uris":[""]},{"id":"ITEM-2","itemData":{"DOI":"10.1016/j.revpalbo.2008.06.007","ISSN":"0034-6667","abstract":"Accurate differentiation between tropical forest and savannah ecosystems in the fossil pollen record is hampered by the combination of: i) poor taxonomic resolution in pollen identification, and ii) the high species diversity of many lowland tropical families, i.e. with many different growth forms living in numerous environmental settings. These barriers to interpreting the fossil record hinder our understanding of the past distributions of different Neotropical ecosystems and consequently cloud our knowledge of past climatic, biodiversity and carbon storage patterns. Modern pollen studies facilitate an improved understanding of how ecosystems are represented by the pollen their plants produce and therefore aid interpretation of fossil pollen records. To understand how to differentiate ecosystems palynologically, it is essential that a consistent sampling method is used across ecosystems. However, to date, modern pollen studies from tropical South America have employed a variety of methodologies (e.g. pollen traps, moss polsters, soil samples). In this paper, we present the first modern pollen study from the Neotropics to examine the modern pollen rain from moist evergreen tropical forest (METF), semi-deciduous dry tropical forest (SDTF) and wooded savannah (cerrad?o) using a consistent sampling methodology (pollen traps). Pollen rain was sampled annually in September for the years 1999–2001 from within permanent vegetation study plots in, or near, the Noel Kempff Mercado National Park (NKMNP), Bolivia. Comparison of the modern pollen rain within these plots with detailed floristic inventories allowed estimates of the relative pollen productivity and dispersal for individual taxa to be made (% pollen/% vegetation or ‘p/v’). The applicability of these data to interpreting fossil records from lake sediments was then explored by comparison with pollen assemblages obtained from five lake surface samples. Pollen productivity is demonstrated to vary inter-annually and conforms to a consistent hierarchy for any given year: METF > SDTF > cerrad?o. This suggests an association between pollen productivity and basic structural characteristics of the ecosystem, i.e. closed canopy vs. open canopy vs. savannah. Comparison of modern pollen and vegetation revealed that some important floristic elements were completely absent from the pollen: Qualea and Erisma (METF), Bauhinia, Simira and Guazuma (SDTF), and Pouteria and Caryocar (cerrad?o). Anadenanthera was foun…","author":[{"dropping-particle":"","family":"Gosling","given":"William D.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Tate","given":"Nicholas J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Review of Palaeobotany and Palynology","id":"ITEM-2","issue":"1–2","issued":{"date-parts":[["2009","1"]]},"note":"Uses a surface sample from Mandiore = 51.6% Poaceae pollen at 0-1cm depth.","page":"70-85","title":"Differentiation between Neotropical rainforest, dry forest, and savannah ecosystems by their modern pollen spectra and implications for the fossil pollen record","type":"article-journal","volume":"153"},"uris":[""]},{"id":"ITEM-3","itemData":{"DOI":"10.1016/j.palaeo.2010.05.009","ISSN":"0031-0182","abstract":"An ongoing controversy in Amazonian palaeoecology is the manner in which Amazonian rainforest communities have responded to environmental change over the last glacial–interglacial cycle. Much of this controversy results from an inability to identify the floristic heterogeneity exhibited by rainforest communities within fossil pollen records. We apply multivariate (Principal Components Analysis) and classification (Unweighted Pair Group with Arithmetic Mean Agglomerative Classification) techniques to floral-biometric, modern pollen trap and lake sediment pollen data situated within different rainforest communities in the tropical lowlands of Amazonian Bolivia. Modern pollen rain analyses from artificial pollen traps show that evergreen terra firme (well-drained), evergreen terra firme liana, evergreen seasonally inundated, and evergreen riparian rainforests may be readily differentiated, floristically and palynologically. Analogue matching techniques, based on Euclidean distance measures, are employed to compare these pollen signatures with surface sediment pollen assemblages from five lakes: Laguna Bella Vista, Laguna Chaplin, and Laguna Huachi situated within the Madeira-Tapajós moist forest ecoregion, and Laguna Isirere and Laguna Loma Suarez, which are situated within forest patches in the Beni savanna ecoregion. The same numerical techniques are used to compare rainforest pollen trap signatures with the fossil pollen record of Laguna Chaplin.\n\nPollen assemblages of pollen traps situated within riparian forest communities are most similar to surface sediment samples from Lagunas Bella Vista and Chaplin. Pollen derived from terra firme forests also comprises a significant proportion of these assemblages. Together, these pollen spectra successfully identify riparian and terra firme rainforest communities surrounding the two lakes today. Close similarity between terra firme liana pollen trap assemblages and surface samples obtained from Laguna Huachi, a lake surrounded by relatively undisturbed liana forest, suggests liana forest pollen rain may also be identified within lake sediment records. Pollen spectra obtained from surface sediment samples from lakes situated within gallery forests of the Beni savanna ecoregion are significantly different to those obtained from the Madeira-Tapajós ecoregion, reflecting their different floristic compositions. By applying our findings to the previously published Laguna Chaplin Quaternary pollen record, we show that…","author":[{"dropping-particle":"","family":"Burn","given":"Michael J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Palaeogeography, Palaeoclimatology, Palaeoecology","id":"ITEM-3","issue":"1–2","issued":{"date-parts":[["2010","9"]]},"page":"1-18","title":"Pollen-based differentiation of Amazonian rainforest communities and implications for lowland palaeoecology in tropical South America","type":"article-journal","volume":"295"},"uris":[""]}],"mendeley":{"formattedCitation":"(Burn et al., 2010; Gosling et al., 2005, 2009)","plainTextFormattedCitation":"(Burn et al., 2010; Gosling et al., 2005, 2009)","previouslyFormattedCitation":"(Burn et al., 2010; Gosling et al., 2005, 2009)"},"properties":{"noteIndex":0},"schema":""}(Burn et al., 2010; Gosling et al., 2005, 2009). These inventories do not capture understorey grasses, vines or small lianas. For the savannah plots, horizontal ground cover of each plant along line transects were recorded, full details on the SALVIAS and RAINFOR databases ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"URL":"","accessed":{"date-parts":[["2018","10","25"]]},"author":[{"dropping-particle":"","family":"SALVIAS","given":"","non-dropping-particle":"","parse-names":false,"suffix":""}],"id":"ITEM-1","issued":{"date-parts":[["2004"]]},"title":"SALVIAS: Synthesis and Analysis of Local Vegetation Inventories Across Scales","type":"webpage"},"uris":[""]},{"id":"ITEM-2","itemData":{"author":[{"dropping-particle":"","family":"Malhi","given":"Y.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Phillips","given":"O.L.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Llloyd","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Baker","given":"T.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Wright","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Almedia","given":"S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Arroyo","given":"L.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Journal of Vegetation Science","id":"ITEM-2","issue":"3","issued":{"date-parts":[["2002"]]},"page":"439-450","title":"An international network to monitor the structure, composition and dynamics of Amazonian forests (RAINFOR)","type":"article-journal","volume":"13"},"uris":[""]}],"mendeley":{"formattedCitation":"(Malhi et al., 2002; SALVIAS, 2004)","plainTextFormattedCitation":"(Malhi et al., 2002; SALVIAS, 2004)","previouslyFormattedCitation":"(Malhi et al., 2002; SALVIAS, 2004)"},"properties":{"noteIndex":0},"schema":""}(Malhi et al., 2002; SALVIAS, 2004). 3.2 Laboratory methodsPollen trap samples were processed using the ‘wash’ methodology and prepared for microscopy. Details of the methodology are available in the original publications ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.revpalbo.2008.06.007","ISSN":"0034-6667","abstract":"Accurate differentiation between tropical forest and savannah ecosystems in the fossil pollen record is hampered by the combination of: i) poor taxonomic resolution in pollen identification, and ii) the high species diversity of many lowland tropical families, i.e. with many different growth forms living in numerous environmental settings. These barriers to interpreting the fossil record hinder our understanding of the past distributions of different Neotropical ecosystems and consequently cloud our knowledge of past climatic, biodiversity and carbon storage patterns. Modern pollen studies facilitate an improved understanding of how ecosystems are represented by the pollen their plants produce and therefore aid interpretation of fossil pollen records. To understand how to differentiate ecosystems palynologically, it is essential that a consistent sampling method is used across ecosystems. However, to date, modern pollen studies from tropical South America have employed a variety of methodologies (e.g. pollen traps, moss polsters, soil samples). In this paper, we present the first modern pollen study from the Neotropics to examine the modern pollen rain from moist evergreen tropical forest (METF), semi-deciduous dry tropical forest (SDTF) and wooded savannah (cerrad?o) using a consistent sampling methodology (pollen traps). Pollen rain was sampled annually in September for the years 1999–2001 from within permanent vegetation study plots in, or near, the Noel Kempff Mercado National Park (NKMNP), Bolivia. Comparison of the modern pollen rain within these plots with detailed floristic inventories allowed estimates of the relative pollen productivity and dispersal for individual taxa to be made (% pollen/% vegetation or ‘p/v’). The applicability of these data to interpreting fossil records from lake sediments was then explored by comparison with pollen assemblages obtained from five lake surface samples. Pollen productivity is demonstrated to vary inter-annually and conforms to a consistent hierarchy for any given year: METF > SDTF > cerrad?o. This suggests an association between pollen productivity and basic structural characteristics of the ecosystem, i.e. closed canopy vs. open canopy vs. savannah. Comparison of modern pollen and vegetation revealed that some important floristic elements were completely absent from the pollen: Qualea and Erisma (METF), Bauhinia, Simira and Guazuma (SDTF), and Pouteria and Caryocar (cerrad?o). Anadenanthera was foun…","author":[{"dropping-particle":"","family":"Gosling","given":"William D.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Tate","given":"Nicholas J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Review of Palaeobotany and Palynology","id":"ITEM-1","issue":"1–2","issued":{"date-parts":[["2009","1"]]},"note":"Uses a surface sample from Mandiore = 51.6% Poaceae pollen at 0-1cm depth.","page":"70-85","title":"Differentiation between Neotropical rainforest, dry forest, and savannah ecosystems by their modern pollen spectra and implications for the fossil pollen record","type":"article-journal","volume":"153"},"uris":[""]},{"id":"ITEM-2","itemData":{"DOI":"10.1016/j.palaeo.2010.05.009","ISSN":"0031-0182","abstract":"An ongoing controversy in Amazonian palaeoecology is the manner in which Amazonian rainforest communities have responded to environmental change over the last glacial–interglacial cycle. Much of this controversy results from an inability to identify the floristic heterogeneity exhibited by rainforest communities within fossil pollen records. We apply multivariate (Principal Components Analysis) and classification (Unweighted Pair Group with Arithmetic Mean Agglomerative Classification) techniques to floral-biometric, modern pollen trap and lake sediment pollen data situated within different rainforest communities in the tropical lowlands of Amazonian Bolivia. Modern pollen rain analyses from artificial pollen traps show that evergreen terra firme (well-drained), evergreen terra firme liana, evergreen seasonally inundated, and evergreen riparian rainforests may be readily differentiated, floristically and palynologically. Analogue matching techniques, based on Euclidean distance measures, are employed to compare these pollen signatures with surface sediment pollen assemblages from five lakes: Laguna Bella Vista, Laguna Chaplin, and Laguna Huachi situated within the Madeira-Tapajós moist forest ecoregion, and Laguna Isirere and Laguna Loma Suarez, which are situated within forest patches in the Beni savanna ecoregion. The same numerical techniques are used to compare rainforest pollen trap signatures with the fossil pollen record of Laguna Chaplin.\n\nPollen assemblages of pollen traps situated within riparian forest communities are most similar to surface sediment samples from Lagunas Bella Vista and Chaplin. Pollen derived from terra firme forests also comprises a significant proportion of these assemblages. Together, these pollen spectra successfully identify riparian and terra firme rainforest communities surrounding the two lakes today. Close similarity between terra firme liana pollen trap assemblages and surface samples obtained from Laguna Huachi, a lake surrounded by relatively undisturbed liana forest, suggests liana forest pollen rain may also be identified within lake sediment records. Pollen spectra obtained from surface sediment samples from lakes situated within gallery forests of the Beni savanna ecoregion are significantly different to those obtained from the Madeira-Tapajós ecoregion, reflecting their different floristic compositions. By applying our findings to the previously published Laguna Chaplin Quaternary pollen record, we show that…","author":[{"dropping-particle":"","family":"Burn","given":"Michael J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Palaeogeography, Palaeoclimatology, Palaeoecology","id":"ITEM-2","issue":"1–2","issued":{"date-parts":[["2010","9"]]},"page":"1-18","title":"Pollen-based differentiation of Amazonian rainforest communities and implications for lowland palaeoecology in tropical South America","type":"article-journal","volume":"295"},"uris":[""]},{"id":"ITEM-3","itemData":{"DOI":"10.1016/j.revpalbo.2011.01.001","ISSN":"00346667","abstract":"The majority of vegetation reconstructions from the Neotropics are derived from fossil pollen records extracted from lake sediments. However, the interpretation of these records is restricted by limited knowledge of the contemporary relationships between the vegetation and pollen rain of Neotropical ecosystems, especially for more open vegetation such as savannas. This research aims to improve the interpretation of these records by investigating the vegetation and modern pollen rain of different savanna ecosystems in Bolivia using vegetation inventories, artificial pollen traps and surface lake sediments. Two types of savanna were studied, upland savannas (cerrado), occurring on well drained soils, and seasonally-inundated savannas occurring on seasonally water-logged soils. Quantitative vegetation data are used to identify taxa that are floristically important in the different savanna types and to allow modern pollen/vegetation ratios to be calculated. Artificial pollen traps from the upland savanna site are dominated by Moraceae (35%), Poaceae (30%), Alchornea (6%) and Cecropia (4%). The two seasonally-inundated savanna sites are dominated by Moraceae (37%), Poaceae (20%), Alchornea (8%) and Cecropia (7%), and Moraceae (25%), Cyperaceae (22%), Poaceae (19%) and Cecropia (9%), respectively. The modern pollen rain of seasonally-inundated savannas from surface lake sediments is dominated by Cyperaceae (35%), Poaceae (33%), Moraceae (9%) and Asteraceae (5%). Upland and seasonally-flooded savannas were found to be only subtly distinct from each other palynologically. All sites have a high proportion of Moraceae pollen due to effective wind dispersal of this pollen type from areas of evergreen forest close to the study sites. Modern pollen/vegetation ratios show that many key woody plant taxa are absent/under-represented in the modern pollen rain (e.g., Caryocar and Tabebuia). The lower-than-expected percentages of Poaceae pollen, and the scarcity of savanna indicators, in the modern pollen rain of these ecosystems mean that savannas could potentially be overlooked in fossil pollen records without consideration of the full pollen spectrum available.","author":[{"dropping-particle":"","family":"Jones","given":"Huw T.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Pennington","given":"R. Toby","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Review of Palaeobotany and Palynology","id":"ITEM-3","issue":"3-4","issued":{"date-parts":[["2011","4"]]},"page":"223-237","title":"Characterisation of Bolivian savanna ecosystems by their modern pollen rain and implications for fossil pollen records","type":"article-journal","volume":"164"},"uris":[""]}],"mendeley":{"formattedCitation":"(Burn et al., 2010; Gosling et al., 2009; Jones et al., 2011)","plainTextFormattedCitation":"(Burn et al., 2010; Gosling et al., 2009; Jones et al., 2011)","previouslyFormattedCitation":"(Burn et al., 2010; Gosling et al., 2009; Jones et al., 2011)"},"properties":{"noteIndex":0},"schema":""}(Burn et al., 2010; Gosling et al., 2009; Jones et al., 2011). Pollen identification was carried out using the Neotropical pollen reference collection of over 1000 species held at the Universities of Leicester and Edinburgh and published pollen atlases ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"author":[{"dropping-particle":"","family":"Roubik","given":"DW","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Moreno","given":"PJE","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Monographs in systematic botany, Missouri Botanical Garden, St. Louis","id":"ITEM-1","issued":{"date-parts":[["1991"]]},"title":"Pollen and spores of Barro Colorado Island","type":"article-journal","volume":"36"},"uris":[""]},{"id":"ITEM-2","itemData":{"author":[{"dropping-particle":"","family":"Colinvaux","given":"Paul A.