Special feature: measuring components of ecological resilience in long-term ecological datasets

Ecological resilience has become a focal concept in ecosystem management. Palaeoecological records (i.e. the sub-fossil remains preserved in sediments) are useful archives to address ecological resilience since they can be used to reconstruct long-term temporal variations in ecosystem properties. The special feature presented here includes nine new papers from members and associates of the PAGES EcoRe3 community. The papers build on previous work in palaeoecology to investigate, identify and compare components of ecosystem resilience on centennial to millennial timescales. There are four key messages that can be summarized from the findings of papers within the special feature: (i) multi-proxy studies reveal insights into the presence and mechanisms of alternative states; (ii) transitions between alternative states may not necessarily be abrupt; (iii) components of ecological resilience can be identified in long-term ecological data and (iv) the palaeoecological record can also provide insights into factors influencing the resilience of ecosystem functioning. Overall, these papers demonstrate the importance of using long-term ecological records for addressing questions related to the theoretical framework provided by ecological resilience.publishedVersio

Vegetation- memory effects and their association with vegetation resilience in global drylands

Vegetation memory describes the effect of antecedent environmental and ecological conditions on the present ecosystem state and has been proposed as an important proxy for vegetation resilience. In particular, strong vegetation memory has been identified in dryland regions, but the factors underlying the spatial patterns of vegetation memory remain unknown. We aim to map the components and drivers of vegetation memory in dryland regions using state-of-the-art climate reanalysis data and refined approaches to identify vegetation-memory characteristics across dryland regions worldwide. Using a framework which distinguishes between intrinsic and extrinsic ecological memory, we show that (i) intrinsic memory is a much stronger component than extrinsic memory in the majority of dryland regions and (ii) climate reanalysis datasets change the detection of extrinsic vegetation memory in some global dryland regions. Synthesis. Our study offers a global picture of the vegetation response to two climate variables using satellite data, information which is potentially relevant for mapping components and properties of vegetation responses worldwide. However, the large differences in the spatial patterns in intrinsic vegetation memory in our study compared to previous analyses show the overall sensitivity of this component to the initial choice of extrinsic predictor variables. As a result, we caution against using the oversimplified link between intrinsic vegetation-memory and vegetation recovery rates at large spatial scales.publishedVersio

Fossil pollen and spores as a tool for reconstructing ancient solar-ultraviolet irradiance received by plants : an assessment of prospects and challenges using proxy-system modelling

Ultraviolet-B radiation (UV-B, 280-315 nm) constitutes less than 1% of the total solar radiation that reaches the Earth's surface but has a disproportional impact on biological and ecological processes from the individual to the ecosystem level. Absorption of UV-B by ozone is also one of the primary heat sources to the stratosphere, so variations in UV-B have important relationships to the Earth's radiation budget. Yet despite its importance for understanding atmospheric and ecological processes, there is limited understanding about the changes in UV-B radiation in the geological past. This is because systematic measurements of total ozone and surface UV-B only exist since the 1970s, so biological or geochemical proxies from sediment archives are needed to reconstruct UV-B irradiance received at the Earth surface beyond the experimental record. Recent developments have shown that the quantification of UV-B-absorbing compounds in pollen and spores have the potential to provide a continuous record of the solar-ultraviolet radiation received by plants. There is increasing interest in developing this proxy in palaeoclimatic and palaeoecological research. However, differences in interpretation exist between palaeoecologists, who are beginning to apply the proxy under various geological settings, and UV-B ecologists, who question whether a causal dose-response relationship of pollen and spore chemistry to UV-B irradiance has really been established. Here, we use a proxy-system modelling approach to systematically assess components of the pollen-and spore-based UV-B-irradiance proxy to ask how these differences can be resolved. We identify key unknowns and uncertainties in making inferences about past UV-B irradiance, from the pollen sensor, the sedimentary archive, and through the laboratory and experimental procedures in order to target priority areas of future work. We argue that an interdisciplinary approach, modifying methods used by plant ecologists studying contemporary responses to solar-UV-B radiation specifically to suit the needs of palaeoecological analyses, provides a way forward in developing the most reliable reconstructions for the UV-B irradiance received by plants across a range of timescales.Peer reviewe

