Pollen-based differentiation of Amazonian rainforest communities and implications for lowland palaeoecology in tropical South America

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.

A review of Holocene rainforest ecotonal dynamics at opposite ends of the Amazon - Bolivia versus Colombia

In this review paper, the aim is to compare and contrast fossil pollen evidence for Holocene rainforest ecotonal dynamics at opposite ends of the Amazon basin – the southern ecotone in NE lowland Bolivia versus the northern ecotone in lowland Colombia. During the Holocene, tropical South America experienced major changes in precipitation (Silva Dias et al. 2009). Consideration of Amazonian rainforest dynamics over this time-frame may therefore provide important insights into rainforest responsiveness to climate change.

Palynological differentiation between genera of the Moraceae family and implications for Amazonian palaeoecology

The Moraceae family is one of the most abundant and ecologically important families in Neotropical rainforests and is very well-represented in Amazonian fossil pollen records. However, difficulty in differentiating palynologically between the genera within this family, or between the Moraceae and Urticaceae families, has limited the amount of palaeoecological information that can be extracted from these records. The aim of this paper is to analyse the morphological properties of pollen from Amazonian species of Moraceae in order to determine whether the pollen taxonomy of this family can be improved. Descriptive and morphometric methods are used to identify and differentiate key pollen types of the Moraceae (mulberry) and Urticaceae (nettle) families which are represented in Amazonian rainforest communities of Noel Kempff Mercado National Park (NKMNP), Northeast Bolivia. We demonstrate that Helicostylis, Brosimum, Pseudolmedia, Sorocea and Pourouma pollen can be identified in tropical pollen assemblages and present digital images of, and a taxonomic key to, the Moraceae pollen types of NKMNP. Indicator species, Maquira coriacea (riparian evergreen forest) and Brosimum gaudichaudii (open woodland and upland savanna communities), also exhibit unique pollen morphologies. The ability to recognise these ecologically important taxa in pollen records provides the potential for much more detailed and reliable Neotropical palaeovegetation reconstructions than have hitherto been possible. In particular, this improved taxonomic resolution holds promise for resolving long-standing controversies over the interpretation of key Amazonian Quaternary pollen records.

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.

Classification of seed storage behaviour of 67 Amazonian tree species

Information was collated on the seed storage behaviour of 67 tree species native to the Amazon rainforest of Brazil; 38 appeared to show orthodox, 23 recalcitrant and six intermediate seed storage behaviour. A double-criteria key based on thousand-seed weight and seed moisture content at shedding to estimate likely seed storage behaviour, developed previously, showed good agreement with the above classifications. The key can aid seed storage behaviour identification considerably.

NOx and O3 above a tropical rainforest: an analysis with a global and box model

A cross-platform field campaign, OP3, was conducted in the state of Sabah in Malaysian Borneo between April and July of 2008. Among the suite of observations recorded, the campaign included measurements of NOx and O3 – crucial outputs of any model chemistry mechanism. We describe the measurements of these species made from both the ground site and aircraft. We then use the output from two resolutions of the chemistry transport model p-TOMCAT to illustrate the ability of a global model chemical mechanism to capture the chemistry at the rainforest site. The basic model performance is good for NOx and poor for ozone. A box model containing the same chemical mechanism is used to explore the results of the global model in more depth and make comparisons between the two. Without some parameterization of the nighttime boundary layer – free troposphere mixing (i.e. the use of a dilution parameter), the box model does not reproduce the observations, pointing to the importance of adequately representing physical processes for comparisons with surface measurements. We conclude with a discussion of box model budget calculations of chemical reaction fluxes, deposition and mixing, and compare these results to output from p-TOMCAT. These show the same chemical mechanism behaves similarly in both models, but that emissions and advection play particularly strong roles in influencing the comparison to surface measurements.

Fire-free land use in pre-1492 Amazonian savannas

The nature and scale of pre-Columbian land use and the consequences of the 1492 “Columbian Encounter” (CE) on Amazonia are among the more debated topics in New World archaeology and paleoecology. However, pre-Columbian human impact in Amazonian savannas remains poorly understood. Most paleoecological studies have been conducted in neotropical forest contexts. Of studies done in Amazonian savannas, none has the temporal resolution needed to detect changes induced by either climate or humans before and after A.D. 1492, and only a few closely integrate paleoecological and archaeological data. We report a high-resolution 2,150-y paleoecological record from a French Guianan coastal savanna that forces reconsideration of how pre-Columbian savanna peoples practiced raised-field agriculture and how the CE impacted these societies and environments. Our combined pollen, phytolith, and charcoal analyses reveal unexpectedly low levels of biomass burning associated with pre-A.D. 1492 savanna raised-field agriculture and a sharp increase in fires following the arrival of Europeans. We show that pre-Columbian raised-field farmers limited burning to improve agricultural production, contrasting with extensive use of fire in pre-Columbian tropical forest and Central American savanna environments, as well as in present-day savannas. The charcoal record indicates that extensive fires in the seasonally flooded savannas of French Guiana are a post-Columbian phenomenon, postdating the collapse of indigenous populations. The discovery that pre-Columbian farmers practiced fire-free savanna management calls into question the widely held assumption that pre-Columbian Amazonian farmers pervasively used fire to manage and alter ecosystems and offers fresh perspectives on an emerging alternative approach to savanna land use and conservation that can help reduce carbon emissions.