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Oliveira","given":"Paulo E.","non-dropping-particle":"de","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Pati?o","given":"Jorge E. M.","non-dropping-particle":"","parse-names":false,"suffix":""}],"id":"ITEM-2","issued":{"date-parts":[["1999"]]},"publisher":"Harwood Academic Publishers","publisher-place":"Amsterdam","title":"Amazon Pollen Manual and Atlas","type":"book"},"uris":[""]}],"mendeley":{"formattedCitation":"(Colinvaux et al., 1999; Roubik and Moreno, 1991)","plainTextFormattedCitation":"(Colinvaux et al., 1999; Roubik and Moreno, 1991)","previouslyFormattedCitation":"(Colinvaux et al., 1999; Roubik and Moreno, 1991)"},"properties":{"noteIndex":0},"schema":""}(Colinvaux et al., 1999; Roubik and Moreno, 1991). Phytoliths were extracted from 100ml of soil using the wet oxidation protocol ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"author":[{"dropping-particle":"","family":"Piperno","given":"Dolores R.","non-dropping-particle":"","parse-names":false,"suffix":""}],"id":"ITEM-1","issued":{"date-parts":[["2006"]]},"publisher":"Altamire Press","title":"Phytoliths: A comprehensive guide for Archeologists and Paleoecologists","type":"book"},"uris":[""]}],"mendeley":{"formattedCitation":"(Piperno, 2006)","plainTextFormattedCitation":"(Piperno, 2006)","previouslyFormattedCitation":"(Piperno, 2006)"},"properties":{"noteIndex":0},"schema":""}(Piperno, 2006). Phytolith identification was carried out using published references and the Neotropical phytolith collection of over 500 species held at the University of Exeter Archaeobotany Laboratory ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.revpalbo.2013.01.004","ISSN":"00346667","abstract":"The interpretation of Neotropical fossil phytolith assemblages for palaeoenvironmental and archaeological reconstructions relies on the development of appropriate modern analogues. We analyzed modern phytolith assemblages from the soils of ten distinctive tropical vegetation communities in eastern lowland Bolivia, ranging from terra firme humid evergreen forest to seasonally-inundated savannah. Results show that broad ecosystems – evergreen tropical forest, semi-deciduous dry tropical forest, and savannah – can be clearly differentiated by examination of their phytolith spectra and the application of Principal Component Analysis (PCA). Differences in phytolith assemblages between particular vegetation communities within each of these ecosystems are more subtle, but can still be identified. Comparison of phytolith assemblages with pollen rain data and stable carbon isotope analyses from the same vegetation plots show that these proxies are not only complementary, but significantly improve taxonomic and ecosystem resolution, and therefore our ability to interpret palaeoenvironmental and archaeological records. Our data underline the utility of phytolith analyses for reconstructing Amazon Holocene vegetation histories and pre-Columbian land use, particularly the high spatial resolution possible with terrestrial soil-based phytolith studies.","author":[{"dropping-particle":"","family":"Dickau","given":"Ruth","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Whitney","given":"Bronwen S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"José","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Soto","given":"J. Daniel","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Metcalfe","given":"Phil","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Street-Perrott","given":"F. Alayne","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Loader","given":"Neil J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Ficken","given":"Katherine J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Review of Palaeobotany and Palynology","id":"ITEM-1","issued":{"date-parts":[["2013","6"]]},"page":"15-37","publisher":"Elsevier B.V.","title":"Differentiation of neotropical ecosystems by modern soil phytolith assemblages and its implications for palaeoenvironmental and archaeological reconstructions","type":"article-journal","volume":"193"},"uris":[""]}],"mendeley":{"formattedCitation":"(Dickau et al., 2013)","plainTextFormattedCitation":"(Dickau et al., 2013)","previouslyFormattedCitation":"(Dickau et al., 2013)"},"properties":{"noteIndex":0},"schema":""}(Dickau et al., 2013). 3.3 Numerical analysisThe data were summarised to ecosystem level, i.e. evergreen forest, dry forest and savannah, as these are the most useful vegetation classifications for application to the palaeo-record. 12 samples from each ecosystem were selected from the full published datasets to ensure a consistent number of comparisons between assemblages within each ecosystem (Table 1). Taxonomic classifications in the pollen data were harmonised between datasets from different authors to a common standard. For example, all Moraceae and Urticaceae genera were grouped into one category, as some datasets did not differentiate the constituent genera. For the phytolith data, only A-fraction counts were used to enable comparison to samples from lake sediment where there is often insufficient material available to fractionate the samples into different size classes during preparation. The calculations of variability in these assemblages are therefore conservative, as variability in palaeo-records is likely to be higher given the higher diversity of classifications.The Hellinger transformation was applied to harmonised pollen and phytolith data before Euclidean distances between all pollen assemblages and between all phytolith assemblages were calculated using vegan 2.5-2 in R 3.5.0 (Oksanen et al. 2018). This method is equivalent to calculation of Hellinger distances, which reduce the influence of zero abundance samples ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1007/978-94-007-2745-8","ISBN":"9789400727441","ISSN":"00134651","abstract":"First book covering full range of modern data-analytical and statistical techniques used in paleolimnology and paleoecology Features exploratory data analysis, error estimation, clustering, ordination and modern statistical learning techniques Includes case studies on human impact, lake development and climate change ? Numerical and statistical methods have rapidly become part of a palaeolimnologist’s tool-kit. They are used to explore and summarise complex data, reconstruct past environmental variables from fossil assemblages, and test competing hypotheses about the causes of observed changes in lake biota through history. This book brings together a wide array of numerical and statistical techniques currently available for use in palaeolimnology and other branches of palaeoecology.","author":[{"dropping-particle":"","family":"Legendre","given":"Pierre","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Birks","given":"H John B","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Tracking Environmental Change using Lakes Sediment: Volume 5 Data Handling and Numerical Techniques","editor":[{"dropping-particle":"","family":"Birks","given":"H John B","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Lotter","given":"Andre F","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Juggins","given":"Steve","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Smol","given":"John P","non-dropping-particle":"","parse-names":false,"suffix":""}],"id":"ITEM-1","issued":{"date-parts":[["2012"]]},"note":"Section on distances methods","page":"201-248","publisher":"Springer","title":"Chapter 8: From Classical to Canonical Ordination","type":"chapter"},"uris":[""]},{"id":"ITEM-2","itemData":{"DOI":"10.1007/s004420100716","ISSN":"0029-8549","author":[{"dropping-particle":"","family":"Legendre","given":"Pierre","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Gallagher","given":"Eugene D.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Oecologia","id":"ITEM-2","issue":"2","issued":{"date-parts":[["2001","10","1"]]},"page":"271-280","publisher":"Springer Berlin Heidelberg","title":"Ecologically meaningful transformations for ordination of species data","type":"article-journal","volume":"129"},"uris":[""]}],"mendeley":{"formattedCitation":"(Legendre and Birks, 2012; Legendre and Gallagher, 2001)","plainTextFormattedCitation":"(Legendre and Birks, 2012; Legendre and Gallagher, 2001)","previouslyFormattedCitation":"(Legendre and Birks, 2012; Legendre and Gallagher, 2001)"},"properties":{"noteIndex":0},"schema":""}(Legendre and Birks, 2012; Legendre and Gallagher, 2001). Boxplots of distances within each plot and within each ecosystem (including distances within plots and between plots in the same ecosystem) and between ecosystems for pollen and phytoliths were created using the base boxplot function in R 3.5.0. A one-way ANOVA (Distance ~ Ecosystem) was used to determine if the distances within ecosystems were significantly different between the ecosystems. TukeysHSD was used to determine which ecosystems were significantly different to each other. Principal Component Analysis (PCA) was carried out on the fossil and modern datasets to assess if the modern samples are suitable analogs for the fossil samples. PCAs were produced using vegan 2.5-2 in R 3.5.0 (Oksanen et al. 2018).Lower quartile (Q1), median (Q2) and upper quartile (Q3) of the Hellinger distances within each ecosystem for each proxy were calculated to use as thresholds for floristic turnover within ecosystems (Table 2). The three quartiles were also calculated for distances between ecosystems, to use as thresholds for ecosystem turnover (Table 3). These thresholds reflect different levels of confidence that turnover is taking place: Q1 is a weak indication of ecosystem turnover, Q2 is a good indication of ecosystem turnover, and Q3 is a strong indication of ecosystem turnover. Hellinger distances were calculated between all samples at all depths for pollen and phytolith data from the palaeo-record at Laguna Granja and the pollen data from Laguna Chaplin, for comparison to the thresholds identified for ecosystem turnover.3.4 MapThe map of Noel Kempff Mercado National Park and surrounding ecosystems (Fig. 1) was created using ArcGIS 10.4. A vegetation classification of our study area, based on supervised Landsat imagery, was generously provided by the Museo de Historia Natural ‘Noel Kempff Mercado’, Santa Cruz, Bolivia in 2015.4. Results[Figure 2 – Boxplots of Hellinger distances. Plot A and B show distances within individual study plots for phytoliths and pollen respectively. CH – terra firme evergreen liana forest, LF – terra firme evergreen forest, AC – terra firme semi-deciduous dry forest, CP – seasonally-inundated semi-deciduous dry forest, MV – seasonally-inundated semi-deciduous dry forest, FC – terra firme savannah, FT – seasonally-inundated savannah, TO – seasonally-inundated savannah. Plots C and D show distances within ecosystems, plots E and F show distances between ecosystems within biomes.]4.1 Variation within plotsThe level of variation between phytolith assemblages within the two plots in the evergreen forest ecosystem class (CH, LF) are similar, both in terms of median distance (~5) and range of distance (~3-7) (Fig. 2A). Within the dry forest plots (AC, CP, MV) the terra firme plots (AC and CP) have much higher variability than the seasonally inundated plot. In the savannah plots (FC, FT, TO) the variation between assemblages within the plots is generally lower than for the evergreen and dry forest plots. The two evergreen plots (CH, LF) also show similar levels of variation between pollen assemblages (Fig. 2B). The dry forest plots show a different pattern however, with all three plots (AC, CP, MV), both terra firme and seasonally inundated, showing high levels of variation, although the seasonally inundated plot (MV) does show a greater range of variability than the two terra firme plots (AC, CP). The terra firme savannah plot (FC) shows particularly low variability in pollen assemblages, while the two seasonally inundated savannah plots (FT, TO) show greater variability and a greater range of variability. 4.2 Variation within ecosystemsThe level of variability overall is higher in the pollen assemblages than the phytolith assemblages (Fig 2C, D). For phytoliths, the dry forest ecosystem shows the highest variability and the greatest range of variability. The evergreen forest and savannah ecosystems show similar, lower levels of variability and smaller ranges (Fig. 2C). For pollen, the dry forest ecosystem also contains the highest variability in pollen assemblages, followed by evergreen forest, with savannas containing the lowest variability. The range of variability for pollen is similar across all three ecosystems (Fig. 2D).4.3 Variation between ecosystemsFor phytoliths, the highest variation is between evergreen forest and savannah assemblages, followed by between dry forest and savannah assemblages (Fig. 2E). The lowest variation in phytolith assemblages is between the two forested ecosystems, dry forest and evergreen forest. For pollen, all three comparisons between ecosystems show similar levels of variation between assemblages (Fig. 2F).A one-way ANOVA (Distance ~ Ecosystem) shows that ecosystem type is a statistically significant factor in the distribution of differences (p=0.001) for both pollen and phytoliths (Tables S3 and S4). TukeysHSD post-hoc test shows that for phytoliths dry forest is significantly different to evergreen forest and savannah, but that evergreen forest and savannah are not significantly different from each other. For pollen, TukeysHSD showed that the level of variation is statistically different between all three ecosystems. 4.4 PCA results[Figure 3 – Principal Component Analysis (PCA) biplots for modern and fossil samples. Panel A – modern pollen samples from NKMNP plots, fossil pollen samples from Laguna Granja and Laguna Chaplin. Panel B – modern phytolith samples from NKMNP plots and fossil phytolith samples from Laguna Granja. Colours represent sample location: evergreen forest plots from NKMNP are coloured dark green; dry forest plots from NKMNP are light green; savannah plots from NKMNP are yellow; fossil samples from Laguna Chaplin are violet; fossil samples from Laguna Granja are dark blue.]PCA results for both the modern pollen and phytoliths (Fig. 3) show differentiation of savannah (yellow), dry forest (light green) and evergreen forest (dark green) samples along PCA axis 1 with some overlap between dry and evergreen forest samples. PCA axis 1 may represent a gradient of arboreal cover, from savannah to evergreen forest. The fossil pollen samples (Fig. 3A) show differentiation of Lagunas Granja (blue) and Chaplin (violet). Laguna Chaplin samples are clustered into three time periods: the basal samples plot negatively on PCA axis 2; the intermediate samples plot towards the centre of the PCA; the most recent samples plot positively on PCA axis 1, close to the modern dry and evergreen forest samples. The fossil phytolith samples from Laguna Granja (Fig. 3B) show overlap with modern savannah and dry forest samples in the centre of the PCA. 4.5 Thresholds for floristic and ecosystem turnoverTable 2 – Thresholds for floristic change within ecosystems. Lower quartile, median and upper quartile of Hellinger distances between proxy assemblages within evergreen forest, dry forest and savannah from modern samples at Noel Kempff Mercado National Park (NKMNP). The lower quartile (Q1) represents a weak indication of floristic change within the ecosystem, the median (Q2) represents a good indication, and the upper quartile (Q3) represents a strong indication. Thresholds are given to 3 significant figures.ProxyEcosystemLower quartile (Q1)Median (Q2)Upper quartile (Q3)PhytolithEvergreen forest0.4170.5010.551Dry forest0.5160.8210.995Savannah0.3130.4710.540PollenEvergreen forest0.5860.6660.800Dry forest0.6980.8590.967Savannah0.4170.5280.688The range of thresholds identified for floristic turnover are the lower quartile (Q1), median (Q2) and upper quartile (Q3) of the Hellinger distances within each ecosystem, for each proxy (Table 2). These thresholds reflect the patterns of variability in the boxplots at ecosystem level (Fig. 2C, D). This pattern is generally similar for both pollen and phytoliths; the highest thresholds are in dry forest as these ecosystems presented the highest variability, while the thresholds for evergreen forest and savannah are similar and lower. The absolute values of the thresholds are slightly higher for pollen compared to phytoliths. Table 3 – Thresholds for ecosystem turnover. Lower quartile, median and upper quartile of Hellinger distances between proxy assemblages between evergreen forest, dry forest and savannah from modern samples at Noel Kempff Mercado National Park (NKMNP). The lower quartile (Q1) represents a weak indication of ecosystem turnover, the median (Q2) represents a good indication, and the upper quartile (Q3) represents a strong indication. Thresholds are given to 3 significant figures.ProxyEcosystemLower quartile (Q1)Median (Q2)Upper quartile (Q3)PhytolithsDry forest - evergreen forest0.7110.8200.938Dry forest - savannah0.8430.9621.06Evergreen forest - savannah0.9861.151.22PollenDry forest - evergreen forest0.7840.8730.967Dry forest - savannah0.7790.8920.975Evergreen forest - savannah0.7810.8350.898The ecosystem turnover thresholds are the lower (Q1), median (Q2) and upper quartiles (Q3) for between ecosystem distances (Table 3). The highest thresholds for phytoliths are for turnover between evergreen forest and savannah, followed by dry forest and savannah. Turnover between the two forested ecosystems has the lowest threshold. For pollen, the threshold is similar for turnover between all three ecosystems. 