There is a close correspondence between regions with globally disappearing climates and previously identified biodiversity hotspots; for these regions, standard conservation solutions (e.g., assisted migration and networked reserves) may be insufficient to preserve biodiversity. biodiversity hotspots ͉ climate change ͉ dispersal limitations ͉ global-change ecology ͉ ecological surprises

Localized ecological systems are known to shift abruptly and irreversibly from one state to another when they are forced across critical thresholds. Here we review evidence that the global ecosystem as a whole can react in the same way and is approaching a planetary-scale critical transition as a result of human influence. The plausibility of a planetary-scale 'tipping point' highlights the need to improve biological forecasting by detecting early warning signs of critical transitions on global as well as local scales, and by detecting feedbacks that promote such transitions. It is also necessary to address root causes of how humans are forcing biological changes.

w ww ww w. .f fr ro on nt ti ie er rs si in ne ec co ol lo og gy y. .o or rg g H ow do you study an ecosystem no ecologist has ever seen? This is a problem for both paleoecologists and global-change ecologists, who seek to understand ecological systems for time periods outside the realm of modern observations. One group looks to the past and the other to the future, but both use our understanding of extant ecosystems and processes as a common starting point for scientific inference. This is familiar to paleoecologists as the principle of uniformitarianism (ie "the present is the key to the past"), whereby understanding modern processes aids interpretation of fossil records. Similarly, global-change ecologists apply a forward-projected form of uniformitarianism, using models based on present-day ecological patterns and processes to forecast ecological responses to future change. Thus, both paleoecology and global-change ecology are inextricably rooted in the current, and research into long-term ecological dynamics, past or future, is heavily conditioned by our current observations and personal experience.The further our explorations carry us from the present, the murkier our vision becomes. This is not just because fossil archives become sparser as we look deeper into the past, nor because the chains of future contingency become increasingly long. Rather, the further we move from the present, the more it becomes an inadequate model for past and future system behavior. The current state of the Earth system, and its constituent ecosystems, is just one of many possible states, and both past and future system states may differ fundamentally from the present. The more that environments, past or future, differ from the present, the more our understanding of ecological patterns and processes will be incomplete and the less accurately will our models predict key ecological phenomena such as species distributions, community composition, species interactions, and biogeochemicalprocess rates.Here, we focus on "no-analog" plant communities (Panel 1), their relationship to climate, and the challenges they pose to predictive ecological models. We briefly summarize a niche-based, conceptual framework explaining how no-analog communities arise (Jackson and Overpeck 2000). We discuss past no-analog communities, using the well documented late-glacial communities as a detailed case study (Jackson and Williams 2004), and argue that these communities were shaped by environmental conditions also without modern counterpart (Williams et al. 2001). We then turn to the future, identifying regions of the world at risk of developing future novel climates (Williams et al. 2007). Finally, we discuss the implications for global-change ecology, including the risk of future novel ecosystems (Hobbs et al. 2006) and the challenges posed for ecological forecasting. I In n a a n nu ut ts sh he el ll l: :• Many past ecological communities were compositionally unlike modern communities • The formation and dissolution of these past "no-analog" communit...

Subfossil pollen and plant macrofossil data derived from 14 C-dated sediment profiles can provide quantitative information on glacial and interglacial climates. The data allow climate variables related to growingseason warmth, winter cold, and plant-available moisture to be reconstructed. 123Clim Dyn (2011) 37:775-802 DOI 10.1007 surface-pollen assemblages are shown to be accurate and unbiased. Reconstructed LGM and MH climate anomaly patterns are coherent, consistent between variables, and robust with respect to the choice of technique. They support a conceptual model of the controls of Late Quaternary climate change whereby the first-order effects of orbital variations and greenhouse forcing on the seasonal cycle of temperature are predictably modified by responses of the atmospheric circulation and surface energy balance.

