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The impact of climate change on the structure of Pleistocene food webs across the mammoth steppe

Species interactions form food webs, impacting community structure and, potentially, ecological dynamics. It is likely that global climatic perturbations that occur over long periods of time have a significant influence on species interaction patterns. Here, we integrate stable isotope analysis and network theory to reconstruct patterns of trophic interactions for six independent mammalian communities that inhabited mammoth steppe environments spanning western Europe to eastern Alaska (Beringia) during the Late Pleis- tocene. We use a Bayesian mixing model to quantify the contribution of prey to the diets of local predators, and assess how the structure of trophic inter- actions changed across space and the Last Glacial Maximum (LGM), a global climatic event that severely impacted mammoth steppe communities. We find that large felids had diets that were more constrained than those of co-occurring predators, and largely influenced by an increase in Rangifer abun- dance after the LGM. Moreover, the structural organization of Beringian and European communities strongly differed: compared with Europe, species inter- actions in Beringian communities before—and possibly after—the LGM were highly modular. We suggest that this difference in modularity may have been driven by the geographical insularity of Beringian communities.

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Why a collapse of global civilization will be avoided: a comment on Ehrlich & Ehrlich

1st paragraph: Ehrlich FRS & Ehrlich [1] claim that over-population, over-consumption and the future climate mean that ‘preventing a global collapse of civilization is perhaps the foremost challenge confronting humanity’. What is missing from the well- referenced perspective of the potential downsides for the future of humanity is any balancing assessment of the progress being made on these three chal- lenges (and the many others they cite by way of detail) that suggests that the problems are being dealt with in a way that will not require a major disruption to the human condition or society. Earlier dire predictions have been made in the same mode by Malthus FRS [2] on food security, Jevons FRS [3] on coal exhaustion, King FRS & Murray [4] on peak oil, and by many others. They have all been overcome by the exercise of human ingenuity just as the doom was being prophesied with the deployment of steam engines to greatly improve agricultural efficiency, and the discoveries of oil and of fracking oil and gas, respectively, for the three examples given. It is incumbent on those who would continue to predict gloom to learn from history and make a comprehen- sive review of human progress before coming to their conclusions. The problems as perceived today by Ehrlich FRS and Ehrlich will be similarly seen off by work in progress by scientists and engineers. My comment is intended to summarize and reference the potential upsides being produced by today’s human ingenuity, and I leave the reader to weigh the balance for the future, taking into account the lessons of recent history.

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Future collapse: how optimistic should we be?

1st paragraph: Prof. Kelly FRS is optimistic about the chances of avoiding a collapse, but sadly we find his arguments entirely unpersuasive. For example, have Malthus (or we) really been wrong about food security? Roughly 850 million people are seriously undernourished (lacking sufficient calories) today, and perhaps 2 billion are malnourished (lacking one or more essential nutrients) [1]. When Malthus lived, there were only about 1 billion people on the planet. We agree that there are many things that could be done to feed today’s population of 7.1 billion, or even perhaps over 9 billion in 2050. Many of them (e.g. limiting waste) have been discussed for 50 years with little sign of progress. We do not think any serious analyst doubts that, if it were equitably distributed, today’s food production could nourish everyone adequately. Equally, we know of no serious analyst who believes such distribution is likely in the future. The concern is that climate disruption combined with other problems with the agricultural system will make it impossible to feed an ever larger future population, even if equal distribution were achieved. That concern is reinforced by the recent observation that, even before the likely heavy impacts of climate disruption on agriculture appear, production is failing to keep pace with projected needs [2].