4.4.1 Application of thresholds to L. GranjaThe original interpretation of the pollen and phytolith records ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.yqres.2003.12.004","ISSN":"00335894","abstract":"Pollen and charcoal records from two large, shallow lakes reveal that throughout most of the past 50,000 yr Noel Kempff Mercado National Park, in northeastern lowland Bolivia (southwestern Amazon Basin), was predominantly covered by savannas and seasonally dry semideciduous forests. Lowered atmospheric CO2 concentrations, in combination with a longer dry season, caused expansion of dry forests and savannas during the last glacial period, especially at the last glacial maximum. These ecosystems persisted until the mid-Holocene, although they underwent significant species reassortment. Forest communities containing a mixture of evergreen and semideciduous species began to expand between 6000 and 3000 14C yr B.P. Humid evergreen rain forests expanded to cover most of the area within the past 2000 14C yr B.P., coincident with a reduction in fire frequencies. Comparisons between modern pollen spectra and vegetation reveal that the Moraceae-dominated rain forest pollen spectra likely have a regional source area at least 2–3 km beyond the lake shore, whereas the grass- and sedge-dominated savanna pollen spectra likely have a predominantly local source area. The Holocene vegetation changes are consistent with independent paleoprecipitation records from the Bolivian Altiplano and paleovegetation records from other parts of southwestern Amazonia. The progressive expansion in rain forests through the Holocene can be largely attributed to enhanced convective activity over Amazonia, due to greater seasonality of insolation in the Southern Hemisphere tropics driven by the precession cycle according to the Milankovitch Astronomical Theory.","author":[{"dropping-particle":"","family":"Burbridge","given":"Rachel E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Quaternary Research","id":"ITEM-1","issue":"2","issued":{"date-parts":[["2004","3"]]},"note":"chapling and Bella Vista records - dry forest/savanna until 2000 bp when precip increased - humid evergreen","page":"215-230","title":"Fifty-thousand-year vegetation and climate history of Noel Kempff Mercado National Park, Bolivian Amazon","type":"article-journal","volume":"61"},"uris":[""]},{"id":"ITEM-2","itemData":{"DOI":"10.1177/0959683615581204","ISBN":"0959-6836","ISSN":"0959-6836","abstract":"The nature and extent of pre-Columbian (pre-AD 1492) human impact in Amazonia is a contentious issue. The Bolivian Amazon has yielded some of the most impressive evidence for large and complex pre-Columbian societies in the Amazon basin, yet there remains relatively little data concerning the land use of these societies over time. Palaeoecology, when integrated with archaeological data, has the potential to fill these gaps in our knowledge. We present a 6000-year record of anthropogenic burning, agriculture and vegetation change, from an oxbow lake located adjacent to a pre-Columbian ring ditch in north-east Bolivia (13°15′44″S, 63°42′37″W). Human occupation around the lake site is inferred from pollen and phytoliths of maize (Zea mays L.) and macroscopic charcoal evidence of anthropogenic burning. First occupation around the lake was radiocarbon dated to ~2500 calibrated years before present (BP). The persistence of maize in the record from ~1850 BP suggests that it was an important crop grown in the ring-ditch region in pre-Columbian times, and abundant macroscopic charcoal suggests that pre-Columbian land management entailed more extensive burning of the landscape than the slash-and-burn agriculture practised around the site today. The site was occupied continuously until near-modern times, although there is evidence for a decline in agricultural intensity or change in land-use strategy, and possible population decline, from ~600–500 BP. The long and continuous occupation, which predates the establishment of rainforest in the region, suggests that pre-Columbian land use may have had a significant influence on ecosystem development at this site over the last ~2000 years.","author":[{"dropping-particle":"","family":"Carson","given":"J. F.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Watling","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"F. E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Whitney","given":"B. S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Prumers","given":"H.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Soto","given":"J. D.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"The Holocene","id":"ITEM-2","issue":"8","issued":{"date-parts":[["2015","5","1"]]},"note":"From Duplicate 2 (Pre-Columbian land use in the ring-ditch region of the Bolivian Amazon - Carson, J. F.; Watling, J.; Mayle, F. E.; Whitney, B. S.; Iriarte, J.; Prumers, H.; Soto, J. D.)\n\nUsed phytoliths and pollen from oxbow lake in NE Bolivia - oxbow 6000 yrs old.\n\nNice passage about phytoliths from lakes and their source area compared to pollen - assumes local representation due to modes of transport available.","page":"1285-1300","title":"Pre-Columbian land use in the ring-ditch region of the Bolivian Amazon","type":"article-journal","volume":"25"},"uris":[""]}],"mendeley":{"formattedCitation":"(Burbridge et al., 2004; Carson et al., 2015)","plainTextFormattedCitation":"(Burbridge et al., 2004; Carson et al., 2015)","previouslyFormattedCitation":"(Burbridge et al., 2004; Carson et al., 2015)"},"properties":{"noteIndex":0},"schema":""}(Carson et al., 2015) were used to identify the direction of change between ecosystems and select the appropriate thresholds to apply to pairwise comparisons of the pollen and phytoliths records down-core at L. Granja and Chaplin.[Figure 4 – Stratigraphic plots of Hellinger distances between pairwise assemblages downcore at Laguna Granja (phytoliths and pollen) and Laguna Chaplin (pollen). Age-depth models taken from original papers ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.yqres.2003.12.004","ISSN":"00335894","abstract":"Pollen and charcoal records from two large, shallow lakes reveal that throughout most of the past 50,000 yr Noel Kempff Mercado National Park, in northeastern lowland Bolivia (southwestern Amazon Basin), was predominantly covered by savannas and seasonally dry semideciduous forests. Lowered atmospheric CO2 concentrations, in combination with a longer dry season, caused expansion of dry forests and savannas during the last glacial period, especially at the last glacial maximum. These ecosystems persisted until the mid-Holocene, although they underwent significant species reassortment. Forest communities containing a mixture of evergreen and semideciduous species began to expand between 6000 and 3000 14C yr B.P. Humid evergreen rain forests expanded to cover most of the area within the past 2000 14C yr B.P., coincident with a reduction in fire frequencies. Comparisons between modern pollen spectra and vegetation reveal that the Moraceae-dominated rain forest pollen spectra likely have a regional source area at least 2–3 km beyond the lake shore, whereas the grass- and sedge-dominated savanna pollen spectra likely have a predominantly local source area. The Holocene vegetation changes are consistent with independent paleoprecipitation records from the Bolivian Altiplano and paleovegetation records from other parts of southwestern Amazonia. The progressive expansion in rain forests through the Holocene can be largely attributed to enhanced convective activity over Amazonia, due to greater seasonality of insolation in the Southern Hemisphere tropics driven by the precession cycle according to the Milankovitch Astronomical Theory.","author":[{"dropping-particle":"","family":"Burbridge","given":"Rachel E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Quaternary Research","id":"ITEM-1","issue":"2","issued":{"date-parts":[["2004","3"]]},"note":"chapling and Bella Vista records - dry forest/savanna until 2000 bp when precip increased - humid evergreen","page":"215-230","title":"Fifty-thousand-year vegetation and climate history of Noel Kempff Mercado National Park, Bolivian Amazon","type":"article-journal","volume":"61"},"uris":[""]},{"id":"ITEM-2","itemData":{"DOI":"10.1177/0959683615581204","ISBN":"0959-6836","ISSN":"0959-6836","abstract":"The nature and extent of pre-Columbian (pre-AD 1492) human impact in Amazonia is a contentious issue. The Bolivian Amazon has yielded some of the most impressive evidence for large and complex pre-Columbian societies in the Amazon basin, yet there remains relatively little data concerning the land use of these societies over time. Palaeoecology, when integrated with archaeological data, has the potential to fill these gaps in our knowledge. We present a 6000-year record of anthropogenic burning, agriculture and vegetation change, from an oxbow lake located adjacent to a pre-Columbian ring ditch in north-east Bolivia (13°15′44″S, 63°42′37″W). Human occupation around the lake site is inferred from pollen and phytoliths of maize (Zea mays L.) and macroscopic charcoal evidence of anthropogenic burning. First occupation around the lake was radiocarbon dated to ~2500 calibrated years before present (BP). The persistence of maize in the record from ~1850 BP suggests that it was an important crop grown in the ring-ditch region in pre-Columbian times, and abundant macroscopic charcoal suggests that pre-Columbian land management entailed more extensive burning of the landscape than the slash-and-burn agriculture practised around the site today. The site was occupied continuously until near-modern times, although there is evidence for a decline in agricultural intensity or change in land-use strategy, and possible population decline, from ~600–500 BP. The long and continuous occupation, which predates the establishment of rainforest in the region, suggests that pre-Columbian land use may have had a significant influence on ecosystem development at this site over the last ~2000 years.","author":[{"dropping-particle":"","family":"Carson","given":"J. F.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Watling","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"F. E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Whitney","given":"B. S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Prumers","given":"H.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Soto","given":"J. D.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"The Holocene","id":"ITEM-2","issue":"8","issued":{"date-parts":[["2015","5","1"]]},"note":"From Duplicate 2 (Pre-Columbian land use in the ring-ditch region of the Bolivian Amazon - Carson, J. F.; Watling, J.; Mayle, F. E.; Whitney, B. S.; Iriarte, J.; Prumers, H.; Soto, J. D.)\n\nUsed phytoliths and pollen from oxbow lake in NE Bolivia - oxbow 6000 yrs old.\n\nNice passage about phytoliths from lakes and their source area compared to pollen - assumes local representation due to modes of transport available.","page":"1285-1300","title":"Pre-Columbian land use in the ring-ditch region of the Bolivian Amazon","type":"article-journal","volume":"25"},"uris":[""]}],"mendeley":{"formattedCitation":"(Burbridge et al., 2004; Carson et al., 2015)","plainTextFormattedCitation":"(Burbridge et al., 2004; Carson et al., 2015)","previouslyFormattedCitation":"(Burbridge et al., 2004; Carson et al., 2015)"},"properties":{"noteIndex":0},"schema":""}(Burbridge et al., 2004; Carson et al., 2015). Red dashed lines represent thresholds for floristic change within the ecosystem: lower quartile (Q1), median (Q2) and upper quartile (Q3) of Hellinger distances for the appropriate ecosystem and proxy, calculated based on modern distances between samples in vegetation study plots at Noel Kempff Mercado National Park (NKMNP) (Table 2). For Granja phytoliths, the thresholds for evergreen forest are plotted. For Granja and Chaplin pollen, the thresholds for savannah are plotted. Green dashed lines represent thresholds for ecosystem turnover: lower quartile (Q1), median (Q2) and upper quartile (Q3) of Hellinger distances for the turnover between evergreen forest and savannah, for the appropriate proxy (Table 3).]ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1177/0959683615581204","ISBN":"0959-6836","ISSN":"0959-6836","abstract":"The nature and extent of pre-Columbian (pre-AD 1492) human impact in Amazonia is a contentious issue. The Bolivian Amazon has yielded some of the most impressive evidence for large and complex pre-Columbian societies in the Amazon basin, yet there remains relatively little data concerning the land use of these societies over time. Palaeoecology, when integrated with archaeological data, has the potential to fill these gaps in our knowledge. We present a 6000-year record of anthropogenic burning, agriculture and vegetation change, from an oxbow lake located adjacent to a pre-Columbian ring ditch in north-east Bolivia (13°15′44″S, 63°42′37″W). Human occupation around the lake site is inferred from pollen and phytoliths of maize (Zea mays L.) and macroscopic charcoal evidence of anthropogenic burning. First occupation around the lake was radiocarbon dated to ~2500 calibrated years before present (BP). The persistence of maize in the record from ~1850 BP suggests that it was an important crop grown in the ring-ditch region in pre-Columbian times, and abundant macroscopic charcoal suggests that pre-Columbian land management entailed more extensive burning of the landscape than the slash-and-burn agriculture practised around the site today. The site was occupied continuously until near-modern times, although there is evidence for a decline in agricultural intensity or change in land-use strategy, and possible population decline, from ~600–500 BP. The long and continuous occupation, which predates the establishment of rainforest in the region, suggests that pre-Columbian land use may have had a significant influence on ecosystem development at this site over the last ~2000 years.","author":[{"dropping-particle":"","family":"Carson","given":"J. F.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Watling","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"F. E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Whitney","given":"B. S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Prumers","given":"H.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Soto","given":"J. D.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"The Holocene","id":"ITEM-1","issue":"8","issued":{"date-parts":[["2015","5","1"]]},"note":"From Duplicate 2 (Pre-Columbian land use in the ring-ditch region of the Bolivian Amazon - Carson, J. F.; Watling, J.; Mayle, F. E.; Whitney, B. S.; Iriarte, J.; Prumers, H.; Soto, J. D.)\n\nUsed phytoliths and pollen from oxbow lake in NE Bolivia - oxbow 6000 yrs old.\n\nNice passage about phytoliths from lakes and their source area compared to pollen - assumes local representation due to modes of transport available.","page":"1285-1300","title":"Pre-Columbian land use in the ring-ditch region of the Bolivian Amazon","type":"article-journal","volume":"25"},"uris":[""]}],"mendeley":{"formattedCitation":"(Carson et al., 2015)","manualFormatting":"Carson et al., (2015)","plainTextFormattedCitation":"(Carson et al., 2015)","previouslyFormattedCitation":"(Carson et al., 2015)"},"properties":{"noteIndex":0},"schema":""}Carson et al., (2015) interpret the phytolith record at L. Granja to represent local riparian gallery forest around the edges of the lake from 6,000 – 2,500 yBP, followed by an opening up of the forest from 2,500-500 yBP, and then expansion of the gallery forest within the last 500 years (Fig. S5). Therefore, the phytolith thresholds for evergreen forest (Table 2) were selected to compare to the pairwise distances down-core in the phytolith record. The three basal phytolith pairwise distances meet or exceed the Q3 threshold indicating strong evidence of floristic turnover within the ecosystem (Fig. 4). Throughout the middle of the core, pairwise distances meet the Q1 threshold indicating weak evidence for floristic turnover within the ecosystem. In the top section of the core, pairwise distances fluctuate below the Q1 threshold, only reaching the Q1 threshold three further times (Fig. 4). The thresholds for ecosystem turnover (Table 3) are not met throughout the core.ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1177/0959683615581204","ISBN":"0959-6836","ISSN":"0959-6836","abstract":"The nature and extent of pre-Columbian (pre-AD 1492) human impact in Amazonia is a contentious issue. The Bolivian Amazon has yielded some of the most impressive evidence for large and complex pre-Columbian societies in the Amazon basin, yet there remains relatively little data concerning the land use of these societies over time. Palaeoecology, when integrated with archaeological data, has the potential to fill these gaps in our knowledge. We present a 6000-year record of anthropogenic burning, agriculture and vegetation change, from an oxbow lake located adjacent to a pre-Columbian ring ditch in north-east Bolivia (13°15′44″S, 63°42′37″W). Human occupation around the lake site is inferred from pollen and phytoliths of maize (Zea mays L.) and macroscopic charcoal evidence of anthropogenic burning. First occupation around the lake was radiocarbon dated to ~2500 calibrated years before present (BP). The persistence of maize in the record from ~1850 BP suggests that it was an important crop grown in the ring-ditch region in pre-Columbian times, and abundant macroscopic charcoal suggests that pre-Columbian land management entailed more extensive burning of the landscape than the slash-and-burn agriculture practised around the site today. The site was occupied continuously until near-modern times, although there is evidence for a decline in agricultural intensity or change in land-use strategy, and possible population decline, from ~600–500 BP. The long and continuous occupation, which predates the establishment of rainforest in the region, suggests that pre-Columbian land use may have had a significant influence on ecosystem development at this site over the last ~2000 years.","author":[{"dropping-particle":"","family":"Carson","given":"J. F.