Differentiation of neotropical ecosystems by modern soil phytolith assemblages and its implications for palaeoenvironmental and archaeological reconstructions

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.

Differentiation between Neotropical rainforest, dry forest and savannah ecosystems by their modern pollen spectra and implications for the fossil pollen record

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 (cerradão) 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.

Impact of a drier Early-Mid-Holocene climate upon Amazonian forests

This paper uses a palaeoecological approach to examine the impact of drier climatic conditions of the Early-Mid-Holocene (ca 8000-4000 years ago) upon Amazonia's forests and their fire regimes. Palaeovegetation (pollen data) and palaeofire (charcoal) records are synthesized from 20 sites within the present tropical forest biome, and the underlying causes of any emergent patterns or changes are explored by reference to independent palaeoclimate data and present-day patterns of precipitation, forest cover and fire activity across Amazonia. During the Early-Mid-Holocene, Andean cloud forest taxa were replaced by lowland tree taxa as the cloud base rose while lowland ecotonal areas, which are presently covered by evergreen rainforest, were instead dominated by savannahs and/or semi-deciduous dry forests. Elsewhere in the Amazon Basin there is considerable spatial and temporal variation in patterns of vegetation disturbance and fire, which probably reflects the complex heterogeneous patterns in precipitation and seasonality across the basin, and the interactions between climate change, drought- and fire susceptibility of the forests, and Palaeo-Indian land use. Our analysis shows that the forest biome in most parts of Amazonia appears to have been remarkably resilient to climatic conditions significantly drier than those of today, despite widespread evidence of forest burning. Only in ecotonal areas is there evidence of biome replacement in the Holocene. From this palaeoecological perspective, we argue against the Amazon forest 'dieback' scenario simulated for the future.

The surface urban energy and water balance scheme (SUEWS): evaluation in Los Angeles and Vancouver

An urban energy and water balance model is presented which uses a small number of commonly measured meteorological variables and information about the surface cover. Rates of evaporation-interception for a single layer with multiple surface types (paved, buildings, coniferous trees and/or shrubs, deciduous trees and/or shrubs, irrigated grass, non-irrigated grass and water) are calculated. Below each surface type, except water, there is a single soil layer. At each time step the moisture state of each surface is calculated. Horizontal water movements at the surface and in the soil are incorporated. Particular attention is given to the surface conductance used to model evaporation and its parameters. The model is tested against direct flux measurements carried out over a number of years in Vancouver, Canada and Los Angeles, USA. At all measurement sites the model is able to simulate the net all-wave radiation and turbulent sensible and latent heat well (RMSE = 25–47 W m−2, 30–64 and 20–56 W m−2, respectively). The model reproduces the diurnal cycle of the turbulent fluxes but typically underestimates latent heat flux and overestimates sensible heat flux in the day time. The model tracks measured surface wetness and simulates the variations in soil moisture content. It is able to respond correctly to short-term events as well as annual changes. The largest uncertainty relates to the determination of surface conductance. The model has the potential be used for multiple applications; for example, to predict effects of regulation on urban water use, landscaping and planning scenarios, or to assess climate mitigation strategies.

New age estimates for the Palaeolithic assemblages and Pleistocene succession of Casablanca, Morocco