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Life history predicts risk of species decline in a stochastic world

Understanding what traits determine the extinction risk of species has been a long-standing challenge. Natural populations increasingly experience reductions in habitat and population size concurrent with increasing novel environmental variation owing to anthropogenic disturbance and climate change. Recent studies show that a species risk of decline towards extinction is often non-random across species with differ- ent life histories. We propose that species with life histories in which all stage-specific vital rates are more evenly important to population growth rate may be less likely to decline towards extinction under these pressures. To test our prediction, we modelled declines in population growth rates under simulated stochas- tic disturbance to the vital rates of 105 species taken from the literature. Populations with more equally important vital rates, determined using elasticity analysis, declined more slowly across a gradient of increas- ing simulated environmental variation. Furthermore, higher evenness of elasticity was significantly correlated with a reduced chance of listing as Threatened on the International Union for Conservation of Nature Red List. The relative importance of life-history traits of diverse species can help us infer how natural assemblages will be affected by novel anthropogenic and climatic disturbances. Keywords: International Union for Conservation of Nature Red List; extinction; life history; stage-based; elasticity; stochasticity

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On a collision course: competition and dispersal differences create no-analogue communities and cause extinctions during climate change

Most climate change predictions omit species interactions and interspecific variation in dispersal. Here, we develop a model of multiple competing species along a warming climatic gradient that includes temperature- dependent competition, differences in niche breadth and interspecific differences in dispersal ability. Competition and dispersal differences decreased diversity and produced so-called ‘no-analogue’ commu- nities, defined as a novel combination of species that does not currently co-occur. Climate change altered community richness the most when species had narrow niches, when mean community-wide dispersal rates were low and when species differed in dispersal abilities. With high interspecific dispersal variance, the best dispersers tracked climate change, out-competed slower dispersers and caused their extinction. Overall, competition slowed the advance of colonists into newly suitable habitats, creating lags in climate tracking. We predict that climate change will most threaten communities of species that have narrow niches (e.g. tropics), vary in dispersal (most communities) and compete strongly. Current forecasts probably underestimate climate change impacts on biodiversity by neglecting competition and dispersal differences. Keywords: climate change; competition; dispersal; community ecology; movement ecology; thermal performance breadth

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Genetic change for earlier migration timing in a pink salmon population

To predict how climate change will influence populations, it is necessary to understand the mechanisms, particularly microevolution and phenotypic plasticity, that allow populations to persist in novel environmental conditions. Although evidence for climate-induced phenotypic change in populations is widespread, evidence documenting that these phenotypic changes are due to microevolution is exceed- ingly rare. In this study, we use 32 years of genetic data (17 complete generations) to determine whether there has been a genetic change towards earlier migration timing in a population of pink salmon that shows phenotypic change; average migration time occurs nearly two weeks earlier than it did 40 years ago. Experimental genetic data support the hypothesis that there has been directional selection for earlier migration timing, resulting in a substantial decrease in the late-migrating phenotype (from more than 30% to less than 10% of the total abundance). From 1983 to 2011, there was a significant decrease—over threefold—in the frequency of a genetic marker for late-migration timing, but there were minimal changes in allele frequencies at other neutral loci. These results demonstrate that there has been rapid microevolution for earlier migration timing in this population. Circadian rhythm genes, however, did not show any evidence for selective changes from 1993 to 2009. Keywords: microevolution; genetic change; salmon; circadian rhythms; climate change; migration timing

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How the type of anthropogenic change alters the consequences of ecological traps

Understanding altered ecological and evolutionary dynamics in novel environments is vital for predicting species responses to rapid environmental change. One fundamental concept relevant to such dynamics is the ecological trap, which arises from rapid anthropogenic change and can facilitate extinction. Ecological traps occur when formerly adaptive habitat preferences become maladaptive because the cues individuals preferentially use in selecting habitats lead to lower fitness than other alternatives. While it has been emphasized that traps can arise from different types of anthropogenic change, the resulting consequences of these different types of traps remain unknown. Using a novel model framework that builds upon the Price equation from evolutionary genetics, we provide the first analysis that contrasts the ecological and evolutionary consequences of ecological traps arising from two general types of perturbations known to trigger traps. Our model suggests that traps arising from degradation of existing habitats are more likely to facilitate extinction than those arising from the addition of novel trap habitat. Importantly, our framework reveals the mechanisms of these outcomes and the substantial scope for persistence via rapid evolution that may buffer many populations from extinction, helping to resolve the paradox of continued persistence of many species in dramatically altered landscapes. Keywords: attractive sink; evolutionary trap; habitat selection; maladaptation; Price equation; rapid evolution

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How does climate change cause extinction?