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Watling","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"F. E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Whitney","given":"B. S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Prumers","given":"H.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Soto","given":"J. D.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"The Holocene","id":"ITEM-1","issue":"8","issued":{"date-parts":[["2015","5","1"]]},"note":"From Duplicate 2 (Pre-Columbian land use in the ring-ditch region of the Bolivian Amazon - Carson, J. F.; Watling, J.; Mayle, F. E.; Whitney, B. S.; Iriarte, J.; Prumers, H.; Soto, J. D.)\n\nUsed phytoliths and pollen from oxbow lake in NE Bolivia - oxbow 6000 yrs old.\n\nNice passage about phytoliths from lakes and their source area compared to pollen - assumes local representation due to modes of transport available.","page":"1285-1300","title":"Pre-Columbian land use in the ring-ditch region of the Bolivian Amazon","type":"article-journal","volume":"25"},"uris":[""]}],"mendeley":{"formattedCitation":"(Carson et al., 2015)","manualFormatting":"Carson et al., (2015)","plainTextFormattedCitation":"(Carson et al., 2015)","previouslyFormattedCitation":"(Carson et al., 2015)"},"properties":{"noteIndex":0},"schema":""}Carson et al., (2015) interpret the pollen record at L. Granja to represent open, terra firme savannah from 6,000 yBP to 500 yBP when terra firme evergreen forest expands into the surrounding landscape (Fig. S6). Therefore, the pollen thresholds for savannah (Table 2) were selected to compare to the pairwise distances down-core in the pollen record. Almost all pairwise distances from the pollen record fluctuate between the Q1 and Q3 thresholds for floristic turnover (Fig. 4). From 6,000 calyrBP to 2,500 calyrBP most distances meet the Q2 threshold, indicating good evidence for floristic turnover. From 2,500 calyrBP to 1,500 calyrBP distances are closer to the Q1 threshold for weak indication of floristic turnover. From 1,500 calyrBP to present, distances are close to the Q2 threshold, showing good evidence of floristic turnover. The thresholds for ecosystem turnover (Table 3) are not met throughout the core. In addition to pairwise distances down-core, the distances between all samples at all depths were also calculated. To test whether the high levels of variation between the basal three phytolith samples at L. Granja was due to the low sampling resolution (500 years c.f. <100 years near the top of the core), the thresholds for floristic and ecosystem turnover were compared to distances between all samples. In the phytolith record from L. Granja, Q1 threshold for weak indication of floristic turnover is met by comparison of the top section from 600 calyrBP to present with the rest of the core, matching the timing of gallery forest expansion interpreted by ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1177/0959683615581204","ISBN":"0959-6836","ISSN":"0959-6836","abstract":"The nature and extent of pre-Columbian (pre-AD 1492) human impact in Amazonia is a contentious issue. The Bolivian Amazon has yielded some of the most impressive evidence for large and complex pre-Columbian societies in the Amazon basin, yet there remains relatively little data concerning the land use of these societies over time. Palaeoecology, when integrated with archaeological data, has the potential to fill these gaps in our knowledge. We present a 6000-year record of anthropogenic burning, agriculture and vegetation change, from an oxbow lake located adjacent to a pre-Columbian ring ditch in north-east Bolivia (13°15′44″S, 63°42′37″W). Human occupation around the lake site is inferred from pollen and phytoliths of maize (Zea mays L.) and macroscopic charcoal evidence of anthropogenic burning. First occupation around the lake was radiocarbon dated to ~2500 calibrated years before present (BP). The persistence of maize in the record from ~1850 BP suggests that it was an important crop grown in the ring-ditch region in pre-Columbian times, and abundant macroscopic charcoal suggests that pre-Columbian land management entailed more extensive burning of the landscape than the slash-and-burn agriculture practised around the site today. The site was occupied continuously until near-modern times, although there is evidence for a decline in agricultural intensity or change in land-use strategy, and possible population decline, from ~600–500 BP. The long and continuous occupation, which predates the establishment of rainforest in the region, suggests that pre-Columbian land use may have had a significant influence on ecosystem development at this site over the last ~2000 years.","author":[{"dropping-particle":"","family":"Carson","given":"J. F.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Watling","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"F. E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Whitney","given":"B. S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Prumers","given":"H.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Soto","given":"J. D.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"The Holocene","id":"ITEM-1","issue":"8","issued":{"date-parts":[["2015","5","1"]]},"note":"From Duplicate 2 (Pre-Columbian land use in the ring-ditch region of the Bolivian Amazon - Carson, J. F.; Watling, J.; Mayle, F. E.; Whitney, B. S.; Iriarte, J.; Prumers, H.; Soto, J. D.)\n\nUsed phytoliths and pollen from oxbow lake in NE Bolivia - oxbow 6000 yrs old.\n\nNice passage about phytoliths from lakes and their source area compared to pollen - assumes local representation due to modes of transport available.","page":"1285-1300","title":"Pre-Columbian land use in the ring-ditch region of the Bolivian Amazon","type":"article-journal","volume":"25"},"uris":[""]}],"mendeley":{"formattedCitation":"(Carson et al., 2015)","manualFormatting":"Carson et al., (2015)","plainTextFormattedCitation":"(Carson et al., 2015)","previouslyFormattedCitation":"(Carson et al., 2015)"},"properties":{"noteIndex":0},"schema":""}Carson et al., (2015). The Q2 and Q3 thresholds for good and strong indication of floristic turnover are only met by the three basal samples identified by pairwise comparison downcore. The three thresholds for ecosystem turnover are only met by isolated comparisons of specific samples. In the pollen record from L. Granja, the Q2 threshold for good indication of floristic turnover is met by comparison of the top section from 500 calyrBP to present with the rest of the core, matching the timing of ecosystem turnover from savannah to evergreen forest identified by ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1177/0959683615581204","ISBN":"0959-6836","ISSN":"0959-6836","abstract":"The nature and extent of pre-Columbian (pre-AD 1492) human impact in Amazonia is a contentious issue. The Bolivian Amazon has yielded some of the most impressive evidence for large and complex pre-Columbian societies in the Amazon basin, yet there remains relatively little data concerning the land use of these societies over time. Palaeoecology, when integrated with archaeological data, has the potential to fill these gaps in our knowledge. We present a 6000-year record of anthropogenic burning, agriculture and vegetation change, from an oxbow lake located adjacent to a pre-Columbian ring ditch in north-east Bolivia (13°15′44″S, 63°42′37″W). Human occupation around the lake site is inferred from pollen and phytoliths of maize (Zea mays L.) and macroscopic charcoal evidence of anthropogenic burning. First occupation around the lake was radiocarbon dated to ~2500 calibrated years before present (BP). The persistence of maize in the record from ~1850 BP suggests that it was an important crop grown in the ring-ditch region in pre-Columbian times, and abundant macroscopic charcoal suggests that pre-Columbian land management entailed more extensive burning of the landscape than the slash-and-burn agriculture practised around the site today. The site was occupied continuously until near-modern times, although there is evidence for a decline in agricultural intensity or change in land-use strategy, and possible population decline, from ~600–500 BP. The long and continuous occupation, which predates the establishment of rainforest in the region, suggests that pre-Columbian land use may have had a significant influence on ecosystem development at this site over the last ~2000 years.","author":[{"dropping-particle":"","family":"Carson","given":"J. F.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Watling","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"F. E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Whitney","given":"B. S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Prumers","given":"H.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Soto","given":"J. D.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"The Holocene","id":"ITEM-1","issue":"8","issued":{"date-parts":[["2015","5","1"]]},"note":"From Duplicate 2 (Pre-Columbian land use in the ring-ditch region of the Bolivian Amazon - Carson, J. F.; Watling, J.; Mayle, F. E.; Whitney, B. S.; Iriarte, J.; Prumers, H.; Soto, J. D.)\n\nUsed phytoliths and pollen from oxbow lake in NE Bolivia - oxbow 6000 yrs old.\n\nNice passage about phytoliths from lakes and their source area compared to pollen - assumes local representation due to modes of transport available.","page":"1285-1300","title":"Pre-Columbian land use in the ring-ditch region of the Bolivian Amazon","type":"article-journal","volume":"25"},"uris":[""]}],"mendeley":{"formattedCitation":"(Carson et al., 2015)","manualFormatting":"Carson et al., (2015)","plainTextFormattedCitation":"(Carson et al., 2015)","previouslyFormattedCitation":"(Carson et al., 2015)"},"properties":{"noteIndex":0},"schema":""}Carson et al., (2015). The Q3 threshold for floristic turnover and all three thresholds for ecosystem turnover are only met by isolated comparisons of specific samples. 4.4.2 Application of thresholds to L. ChaplinADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1126/science.290.5500.2291","ISSN":"0036-8075","PMID":"11125139","abstract":"Amazonian rain forest-savanna boundaries are highly sensitive to climatic change and may also play an important role in rain forest speciation. However, their dynamics over millennial time scales are poorly understood. Here, we present late Quaternary pollen records from the southern margin of Amazonia, which show that the humid evergreen rain forests of eastern Bolivia have been expanding southward over the past 3000 years and that their present-day limit represents the southernmost extent of Amazonian rain forest over at least the past 50,000 years. This rain forest expansion is attributed to increased seasonal latitudinal migration of the Intertropical Convergence Zone, which can in turn be explained by Milankovitch astronomic forcing.","author":[{"dropping-particle":"","family":"Mayle","given":"F E","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Burbridge","given":"R","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"T J","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Science (New York, N.Y.)","id":"ITEM-1","issue":"2000","issued":{"date-parts":[["2000"]]},"page":"2291-2294","title":"Millennial-scale dynamics of southern Amazonian rain forests.","type":"article-journal","volume":"290"},"uris":[""]}],"mendeley":{"formattedCitation":"(Mayle et al., 2000)","manualFormatting":"Mayle et al., (2000)","plainTextFormattedCitation":"(Mayle et al., 2000)","previouslyFormattedCitation":"(Mayle et al., 2000)"},"properties":{"noteIndex":0},"schema":""}Mayle et al., (2000) and ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.yqres.2003.12.004","ISSN":"00335894","abstract":"Pollen and charcoal records from two large, shallow lakes reveal that throughout most of the past 50,000 yr Noel Kempff Mercado National Park, in northeastern lowland Bolivia (southwestern Amazon Basin), was predominantly covered by savannas and seasonally dry semideciduous forests. Lowered atmospheric CO2 concentrations, in combination with a longer dry season, caused expansion of dry forests and savannas during the last glacial period, especially at the last glacial maximum. These ecosystems persisted until the mid-Holocene, although they underwent significant species reassortment. Forest communities containing a mixture of evergreen and semideciduous species began to expand between 6000 and 3000 14C yr B.P. Humid evergreen rain forests expanded to cover most of the area within the past 2000 14C yr B.P., coincident with a reduction in fire frequencies. Comparisons between modern pollen spectra and vegetation reveal that the Moraceae-dominated rain forest pollen spectra likely have a regional source area at least 2–3 km beyond the lake shore, whereas the grass- and sedge-dominated savanna pollen spectra likely have a predominantly local source area. The Holocene vegetation changes are consistent with independent paleoprecipitation records from the Bolivian Altiplano and paleovegetation records from other parts of southwestern Amazonia. The progressive expansion in rain forests through the Holocene can be largely attributed to enhanced convective activity over Amazonia, due to greater seasonality of insolation in the Southern Hemisphere tropics driven by the precession cycle according to the Milankovitch Astronomical Theory.","author":[{"dropping-particle":"","family":"Burbridge","given":"Rachel E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Quaternary Research","id":"ITEM-1","issue":"2","issued":{"date-parts":[["2004","3"]]},"note":"chapling and Bella Vista records - dry forest/savanna until 2000 bp when precip increased - humid evergreen","page":"215-230","title":"Fifty-thousand-year vegetation and climate history of Noel Kempff Mercado National Park, Bolivian Amazon","type":"article-journal","volume":"61"},"uris":[""]}],"mendeley":{"formattedCitation":"(Burbridge et al., 2004)","manualFormatting":"Burbridge et al., (2004)","plainTextFormattedCitation":"(Burbridge et al., 2004)","previouslyFormattedCitation":"(Burbridge et al., 2004)"},"properties":{"noteIndex":0},"schema":""}Burbridge et al., (2004) interpret the pollen record at L. Chaplin to represent a palm savannah marsh from ~51,000 to 48,000 yBP, then a seasonally inundated savannah with areas of semi-deciduous dry forest from ~48,000 to 2000 yBP, after which ecosystem turnover occurred and humid evergreen forest expanded to dominate the lake catchment (Fig. S7). Therefore, the pollen thresholds for savannah (Table 2) were selected to compare to the pairwise distances down-core. The majority of pairwise distances are close to the Q1 threshold, indicating weak evidence for floristic turnover (Fig. 4). One distance meets the Q2 threshold, indicating good evidence for floristic turnover, at 48,000 calyrBP coincident with the change from palm savannah marsh to seasonally inundated savannah ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.yqres.2003.12.004","ISSN":"00335894","abstract":"Pollen and charcoal records from two large, shallow lakes reveal that throughout most of the past 50,000 yr Noel Kempff Mercado National Park, in northeastern lowland Bolivia (southwestern Amazon Basin), was predominantly covered by savannas and seasonally dry semideciduous forests. Lowered atmospheric CO2 concentrations, in combination with a longer dry season, caused expansion of dry forests and savannas during the last glacial period, especially at the last glacial maximum. These ecosystems persisted until the mid-Holocene, although they underwent significant species reassortment. Forest communities containing a mixture of evergreen and semideciduous species began to expand between 6000 and 3000 14C yr B.P. Humid evergreen rain forests expanded to cover most of the area within the past 2000 14C yr B.P., coincident with a reduction in fire frequencies. Comparisons between modern pollen spectra and vegetation reveal that the Moraceae-dominated rain forest pollen spectra likely have a regional source area at least 2–3 km beyond the lake shore, whereas the grass- and sedge-dominated savanna pollen spectra likely have a predominantly local source area. The Holocene vegetation changes are consistent with independent paleoprecipitation records from the Bolivian Altiplano and paleovegetation records from other parts of southwestern Amazonia. The progressive expansion in rain forests through the Holocene can be largely attributed to enhanced convective activity over Amazonia, due to greater seasonality of insolation in the Southern Hemisphere tropics driven by the precession cycle according to the Milankovitch Astronomical Theory.","author":[{"dropping-particle":"","family":"Burbridge","given":"Rachel E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Quaternary Research","id":"ITEM-1","issue":"2","issued":{"date-parts":[["2004","3"]]},"note":"chapling and Bella Vista records - dry forest/savanna until 2000 bp when precip increased - humid evergreen","page":"215-230","title":"Fifty-thousand-year vegetation and climate history of Noel Kempff Mercado National Park, Bolivian Amazon","type":"article-journal","volume":"61"},"uris":[""]}],"mendeley":{"formattedCitation":"(Burbridge et al., 2004)","plainTextFormattedCitation":"(Burbridge et al., 2004)","previouslyFormattedCitation":"(Burbridge et al., 2004)"},"properties":{"noteIndex":0},"schema":""}(Burbridge et al., 2004). The ecosystem turnover identified by the authors at 2,000 calyrBP does not meet the Q2 or Q3 thresholds for floristic change. The thresholds for ecosystem turnover are not met throughout the core (Fig. 4).However, when comparing the full matrix of sample depths, the Q3 threshold for strong indication of floristic turnover is reached by comparisons between the top section of the core (2,500 calyrBP to present) with the majority of the rest of core. Furthermore, the Q2 threshold for good indication of ecosystem turnover is met by comparisons between the top 3,000 calyrBP to present section with the majority of the rest of the core. This broadly matches the interpretation of ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.yqres.2003.12.004","ISSN":"00335894","abstract":"Pollen