Sensitivity of global terrestrial ecosystems to climate variability

The identification of properties that contribute to the persistence and resilience of ecosystems despite climate change constitutes a research priority of global relevance1. Here we present a novel, empirical approach to assess the relative sensitivity of ecosystems to climate variability, one property of resilience that builds on theoretical modelling work recognizing that systems closer to critical thresholds respond more sensitively to external perturbations2. We develop a new metric, the vegetation sensitivity index, that identifies areas sensitive to climate variability over the past 14 years. The metric uses time series data derived from the moderate-resolution imaging spectroradiometer (MODIS) enhanced vegetation index3, and three climatic variables that drive vegetation productivity4 (air temperature, water availability and cloud cover). Underlying the analysis is an autoregressive modelling approach used to identify climate drivers of vegetation productivity on monthly timescales, in addition to regions with memory effects and reduced response rates to external forcing5. We find ecologically sensitive regions with amplified responses to climate variability in the Arctic tundra, parts of the boreal forest belt, the tropical rainforest, alpine regions worldwide, steppe and prairie regions of central Asia and North and South America, the Caatinga deciduous forest in eastern South America, and eastern areas of Australia. Our study provides a quantitative methodology for assessing the relative response rate of ecosystems—be they natural or with a strong anthropogenic signature—to environmental variability, which is the first step towards addressing why some regions appear to be more sensitive than others, and what impact this has on the resilience of ecosystem service provision and human well-being.acceptedVersio

Chemical variations in Quercus pollen as a tool for taxonomic identification: Implications for long-term ecological and biogeographical research

Aim Fossil pollen is an important tool for understanding biogeographical patterns in the past, but the taxonomic resolution of the fossil‐pollen record may be limited to genus or even family level. Chemical analysis of pollen grains has the potential to increase the taxonomic resolution of pollen analysis, but present‐day chemical variability is poorly understood. This study aims to investigate whether a phylogenetic signal is present in the chemical variations of Quercus L. pollen and to assess the prospects of chemical techniques for identification in biogeographical research. Location Portugal. Taxon Six taxa (five species, one subspecies) of Quercus L., Q. faginea, Q. robur, Q. robur ssp. estremadurensis, Q. coccifera, Q. rotundifolia and Q. suber belonging to three sections: Cerris, Ilex and Quercus (Denk, Grimm, Manos, Deng, & Hipp, 2017). Methods We collected pollen samples from 297 individual Quercus trees across a 4° (~450 km) latitudinal gradient and determined chemical differences using Fourier‐transform infrared spectroscopy (FTIR). We used canonical powered partial least squares regression (CPPLS) and discriminant analysis to describe within‐ and between‐species chemical variability. Results We find clear differences in the FTIR spectra from Quercus pollen at the section level (Cerris: ~98%; Ilex: ~100%; Quercus: ~97%). Successful discrimination is based on spectral signals related to lipids and sporopollenins. However, discrimination of species within individual Quercus sections is more challenging: overall, species recall is ~76% and species misidentifications within sections lie between 18% and 31% of the test set. Main Conclusions Our results demonstrate that subgenus level differentiation of Quercus pollen is possible using FTIR methods, with successful classification at the section level. This indicates that operator‐independent FTIR approaches can surpass traditional morphological techniques using light microscopy. Our results have implications both for providing new insights into past colonization pathways of Quercus, and likewise for forecasting future responses to climate change. However, before FTIR techniques can be applied more broadly across palaeoecology and biogeography, our results also highlight a number of research challenges that still need to be addressed, including developing sporopollenin‐specific taxonomic discriminators and determining a more complete understanding of the effects of environmental variation on pollen‐chemical signatures in Quercus.publishedVersio

Long-term ecological responses of a lowland dipterocarp forest to climate changes and nutrient availability

Understanding the long-term impact of projected climate change on tropical rainforests is critical given their central role in the Earth's system. Palaeoecological records can provide a valuable perspective on this problem. Here, we examine the effects of past climatic changes on the dominant forest type of Southeast Asia – lowland dipterocarp forest. We use a range of proxies extracted from a 1400-yr-old lacustrine sedimentary sequence from north-eastern Philippines to determine long-term vegetation responses of lowland dipterocarp forest, including its dominant tree group dipterocarps, to changes in precipitation, fire and nutrient availability over time. Our results show a positive relationship between dipterocarp pollen accumulation rates (PARs) and leaf wax hydrogen isotope values, which suggests a negative effect of drier conditions on dipterocarp abundance. Furthermore, we find a positive relationship between dipterocarp PARs and the proxy for phosphorus availability, which suggests phosphorus controls the productivity of these keystone trees on longer time scales. Other pollen taxa show widely varying relationships with the abiotic factors, demonstrating a high diversity of plant functional responses. Our findings provide novel insights into lowland dipterocarp forest responses to changing climatic conditions in the past and highlight potential impacts of future climate change on this globally important ecosystem.publishedVersio