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.

Quantitative estimates of glacial and Holocene temperature and precipitation change in lowland Amazonian Bolivia

Quantitative estimates of temperature and precipitation change during the late Pleistocene and Holocene have been difficult to obtain for much of the lowland Neotropics. Using two published lacustrine pollen records and a climate-vegetation model based on the modern abundance distributions of 154 Neotropical plant families, we demonstrate how family-level counts of fossil pollen can be used to quantitatively reconstruct tropical paleoclimate and provide needed information on historic patterns of climatic change. With this family-level analysis, we show that one area of the lowland tropics, northeastern Bolivia, experienced cooling (1–3 °C) and drying (400 mm/yr), relative to present, during the late Pleistocene (50,000–12,000 calendar years before present [cal. yr B.P.]). Immediately prior to the Last Glacial Maximum (LGM, ca. 21,000 cal. yr B.P.), we observe a distinct transition from cooler temperatures and variable precipitation to a period of warmer temperatures and relative dryness that extends to the middle Holocene (5000–3000 cal. yr B.P.). This prolonged reduction in precipitation occurs against the backdrop of increasing atmospheric CO2 concentrations, indicating that the presence of mixed savanna and dry-forest communities in northeastern Bolivia durng the LGM was not solely the result of low CO2 levels, as suggested previously, but also lower precipitation. The results of our analysis demonstrate the potential for using the distribution and abundance structure of modern Neotropical plant families to infer paleoclimate from the fossil pollen record.

Environmental impact of geometric earthwork construction in pre-Columbian Amazonia

There is considerable controversy over whether pre-Columbian (pre-A.D. 1492) Amazonia was largely “pristine” and sparsely populated by slash-and-burn agriculturists, or instead a densely populated, domesticated landscape, heavily altered by extensive deforestation and anthropogenic burning. The discovery of hundreds of large geometric earthworks beneath intact rainforest across southern Amazonia challenges its status as a pristine landscape, and has been assumed to indicate extensive pre-Columbian deforestation by large populations. We tested these assumptions using coupled local- and regional-scale paleoecological records to reconstruct land use on an earthwork site in northeast Bolivia within the context of regional, climate-driven biome changes. This approach revealed evidence for an alternative scenario of Amazonian land use, which did not necessitate labor-intensive rainforest clearance for earthwork construction. Instead, we show that the inhabitants exploited a naturally open savanna landscape that they maintained around their settlement despite the climatically driven rainforest expansion that began ∼2,000 y ago across the region. Earthwork construction and agriculture on terra firme landscapes currently occupied by the seasonal rainforests of southern Amazonia may therefore not have necessitated large-scale deforestation using stone tools. This finding implies far less labor—and potentially lower population density—than previously supposed. Our findings demonstrate that current debates over the magnitude and nature of pre-Columbian Amazonian land use, and its impact on global biogeochemical cycling, are potentially flawed because they do not consider this land use in the context of climate-driven forest–savanna biome shifts through the mid-to-late Holocene.

Differentiation of neotropical ecosystems by modern soil phytolith assemblages and its implications for palaeoenvironmental and archaeological reconstructions II: Southwestern Amazonian forests

Accurate archaeological and palaeoenvironmental reconstructions using phytoliths relies on the study of modern reference material. In eastern Acre, Brazil, we examined whether the five most common forest types present today were 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 forest with 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.

Temporal patterns of nutrient availability around nests of leaf-cutting ants (Atta colombica) in secondary moist tropical forest

Leaf-cutting ants consume up to 10% of canopy leaves in the foraging area of their colony and therefore represent a key perturbation in the nutrient cycle of tropical forests. We used a chronosequence of nest sites on Barro, Colorado Island, Panama, to assess the influence of leaf-cutting ants (Atta colombica) on nutrient availability in a neotropical rainforest. Twelve nest sites were sampled, including active nests, recently abandoned nests (<1 year) and long-abandoned nests (>1 year). Waste material discarded by the ants down-slope from the nests contained large concentrations of nitrogen and phosphorus in both total and soluble forms, but decomposed within one year after the nests were abandoned. Despite this, soil under the waste material contained high concentrations of nitrate and ammonium that persisted after the disappearance of the waste, although soluble phosphate returned to background concentrations within one year of nest abandonment. Fine roots were more abundant in soil under waste than control soils up to one year after nest abandonment, but were not significantly different for older sites. In contrast to the waste dumps, soil above the underground nest chambers consistently contained lower nutrient concentrations than control soils, although this was not statistically significant. We conclude that the 'islands of fertility' created by leaf-cutting ants provide a nutritional benefit to nearby plants for less than one year after nest abandonment in the moist tropical environment of Barro Colorado Island. Published by Elsevier Ltd.