Anthropogenic climate change is predicted to be a major cause of species extinctions in the next 100 years. But what will actually cause these extinctions? For example, will it be limited physiological tolerance to high temperatures, changing biotic interactions or other factors? Here, we systematically review the proximate causes of climate-change related extinctions and their empirical support. We find 136 case studies of climatic impacts that are potentially relevant to this topic. However, only seven ident- ified proximate causes of demonstrated local extinctions due to anthropogenic climate change. Among these seven studies, the proximate causes vary widely. Surprisingly, none show a straightforward relation- ship between local extinction and limited tolerances to high temperature. Instead, many studies implicate species interactions as an important proximate cause, especially decreases in food availability. We find very similar patterns in studies showing decreases in abundance associated with climate change, and in those studies showing impacts of climatic oscillations. Collectively, these results highlight our disturbingly limited knowledge of this crucial issue but also support the idea that changing species interactions are an important cause of documented population declines and extinctions related to climate change. Finally, we briefly outline general research strategies for identifying these proximate causes in future studies. Keywords: climate change; extinction; physiological tolerances; species interactions

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Genetic consequences of climate change for northern plants

Climate change will lead to loss of range for many species, and thus to loss of genetic diversity crucial for their long-term persistence. We analysed range-wide genetic diversity (amplified fragment length poly- morphisms) in 9581 samples from 1200 populations of 27 northern plant species, to assess genetic consequences of range reduction and potential association with species traits. We used species distri- bution modelling (SDM, eight techniques, two global circulation models and two emission scenarios) to predict loss of range and genetic diversity by 2080. Loss of genetic diversity varied considerably among species, and this variation could be explained by dispersal adaptation (up to 57%) and by genetic differentiation among populations (FST; up to 61%). Herbs lacking adaptations for long-distance disper- sal were estimated to lose genetic diversity at higher rate than dwarf shrubs adapted to long-distance dispersal. The expected range reduction in these 27 northern species was larger than reported for tem- perate plants, and all were predicted to lose genetic diversity according to at least one scenario. SDM combined with FST estimates and/or with species trait information thus allows the prediction of species’ vulnerability to climate change, aiding rational prioritization of conservation efforts. Keywords: conservation genetics; FST; genetic diversity; range reduction; species distribution model; species traits

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Rising atmospheric carbon dioxide concentration and the future of C 4 crops for food and fuel

Crops with the C4 photosynthetic pathway are vital to global food supply, particularly in the tropical regions where human well-being and agricultural productivity are most closely linked. While rising atmospheric [CO2 ] is the driving force behind the greater temperatures and water stress, which threaten to reduce future crop yields, it also has the potential to directly benefit crop physiology. The nature of C4 plant responses to elevated [CO2 ] has been controversial. Recent evidence from free-air CO2 enrichment (FACE) experiments suggests that elevated [CO2] does not directly stimulate C4 photosynthesis. Nonetheless, drought stress can be ameliorated at elevated [CO2] as a result of lower stomatal conductance and greater intercellular [CO2]. Therefore, unlike C3 crops for which there is a direct enhancement of photosynthesis by elevated [CO2 ], C4 crops will only benefit from elevated [CO2 ] in times and places of drought stress. Current projections of future crop yields have assumed that rising [CO2] will directly enhance photosynthesis in all situations and, therefore, are likely to be overly optimistic. Additional experiments are needed to evaluate the extent to which amelioration of drought stress by elevated [CO2 ] will improve C4 crop yields for food and fuel over the range of C4 crop growing conditions and genotypes.