and charcoal records from two large, shallow lakes reveal that throughout most of the past 50,000 yr Noel Kempff Mercado National Park, in northeastern lowland Bolivia (southwestern Amazon Basin), was predominantly covered by savannas and seasonally dry semideciduous forests. Lowered atmospheric CO2 concentrations, in combination with a longer dry season, caused expansion of dry forests and savannas during the last glacial period, especially at the last glacial maximum. These ecosystems persisted until the mid-Holocene, although they underwent significant species reassortment. Forest communities containing a mixture of evergreen and semideciduous species began to expand between 6000 and 3000 14C yr B.P. Humid evergreen rain forests expanded to cover most of the area within the past 2000 14C yr B.P., coincident with a reduction in fire frequencies. Comparisons between modern pollen spectra and vegetation reveal that the Moraceae-dominated rain forest pollen spectra likely have a regional source area at least 2–3 km beyond the lake shore, whereas the grass- and sedge-dominated savanna pollen spectra likely have a predominantly local source area. The Holocene vegetation changes are consistent with independent paleoprecipitation records from the Bolivian Altiplano and paleovegetation records from other parts of southwestern Amazonia. The progressive expansion in rain forests through the Holocene can be largely attributed to enhanced convective activity over Amazonia, due to greater seasonality of insolation in the Southern Hemisphere tropics driven by the precession cycle according to the Milankovitch Astronomical Theory.","author":[{"dropping-particle":"","family":"Burbridge","given":"Rachel E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Quaternary Research","id":"ITEM-1","issue":"2","issued":{"date-parts":[["2004","3"]]},"note":"chapling and Bella Vista records - dry forest/savanna until 2000 bp when precip increased - humid evergreen","page":"215-230","title":"Fifty-thousand-year vegetation and climate history of Noel Kempff Mercado National Park, Bolivian Amazon","type":"article-journal","volume":"61"},"uris":[""]},{"id":"ITEM-2","itemData":{"DOI":"10.1126/science.290.5500.2291","ISSN":"0036-8075","PMID":"11125139","abstract":"Amazonian rain forest-savanna boundaries are highly sensitive to climatic change and may also play an important role in rain forest speciation. However, their dynamics over millennial time scales are poorly understood. Here, we present late Quaternary pollen records from the southern margin of Amazonia, which show that the humid evergreen rain forests of eastern Bolivia have been expanding southward over the past 3000 years and that their present-day limit represents the southernmost extent of Amazonian rain forest over at least the past 50,000 years. This rain forest expansion is attributed to increased seasonal latitudinal migration of the Intertropical Convergence Zone, which can in turn be explained by Milankovitch astronomic forcing.","author":[{"dropping-particle":"","family":"Mayle","given":"F E","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Burbridge","given":"R","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"T J","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Science (New York, N.Y.)","id":"ITEM-2","issue":"2000","issued":{"date-parts":[["2000"]]},"page":"2291-2294","title":"Millennial-scale dynamics of southern Amazonian rain forests.","type":"article-journal","volume":"290"},"uris":[""]}],"mendeley":{"formattedCitation":"(Burbridge et al., 2004; Mayle et al., 2000)","plainTextFormattedCitation":"(Burbridge et al., 2004; Mayle et al., 2000)","previouslyFormattedCitation":"(Burbridge et al., 2004; Mayle et al., 2000)"},"properties":{"noteIndex":0},"schema":""}(Burbridge et al., 2004; Mayle et al., 2000) that ecosystem turnover occurred around 2,000 calyrBP from a savannah-forest community to an evergreen forest community. 5. Discussion5.1 Variability in pollen and phytolith assemblagesThe overall magnitude of variability is higher in the pollen assemblages than the phytolith assemblages (Fig 2C, D). This reflects the higher diversity of pollen types within tropical ecosystems, which can be >100 from one site, compared to phytolith types which do not normally exceed 50. At the plot level, particularly high levels of variability were found within both humid evergreen forest plots and individual semi-deciduous dry forest plots (Fig. 2A, B). The high variability within these 1-hectare plots has implications for small-scale palaeo-ecological studies such as from soil pits or peat cores, as it shows that a significant amount of variability in the phytolith or pollen assemblage is consistent with samples from within one modern plant community. Caution is therefore required in interpretation of floristic changes at the scale of a single soil pit or peat core, as small changes in the pollen or phytolith records may only reflect proxy variability within that ecosystem. For example, increases in phytolith palm abundance within evergreen forest soil pit samples has been interpreted as a record of human influence in Acre state, Brazil ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1073/pnas.1614359114","ISSN":"1091-6490","PMID":"28167791","abstract":"Over 450 pre-Columbian (pre-AD 1492) geometric ditched enclosures (\"geoglyphs\") occupy ～13,000 km(2) of Acre state, Brazil, representing a key discovery of Amazonian archaeology. These huge earthworks were concealed for centuries under terra firme (upland interfluvial) rainforest, directly challenging the \"pristine\" status of this ecosystem and its perceived vulnerability to human impacts. We reconstruct the environmental context of geoglyph construction and the nature, extent, and legacy of associated human impacts. We show that bamboo forest dominated the region for ≥6,000 y and that only small, temporary clearings were made to build the geoglyphs; however, construction occurred within anthropogenic forest that had been actively managed for millennia. In the absence of widespread deforestation, exploitation of forest products shaped a largely forested landscape that survived intact until the late 20th century.","author":[{"dropping-particle":"","family":"Watling","given":"Jennifer","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"José","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Schaan","given":"Denise","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Pessenda","given":"Luiz C R","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Loader","given":"Neil J","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Street-Perrott","given":"F Alayne","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Dickau","given":"Ruth E","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Damasceno","given":"Antonia","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Ranzi","given":"Alceu","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Proceedings of the National Academy of Sciences of the United States of America","id":"ITEM-1","issue":"8","issued":{"date-parts":[["2017","2","21"]]},"page":"1868-1873","publisher":"National Academy of Sciences","title":"Impact of pre-Columbian \"geoglyph\" builders on Amazonian forests.","type":"article-journal","volume":"114"},"uris":[""]}],"mendeley":{"formattedCitation":"(Watling et al., 2017)","plainTextFormattedCitation":"(Watling et al., 2017)","previouslyFormattedCitation":"(Watling et al., 2017)"},"properties":{"noteIndex":0},"schema":""}(Watling et al., 2017) but this may instead reflect the high variability in phytolith assemblages from evergreen forest soil samples. The down-profile variation seen in the soil phytolith palaeo-record may therefore not reflect significant floristic changes in the vegetation. This demonstrates the importance of using a modern training dataset to define the level of variation in modern assemblages from within a plant community to enable the differentiation of proxy noise from significant vegetation change. At the ecosystem level, the highest variability in both pollen and phytolith assemblages was found in dry forest (Fig 2C, D). This is likely to be partly due to the differences between the two dry forest types included in this classification. The dry forest ecosystem class contains both seasonally inundated and terra firme dry forest, which the vegetation inventories show to be floristically distinct in terms of the most dominant tree species ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"URL":"","accessed":{"date-parts":[["2018","10","25"]]},"author":[{"dropping-particle":"","family":"SALVIAS","given":"","non-dropping-particle":"","parse-names":false,"suffix":""}],"id":"ITEM-1","issued":{"date-parts":[["2004"]]},"title":"SALVIAS: Synthesis and Analysis of Local Vegetation Inventories Across Scales","type":"webpage"},"uris":[""]}],"mendeley":{"formattedCitation":"(SALVIAS, 2004)","plainTextFormattedCitation":"(SALVIAS, 2004)","previouslyFormattedCitation":"(SALVIAS, 2004)"},"properties":{"noteIndex":0},"schema":""}(SALVIAS, 2004). For example, in the terra firme plots, the most common taxa are Caesalpinia floribunda, Tabebuia roseo-alba, and Anadenanthera colubrina whereas in the seasonally inundated plots the most common taxa are Inga ingoides, Physocalymma scaberimum and Hirtella gracilipes ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.revpalbo.2013.01.004","ISSN":"00346667","abstract":"The interpretation of Neotropical fossil phytolith assemblages for palaeoenvironmental and archaeological reconstructions relies on the development of appropriate modern analogues. We analyzed modern phytolith assemblages from the soils of ten distinctive tropical vegetation communities in eastern lowland Bolivia, ranging from terra firme humid evergreen forest to seasonally-inundated savannah. Results show that broad ecosystems – evergreen tropical forest, semi-deciduous dry tropical forest, and savannah – can be clearly differentiated by examination of their phytolith spectra and the application of Principal Component Analysis (PCA). Differences in phytolith assemblages between particular vegetation communities within each of these ecosystems are more subtle, but can still be identified. Comparison of phytolith assemblages with pollen rain data and stable carbon isotope analyses from the same vegetation plots show that these proxies are not only complementary, but significantly improve taxonomic and ecosystem resolution, and therefore our ability to interpret palaeoenvironmental and archaeological records. Our data underline the utility of phytolith analyses for reconstructing Amazon Holocene vegetation histories and pre-Columbian land use, particularly the high spatial resolution possible with terrestrial soil-based phytolith studies.","author":[{"dropping-particle":"","family":"Dickau","given":"Ruth","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Whitney","given":"Bronwen S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"José","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Soto","given":"J. Daniel","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Metcalfe","given":"Phil","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Street-Perrott","given":"F. Alayne","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Loader","given":"Neil J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Ficken","given":"Katherine J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Review of Palaeobotany and Palynology","id":"ITEM-1","issued":{"date-parts":[["2013","6"]]},"page":"15-37","publisher":"Elsevier B.V.","title":"Differentiation of neotropical ecosystems by modern soil phytolith assemblages and its implications for palaeoenvironmental and archaeological reconstructions","type":"article-journal","volume":"193"},"uris":[""]}],"mendeley":{"formattedCitation":"(Dickau et al., 2013)","plainTextFormattedCitation":"(Dickau et al., 2013)","previouslyFormattedCitation":"(Dickau et al., 2013)"},"properties":{"noteIndex":0},"schema":""}(Dickau et al., 2013). This floristic difference in tree species explains the different levels of variation in the pollen assemblage between the two types of dry forest. The more significant difference between the two forest types in the phytolith assemblages (Fig. 2A) suggests that there is more significant variation in the understorey vegetation of these forests, as phytoliths are well suited for differentiation of herbaceous taxa ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"author":[{"dropping-particle":"","family":"Piperno","given":"Dolores R.","non-dropping-particle":"","parse-names":false,"suffix":""}],"id":"ITEM-1","issued":{"date-parts":[["2006"]]},"publisher":"Altamire Press","title":"Phytoliths: A comprehensive guide for Archeologists and Paleoecologists","type":"book"},"uris":[""]}],"mendeley":{"formattedCitation":"(Piperno, 2006)","manualFormatting":"(Piperno, 2006; Plumpton et al., in submission)","plainTextFormattedCitation":"(Piperno, 2006)","previouslyFormattedCitation":"(Piperno, 2006)"},"properties":{"noteIndex":0},"schema":""}(Piperno, 2006). Given the variation in inundation regime, variation in understorey vegetation would be expected. Unfortunately, the vegetation inventories for forest plots did not sample herbaceous taxa so this cannot be empirically tested. However, these results suggest that phytoliths may be more sensitive to the variability between different dry forest types, due to their ability to detect variability in the understorey vegetation. Therefore, to detect changes in inundation regime within dry forest ecosystems, phytoliths may be a better proxy choice than pollen. Within evergreen forest, pollen assemblages show higher variability than phytolith assemblages (Fig 2C, D). This likely reflects the ability of pollen to differentiate a wide range of arboreal taxa common to these plant communities, such as Moraceae, Alchornea and Melastomataceae/Combretaceae. As phytoliths generally only produce general arboreal indicators, such as granulate globulars, they are not able to pick up the spatial variability in tree species within these plots. In contrast to the dry forest, evergreen forests tend to have dense canopies leading to less well-developed understorey vegetation ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"author":[{"dropping-particle":"","family":"Killeen","given":"T.J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Schulenberg","given":"T.S.","non-dropping-particle":"","parse-names":false,"suffix":""}],"id":"ITEM-1","issued":{"date-parts":[["1998"]]},"publisher":"Conservation International","publisher-place":"Washington, DC","title":"A biological assessment of Parc Nacional Noel Kempff Mercado, Bolivia, RAP Working Papers 10","type":"book"},"uris":[""]}],"mendeley":{"formattedCitation":"(Killeen and Schulenberg, 1998)","plainTextFormattedCitation":"(Killeen and Schulenberg, 1998)","previouslyFormattedCitation":"(Killeen and Schulenberg, 1998)"},"properties":{"noteIndex":0},"schema":""}(Killeen and Schulenberg, 1998). Therefore, there is scarce herbaceous variability for the phytolith record to record within evergreen forest. These results show that pollen is more sensitive to variability within evergreen forest than phytoliths, due to their ability to detect changes in arboreal floristic composition. Within the savannah communities, pollen and phytolith assemblages show similar levels of variability (Fig 2C, D), although phytoliths are more sensitive to variation in terra firme upland savannah, and pollen more sensitive to variation in seasonally-inundated savannah (Fig. 2A, B). This reflects the complex mixture of both arboreal and herbaceous vegetation which forms the majority of savannah plant communities in south-west Amazonia. Cerrado savannah physiognomies range from closed woodland (cerrad?o) to open grassland with no trees or shrubs (campo limpo) ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"author":[{"dropping-particle":"","family":"Oliveira Filho","given":"A.T.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Ratter","given":"J.A.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"The cerrados of Brazil. Ecology an natural history of a Neotropical savanna (P.S. Oliveira & R.J. Marquis, eds.). Columbia University Press, New York.","id":"ITEM-1","issued":{"date-parts":[["2002"]]},"page":"91-120","title":"Vegetation physiognomies and woody flora of the cerrado biome.","type":"chapter"},"uris":[""]}],"mendeley":{"formattedCitation":"(Oliveira Filho and Ratter, 2002)","plainTextFormattedCitation":"(Oliveira Filho and Ratter, 2002)","previouslyFormattedCitation":"(Oliveira Filho and Ratter, 2002)"},"properties":{"noteIndex":0},"schema":""}(Oliveira Filho and Ratter, 2002). The cerrado savannah plot here represents open woodland savannah (campo cerrado) ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.revpalbo.2011.01.001","ISSN":"00346667","abstract":"The majority of vegetation reconstructions from the Neotropics are derived from fossil pollen records extracted from lake sediments. However, the interpretation of these records is restricted by limited knowledge of the contemporary relationships between the vegetation and pollen rain of Neotropical ecosystems, especially for more open vegetation such as savannas. This research aims to improve the interpretation of these records by investigating the vegetation and modern pollen rain of different savanna ecosystems in Bolivia using vegetation inventories, artificial pollen traps and surface lake sediments. Two types of savanna were studied, upland savannas (cerrado), occurring on well drained soils, and seasonally-inundated savannas occurring on seasonally water-logged soils. Quantitative vegetation data are used to identify taxa that are floristically important in the different savanna types and to allow modern pollen/vegetation ratios to be calculated. Artificial pollen traps from the upland savanna site are dominated by Moraceae (35%), Poaceae (30%), Alchornea (6%) and Cecropia (4%). The two seasonally-inundated savanna sites are dominated by Moraceae (37%), Poaceae (20%), Alchornea (8%) and Cecropia (7%), and Moraceae (25%), Cyperaceae (22%), Poaceae (19%) and Cecropia (9%), respectively. The modern pollen rain of seasonally-inundated savannas from surface lake sediments is dominated by Cyperaceae (35%), Poaceae (33%), Moraceae (9%) and Asteraceae (5%). Upland and seasonally-flooded savannas were found to be only subtly distinct from each other palynologically. All sites have a high proportion of Moraceae pollen due to effective wind dispersal of this pollen type from areas of evergreen forest close to the study sites. Modern pollen/vegetation ratios show that many key woody plant taxa are absent/under-represented in the modern pollen rain (e.g., Caryocar and Tabebuia). The lower-than-expected percentages of Poaceae pollen, and the scarcity of savanna indicators, in the modern pollen rain of these ecosystems mean that savannas could potentially be overlooked in fossil pollen records without consideration of the full pollen spectrum available.","author":[{"dropping-particle":"","family":"Jones","given":"Huw T.