Marine and aeolian Quaternary sediments from Casablanca, Morocco were dated using the optically stimulated luminescence (OSL) signal of quartz grains. These sediments form part of an extensive succession spanning the Pleistocene, and contain a rich faunal and archaeological record, including an Acheulian lithic assemblage from before the Brunhes–Matayama boundary, and a Homo erectus jaw from younger cave deposits. Sediment samples from the sites of Reddad Ben Ali, Oulad J’mel, Sidi Abderhamane and Thomas Quarries have been dated, in order to assess the upper limits of OSL. The revision of previously measured mammalian tooth enamel electron spin resonance (ESR) dates from the Grotte des Rhinocéros, Oulad Hamida Quarry 1, incorporating updated environmental dose rate measurements and attenuation calculations, also provide chronological constraint for the archaeological material preserved at Thomas Quarries. Several OSL age estimates extend back to around 500,000 years, with a single sample providing an OSL age close to 1 Ma in magnetically reversed sediments. These luminescence dates are some of the oldest determined, and their reliability is assessed using both internal criteria based on stratigraphic consistency, and external lithostratigraphic, morphostratigraphic and independent chronological constraints. For most samples, good internal agreement is observed using single aliquot regenerative-dose OSL measurements, while multiple aliquot additive-dose measurements generally have poorer resolution and consistency. Novel slow-component and component-resolved OSL approaches applied to four samples provide significantly enhanced dating precision, and an examination of the degree of signal zeroing at deposition. A comparison of the OSL age estimates with the updated ESR dates and one U-series date demonstrate that this method has great potential for providing reliable age estimates for sediments of this antiquity. We consider the cause of some slight age inversion observed at Thomas Quarries, and provide recommendations for further luminescence dating within this succession.

Palaeovegetation in China during the late Quaternary: Biome reconstructions based on a global scheme of plant functional types

Two previous reconstructions of palaeovegetation across the whole of China were performed using a simple classification of plant functional types (PFTs). Now a more explicit, global PFT classification scheme has been developed, and a substantial number of additional pollen records have become available. Here we apply the global scheme of PFTs to a comprehensive set of pollen records available from China to test the applicability of the global scheme of PFTs in China, and to obtain a well-founded reconstruction of changing palaeovegetation patterns. A total of 806 pollen surface samples, 188 mid-Holocene (MH, 6000 14C yr BP) and 50 last glacial maximum (LGM, 18,000 14C yr BP) pollen records were used to reconstruct vegetation patterns in China, based on a new global classification system of PFTs and a standard numerical technique for biome assignment (biomization). The biome reconstruction based on pollen surface samples showed convincing agreement with present potential natural vegetation. Coherent patterns of change in biome distribution between MH, LGM and present are observed. In the MH, cold and cool-temperate evergreen needleleaf forests and mixed forests, temperate deciduous broadleaf forest, and warm-temperate evergreen broadleaf and mixed forest in eastern China were shifted northward by 200–500 km. Cold-deciduous forest in northeastern China was replaced by cold evergreen needleleaf forest while in central northern China, cold-deciduous forest was present at some sites now occupied by temperate grassland and desert. The forest–grassland boundary was 200–300 km west of its present position. Temperate xerophytic shrubland, temperate grassland and desert covered a large area on the Tibetan Plateau, but the area of tundra was reduced. Treeline was 300–500 m higher than present in Tibet. These changes imply generally warmer winters, longer growing seasons and more precipitation during the MH. Westward shifts of the forest–shrubland–grassland and grassland–desert boundaries imply greater moisture availability in the MH, consistent with a stronger summer monsoon. During the LGM, in contrast, cold-deciduous forest, cool-temperate evergreen needleleaf forest, cool mixed forests, warm-temperate evergreen broadleaf and mixed forest in eastern China were displaced to the south by 300–1000 km, while temperate deciduous broadleaf forest, pure warm-temperate evergreen forest, tropical semi-evergreen and evergreen broadleaf forests were restricted or absent from the mainland of southern China, implying colder winters than present. Strong shifts of temperate xerophytic shrubland, temperate grassland and desert to the south and east in northern and western China and on the Tibetan Plateau imply drier conditions than present.

Pollen-based reconstructions of biome distributions for Australia, Southeast Asia and the Pacific (SEAPAC region) at 0,6000 and 18,000 14C yr B.P.

