Functional traits-not nativeness-shape the effects of large mammalian herbivores on plant communities.

Large mammalian herbivores (megafauna) have experienced extinctions and declines since prehistory. Introduced megafauna have partly counteracted these losses yet are thought to have unusually negative effects on plants compared with native megafauna. Using a meta-analysis of 3995 plot-scale plant abundance and diversity responses from 221 studies, we found no evidence that megafauna impacts were shaped by nativeness, "invasiveness," "feralness," coevolutionary history, or functional and phylogenetic novelty. Nor was there evidence that introduced megafauna facilitate introduced plants more than native megafauna. Instead, we found strong evidence that functional traits shaped megafauna impacts, with larger-bodied and bulk-feeding megafauna promoting plant diversity. Our work suggests that trait-based ecology provides better insight into interactions between megafauna and plants than do concepts of nativeness.

Worldwide Late Pleistocene and Early Holocene population declines in extant megafauna are associated with Homo sapiens expansion rather than climate change.

The worldwide extinction of megafauna during the Late Pleistocene and Early Holocene is evident from the fossil record, with dominant theories suggesting a climate, human or combined impact cause. Consequently, two disparate scenarios are possible for the surviving megafauna during this time period - they could have declined due to similar pressures, or increased in population size due to reductions in competition or other biotic pressures. We therefore infer population histories of 139 extant megafauna species using genomic data which reveal population declines in 91% of species throughout the Quaternary period, with larger species experiencing the strongest decreases. Declines become ubiquitous 32-76 kya across all landmasses, a pattern better explained by worldwide Homo sapiens expansion than by changes in climate. We estimate that, in consequence, total megafauna abundance, biomass, and energy turnover decreased by 92-95% over the past 50,000 years, implying major human-driven ecosystem restructuring at a global scale.

The effect of past defaunation on ranges, niches, and future biodiversity forecasts.

Humans have reshaped the distribution of biodiversity across the globe, extirpating species from regions otherwise suitable and restricting populations to a subset of their original ranges. Here, we ask if anthropogenic range contractions since the Late Pleistocene led to an under-representation of the realized niches for megafauna, an emblematic group of taxa often targeted for restoration actions. Using reconstructions of past geographic distributions (i.e., natural ranges) for 146 extant terrestrial large-bodied (>44 kg) mammals, we estimate their climatic niches as if they had retained their original distributions and evaluate their observed niche dynamics. We found that range contractions led to a sizeable under-representation of the realized niches of several species (i.e., niche unfilling). For 29 species, more than 10% of the environmental space once seen in their natural ranges has been lost due to anthropogenic activity, with at least 12 species undergoing reductions of more than 50% of their realized niches. Eighteen species may now be confined to low-suitability locations, where fitness and abundance are likely diminished; we consider these taxa 'climatic refugees'. For those species, conservation strategies supported by current ranges risk being misguided if current, suboptimal habitats are considered baseline for future restoration actions. Because most climate-based biodiversity forecasts rely exclusively on current occurrence records, we went on to test the effect of neglecting historical information on estimates of species' potential distribution - as a proxy of sensitivity to climate change. We found that niche unfilling driven by past range contraction leads to an overestimation of sensitivity to future climatic change, resulting in 50% higher rates of global extinction, and underestimating the potential for megafauna conservation and restoration under future climate change. In conclusion, range contractions since the Late Pleistocene have also left imprints on megafauna realized climatic niches. Therefore, niche truncation driven by defaunation can directly affect climate and habitat-based conservation strategies.

Anthropocene refugia: integrating history and predictive modelling to assess the space available for biodiversity in a human-dominated world.

During periods of strong environmental change, some areas may serve as refugia, where components of biodiversity can find protection, persist and potentially expand from should conditions again become favourable. The refugia concept has previously been used in the context of climatic change, to describe climatically stable areas in which taxa survived past Quaternary glacial-interglacial oscillations, or where they might persist in the future under anthropogenic climate change. However, with the recognition that Earth has entered the Anthropocene, an era in which human activities are the dominant driving force on ecosystems, it is critical to also consider human pressures on the environment as factors limiting species distributions. Here, we present a novel concept, Anthropocene refugia, to refer to areas that provide spatial and temporal protection from human activities and that will remain suitable for a given taxonomic unit in the long-term. It integrates a deep-time perspective on species biogeography that provides information on the natural rather than current-day relictual distribution of species, with spatial information on modern and future anthropogenic threats. We define the concept and propose a methodology to effectively identify and map realized and potential current and future refugia, using examples for two megafaunal species as a proof of concept. We argue that identifying Anthropocene refugia will improve biodiversity conservation and restoration by allowing better prediction of key areas for conservation and potential for re-expansions today and in the future. More generally, it forms a new conceptual framework to assess and manage the impact of anthropogenic activities on past, current and future patterns of species distributions. This article is part of a discussion meeting issue 'The past is a foreign country: how much can the fossil record actually inform conservation?'

Shifted distribution baselines: neglecting long-term biodiversity records risks overlooking potentially suitable habitat for conservation management.

