Rate-of-change analysis in palaeoecology revisited: a new approach

Dynamics in the rate of compositional change (rate-of-change; RoC), preserved in paleoecological sequences, are thought to reflect changes due to exogenous (climate and human forcing) or endogenous (local dynamics and biotic interactions) drivers. However, changes in sedimentation rates and sampling strategies can result in an uneven distribution of time intervals and are known to affect RoC estimates. Furthermore, there has been relatively little exploration of the implications of these challenges in quantifying RoC in paleoecology. Here, we introduce R-Ratepol – an easy-to-use R package – that provides a robust numerical technique for detecting and summarizing RoC patterns in complex multivariate time-ordered stratigraphical sequences. First, we compare the performance of common methods of estimating RoC and detecting periods of high RoC (peak-point) using simulated pollen-stratigraphical data with known patterns of compositional change and temporal resolution. In addition, we propose a new method of binning with a moving window, which shows a more than 5-fold increase in the correct detection of peak-points compared to the more traditional way of using individual levels. Next, we apply our new methodology to four representative European pollen sequences and show that our approach also performs well in detecting periods of significant compositional change during known onsets of human activity, early land-use transformation, and changes in fire frequency. Expanding the approach using R-Ratepol to open-access paleoecological datasets in global databases, such as Neotoma, will allow future paleoecological and macroecological studies to quantify major changes in biotic composition or in sets of abiotic variables across broad spatiotemporal scales.publishedVersio

A guide to the processing and standardization of global palaeoecological data for large-scale syntheses using fossil pollen

Aim: Palaeoecological data are crucial for comprehending large-scale biodiversity patterns and the natural and anthropogenic drivers that influence them over time. Over the last decade, the availability of open-access research databases of palaeoecological proxies has substantially increased. These databases open the door to research questions needing advanced numerical analyses and modelling based on big-data compilations. However, compiling and analysing palaeoecological data pose unique challenges that require a guide for producing standardized and reproducible compilations. Innovation: We present a step-by-step guide of how to process fossil pollen data into a standardized dataset compilation ready for macroecological and palaeoecological analyses. We describe successive criteria that will enhance the quality of the compilations. Though these criteria are project and research question-dependent, we discuss the most important assumptions that should be considered and adjusted accordingly. Our guide is accompanied by an R-workflow—called FOSSILPOL—and corresponding R-package—called R-Fossilpol—that provide a detailed protocol ready for interdisciplinary users. We illustrate the workflow by sourcing and processing Scandinavian fossil pollen datasets and show the reproducibility of continental-scale data processing. Main Conclusions: The study of biodiversity and macroecological patterns through time and space requires large-scale syntheses of palaeoecological datasets. The data preparation for such syntheses must be transparent and reproducible. With our FOSSILPOL workflow and R-package, we provide a protocol for optimal handling of large compilations of fossil pollen datasets and workflow reproducibility. Our workflow is also relevant for the compilation and synthesis of other palaeoecological proxies and as such offers a guide for synthetic and cross-disciplinary analyses with macroecological, biogeographical and palaeoecological perspectives. However, we emphasize that expertise and informed decisions based on palaeoecological knowledge remain crucial for high-quality data syntheses and should be strongly embedded in studies that rely on the increasing amount of open-access palaeoecological data.publishedVersio

Exploring spatio-temporal patterns of palynological changes in Asia during the Holocene