Aim  This paper documents reconstructions of the vegetation patterns in Australia, Southeast Asia and the Pacific (SEAPAC region) in the mid-Holocene and at the last glacial maximum (LGM). Methods  Vegetation patterns were reconstructed from pollen data using an objective biomization scheme based on plant functional types. The biomization scheme was first tested using 535 modern pollen samples from 377 sites, and then applied unchanged to fossil pollen samples dating to 6000 ± 500 or 18,000 ± 1000 14C yr bp. Results  1. Tests using surface pollen sample sites showed that the biomization scheme is capable of reproducing the modern broad-scale patterns of vegetation distribution. The north–south gradient in temperature, reflected in transitions from cool evergreen needleleaf forest in the extreme south through temperate rain forest or wet sclerophyll forest (WSFW) and into tropical forests, is well reconstructed. The transitions from xerophytic through sclerophyll woodlands and open forests to closed-canopy forests, which reflect the gradient in plant available moisture from the continental interior towards the coast, are reconstructed with less geographical precision but nevertheless the broad-scale pattern emerges. 2. Differences between the modern and mid-Holocene vegetation patterns in mainland Australia are comparatively small and reflect changes in moisture availability rather than temperature. In south-eastern Australia some sites show a shift towards more moisture-stressed vegetation in the mid-Holocene with xerophytic woods/scrub and temperate sclerophyll woodland and shrubland at sites characterized today by WSFW or warm-temperate rain forest (WTRF). However, sites in the Snowy Mountains, on the Southern Tablelands and east of the Great Dividing Range have more moisture-demanding vegetation in the mid-Holocene than today. South-western Australia was slightly drier than today. The single site in north-western Australia also shows conditions drier than today in the mid-Holocene. Changes in the tropics are also comparatively small, but the presence of WTRF and tropical deciduous broadleaf forest and woodland in the mid-Holocene, in sites occupied today by cool-temperate rain forest, indicate warmer conditions. 3. Expansion of xerophytic vegetation in the south and tropical deciduous broadleaf forest and woodland in the north indicate drier conditions across mainland Australia at the LGM. None of these changes are informative about the degree of cooling. However the evidence from the tropics, showing lowering of the treeline and forest belts, indicates that conditions were between 1 and 9 °C (depending on elevation) colder. The encroachment of tropical deciduous broadleaf forest and woodland into lowland evergreen broadleaf forest implies greater aridity. Main conclusions  This study provides the first continental-scale reconstruction of mid-Holocene and LGM vegetation patterns from Australia, Southeast Asia and the Pacific (SEAPAC region) using an objective biomization scheme. These data will provide a benchmark for evaluation of palaeoclimate simulations within the framework of the Palaeoclimate Modelling Intercomparison Project.

Palaeovegetation of China: a pollen data-based synthesis for the mid-Holocene and last glacial maximum.

Pollen data from China for 6000 and 18,000 14C yr bp were compiled and used to reconstruct palaeovegetation patterns, using complete taxon lists where possible and a biomization procedure that entailed the assignment of 645 pollen taxa to plant functional types. A set of 658 modern pollen samples spanning all biomes and regions provided a comprehensive test for this procedure and showed convincing agreement between reconstructed biomes and present natural vegetation types, both geographically and in terms of the elevation gradients in mountain regions of north-eastern and south-western China. The 6000 14C yr bp map confirms earlier studies in showing that the forest biomes in eastern China were systematically shifted northwards and extended westwards during the mid-Holocene. Tropical rain forest occurred on mainland China at sites characterized today by either tropical seasonal or broadleaved evergreen/warm mixed forest. Broadleaved evergreen/warm mixed forest occurred further north than today, and at higher elevation sites within the modern latitudinal range of this biome. The northern limit of temperate deciduous forest was shifted c. 800 km north relative to today. The 18,000 14C yr bp map shows that steppe and even desert vegetation extended to the modern coast of eastern China at the last glacial maximum, replacing today’s temperate deciduous forest. Tropical forests were excluded from China and broadleaved evergreen/warm mixed forest had retreated to tropical latitudes, while taiga extended southwards to c. 43°N.

Mid-Holocene and glacial-maximum vegetation geography of the northern continents and Africa.