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Slow Recovery from Perturbations as a Generic Indicator of a Nearby Catastrophic Shift

The size of the basin of attraction in ecosystems with alternative stable states is often referred to as “ecological resilience.” Ecosystems with a low ecological resilience may easily be tipped into an alternative basin of attraction by a stochastic event. Unfortunately, it is very difficult to measure ecological resilience in practice. Here we show that the rate of recovery from small perturbations (some- times called “engineering resilience”) is a remarkably good indicator of ecological resilience. Such recovery rates decrease as a catastrophic regime shift is approached, a phenomenon known in physics as “crit- ical slowing down.” We demonstrate the robust occurrence of critical slowing down in six ecological models and outline a possible ex- perimental approach to quantify differences in recovery rates. In all the models we analyzed, critical slowing down becomes apparent quite far from a threshold point, suggesting that it may indeed be of practical use as an early warning signal. Despite the fact that critical slowing down could also indicate other critical transitions, such as a stable system becoming oscillatory, the robustness of the phenomenon makes it a promising indicator of loss of resilience and the risk of upcoming regime shifts in a system. Keywords: alternative stable states, catastrophic bifurcations, critical slowing down, early warning signals, resilience, return time.

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The beaver meadow complex revisited – the role of beavers in post-glacial floodplain development

We evaluate the validity of the beaver-meadow complex hypothesis, used to explain the deposition of extensive fine sediment in broad, low-gradient valleys. Previous work establishes that beaver damming forms wet meadows with multi-thread channels and enhanced sediment storage, but the long-term geomorphic effects of beaver are unclear. We focus on two low-gradient broad valleys, Beaver Meadows and Moraine Park, in Rocky Mountain National Park (Colorado, USA). Both valleys experienced a dramatic decrease in beaver population in the past century and provide an ideal setting for determining whether contemporary geomorphic conditions and sedimentation are within the historical range of variability of valley bottom processes. We examine the geomorphic significance of beaver-pond sediment by determining the rates and types of sedimentation since the middle Holocene and the role of beaver in driving floodplain evolution through increased channel complexity and fine sediment deposition. Sediment analyses from cores and cutbanks indicate that 33–50% of the alluvial sediment in Beaver Meadows is ponded and 28–40% was deposited in-channel; in Moraine Park 32–41% is ponded sediment and 40–52% was deposited in-channel. Radiocar- bon ages spanning 4300 years indicate long-term aggradation rates of ~0.05 cm yr-1. The observed highly variable short-term rates indicate temporal heterogeneity in aggradation, which in turn reflects spatial heterogeneity in processes at any point in time. Channel complexity increases directly downstream of beaver dams. The increased complexity forms a positive feedback for beaver-induced sedimentation; the multi-thread channel increases potential channel length for further damming, which increases the potential area occupied by beaver ponds and the volume of fine sediment trapped. Channel complexity decreased significantly as surveyed beaver population decreased. Beaver Meadows and Moraine Park represent settings where beaver substantially influence post-glacial floodplain aggradation. These findings underscore the importance of understanding the historical range of variability of valley bottom processes, and implications for environmental restoration. Copyright © 2011 John Wiley & Sons, Ltd. KEYWORDS: floodplain; sedimentation; beaver; Holocene

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Global imprint of climate change on marine life

Past meta-analyses of the response of marine organisms to climate change have examined a limited range of locations1,2, taxonomic groups2–4 and/or biological responses5,6. This has precluded a robust overview of the effect of climate change in the global ocean. Here, we synthesized all available studies of the consistency of marine ecological observations with expectations under climate change. This yielded a meta- database of 1,735 marine biological responses for which either regional or global climate change was considered as a driver. Included were instances of marine taxa responding as expected, in a manner inconsistent with expectations, and taxa demonstrating no response. From this database, 81–83% of all observations for distribution, phenology, community composition, abundance, demography and calcification across taxa and ocean basins were consistent with the expected impacts of climate change. Of the species responding to climate change, rates of distribution shifts were, on average, consistent with those required to track ocean surface temperature changes. Conversely, we did not find a relationship between regional shifts in spring phenology and the seasonality of temperature. Rates of observed shifts in species’ distributions and phenology are comparable to, or greater, than those for terrestrial systems.