Historical legacies influence present-day ecosystem composition and dynamics. It is therefore important to understand the long-term dynamics of ecosystems and their properties. Analysis of ecosystem properties during the Holocene using fossil pollen assemblages provides valuable insights into past ecosystem dynamics by summarising so-called pollen-assemblage properties (PAPs). Using 205 fossil pollen data-sets (records), we quantify eight PAPs [pollen-taxonomic richness, diversity, evenness, pollen-compositional turnover, pollen-compositional change, and rate of pollen-compositional change (RoC)] for the Asian continent at different spatial scales (in individual records, within and across climate-zones, and within the continent) and time (temporal patterns over the past 12,000 years). Regression tree (RT) partitioning of the PAP-estimates using sample-age as a sole predictor revealed the “change-point(s)” (time or sample-age of major change in a PAP). We estimated the density of RT and multivariate regression tree (MRT) change-points in 1,000-year time bins during the Holocene. Pollen-compositional turnover (range of sample scores along the first DCCA axis) and change (number of MRT partitions) in each record reveal gradual spatial variation across latitude and a decline with longitude eastward. Temporally, compositional turnover declines linearly throughout the Holocene at all spatial scales. Other PAPs are heterogeneous across and within spatial scales, being more detectable at coarser scales. RT and MRT change-point density is broadly consistent in climate-zones and the continent, increasing from the early- to mid-Holocene, and mostly decrease from the mid-Holocene to the present for all PAPs. The heterogenous patterns in PAPs across the scales of study most likely reflect responses to variations in regional environmental conditions, anthropogenic land-use, and their interactions over space and time. Patterns at the climate-zone and continental scales indicate a gradual but congruent decline in major PAPs such as compositional turnover, rate of compositional change, and major temporal compositional changes (MRT) during the Holocene, especially during recent millennia, suggesting that vegetation in Asia has become progressively more homogenous. Data properties (e.g., spatial distribution of the records, distribution of samples within the records, and data-standardisation and analytical approaches) may also have partly influenced the results. It is critically important to evaluate the data properties and the approaches to data standardisation and summarisation.publishedVersio

Approaches to pollen taxonomic harmonisation in Quaternary palynology

Pollen taxonomic harmonisation involves the standardisation of the nomenclature of pollen and fern spores with similar morphotypes at the determination level that is common to all grains or spores with that morphotype within the pollen sequence(s) of interest. Such harmonisation is required prior to subsequent investigations such as numerical analysis, comparing, mapping, synthesis, and environmental reconstruction involving several pollen sequences. Here we present two approaches to harmonisation. These are a ‘top-down’ and a ‘bottom-up’ approach. The bottom-up approach is preferred. It is based on the concept of the regional pollen flora for the sequence(s) in the spatial area(s) of study. We present bottom-up harmonisation tables for the continental or sub-continental scales developed for the Humans on Planet Earth (HOPE) project. The tables are for North America, Latin America, Europe, Asia (three parts), and Indo-Pacific. These harmonisations are project-specific and sequence-specific, relating to the geographical area and to the sequences in the area under consideration, both of which are linked to the research questions being addressed. A new bottom-up harmonisation with a consistent taxonomic level and nomenclature is needed when additional sequences or areas are added. However, the HOPE tables can serve as a starting point for further research involving multi-sequence analyses or syntheses.publishedVersio

Rapid assessment of the three-dimensional distribution of dominant arboreal ants in tropical forests

Abstract Ants are omnipresent in tropical forests, especially territorially dominant arboreal ants whose territories are spatially segregated forming ‘ant mosaics’. These ecologically important species are rarely used in conservation monitoring because of the difficulty in collecting them. We developed a standardised baitline protocol to study the distribution of dominant ants on canopy trees and also a procedure to objectively define species dominance, even in unknown ant assemblages. Besides eliminating the need to climb trees, this protocol allows live arboreal ant specimens to be sampled at different heights. Behavioural aggressiveness assays between the collected workers provide data on the three-dimensional distribution of colonies and on interactions between species. We compared the results of the behavioural tests to those from null models. In the New Guinean lowland forest studied, we show that the canopy was either shared by multiple territorial species or inhabited by a single species with a large territory. The baitline protocol collected up to half of the arboreal ant species found in a felling census. However, the proportion of species collected at baits decreased with the increasing spatial dominance of single territorial species. Behavioural observations used in the protocol allowed a more efficient detection of ant mosaics than null models. Territorially dominant ants were active on both understorey and canopy trees. The protocol is fast and easy to replicate. It is a potential tool for understanding and monitoring the spatiotemporal dynamics of arboreal ant assemblages and can detect populous colonies, including those of invasive species

Inter-specific aggression generates ant mosaics in canopies of primary tropical rainforest