Setting appropriate conservation measures to halt the loss of biodiversity requires a good understanding of species' habitat requirements and potential distribution. Recent (past few decades) ecological data are typically used to estimate and understand species' ecological niches. However, historical local extinctions may have truncated species-environment relationships, resulting in a biased perception of species' habitat preferences. This may result in incorrect assessments of the area potentially available for their conservation. Incorporating long-term (centuries-old) occurrence records with recent records may provide better information on species-environment relationships and improve the modelling and understanding of habitat suitability. We test whether neglecting long-term occurrence records leads to an underestimation of species' historical niche and potential distribution and identify which species are more vulnerable to this effect. We compare outputs of species distribution models and niche hypervolumes built using recent records only with those built using both recent and long-term (post-1500) records, for a set of 34 large mammal species in South Africa. We find that, while using recent records only is adequate for some species, adding historical records in the analyses impacts estimates of the niche and habitat suitability for 12 species (34%) in our dataset, and that this effect is significantly higher for carnivores. These results show that neglecting long-term biodiversity records in spatial analyses risks misunderstanding, and generally underestimating, species' niches, which in turn may lead to ill-informed management decisions, with significant implications for the effectiveness of conservation efforts. This article is part of a discussion meeting issue 'The past is a foreign country: how much can the fossil record actually inform conservation?'

Unshifting the baseline: a framework for documenting historical population changes and assessing long-term anthropogenic impacts.

Ecological baselines-reference states of species' distributions and abundances-are key to the scientific arguments underpinning many conservation and management interventions, as well as to the public support to such interventions. Yet societal as well as scientific perceptions of these baselines are often based on ecosystems that have been deeply transformed by human actions. Despite increased awareness about the pervasiveness and implications of this shifting baseline syndrome, ongoing global assessments of the state of biodiversity do not take into account the long-term, cumulative, anthropogenic impacts on biodiversity. Here, we propose a new framework for documenting such impacts, by classifying populations according to the extent to which they deviate from a baseline in the absence of human actions. We apply this framework to the bowhead whale (Balaena mysticetus) to illustrate how it can be used to assess populations with different geographies and timelines of known or suspected impacts. Through other examples, we discuss how the framework can be applied to populations for which there is a wide diversity of existing knowledge, by making the best use of the available ecological, historical and archaeological data. Combined across multiple populations, this framework provides a standard for assessing cumulative anthropogenic impacts on biodiversity. This article is part of a discussion meeting issue 'The past is a foreign country: how much can the fossil record actually inform conservation?'

A spatially explicit estimate of the prewhaling abundance of the endangered North Atlantic right whale.

The North Atlantic right whale (NARW) (Eubalaena glacialis) is one of the world's most threatened whales. It came close to extinction after nearly a millennium of exploitation and currently persists as a population of only approximately 500 individuals. Setting appropriate conservation targets for this species requires an understanding of its historical population size, as a baseline for measuring levels of depletion and progress toward recovery. This is made difficult by the scarcity of records over this species' long whaling history. We sought to estimate the preexploitation population size of the North Atlantic right whale and understand how this species was distributed across its range. We used a spatially explicit data set on historical catches of North Pacific right whales (NPRWs) (Eubalaena japonica) to model the relationship between right whale relative density and the environment during the summer feeding season. Assuming the 2 right whale species select similar environments, we projected this model to the North Atlantic to predict how the relative abundance of NARWs varied across their range. We calibrated these relative abundances with estimates of the NPRW total prewhaling population size to obtain high and low estimates for the overall NARW population size prior to exploitation. The model predicted 9,075-21,328 right whales in the North Atlantic. The current NARW population is thus <6% of the historical North Atlantic carrying capacity and has enormous potential for recovery. According to the model, in June-September NARWs concentrated in 2 main feeding areas: east of the Grand Banks of Newfoundland and in the Norwegian Sea. These 2 areas may become important in the future as feeding grounds and may already be used more regularly by this endangered species than is thought.

How predictability of feeding patches affects home range and foraging habitat selection in avian social scavengers?

Feeding stations are commonly used to sustain conservation programs of scavengers but their impact on behaviour is still debated. They increase the temporal and spatial predictability of food resources while scavengers have supposedly evolved to search for unpredictable resources. In the Grands Causses (France), a reintroduced population of Griffon vultures Gyps fulvus can find carcasses at three types of sites: 1. "light feeding stations", where farmers can drop carcasses at their farm (spatially predictable), 2. "heavy feeding stations", where carcasses from nearby farms are concentrated (spatially and temporally predictable) and 3. open grasslands, where resources are randomly distributed (unpredictable). The impact of feeding stations on vulture's foraging behaviour was investigated using 28 GPS-tracked vultures. The average home range size was maximal in spring (1272 ± 752 km(2)) and minimal in winter (473 ± 237 km(2)) and was highly variable among individuals. Analyses of home range characteristics and feeding habitat selection via compositional analysis showed that feeding stations were always preferred compared to the rest of the habitat where vultures can find unpredictable resources. Feeding stations were particularly used when resources were scarce (summer) or when flight conditions were poor (winter), limiting long-ranging movements. However, when flight conditions were optimal, home ranges also encompassed large areas of grassland where vultures could find unpredictable resources, suggesting that vultures did not lose their natural ability to forage on unpredictable resources, even when feeding stations were available. However during seasons when food abundance and flight conditions were not limited, vultures seemed to favour light over heavy feeding stations, probably because of the reduced intraspecific competition and a pattern closer to the natural dispersion of resources in the landscape. Light feeding stations are interesting tools for managing food resources, but don't prevent vultures to feed at other places with possibly high risk of intoxication (poison).