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Pennington","given":"R. Toby","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Review of Palaeobotany and Palynology","id":"ITEM-1","issue":"3-4","issued":{"date-parts":[["2011","4"]]},"page":"223-237","title":"Characterisation of Bolivian savanna ecosystems by their modern pollen rain and implications for fossil pollen records","type":"article-journal","volume":"164"},"uris":[""]}],"mendeley":{"formattedCitation":"(Jones et al., 2011)","plainTextFormattedCitation":"(Jones et al., 2011)","previouslyFormattedCitation":"(Jones et al., 2011)"},"properties":{"noteIndex":0},"schema":""}(Jones et al., 2011). Within seasonally inundated savannahs, low-elevation areas are almost entirely dominated by grasses, while trees and shrubs are found on raised islands, particularly termite mounds (pampa termitero) in NKMNP ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"author":[{"dropping-particle":"","family":"Killeen","given":"T.J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Schulenberg","given":"T.S.","non-dropping-particle":"","parse-names":false,"suffix":""}],"id":"ITEM-1","issued":{"date-parts":[["1998"]]},"publisher":"Conservation International","publisher-place":"Washington, DC","title":"A biological assessment of Parc Nacional Noel Kempff Mercado, Bolivia, RAP Working Papers 10","type":"book"},"uris":[""]}],"mendeley":{"formattedCitation":"(Killeen and Schulenberg, 1998)","plainTextFormattedCitation":"(Killeen and Schulenberg, 1998)","previouslyFormattedCitation":"(Killeen and Schulenberg, 1998)"},"properties":{"noteIndex":0},"schema":""}(Killeen and Schulenberg, 1998). Pollen may therefore be capturing the variability in savannah trees, both in upland cerrado formations and on islands within seasonally inundated savannah but does not provide the taxonomic resolution to capture variability in grass genera and subfamilies. In contrast, phytoliths will capture the variability in the grasses and herbaceous taxa but not in the majority of the arboreal taxa ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"author":[{"dropping-particle":"","family":"Piperno","given":"Dolores R.","non-dropping-particle":"","parse-names":false,"suffix":""}],"id":"ITEM-1","issued":{"date-parts":[["2006"]]},"publisher":"Altamire Press","title":"Phytoliths: A comprehensive guide for Archeologists and Paleoecologists","type":"book"},"uris":[""]}],"mendeley":{"formattedCitation":"(Piperno, 2006)","plainTextFormattedCitation":"(Piperno, 2006)","previouslyFormattedCitation":"(Piperno, 2006)"},"properties":{"noteIndex":0},"schema":""}(Piperno, 2006). Phytoliths may therefore be well suited to detecting local changes such as the inundation regime in savannas while pollen may be a better choice of proxy for detection of regional climatic changes such as reductions in precipitation ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"author":[{"dropping-particle":"","family":"Piperno","given":"Dolores R.","non-dropping-particle":"","parse-names":false,"suffix":""}],"id":"ITEM-1","issued":{"date-parts":[["2006"]]},"publisher":"Altamire Press","title":"Phytoliths: A comprehensive guide for Archeologists and Paleoecologists","type":"book"},"uris":[""]},{"id":"ITEM-2","itemData":{"DOI":"10.1177/0959683615581204","ISBN":"0959-6836","ISSN":"0959-6836","abstract":"The nature and extent of pre-Columbian (pre-AD 1492) human impact in Amazonia is a contentious issue. The Bolivian Amazon has yielded some of the most impressive evidence for large and complex pre-Columbian societies in the Amazon basin, yet there remains relatively little data concerning the land use of these societies over time. Palaeoecology, when integrated with archaeological data, has the potential to fill these gaps in our knowledge. We present a 6000-year record of anthropogenic burning, agriculture and vegetation change, from an oxbow lake located adjacent to a pre-Columbian ring ditch in north-east Bolivia (13°15′44″S, 63°42′37″W). Human occupation around the lake site is inferred from pollen and phytoliths of maize (Zea mays L.) and macroscopic charcoal evidence of anthropogenic burning. First occupation around the lake was radiocarbon dated to ~2500 calibrated years before present (BP). The persistence of maize in the record from ~1850 BP suggests that it was an important crop grown in the ring-ditch region in pre-Columbian times, and abundant macroscopic charcoal suggests that pre-Columbian land management entailed more extensive burning of the landscape than the slash-and-burn agriculture practised around the site today. The site was occupied continuously until near-modern times, although there is evidence for a decline in agricultural intensity or change in land-use strategy, and possible population decline, from ~600–500 BP. The long and continuous occupation, which predates the establishment of rainforest in the region, suggests that pre-Columbian land use may have had a significant influence on ecosystem development at this site over the last ~2000 years.","author":[{"dropping-particle":"","family":"Carson","given":"J. F.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Watling","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"F. E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Whitney","given":"B. S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Prumers","given":"H.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Soto","given":"J. D.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"The Holocene","id":"ITEM-2","issue":"8","issued":{"date-parts":[["2015","5","1"]]},"note":"From Duplicate 2 (Pre-Columbian land use in the ring-ditch region of the Bolivian Amazon - Carson, J. F.; Watling, J.; Mayle, F. E.; Whitney, B. S.; Iriarte, J.; Prumers, H.; Soto, J. D.)\n\nUsed phytoliths and pollen from oxbow lake in NE Bolivia - oxbow 6000 yrs old.\n\nNice passage about phytoliths from lakes and their source area compared to pollen - assumes local representation due to modes of transport available.","page":"1285-1300","title":"Pre-Columbian land use in the ring-ditch region of the Bolivian Amazon","type":"article-journal","volume":"25"},"uris":[""]},{"id":"ITEM-3","itemData":{"DOI":"10.1016/j.yqres.2013.06.005","ISSN":"00335894","abstract":"We present a multiproxy study of land use by a pre-Columbian earth mounds culture in the Bolivian Amazon. The Monumental Mounds Region (MMR) is an archaeological sub-region characterized by hundreds of pre-Columbian habitation mounds associated with a complex network of canals and causeways, and situated in the forest–savanna mosaic of the Llanos de Moxos. Pollen, phytolith, and charcoal analyses were performed on a sediment core from a large lake (14km2), Laguna San José (14°56.97′S, 64°29.70′W). We found evidence of high levels of anthropogenic burning from AD 400 to AD 1280, corroborating dated occupation layers in two nearby excavated habitation mounds. The charcoal decline pre-dates the arrival of Europeans by at least 100yr, and challenges the notion that the mounds culture declined because of European colonization. We show that the surrounding savanna soils were sufficiently fertile to support crops, and the presence of maize throughout the record shows that the area was continuously cultivated despite land-use change at the end of the earth mounds culture. We suggest that burning was largely confined to the savannas, rather than forests, and that pre-Columbian deforestation was localized to the vicinity of individual habitation mounds, whereas the inter-mound areas remained largely forested.","author":[{"dropping-particle":"","family":"Whitney","given":"Bronwen S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Dickau","given":"Ruth","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Soto","given":"J. Daniel","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"José","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Quaternary Research","id":"ITEM-3","issue":"2","issued":{"date-parts":[["2013"]]},"note":"From Duplicate 1 (Pre-Columbian landscape impact and agriculture in the Monumental Mound region of the Llanos de Moxos, lowland Bolivia - Whitney, Bronwen S.; Dickau, Ruth; Mayle, Francis E.; Soto, J. Daniel; Iriarte, José)\n\nPhytolith-lake paper. San jose 14km2. assume phytolith source area more local than pollen - backed up by comparison of modern veg inventory with surface phytolith assemblage.","page":"207-217","title":"Pre-Columbian landscape impact and agriculture in the Monumental Mound region of the Llanos de Moxos, lowland Bolivia","type":"article-journal","volume":"80"},"uris":[""]}],"mendeley":{"formattedCitation":"(Carson et al., 2015; Piperno, 2006; Whitney et al., 2013)","plainTextFormattedCitation":"(Carson et al., 2015; Piperno, 2006; Whitney et al., 2013)","previouslyFormattedCitation":"(Carson et al., 2015; Piperno, 2006; Whitney et al., 2013)"},"properties":{"noteIndex":0},"schema":""}(Carson et al., 2015; Piperno, 2006; Whitney et al., 2013).. However, to fully investigate savannah dynamics both pollen and phytoliths should be used together to provide taxonomic detail on both the herbaceous and arboreal components of the ecosystem (Dickau et al., 2013).5.2 Modern analogues for fossil samplesThe overlap between the modern and fossil phytolith samples in the PCA (Fig 3B) suggests the modern samples represent good analogues for fossil assemblages, and supports the use of thresholds based on the modern phytolith samples to infer ecosystem turnover in the fossil record. The overlap for modern and fossil pollen samples is less strong (Fig 3A). The modern pollen samples do show good overlap in ordination space with the most recent (2,500 calyrBP to present) fossil samples from Laguna Chaplin, but less overlap with the older Chaplin samples or the Laguna Granja fossil samples. L. Granja is situated ~250km to the north west of NKMNP and L. Chaplin (Fig. 1). L. Granja is situated at the ecotone between humid evergreen forest to the north and the Beni seasonally-inundated savannah- forest mosaic to the south (Fig. 1). The savannah of the Beni basin has a distinct vegetation composition whose pollen rain differs from seasonally-inundated savannah communities in NKMNP: Beni basin pollen rain contains a higher proportion of Poaceae and Cyperaceae and lower Moraceae/Urticaceae than NKMNP pollen rain (Jones et al., 2011). The lack of ordination overlap between L. Granja pollen and modern NKMNP pollen suggests that modern samples specific to the local vegetation of a study site should be used where possible to develop thresholds for floristic and ecosystem turnover, particularly for pollen due to the high diversity of pollen types. For phytoliths, the need for local samples may be less important as there is a lower diversity of phytolith types. 5.3 Thresholds for floristic and ecosystem turnover in the palaeo-recordThe pollen Q1 threshold for weak indication of floristic turnover within savannah was met several times through the pairwise downcore comparisons at L. Chaplin (Fig. 4). However, the pairwise distances downcore do not meet the thresholds for stronger floristic turnover or ecosystem turnover at 2,000 calyrBP, the time at which ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.yqres.2003.12.004","ISSN":"00335894","abstract":"Pollen and charcoal records from two large, shallow lakes reveal that throughout most of the past 50,000 yr Noel Kempff Mercado National Park, in northeastern lowland Bolivia (southwestern Amazon Basin), was predominantly covered by savannas and seasonally dry semideciduous forests. Lowered atmospheric CO2 concentrations, in combination with a longer dry season, caused expansion of dry forests and savannas during the last glacial period, especially at the last glacial maximum. These ecosystems persisted until the mid-Holocene, although they underwent significant species reassortment. Forest communities containing a mixture of evergreen and semideciduous species began to expand between 6000 and 3000 14C yr B.P. Humid evergreen rain forests expanded to cover most of the area within the past 2000 14C yr B.P., coincident with a reduction in fire frequencies. Comparisons between modern pollen spectra and vegetation reveal that the Moraceae-dominated rain forest pollen spectra likely have a regional source area at least 2–3 km beyond the lake shore, whereas the grass- and sedge-dominated savanna pollen spectra likely have a predominantly local source area. The Holocene vegetation changes are consistent with independent paleoprecipitation records from the Bolivian Altiplano and paleovegetation records from other parts of southwestern Amazonia. The progressive expansion in rain forests through the Holocene can be largely attributed to enhanced convective activity over Amazonia, due to greater seasonality of insolation in the Southern Hemisphere tropics driven by the precession cycle according to the Milankovitch Astronomical Theory.","author":[{"dropping-particle":"","family":"Burbridge","given":"Rachel E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Quaternary Research","id":"ITEM-1","issue":"2","issued":{"date-parts":[["2004","3"]]},"note":"chapling and Bella Vista records - dry forest/savanna until 2000 bp when precip increased - humid evergreen","page":"215-230","title":"Fifty-thousand-year vegetation and climate history of Noel Kempff Mercado National Park, Bolivian Amazon","type":"article-journal","volume":"61"},"uris":[""]}],"mendeley":{"formattedCitation":"(Burbridge et al., 2004)","manualFormatting":"Burbridge et al., (2004)","plainTextFormattedCitation":"(Burbridge et al., 2004)","previouslyFormattedCitation":"(Burbridge et al., 2004)"},"properties":{"noteIndex":0},"schema":""}Burbridge et al., (2004) interpret turnover from savannah and dry forest to evergreen forest. This may reflect the fact that the vegetation changes are gradual transitions that are not captured in a single pairwise distance measure. When the full matrix of distances between all samples at all depths was compared, the Q2 threshold for good indication of ecosystem turnover was met by comparison of the top section from 3,000 calyrBP to present with the majority of the rest of the core. This is close to the timing of the ecosystem turnover identified by qualitative interpretation of the pollen record. At L. Granja, the pollen Q2 threshold for good indication of floristic turnover was met by the majority of the pairwise comparisons downcore, indicating evidence of floristic turnover within the savannah ecosystem throughout the majority of the record (Fig. 4). However, the pairwise distances downcore do not meet the thresholds for strong (Q3) floristic or ecosystem turnover at 500 calyrBP when ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1177/0959683615581204","ISBN":"0959-6836","ISSN":"0959-6836","abstract":"The nature and extent of pre-Columbian (pre-AD 1492) human impact in Amazonia is a contentious issue. The Bolivian Amazon has yielded some of the most impressive evidence for large and complex pre-Columbian societies in the Amazon basin, yet there remains relatively little data concerning the land use of these societies over time. Palaeoecology, when integrated with archaeological data, has the potential to fill these gaps in our knowledge. We present a 6000-year record of anthropogenic burning, agriculture and vegetation change, from an oxbow lake located adjacent to a pre-Columbian ring ditch in north-east Bolivia (13°15′44″S, 63°42′37″W). Human occupation around the lake site is inferred from pollen and phytoliths of maize (Zea mays L.) and macroscopic charcoal evidence of anthropogenic burning. First occupation around the lake was radiocarbon dated to ~2500 calibrated years before present (BP). The persistence of maize in the record from ~1850 BP suggests that it was an important crop grown in the ring-ditch region in pre-Columbian times, and abundant macroscopic charcoal suggests that pre-Columbian land management entailed more extensive burning of the landscape than the slash-and-burn agriculture practised around the site today. The site was occupied continuously until near-modern times, although there is evidence for a decline in agricultural intensity or change in land-use strategy, and possible population decline, from ~600–500 BP. The long and continuous occupation, which predates the establishment of rainforest in the region, suggests that pre-Columbian land use may have had a significant influence on ecosystem development at this site over the last ~2000 years.","author":[{"dropping-particle":"","family":"Carson","given":"J. F.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Watling","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"F. E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Whitney","given":"B. S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Prumers","given":"H.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Soto","given":"J. D.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"The Holocene","id":"ITEM-1","issue":"8","issued":{"date-parts":[["2015","5","1"]]},"note":"From Duplicate 2 (Pre-Columbian land use in the ring-ditch region of the Bolivian Amazon - Carson, J. F.; Watling, J.; Mayle, F. E.; Whitney, B. S.; Iriarte, J.; Prumers, H.; Soto, J. D.)\n\nUsed phytoliths and pollen from oxbow lake in NE Bolivia - oxbow 6000 yrs old.