The ant mosaic is a concept of the non-random spatial distribution of individual ant species in trees built upon the assumption of interspecific behavioural associations. However, colony identity and environmental variance may also play a role in species distribution. Here we assess the presence of ant mosaics in a primary forest ecosystem and whether they are structured by species' aggressive behaviours or by habitat filtering. We sampled arboreal ants from vertically stratified baits exposed in 225 canopy trees in a 9-ha plot of primary lowland forest in Papua New Guinea, the largest forest area surveyed to detect ant mosaics. We performed behavioural tests on conspecific ants from adjacent trees to determine the territories of individual colonies. We explored the environmental effects on the ant communities using information on the plot vegetation structure and topography. Furthermore, we created a novel statistical method to test for the community non-random spatial structure across the plot via spatial randomisation of individual colony territories. Finally, we linked spatial segregation among the four most common species to experimentally assessed rates of interspecies aggression. The ant communities comprised 57 species of highly variable abundance and vertical stratification. Ant community composition was spatially dependent, but it was not affected by tree species composition or canopy connectivity. Only local elevation had a significant but rather small effect. Individual colony territories ranged from one tree to 0.7 ha. Species were significantly over-dispersed, with their territory overlap significantly reduced. The level of aggression between pairs of the four most common species was positively correlated with their spatial segregation. Our study demonstrates the presence of ant mosaics in tropical pristine forest, which are maintained by interspecific aggression rather than habitat filtering, with vegetation structure having a rather small and indirect effect, probably linked to microclimate variability

Trait-based approaches as ecological time machines : developing tools for reconstructing long-term variation in ecosystems

Research over the past decade has shown that quantifying spatial variation in ecosystem properties is an effective approach to investigating the effects of environmental change on ecosystems. Yet, current consensus among scientists is that we need a better understanding of short‐ and long‐term (temporal) variation in ecosystem properties to plan effective ecosystem management and predict future ecologies. Trait‐based approaches can be used to reconstruct ecosystem properties from long‐term ecological records and contribute significantly to developing understandings of ecosystem change over decadal to millennial time‐scales. Here, we synthesise current trait‐based approaches and explore how organisms' functional traits (FTs) can be scaled across time and space. We propose a framework for reconstructing long‐term variation in ecosystems by means of analysing FTs derived from palaeoecological datasets. We then summarise challenges that must be overcome to reconcile trait‐based approaches with palaeo‐datasets. Finally, we discuss the benefits and limitations of trait‐based reconstructions of ecosystem temporal dynamics and suggest future directions for research. Reconstructing environmental properties through time vis‐à‐vis FTs can be separated into two parts. The first is to record trait data for organisms present in modern ecosystems, and the second is to reconstruct temporal variability in FTs from palaeoecological datasets, capturing changes in trait composition over time. Translating palaeoecological datasets into FTs is challenging due to taphonomic, taxonomic and chronological uncertainties, as well as uniformitarian assumptions. Explicitly identifying and addressing these challenges is important to effectively calculate changes in FT through time. Palaeo‐trait research offers insights into questions related to short‐ and long‐term ecosystem functioning, environmental change and extinction and community assembly rules across time. As work in this area matures, we expect that trait‐based approaches integrating palaeoecology and neo‐ecology will improve understanding of past ecologies and provide a deeper insight of their implications for present‐day and future ecosystem management and conservation. Read the free Plain Language Summary for this article on the Journal blog

Rate-of-change analysis in palaeoecology revisited: a new approach

Dynamics in the rate of compositional change (rate-of-change; RoC), preserved in paleoecological sequences, are thought to reflect changes due to exogenous (climate and human forcing) or endogenous (local dynamics and biotic interactions) drivers. However, changes in sedimentation rates and sampling strategies can result in an uneven distribution of time intervals and are known to affect RoC estimates. Furthermore, there has been relatively little exploration of the implications of these challenges in quantifying RoC in paleoecology. Here, we introduce R-Ratepol – an easy-to-use R package – that provides a robust numerical technique for detecting and summarizing RoC patterns in complex multivariate time-ordered stratigraphical sequences. First, we compare the performance of common methods of estimating RoC and detecting periods of high RoC (peak-point) using simulated pollen-stratigraphical data with known patterns of compositional change and temporal resolution. In addition, we propose a new method of binning with a moving window, which shows a more than 5-fold increase in the correct detection of peak-points compared to the more traditional way of using individual levels. Next, we apply our new methodology to four representative European pollen sequences and show that our approach also performs well in detecting periods of significant compositional change during known onsets of human activity, early land-use transformation, and changes in fire frequency. Expanding the approach using R-Ratepol to open-access paleoecological datasets in global databases, such as Neotoma, will allow future paleoecological and macroecological studies to quantify major changes in biotic composition or in sets of abiotic variables across broad spatiotemporal scales