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Point of No Return :The massive climate threats we must avoid

The world is quickly reaching a Point of No Return for preventing the worst impacts of climate change. Continuing on the current course will make it difficult, if not impossible, to prevent the widespread and catastrophic impacts of climate change. The costs will be substantial: billions spent to deal with the destruction of extreme weather events, untold human suffering, and the deaths of tens of millions from the impacts by as soon as 2030.

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Responses of wind erosion to climate-induced vegetation changes on the Colorado Plateau

Projected increases in aridity throughout the southwestern United States due to anthropogenic climate change will likely cause reduc- tions in perennial vegetation cover, which leaves soil surfaces exposed to erosion. Accelerated rates of dust emission from wind erosion have large implications for ecosystems and human well- being, yet there is poor understanding of the sources and magni- tude of dust emission in a hotter and drier climate. Here we use a two-stage approach to compare the susceptibility of grasslands and three different shrublands to wind erosion on the Colorado Plateau and demonstrate how climate can indirectly moderate the magnitude of aeolian sediment flux through different responses of dominant plants in these communities. First, using results from 20 y of vegetation monitoring, we found perennial grass cover in grass- lands declined with increasing mean annual temperature in the previous year, whereas shrub cover in shrublands either showed no change or declined as temperature increased, depending on the species. Second, we used these vegetation monitoring results and measurements of soil stability as inputs into a field-validated wind erosion model and found that declines in perennial vegeta- tion cover coupled with disturbance to biological soil crust resulted in an exponential increase in modeled aeolian sediment flux. Thus the effects of increased temperature on perennial plant cover and the correlation of declining plant cover with increased aeolian flux strongly suggest that sustained drought conditions across the southwest will accelerate the likelihood of dust production in the future on disturbed soil surfaces. arid ∣ horizontal flux ∣ land use ∣ national park ∣ threshold shear velocity

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Wilderness and biodiversity conservation

Human pressure threatens many species and ecosystems, so con- servation efforts necessarily prioritize saving them. However, conservation should clearly be proactive wherever possible. In this article, we assess the biodiversity conservation value, and specif- ically the irreplaceability in terms of species endemism, of those of the planet’s ecosystems that remain intact. We find that 24 wil- derness areas, all >1 million hectares, are >70% intact and have human densities of less than or equal to five people per km2. This wilderness covers 44% of all land but is inhabited by only 3% of people. Given this sparse population, wilderness conservation is cost-effective, especially if ecosystem service value is incorporated. Soberingly, however, most wilderness is not speciose: only 18% of plants and 10% of terrestrial vertebrates are endemic to individual wildernesses, the majority restricted to Amazonia, Congo, New Guinea, the Miombo–Mopane woodlands, and the North American deserts. Global conservation strategy must target these five wil- dernesses while continuing to prioritize threatened biodiversity hotspots.

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Flow regime, temperature, and biotic interactions drive differential declines of trout species under climate change

Broad-scale studies of climate change effects on freshwater species have focused mainly on temperature, ignoring critical drivers such as flow regime and biotic interactions. We use downscaled outputs from general circulation models coupled with a hydrologic model to forecast the effects of altered flows and increased temperatures on four interacting species of trout across the interior western United States (1.01 million km2), based on empirical statistical models built from fish surveys at 9,890 sites. Projections under the 2080s A1B emissions scenario forecast a mean 47% decline in total suitable habitat for all trout, a group of fishes of major socioeconomic and ecological significance. We project that native cutthroat trout Oncorhynchus clarkii, already excluded from much of its potential range by nonnative species, will lose a further 58% of habitat due to an increase in temper- atures beyond the species’ physiological optima and continued negative biotic interactions. Habitat for nonnative brook trout Salvelinus fontinalis and brown trout Salmo trutta is predicted to decline by 77% and 48%, respectively, driven by increases in temperature and winter flood frequency caused by warmer, rain- ier winters. Habitat for rainbow trout, Oncorhynchus mykiss, is projected to decline the least (35%) because negative temperature effects are partly offset by flow regime shifts that benefit the species. These results illustrate how drivers other than tempera- ture influence species response to climate change. Despite some uncertainty, large declines in trout habitat are likely, but our find- ings point to opportunities for strategic targeting of mitigation efforts to appropriate stressors and locations. global change | hydrology | invasive species | niche model | distribution modeling