\n\nNice passage about phytoliths from lakes and their source area compared to pollen - assumes local representation due to modes of transport available.","page":"1285-1300","title":"Pre-Columbian land use in the ring-ditch region of the Bolivian Amazon","type":"article-journal","volume":"25"},"uris":[""]}],"mendeley":{"formattedCitation":"(Carson et al., 2015)","manualFormatting":"Carson et al., (2015)","plainTextFormattedCitation":"(Carson et al., 2015)","previouslyFormattedCitation":"(Carson et al., 2015)"},"properties":{"noteIndex":0},"schema":""}Carson et al., (2015) identify ecosystem turnover from savannah to evergreen forest. When the full matrix of distances between all samples at all depths was compared, the Q2 threshold for good indication of floristic turnover within savannah was met by comparison of the last 500 years to present with the rest of the core, which matches the timing of the transition identified from qualitative interpretation of the pollen record. However, the transition at 500 calyrBP did not meet any of the thresholds for ecosystem turnover from savannah to evergreen forest. This may be due to the spatial scale of the change within the catchment of the lake. At present L. Granja is at the very edge of the Amazonian evergreen forest, at the boundary with seasonally-inundated savannah of the Beni basin. Therefore, within the catchment of L. Granja, there would not have been a complete transition from savannah to evergreen forest as the Beni is hydrologically controlled and would have remained savanna-dominated throughout. In comparison, at present L. Chaplin is surrounded almost entirely by evergreen forest, so the transition in the catchment of the lake would have been more significant. These findings demonstrate the importance of considering the local characteristics of the site and heterogeneity of the surrounding landscape in interpretation of the thresholds. Both the spatial and temporal elements of the vegetation change are captured by these thresholds. In the phytolith record at L. Granja, the Q3 threshold for strong indication of floristic turnover was met at the base of the core, not at the depths identified for vegetation change by the authors ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1177/0959683615581204","ISBN":"0959-6836","ISSN":"0959-6836","abstract":"The nature and extent of pre-Columbian (pre-AD 1492) human impact in Amazonia is a contentious issue. The Bolivian Amazon has yielded some of the most impressive evidence for large and complex pre-Columbian societies in the Amazon basin, yet there remains relatively little data concerning the land use of these societies over time. Palaeoecology, when integrated with archaeological data, has the potential to fill these gaps in our knowledge. We present a 6000-year record of anthropogenic burning, agriculture and vegetation change, from an oxbow lake located adjacent to a pre-Columbian ring ditch in north-east Bolivia (13°15′44″S, 63°42′37″W). Human occupation around the lake site is inferred from pollen and phytoliths of maize (Zea mays L.) and macroscopic charcoal evidence of anthropogenic burning. First occupation around the lake was radiocarbon dated to ~2500 calibrated years before present (BP). The persistence of maize in the record from ~1850 BP suggests that it was an important crop grown in the ring-ditch region in pre-Columbian times, and abundant macroscopic charcoal suggests that pre-Columbian land management entailed more extensive burning of the landscape than the slash-and-burn agriculture practised around the site today. The site was occupied continuously until near-modern times, although there is evidence for a decline in agricultural intensity or change in land-use strategy, and possible population decline, from ~600–500 BP. The long and continuous occupation, which predates the establishment of rainforest in the region, suggests that pre-Columbian land use may have had a significant influence on ecosystem development at this site over the last ~2000 years.","author":[{"dropping-particle":"","family":"Carson","given":"J. F.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Watling","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"F. E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Whitney","given":"B. S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Prumers","given":"H.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Soto","given":"J. D.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"The Holocene","id":"ITEM-1","issue":"8","issued":{"date-parts":[["2015","5","1"]]},"note":"From Duplicate 2 (Pre-Columbian land use in the ring-ditch region of the Bolivian Amazon - Carson, J. F.; Watling, J.; Mayle, F. E.; Whitney, B. S.; Iriarte, J.; Prumers, H.; Soto, J. D.)\n\nUsed phytoliths and pollen from oxbow lake in NE Bolivia - oxbow 6000 yrs old.\n\nNice passage about phytoliths from lakes and their source area compared to pollen - assumes local representation due to modes of transport available.","page":"1285-1300","title":"Pre-Columbian land use in the ring-ditch region of the Bolivian Amazon","type":"article-journal","volume":"25"},"uris":[""]}],"mendeley":{"formattedCitation":"(Carson et al., 2015)","plainTextFormattedCitation":"(Carson et al., 2015)","previouslyFormattedCitation":"(Carson et al., 2015)"},"properties":{"noteIndex":0},"schema":""}(Carson et al., 2015). The phytolith record at the depths where the Q3 floristic turnover threshold was reached shows high abundance of Asteraceae phytoliths in particular. This single taxon appears to be driving the large differences between these samples and the rest of the core. This highlights the importance of considering the taxonomic detail of the changes in the record alongside the level of variation in assemblage to identify ecosystem turnover; not all taxa are weighted equally in the qualitative interpretation of palaeo-records. However, when all samples at all depths are compared, the top section of the core from 600 calyrBP to present meets the Q1 threshold for weak indication of floristic turnover compared to the rest of the core. This fits the timing of the closing of the forest canopy around the lake edge identified by ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1177/0959683615581204","ISBN":"0959-6836","ISSN":"0959-6836","abstract":"The nature and extent of pre-Columbian (pre-AD 1492) human impact in Amazonia is a contentious issue. The Bolivian Amazon has yielded some of the most impressive evidence for large and complex pre-Columbian societies in the Amazon basin, yet there remains relatively little data concerning the land use of these societies over time. Palaeoecology, when integrated with archaeological data, has the potential to fill these gaps in our knowledge. We present a 6000-year record of anthropogenic burning, agriculture and vegetation change, from an oxbow lake located adjacent to a pre-Columbian ring ditch in north-east Bolivia (13°15′44″S, 63°42′37″W). Human occupation around the lake site is inferred from pollen and phytoliths of maize (Zea mays L.) and macroscopic charcoal evidence of anthropogenic burning. First occupation around the lake was radiocarbon dated to ~2500 calibrated years before present (BP). The persistence of maize in the record from ~1850 BP suggests that it was an important crop grown in the ring-ditch region in pre-Columbian times, and abundant macroscopic charcoal suggests that pre-Columbian land management entailed more extensive burning of the landscape than the slash-and-burn agriculture practised around the site today. The site was occupied continuously until near-modern times, although there is evidence for a decline in agricultural intensity or change in land-use strategy, and possible population decline, from ~600–500 BP. The long and continuous occupation, which predates the establishment of rainforest in the region, suggests that pre-Columbian land use may have had a significant influence on ecosystem development at this site over the last ~2000 years.","author":[{"dropping-particle":"","family":"Carson","given":"J. F.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Watling","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"F. E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Whitney","given":"B. S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Prumers","given":"H.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Soto","given":"J. D.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"The Holocene","id":"ITEM-1","issue":"8","issued":{"date-parts":[["2015","5","1"]]},"note":"From Duplicate 2 (Pre-Columbian land use in the ring-ditch region of the Bolivian Amazon - Carson, J. F.; Watling, J.; Mayle, F. E.; Whitney, B. S.; Iriarte, J.; Prumers, H.; Soto, J. D.)\n\nUsed phytoliths and pollen from oxbow lake in NE Bolivia - oxbow 6000 yrs old.\n\nNice passage about phytoliths from lakes and their source area compared to pollen - assumes local representation due to modes of transport available.","page":"1285-1300","title":"Pre-Columbian land use in the ring-ditch region of the Bolivian Amazon","type":"article-journal","volume":"25"},"uris":[""]}],"mendeley":{"formattedCitation":"(Carson et al., 2015)","plainTextFormattedCitation":"(Carson et al., 2015)","previouslyFormattedCitation":"(Carson et al., 2015)"},"properties":{"noteIndex":0},"schema":""}(Carson et al., 2015). The lower threshold for floristic change (Q1) is therefore capturing the small scale of vegetation change reflected in the phytolith record.These findings demonstrate the importance of this quantitative approach for assessing the magnitude of vegetation change in the palaeo-record. Use of these thresholds enables differentiation between the magnitude of variation that represents floristic turnover within an ecosystem from that which represents turnover between ecosystems. At L. Chaplin, the vegetation changes between 3,000 and 2,000 calyrBP are of high enough magnitude to be identified as a good indication of ecosystem turnover. However, the changes in the pollen record at L. Granja at 500 calyrBP are only high enough to be identified as a good indication of floristic turnover within an ecosystem. The changes in the phytolith record at L. Granja at 600 calyrBP only represent a weak indication of floristic turnover within an ecosystem. These thresholds therefore provide quantitative distinctions between different scales of turnover in the palaeo-record and enable differentiation of ecosystem turnover from floristic turnover within the ecosystem. However, it is important to consider that these thresholds were only able to capture floristic turnover or ecosystem turnover by comparison of the full matrix of distances between all samples at all depths. Pairwise downcore distances mostly did not meet the thresholds for floristic or ecosystem turnover at the depths that qualitative interpretation of the records suggested turnover was taking place. This may reflect the gradual nature of these vegetation transitions. Previous research has demonstrated that lowland south-western Amazonian sites, such as L. Chaplin, have low rates of ecological change ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1111/geb.12694","ISSN":"1466822X","author":[{"dropping-particle":"","family":"Costa","given":"Gabriel C.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Hampe","given":"Arndt","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Ledru","given":"Marie-Pierre","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Martinez","given":"Pablo A.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mazzochini","given":"Guilherme G.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Shepard","given":"Donald B.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Werneck","given":"Fernanda P.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Moritz","given":"Craig","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Carnaval","given":"Ana Carolina","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Global Ecology and Biogeography","id":"ITEM-1","issue":"3","issued":{"date-parts":[["2018","3","1"]]},"page":"285-297","publisher":"John Wiley & Sons, Ltd (10.1111)","title":"Biome stability in South America over the last 30 kyr: Inferences from long-term vegetation dynamics and habitat modelling","type":"article-journal","volume":"27"},"uris":[""]}],"mendeley":{"formattedCitation":"(Costa et al., 2018)","plainTextFormattedCitation":"(Costa et al., 2018)","previouslyFormattedCitation":"(Costa et al., 2018)"},"properties":{"noteIndex":0},"schema":""}(Costa et al., 2018). Therefore sampling resolution and sedimentation rate will strongly influence the magnitude of change captured by single pairwise distances downcore. Sampling resolution and sedimentation rate can be accounted for by calculating the rate of ecological change for fossil samples ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1007/978-90-481-2672-9_12","abstract":"An analysis of rates of ecological change (RoC) from thirteen pollen records from tropical South America is presented here. The analysis aims to identify the periods of fastest change since the last glacial maximum (LGM) and possible driving mechanisms. Despite rapid cooling periods, region-wide profound droughts, fire and human disturbances, RoC analysis showed that the speed of these climate changes never exceed the species response capabilities. Our results legitimize concerns regarding the resilience of species to accommodate future change and emphasize the urgency for integrative environmental measures.","author":[{"dropping-particle":"","family":"Urrego","given":"Dunia H","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Bush","given":"Mark B","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Silman","given":"Miles R","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Correa-Metrio","given":"Alexander","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Ledru","given":"Marie-Pierre","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Paduano","given":"Gina","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Valencia","given":"Bryan G","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Past climate variability in South America and surrounding regions","id":"ITEM-1","issued":{"date-parts":[["2009"]]},"page":"283-300","title":"Millennial-Scale Ecological Changes in Tropical South America Since the Last Glacial Maximum","type":"chapter"},"uris":[""]}],"mendeley":{"formattedCitation":"(Urrego et al., 2009)","plainTextFormattedCitation":"(Urrego et al., 2009)","previouslyFormattedCitation":"(Urrego et al., 2009)"},"properties":{"noteIndex":0},"schema":""}(Urrego et al., 2009) but this is not possible for modern samples. As the methodology of this paper relies on comparison of modern proxy variability to fossil proxy variability, sampling resolution and sedimentation rate needs to be accounted for by comparison of the full matrix of distances between all samples at all depths in the palaeo-record. Future development of this quantitative methodology for identifying ecosystem turnover in the palaeo-record relies on development and expansion of modern training datasets of pollen and phytolith variability to set thresholds. The NKMNP dataset used in this study is currently the only dataset in tropical South America with both pollen and phytolith data from permanent vegetation study plots within evergreen forest, dry forest and savannah. One outstanding question is whether the thresholds calculated from the NKMNP data are applicable to palaeo-records from other parts of south-western Amazonia. Despite the lack of overlap in ordination space between the modern pollen samples from NKMNP and the fossil samples from L. Granja (Fig. 3A), the thresholds were still able to capture the vegetation turnover at 500 calyrBP in the palaeo-record. Applicability of these thresholds could be further tested through study of palaeo-sites from outside NKMNP. Applying these thresholds to additional sites where ecosystem turnover has been interpreted from the fossil record would also enable a sensitivity analysis of the threshold levels used here. In this pilot investigation, thresholds we set based on the 25th, 50th and 75th percentile of variation in the modern datasets. These threshold levels could be refined through testing on additional palaeo-sites. Applying this approach to numerous palaeo-sites may also improve our understanding of whether different drivers of floristic or ecosystem turnover produce different levels of variability in taxonomic composition. For example, does fire-induced vegetation change produce the same amount of taxonomic variability as flood-induced vegetation change or human-induced vegetation change? The high variability in rates of ecological change found in palaeo-records across tropical South America in the last 3,000 years has been linked to human-induced vegetation change ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1007/978-90-481-2672-9_12","abstract":"An analysis of rates of ecological change (RoC) from thirteen pollen records from tropical South America is presented here. The analysis aims to identify the periods of fastest change since the last glacial maximum (LGM) and possible driving mechanisms. Despite rapid cooling periods, region-wide profound droughts, fire and human disturbances, RoC analysis showed that the speed of these climate changes never exceed the species response capabilities. Our results legitimize concerns regarding the resilience of species to accommodate future change and emphasize the urgency for integrative environmental measures.","author":[{"dropping-particle":"","family":"Urrego","given":"Dunia H","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Bush","given":"Mark B","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Silman","given":"Miles R","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Correa-Metrio","given":"Alexander","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Ledru","given":"Marie-Pierre","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Paduano","given":"Gina","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Valencia","given":"Bryan G","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Past