BIOME 6000 is an international project to map vegetation globally at mid-Holocene (6000 14C yr bp) and last glacial maximum (LGM, 18,000 14C yr bp), with a view to evaluating coupled climate-biosphere model results. Primary palaeoecological data are assigned to biomes using an explicit algorithm based on plant functional types. This paper introduces the second Special Feature on BIOME 6000. Site-based global biome maps are shown with data from North America, Eurasia (except South and Southeast Asia) and Africa at both time periods. A map based on surface samples shows the method’s skill in reconstructing present-day biomes. Cold and dry conditions at LGM favoured extensive tundra and steppe. These biomes intergraded in northern Eurasia. Northern hemisphere forest biomes were displaced southward. Boreal evergreen forests (taiga) and temperate deciduous forests were fragmented, while European and East Asian steppes were greatly extended. Tropical moist forests (i.e. tropical rain forest and tropical seasonal forest) in Africa were reduced. In south-western North America, desert and steppe were replaced by open conifer woodland, opposite to the general arid trend but consistent with modelled southward displacement of the jet stream. The Arctic forest limit was shifted slighly north at 6000 14C yr bp in some sectors, but not in all. Northern temperate forest zones were generally shifted greater distances north. Warmer winters as well as summers in several regions are required to explain these shifts. Temperate deciduous forests in Europe were greatly extended, into the Mediterranean region as well as to the north. Steppe encroached on forest biomes in interior North America, but not in central Asia. Enhanced monsoons extended forest biomes in China inland and Sahelian vegetation into the Sahara while the African tropical rain forest was also reduced, consistent with a modelled northward shift of the ITCZ and a more seasonal climate in the equatorial zone. Palaeobiome maps show the outcome of separate, independent migrations of plant taxa in response to climate change. The average composition of biomes at LGM was often markedly different from today. Refugia for the temperate deciduous and tropical rain forest biomes may have existed offshore at LGM, but their characteristic taxa also persisted as components of other biomes. Examples include temperate deciduous trees that survived in cool mixed forest in eastern Europe, and tropical evergreen trees that survived in tropical seasonal forest in Africa. The sequence of biome shifts during a glacial-interglacial cycle may help account for some disjunct distributions of plant taxa. For example, the now-arid Saharan mountains may have linked Mediterranean and African tropical montane floras during enhanced monsoon regimes. Major changes in physical land-surface conditions, shown by the palaeobiome data, have implications for the global climate. The data can be used directly to evaluate the output of coupled atmosphere-biosphere models. The data could also be objectively generalized to yield realistic gridded land-surface maps, for use in sensitivity experiments with atmospheric models. Recent analyses of vegetation-climate feedbacks have focused on the hypothesized positive feedback effects of climate-induced vegetation changes in the Sahara/Sahel region and the Arctic during the mid-Holocene. However, a far wider spectrum of interactions potentially exists and could be investigated, using these data, both for 6000 14C yr bp and for the LGM.

Present-day and mid-Holocene biomes reconstructed from pollen and plant macrofossil data from the former Soviet Union and Mongolia

Fossil pollen data supplemented by tree macrofossil records were used to reconstruct the vegetation of the Former Soviet Union and Mongolia at 6000 years. Pollen spectra were assigned to biomes using the plant-functional-type method developed by Prentice et al. (1996). Surface pollen data and a modern vegetation map provided a test of the method. This is the first time such a broad-scale vegetation reconstruction for the greater part of northern Eurasia has been attempted with objective techniques. The new results confirm previous regional palaeoenvironmental studies of the mid-Holocene while providing a comprehensive synopsis and firmer conclusions. West of the Ural Mountains temperate deciduous forest extended both northward and southward from its modern range. The northern limits of cool mixed and cool conifer forests were also further north than present. Taiga was reduced in European Russia, but was extended into Yakutia where now there is cold deciduous forest. The northern limit of taiga was extended (as shown by increased Picea pollen percentages, and by tree macrofossil records north of the present-day forest limit) but tundra was still present in north-eastern Siberia. The boundary between forest and steppe in the continental interior did not shift substantially, and dry conditions similar to present existed in western Mongolia and north of the Aral Sea.