Abrupt change in tropical Pacific climate mean state during the Little Ice Age

The mean state of the tropical Pacific ocean-atmosphere climate, in particular its east-west asymmetry, has profound consequences for regional climates and for the El Niño/Southern Oscillation variability. Here we present a new high-resolution paleohydrological record using the stable-hydrogen-isotopic composition of terrestrial-lipid biomarkers (δDwax) from a 1,400-year-old lake sedimentary sequence from northern Philippines. Results show a dramatic and abrupt increase in δDwax values around 1630 AD with sustained high values until around 1900 AD. We interpret this change as a shift to significantly drier conditions in the western tropical Pacific during the second half of the Little Ice Age as a result of a change in tropical Pacific mean state tied to zonal sea surface temperature (SST) gradients. Our findings highlight the prominent role of abrupt shifts in zonal SST gradients on multidecadal to multicentennial timescales in shaping the tropical Pacific hydrology of the last millennium, and demonstrate that a marked transition in the tropical Pacific mean state can occur within a period of a few decades.publishedVersio

Time to better integrate paleoecological research infrastructures with neoecology to improve understanding of biodiversity long-term dynamics and to inform future conservation

Anthropogenic pressures are causing a global decline in biodiversity. Successful attempts at biodiversity conservation requires an understanding of biodiversity patterns as well as the drivers and processes that determine those patterns. To deepen this knowledge, neoecologists have focused on studying present-day or recent historical data, while paleoecologists usually study long-term data through the composition of various biological proxies and environmental indicators. By establishing standard protocols or gathering databases, research infrastructures (RIs) have been instrumental to foster exchange and collaboration among scientists within neoecology (e.g. Global Information Biodiversity Facility or National Ecological Observatory Network) and paleoecology (e.g. Paleobiology Database, Neotoma Paleoecology Database or European Pollen Database). However, these two subdisciplines (and their RIs) have traditionally remained segregated although both provide valuable information that combined can improve our understanding of biodiversity drivers and underlying processes, as well as our predictions of biodiversity responses in the future. For instance, integrative studies between paleo- and neoecology have addressed the global challenge of biodiversity loss by validating climate and ecological models, estimating species fundamental niches, understanding ecological changes and trajectories, or establishing baseline conditions for restoration. Supporting and contributing to research infrastructures from both paleo- and neoecology, as well as their further integration, could boost the amount and improve the quality of such integrative studies. We argue this will enable improved capabilities to anticipate the impacts of global change and biodiversity losses. To boost such integration and illustrate our arguments, we (1) review studies integrating paleo- and neoecology to advance in the light of global changes challenge, (2) describe RIs developed in paleoecology, and (3) discuss opportunities for further integration of RIs from both disciplines (i.e. paleo- and neoecology).publishedVersio

Long-term trends in diatom diversity and palaeoproductivity: a 16 000-year multidecadal record from Lake Baikal, southern Siberia

Biological diversity is inextricably linked to community stability and ecosystem functioning, but our understanding of these relationships in freshwater ecosystems is largely based on short-term observational, experimental, and modelling approaches. Using a multidecadal diatom record for the past ca. 16 000 years from Lake Baikal, we investigate how diversity and palaeoproductivity have responded to climate change during periods of both rapid climate fluctuation and relative climate stability. We show dynamic changes in diatom communities during the past 16 000 years, with decadal shifts in species dominance punctuating millennial-scale seasonal trends. We describe for the first time in Lake Baikal a gradual shift from spring to autumnal diatom communities that started during the Younger Dryas and peaked during the Late Holocene, which likely represents orbitally driven ecosystem responses to long-term changes in seasonality. Using a multivariate classification tree, we show that trends in planktonic and tychoplanktonic diatoms broadly reflect both long-term climatic changes associated with the demise of Northern Hemisphere ice sheets and abrupt climatic changes associated with, for example, the Younger Dryas stadial. Indeed, diatom communities are most different before and after the boundary between the Early and Middle Holocene periods of ca. 8.2 cal kyr BP, associated with the presence and demise of Northern Hemisphere ice sheets respectively. Diatom richness and diversity, estimated using Hill's species numbers, are also shown to be very responsive to periods characterized by abrupt climate change, and using knowledge of diatom autecologies in Lake Baikal, diversity trends are interpreted in terms of resource availability. Using diatom biovolume accumulation rates (BVARs; µm3 cm−2 yr−1), we show that spring diatom crops dominate palaeoproductivity for nearly all of our record, apart from a short period during the Late Holocene, when autumnal productivity dominated between 1.8–1.4 cal kyr BP. Palaeoproductivity was especially unstable during the Younger Dryas, reaching peak rates of 18.3 × 103 µm3 cm−2 yr−1 at ca. 12.3 cal kyr BP. Generalized additive models (GAMs), which explore productivity–diversity relationships (PDRs) during pre-defined climate periods, reveal complex relationships. The strongest statistical evidence for GAMs were found during the Younger Dryas, the Early Holocene, and the Late Holocene, i.e. periods of rapid climate change. We account for these differences in terms of climate-mediated resource availability, and the ability of endemic diatom species in Lake Baikal to adapt to extreme forms of living in this unique ecosystem. Our analyses offer insight into how productivity–diversity relationships may develop in the future under a warming climate