climate variability in South America and surrounding regions","id":"ITEM-1","issued":{"date-parts":[["2009"]]},"page":"283-300","title":"Millennial-Scale Ecological Changes in Tropical South America Since the Last Glacial Maximum","type":"chapter"},"uris":[""]}],"mendeley":{"formattedCitation":"(Urrego et al., 2009)","plainTextFormattedCitation":"(Urrego et al., 2009)"},"properties":{"noteIndex":0},"schema":""}(Urrego et al., 2009). Patterns of variability in the palaeo-record may therefore help distinguish between climate-induced vegetation change, which may show more gradual variation, and human-induced vegetation change.6. ConclusionsThis research aimed to compare the sensitivity of pollen and phytolith assemblages to changes within and between three tropical South American ecosystems: evergreen forest, dry forest and savannah. Pollen is more sensitive than phytoliths to variation in evergreen forest due to pollen’s superior ability to differentiate arboreal taxa which dominate this forest type. Phytoliths are slightly more sensitive than pollen to variation in dry forest due to their greater ability to capture changes in understorey herbaceous taxa associated with differences in canopy density. Phytoliths may be particularly sensitive to changes in inundation regime in dry forests. Both pollen and phytoliths are equally sensitive to variation in savannas due to the complex mixture of arboreal and herbaceous taxa in these communities. Phytoliths may be best suited to investigate changes in inundation regime affecting herbs and grasses in low-elevation areas, while pollen may be best suited to investigate climatic changes affecting arboreal composition within savannas. These findings support the suggestion that the two proxies provide different, complementary taxonomic information and should be used together to improve our understanding of these complex ecosystem dynamics ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.revpalbo.2013.01.004","ISSN":"00346667","abstract":"The interpretation of Neotropical fossil phytolith assemblages for palaeoenvironmental and archaeological reconstructions relies on the development of appropriate modern analogues. We analyzed modern phytolith assemblages from the soils of ten distinctive tropical vegetation communities in eastern lowland Bolivia, ranging from terra firme humid evergreen forest to seasonally-inundated savannah. Results show that broad ecosystems – evergreen tropical forest, semi-deciduous dry tropical forest, and savannah – can be clearly differentiated by examination of their phytolith spectra and the application of Principal Component Analysis (PCA). Differences in phytolith assemblages between particular vegetation communities within each of these ecosystems are more subtle, but can still be identified. Comparison of phytolith assemblages with pollen rain data and stable carbon isotope analyses from the same vegetation plots show that these proxies are not only complementary, but significantly improve taxonomic and ecosystem resolution, and therefore our ability to interpret palaeoenvironmental and archaeological records. Our data underline the utility of phytolith analyses for reconstructing Amazon Holocene vegetation histories and pre-Columbian land use, particularly the high spatial resolution possible with terrestrial soil-based phytolith studies.","author":[{"dropping-particle":"","family":"Dickau","given":"Ruth","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Whitney","given":"Bronwen S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"José","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Soto","given":"J. Daniel","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Metcalfe","given":"Phil","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Street-Perrott","given":"F. Alayne","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Loader","given":"Neil J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Ficken","given":"Katherine J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Review of Palaeobotany and Palynology","id":"ITEM-1","issued":{"date-parts":[["2013","6"]]},"page":"15-37","publisher":"Elsevier B.V.","title":"Differentiation of neotropical ecosystems by modern soil phytolith assemblages and its implications for palaeoenvironmental and archaeological reconstructions","type":"article-journal","volume":"193"},"uris":[""]}],"mendeley":{"formattedCitation":"(Dickau et al., 2013)","manualFormatting":"(Dickau et al., 2013; Plumpton et al., submitted)","plainTextFormattedCitation":"(Dickau et al., 2013)","previouslyFormattedCitation":"(Dickau et al., 2013)"},"properties":{"noteIndex":0},"schema":""}(Dickau et al., 2013; Plumpton et al., submitted). Overall, this analysis shows that the relative sensitivity of the two proxies to changes within an ecosystem depends on the ecosystem type studied. These are important considerations for palaeoecologists selecting proxies for the study of ecosystem turnover in the palaeo-record. This research also aimed to use the variability in the proxy assemblages from evergreen forest, dry forest and savannah as a modern training dataset to define thresholds which could distinguish floristic change within an ecosystem from turnover between ecosystems. When applied to palaeo-records from two test sites, the thresholds reflected the changes identified by the authors of the records and largely differentiated ecosystem turnover from floristic turnover. The thresholds have been shown to provide quantitative distinctions between different scales of turnover in the palaeo-record. These results demonstrate the potential of this quantitative approach for assessing the magnitude of vegetation change in the palaeo-record and differentiating ecosystem turnover from floristic changes within an ecosystem. This quantitative approach is therefore a useful tool for palaeoecologists to apply alongside qualitative interpretation of the palaeo-record for identification of ecosystem turnover. AcknowledgementsFunding for the analysis in this paper was provided by a University of Reading PhD studentship (HP). We thank William Gosling, Michael Burn, Huw Jones and Ruth Dickau for permission to use their pollen and phytolith data from Noel Kempff Mercado National Park, and John Carson and Jenny Watling for permission to use their pollen and phytolith data from Laguna Granja. We also thank the reviewers for their useful comments and suggestions which helped to improve this manuscript.ReferencesADDIN Mendeley Bibliography CSL_BIBLIOGRAPHY Burbridge RE, Mayle FE and Killeen TJ (2004) Fifty-thousand-year vegetation and climate history of Noel Kempff Mercado National Park, Bolivian Amazon. Quaternary Research 61(2): 215–230. DOI: 10.1016/j.yqres.2003.12.004.Burn MJ, Mayle FE and Killeen TJ (2010) Pollen-based differentiation of Amazonian rainforest communities and implications for lowland palaeoecology in tropical South America. Palaeogeography, Palaeoclimatology, Palaeoecology 295(1–2): 1–18. DOI: 10.1016/j.palaeo.2010.05.009.Bush MB, Silman MR and Urrego DH (2004) 48,000 years of Climate and Forest Change in a Biodiversity Hot Spot.Carson JF, Watling J, Mayle FE, et al. (2015) Pre-Columbian land use in the ring-ditch region of the Bolivian Amazon. The Holocene 25(8): 1285–1300. 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(2013) Pre-Columbian landscape impact and agriculture in the Monumental Mound region of the Llanos de Moxos, lowland Bolivia. Quaternary Research 80(2): 207–217. DOI: 10.1016/j.yqres.2013.06.005.List of FiguresFigure 1 - Map of Noel Kempff Mercado National Park (NKMNP) showing major vegetation communities and location of plots MV1, AC2, CP1, LF1, CH1, CH2, FC1, TO1, FT1, and Lagunas Granja and Chaplin. Inset map of South American countries with red-shaded area showing location of NKMNP. Figure 2 – Boxplots of Hellinger distances. Plot A and B show distances within individual study plots for phytoliths and pollen respectively. CH – terra firme evergreen liana forest, LF – terra firme evergreen forest, AC – terra firme semi-deciduous dry forest, CP – seasonally-inundated semi-deciduous dry forest, MV – seasonally-inundated semi-deciduous dry forest, FC – terra firme savannah, FT – seasonally-inundated savannah, TO – seasonally-inundated savannah. Plots C and D show distances within ecosystem classes, plots E and F show distances between ecosystem classes within biomes.Figure 3 – Principal Component Analysis (PCA) biplots for modern and fossil samples. Panel A – modern samples from NKMNP plots, fossil samples from Laguna Granja and Laguna Chaplin. Panel B – modern samples from NKMNP plots and fossil samples from Laguna Granja. Colours represent sample location: evergreen forest plots from NKMNP are coloured dark green; dry forest plots from NKMNP are light green; savannah plots from NKMNP are yellow; fossil samples from Laguna Chaplin are violet; fossil samples from Laguna Granja are dark blue.Figure 4 – Stratigraphic plots of Hellinger distances between pairwise assemblages downcore at Laguna Granja (phytoliths and pollen) and Laguna Chaplin (pollen). Age-depth models taken from original papers ADDIN CSL_CITATION {"citationItems":[{"id":"ITEM-1","itemData":{"DOI":"10.1016/j.yqres.2003.12.004","ISSN":"00335894","abstract":"Pollen and charcoal records from two large, shallow lakes reveal that throughout most of the past 50,000 yr Noel Kempff Mercado National Park, in northeastern lowland Bolivia (southwestern Amazon Basin), was predominantly covered by savannas and seasonally dry semideciduous forests. Lowered atmospheric CO2 concentrations, in combination with a longer dry season, caused expansion of dry forests and savannas during the last glacial period, especially at the last glacial maximum. These ecosystems persisted until the mid-Holocene, although they underwent significant species reassortment. Forest communities containing a mixture of evergreen and semideciduous species began to expand between 6000 and 3000 14C yr B.P. Humid evergreen rain forests expanded to cover most of the area within the past 2000 14C yr B.P., coincident with a reduction in fire frequencies. Comparisons between modern pollen spectra and vegetation reveal that the Moraceae-dominated rain forest pollen spectra likely have a regional source area at least 2–3 km beyond the lake shore, whereas the grass- and sedge-dominated savanna pollen spectra likely have a predominantly local source area. The Holocene vegetation changes are consistent with independent paleoprecipitation records from the Bolivian Altiplano and paleovegetation records from other parts of southwestern Amazonia. The progressive expansion in rain forests through the Holocene can be largely attributed to enhanced convective activity over Amazonia, due to greater seasonality of insolation in the Southern Hemisphere tropics driven by the precession cycle according to the Milankovitch Astronomical Theory.","author":[{"dropping-particle":"","family":"Burbridge","given":"Rachel E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"Francis E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Killeen","given":"Timothy J.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"Quaternary Research","id":"ITEM-1","issue":"2","issued":{"date-parts":[["2004","3"]]},"note":"chapling and Bella Vista records - dry forest/savanna until 2000 bp when precip increased - humid evergreen","page":"215-230","title":"Fifty-thousand-year vegetation and climate history of Noel Kempff Mercado National Park, Bolivian Amazon","type":"article-journal","volume":"61"},"uris":[""]},{"id":"ITEM-2","itemData":{"DOI":"10.1177/0959683615581204","ISBN":"0959-6836","ISSN":"0959-6836","abstract":"The nature and extent of pre-Columbian (pre-AD 1492) human impact in Amazonia is a contentious issue. The Bolivian Amazon has yielded some of the most impressive evidence for large and complex pre-Columbian societies in the Amazon basin, yet there remains relatively little data concerning the land use of these societies over time. Palaeoecology, when integrated with archaeological data, has the potential to fill these gaps in our knowledge. We present a 6000-year record of anthropogenic burning, agriculture and vegetation change, from an oxbow lake located adjacent to a pre-Columbian ring ditch in north-east Bolivia (13°15′44″S, 63°42′37″W). Human occupation around the lake site is inferred from pollen and phytoliths of maize (Zea mays L.) and macroscopic charcoal evidence of anthropogenic burning. First occupation around the lake was radiocarbon dated to ~2500 calibrated years before present (BP). The persistence of maize in the record from ~1850 BP suggests that it was an important crop grown in the ring-ditch region in pre-Columbian times, and abundant macroscopic charcoal suggests that pre-Columbian land management entailed more extensive burning of the landscape than the slash-and-burn agriculture practised around the site today. The site was occupied continuously until near-modern times, although there is evidence for a decline in agricultural intensity or change in land-use strategy, and possible population decline, from ~600–500 BP. The long and continuous occupation, which predates the establishment of rainforest in the region, suggests that pre-Columbian land use may have had a significant influence on ecosystem development at this site over the last ~2000 years.","author":[{"dropping-particle":"","family":"Carson","given":"J. F.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Watling","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Mayle","given":"F. E.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Whitney","given":"B. S.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Iriarte","given":"J.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Prumers","given":"H.","non-dropping-particle":"","parse-names":false,"suffix":""},{"dropping-particle":"","family":"Soto","given":"J. D.","non-dropping-particle":"","parse-names":false,"suffix":""}],"container-title":"The Holocene","id":"ITEM-2","issue":"8","issued":{"date-parts":[["2015","5","1"]]},"note":"From Duplicate 2 (Pre-Columbian land use in the ring-ditch region of the Bolivian Amazon - Carson, J. F.; Watling, J.; Mayle, F. E.; Whitney, B. S.; Iriarte, J.; Prumers, H.; Soto, J. D.)\n\nUsed phytoliths and pollen from oxbow lake in NE Bolivia - oxbow 6000 yrs old.\n\nNice passage about phytoliths from lakes and their source area compared to pollen - assumes local representation due to modes of transport available.","page":"1285-1300","title":"Pre-Columbian land use in the ring-ditch region of the Bolivian Amazon","type":"article-journal","volume":"25"},"uris":[""]}],"mendeley":{"formattedCitation":"(Burbridge et al., 2004; Carson et al., 2015)","plainTextFormattedCitation":"(Burbridge et al., 2004; Carson et al., 2015)","previouslyFormattedCitation":"(Burbridge et al., 2004; Carson et al., 2015)"},"properties":{"noteIndex":0},"schema":""}(Burbridge et al., 2004; Carson et al., 2015). Red dashed lines represent thresholds for floristic change within the ecosystem: lower quartile (Q1), median (Q2) and upper quartile (Q3) of Hellinger distances for the appropriate ecosystem and proxy, calculated based on modern distances between samples in vegetation study plots at Noel Kempff Mercado National Park (NKMNP) (Table 2). For Granja phytoliths, the thresholds for evergreen forest are plotted. For Granja and Chaplin pollen, the thresholds for savannah are plotted. Green dashed lines represent thresholds for ecosystem turnover: lower quartile (Q1), median (Q2) and upper quartile (Q3) of Hellinger distances for the turnover between evergreen forest and savannah, for the appropriate proxy (Table 3).Supplementary Figures S1-7S1 – Summary diagram of modern pollen assemblages from plots within Noel Kempf Mercado National Park (NKMNP). Taxa plotted are those that occur at >1 % abundance in 2 or more samples. X-axis represents percentage abundance. Plots are grouped by ecosystem class: MV, AC, CP – dry forest; LF, CH – evergreen forest; FC, FT, TO – savannah. For full descriptions and analysis of pollen assemblages see original publications (Gosling et al., 2005; Gosling et al., 2009; Jones et al., 2009; Burn et al., 2010; Jones et al., 2011).S2 – Summary diagram of modern phytolith assemblages from plots within Noel Kempf Mercado National Park (NKMNP). Taxa plotted are those that occur at >1 % abundance. X-axis represents percentage abundance.Plots are grouped by ecosystem class: MV, AC, CP – dry forest; LF, CH – evergreen forest; FC, FT, TO – savannah. For full descriptions and analysis of phytolith assemblages see original publication (Dickau et al., 2013).S3 – Table of ANOVA results for Distance~EcosystemClass. Significance codes: 0 ‘***’ 0.001 ‘**’ 0.01 ‘*’ 0.05 ‘.’ 0.1 ‘ ’ 1.DfSum SqSqF ValuePr(>F)Ecosystem Class23.9011.950552.41<2e-16***Residuals1957.2570.0372S4 – Table of Tukeys HSD test results for Distance~EcosystemClass at the 95% confidence level.Ecosystem ClassdifflwruprP adjEvergreen forest-Dry forest-0.2789007-0.3582125-0.199588850.0000000Savannah-Dry forest -0.3135789-0.3928907-0.234267060.0000000Savannah-Evergreen forest -0.0346782-0.1139900 0.044633630.5571915NOVA and Tukeys HSD of the full matrix of distances. See page e discussion to more fully discuss sedimentation rate and smplingS5 – Summary diagram of fossil phytoliths from Laguna Granja, plotted against age of sediment in cal yrBP. Age-depth model taken from original publication (Carson et al., 2015). Taxa plotted are those that occur at >1 % abundance. X-axis represents percentage abundance.S6 – Summary diagram of fossil pollen from Laguna Granja, plotted against age of sediment in cal yrBP. Age-depth model taken from original publication (Carson et al., 2015). Taxa plotted are those that occur at >1 % abundance. X-axis represents percentage abundance.S7 – Summary diagram of fossil pollen from Laguna Chaplin, plotted against age of sediment in cal yrBP. Age-depth model taken from original publication (Burbridge et al., 2004). Taxa plotted are those that occur at >1 % abundance in 2 or more samples. X-axis represents percentage abundance. ................
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