Pollen-based biome reconstructions for China at 0 and 6000 years

Biomization provides an objective and robust method of assigning pollen spectra to biomes so that pollen data can be mapped and compared directly with the output of biomgeographic models. We have tested the applicability of this procedure, originally developed for Europe, to assign modern surface samples from China to biomes. The procedure successfully delineated the major vegetation types of China. When the same procedure was applied to fossil pollen samples for 6000 years ago, the reconstructions showed systematic differences from present, consistent with previous interpretations of vegetation changes since the mid-Holocene. In eastern China, the forest zones were systematically shifted northwards, such that cool mixed forests displaced taiga in northeastern China, while broad-leaved evergreen forest extended c. 300 km and temperate deciduous forestc. 500–600 km beyond their present northern limits. In northwestern China, the area of desert and steppe vegetation was reduced compared to present. On the Tibetan Plateau, forest vegetation extended to higher elevations than today and the area of tundra was reduced. These shifts in biome distributions imply significant changes in climate since 6000 years ago that can be interpreted qualitatively as a response to orbital forcing and its secondary effects on the Asian monsoon.

The climate and biomes of Europe at 6000 yr BP: comparison of model simulations and pollen-based reconstructions

14C-dated pollen and lake-level data from Europe are used to assess the spatial patterns of climate change between 6000 yr BP and present, as simulated by the NCAR CCM1 (National Center for Atmospheric Research, Community Climate Model, version 1) in response to the change in the Earth’s orbital parameters during this perod. First, reconstructed 6000 yr BP values of bioclimate variables obtained from pollen and lake-level data with the constrained-analogue technique are compared with simulated values. Then a 6000 yr BP biome map obtained from pollen data with an objective biome reconstruction (biomization) technique is compared with BIOME model results derived from the same simulation. Data and simulations agree in some features: warmer-than-present growing seasons in N and C Europe allowed forests to extend further north and to higher elevations than today, and warmer winters in C and E Europe prevented boreal conifers from spreading west. More generally, however, the agreement is poor. Predominantly deciduous forest types in Fennoscandia imply warmer winters than the model allows. The model fails to simulate winters cold enough, or summers wet enough, to allow temperate deciduous forests their former extended distribution in S Europe, and it incorrectly simulates a much expanded area of steppe vegetation in SE Europe. Similar errors have also been noted in numerous 6000 yr BP simulations with prescribed modern sea surface temperatures. These errors are evidently not resolved by the inclusion of interactive sea-surface conditions in the CCM1. Accurate representation of mid-Holocene climates in Europe may require the inclusion of dynamical ocean–atmosphere and/or vegetation–atmosphere interactions that most palaeoclimate model simulations have so far disregarded.

Mid-Holocene land-surface conditions in northern Africa and the Arabian peninsula: a data set for the analysis of biogeophysical feedbacks in the climate system

Large changes in the extent of northern subtropical arid regions during the Holocene are attributed to orbitally forced variations in monsoon strength and have been implicated in the regulation of atmospheric trace gas concentrations on millenial timescales. Models that omit biogeophysical feedback, however, are unable to account for the full magnitude of African monsoon amplification and extension during the early to middle Holocene (˜9500–5000 years B.P.). A data set describing land-surface conditions 6000 years B.P. on a 1° × 1° grid across northern Africa and the Arabian Peninsula has been prepared from published maps and other sources of palaeoenvironmental data, with the primary aim of providing a realistic lower boundary condition for atmospheric general circulation model experiments similar to those performed in the Palaeoclimate Modelling Intercomparison Project. The data set includes information on the percentage of each grid cell occupied by specific vegetation types (steppe, savanna, xerophytic woods/scrub, tropical deciduous forest, and tropical montane evergreen forest), open water (lakes), and wetlands, plus information on the flow direction of major drainage channels for use in large-scale palaeohydrological modeling.