No abstract

This paper integrates recent efforts to map the distribution of biomes for the late Quaternary with the detailed evidence that plant species have responded individualistically to climate change at millennial timescales. Using a fossil-pollen data set of over 700 sites, we review late-Quaternary vegetation history in northern and eastern North America across levels of ecological organization from individual taxa to biomes, and apply the insights gained from this review to critically examine the biome maps generated from the pollen data. Higher-order features of the vegetation (e.g., plant associations, physiognomy) emerge from individualistic responses of plant taxa to climate change, and different representations of vegetation history reveal different aspects of vegetation dynamics. Vegetation distribution and composition were relatively stable during full-glacial times (21 000-17 000 yr BP) [calendar years] and during the mid-to late Holocene (7000-500 yr BP), but changed rapidly during the late-glacial period and early Holocene (16 000-8 000 yr BP) and after 500 yr BP. Shifts in plant taxon distributions were characterized by individualistic changes in population abundances and ranges and included large east-west shifts in distribution in addition to the northward redistribution of most taxa. Modern associations such as Fagus-Tsuga and Picea-Alnus-Betula date to the early Holocene, whereas other associations common to the late-glacial period (e.g., Picea-Cyperaceae-Fraxinus-Ostrya/ Carpinus) no longer exist. Biomes are dynamic entities that have changed in distribution, composition, and structure over time. The late-Pleistocene suite of biomes is distinct from those that grew during the Holocene. The pollen-based biome reconstructions are able to capture the major features of late-Quaternary vegetation but downplay the magnitude and variety of vegetational responses to climate change by (1) limiting apparent land-cover change to ecotones, (2) masking internal variations in biome composition, and (3) obscuring the range shifts and changes in abundance among individual taxa. The compositional and structural differences between full-glacial and recent biomes of the same type are similar to or greater than the spatial heterogeneity in the composition and structure of present-day biomes. This spatial and temporal heterogeneity allows biome maps to accommodate individualistic behavior among species but masks climatically important variations in taxonomic composition as well as structural differences between modern biomes and their ancient counterparts.

Demise of the Megafauna Approximately 10,000 years ago, the Pleistocene-Holocene deglaciation in North America produced widespread biotic and environmental change, including extinctions of megafauna, reorganization of plant communities, and increased wildfire. The causal links and sequences of these changes remain unclear. Gill et al. (p. 1100 ; see the Perspective by Johnson ) unravel these connections in an analysis of pollen, charcoal, and the dung fungus Sporormiella from the sediments of Appleman Lake, Indiana. The decline in Pleistocene megafaunal population densities (inferred from fungal spore abundances) preceded both the formation of the lateglacial plant communities and a shift to an enhanced fire regime, thus contradicting hypotheses that invoke habitat change or extraterrestrial impact to explain the megafaunal extinction. The data suggest that population collapse and functional extinction of the megafauna preceded their final extinction by several thousand years.

"Space-for-time" substitution is widely used in biodiversity modeling to infer past or future trajectories of ecological systems from contemporary spatial patterns. However, the foundational assumption-that drivers of spatial gradients of species composition also drive temporal changes in diversity-rarely is tested. Here, we empirically test the space-for-time assumption by constructing orthogonal datasets of compositional turnover of plant taxa and climatic dissimilarity through time and across space from Late Quaternary pollen records in eastern North America, then modeling climatedriven compositional turnover. Predictions relying on space-for-time substitution were ∼72% as accurate as "time-for-time" predictions. However, space-for-time substitution performed poorly during the Holocene when temporal variation in climate was small relative to spatial variation and required subsampling to match the extent of spatial and temporal climatic gradients. Despite this caution, our results generally support the judicious use of space-for-time substitution in modeling community responses to climate change.fossil pollen | global change | paleoecology | generalized dissimilarity modeling V iewed broadly, space-for-time substitution encompasses analyses in which contemporary spatial phenomena are used to understand and model temporal processes that are otherwise unobservable, most notably past and future events. Many fields have developed and debated methods relying on space-for-time substitution, such as ecological chronosequences to study longterm nutrient cycling and plant succession (1-3) and transfer functions for inferring past environmental changes from geological proxies (4, 5). The assumption of space-for-time substitutability has been queried and debated most closely in chronosequence studies, with conclusions ranging from strong support (6) to strong rejection (2) of space-for-time substitution. Increasingly, space-for-time substitution is being applied in biodiversity modeling to project climate-driven changes in species distributions, species richness, and compositional turnover (7-11). Examination of transferability of models for individual species has exposed concerns regarding the projection of these spatial models across time (12-15), and it has been suggested that models based on collective biodiversity properties might be more robust (9,16,17). However, the fundamental assumption that spatial relationships between climate and biodiversity can be used to project temporal trajectories of biodiversity under changing climates remains largely untested (but see refs. 16 and 18).The turnover of species among communities is particularly well suited for testing space-for-time substitution because it can be quantified independently across space or through time and because compositional turnover strongly correlates to climate variations in both space and time (19)(20)(21). However, other factors, such as species history, site history, and species interactions, also influence compositional turnover, independently or...