Responses of leaf traits to climatic gradients: adaptive variation versus compositional shifts

Dynamic global vegetation models (DGVMs) typically rely on plant functional types (PFTs), which are assigned distinct environmental tolerances and replace one another progressively along environmental gradients. Fixed values of traits are assigned to each PFT; modelled trait variation along gradients is thus driven by PFT replacement. But empirical studies have revealed "universal" scaling relationships (quantitative trait variations with climate that are similar within and between species, PFTs and communities); and continuous, adaptive trait variation has been proposed to replace PFTs as the basis for next-generation DGVMs. Here we analyse quantitative leaf-trait variation on long temperature and moisture gradients in China with a view to understanding the relative importance of PFT replacement vs. continuous adaptive variation within PFTs. Leaf area (LA), specific leaf area (SLA), leaf dry matter content (LDMC) and nitrogen content of dry matter were measured on all species at 80 sites ranging from temperate to tropical climates and from dense forests to deserts. Chlorophyll fluorescence traits and carbon, phosphorus and potassium contents were measured at 47 sites. Generalized linear models were used to relate log-transformed trait values to growing-season temperature and moisture indices, with or without PFT identity as a predictor, and to test for differences in trait responses among PFTs. Continuous trait variation was found to be ubiquitous. Responses to moisture availability were generally similar within and between PFTs, but biophysical traits (LA, SLA and LDMC) of forbs and grasses responded differently from woody plants. SLA and LDMC responses to temperature were dominated by the prevalence of evergreen PFTs with thick, dense leaves at the warm end of the gradient. Nutrient (N, P and K) responses to climate gradients were generally similar within all PFTs. Area-based nutrients generally declined with moisture; Narea and Karea declined with temperature, but Parea increased with temperature. Although the adaptive nature of many of these trait-climate relationships is understood qualitatively, a key challenge for modelling is to predict them quantitatively. Models must take into account that community-level responses to climatic gradients can be influenced by shifts in PFT composition, such as the replacement of deciduous by evergreen trees, which may run either parallel or counter to trait variation within PFTs. The importance of PFT shifts varies among traits, being important for biophysical traits but less so for physiological and chemical traits. Finally, models should take account of the diversity of trait values that is found in all sites and PFTs, representing the "pool" of variation that is locally available for the natural adaptation of ecosystem function to environmental change.

Late Würm and Early-Middle Holocene environmental change and human activities in the Northern Apennines, Italy

An enhanced radiocarbon-dated pollen-stratigraphical record from Rovegno (Liguria, 812m asl), northern Apennines (Italy), has provided a history of vegetation succession from before 17,056-16,621 cal yrs BP to the present day. The record indicates the transition from open Pinus woodland to Artemisia dominated grassland, and finally Juniperus shrubland during the late Würm. This is succeeded by Betula and Pinus woodland, and the expansion of thermophilous taxa, namely Abies, Corylus and Quercus during the Late Würm Lateglacial Interstadial. The ‘Younger Dryas’ is possibly represented by an increase in Betula and Artemisia. During the early Holocene, mixed coniferous-deciduous woodland is dominant with Quercus, as well as Abies, Fagus and Corylus. Fagus woodland becomes established sometime before 6488-6318 cal yrs BP, but never becomes a major component of the woodland cover. Throughout the middle Holocene, Abies woodland fl uctuates, with marked declines between 6488-6318 cal yrs BP and 5287-4835 cal yrs BP, although the cause remains uncertain. Finally, the paper evaluates the application of non-pollen palynomorphs, especially coprophilous fungal spores, at Prato Spilla ‘A’ (Emilia Romagna) and concludes that greater caution must be used when interpreting middle Holocene human activity based upon pollen data alone