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Housing growth in and near United States protected areas limits their conservation value

Protected areas are crucial for biodiversity conservation because they provide safe havens for species threatened by land-use change and resulting habitat loss. However, protected areas are only effective when they stop habitat loss within their boundaries, and are connected via corridors to other wild areas. The effectiveness of protected areas is threatened by development; however, the extent of this threat is unknown. We compiled spatially-detailed housing growth data from 1940 to 2030, and quantified growth for each wilderness area, national park, and national forest in the contermi- nous United States. Our findings show that housing development in the United States may severely limit the ability of protected areas to function as a modern “Noah’s Ark.” Between 1940 and 2000, 28 mil- lion housing units were built within 50 km of protected areas, and 940,000 were built within national forests. Housing growth rates during the 1990s within 1 km of protected areas (20% per decade) outpaced the national average (13%). If long-term trends continue, another 17 million housing units will be built within 50 km of pro- tected areas by 2030 (1 million within 1 km), greatly diminishing their conservation value. US protected areas are increasingly iso- lated, housing development in their surroundings is decreasing their effective size, and national forests are even threatened by habitat loss within their administrative boundaries. Protected areas in the United States are thus threatened similarly to those in developing countries. However, housing growth poses the main threat to protected areas in the United States whereas deforestation is the main threat in developing countries. conservation threats | effectiveness | parks | reserves

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Molecular study of worldwide distribution and diversity of soil animals

The global distribution of soil animals and the relationship of below-ground biodiversity to above-ground biodiversity are not well understood. We examined 17,516 environmental 18S rRNA gene sequences representing 20 phyla of soil animals sampled from 11 locations covering a range of biomes and latitudes around the world. No globally cosmopolitan taxa were found and only 14 of 2,259 operational taxonomic units (OTUs) found were common to four or more locations. Half of those were circumpolar and may reflect higher connectivity among circumpolar locations compared with other locations in the study. Even when OTU assembly criteria were relaxed to approximate the family taxo- nomic level, only 34 OTUs were common to four or more locations. A comparison of our diversity and community structure data to environmental factors suggests that below-ground animal diver- sity may be inversely related to above-ground biodiversity. Our data suggest that greater soil inorganic N and lower pH could explain the low below-ground biodiversity found at locations of high above-ground biodiversity. Our locations could also be characterized as being dominated by microarthropods or domi- nated by nematodes. Locations dominated by arthropods were primarily forests with lower soil pH, root biomass, mean annual temperature, low soil inorganic N and higher C:N, litter and moisture compared with nematode-dominated locations, which were mostly grasslands. Overall, our data indicate that small soil animals have distinct biogeographical distributions and provide unique evidence of the link between above-ground and below- ground biodiversity at a global scale. cosmopolitan species | endemism

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Improved probability of detection of ecological “surprises”

Ecological “surprises” are defined as unexpected findings about the natural environment. They are critically important in ecology because they are catalysts for questioning and reformulating views of the natural world, help shape assessments of the veracity of a priori predictions about ecological trends and phenomena, and underpin questioning of effectiveness of resource management. Despite the importance of ecological surprises, major gaps in understanding remain about how studies might be done differently or done better to improve the ability to identify them. We outline the kinds of ecological surprises that have arisen from long-term research programs that we lead in markedly different ecosystems around the world. Based on these case studies, we identify important lessons to guide both existing studies and new investigations to detect ecological surprises more readily, better anticipate unusual ecological phenomena, and take proactive steps to plan for and alleviate “undesirable” ecological surprises. Some of these lessons include: (i) maintain existing, and instigate new, long-term studies; (ii) conduct a range of kinds of parallel and concurrent research in a given target area; (iii) better use past literature and conceptual models of the target ecosystem in posing good questions and developing hypotheses and alternative hypotheses; and (iv) increase the capacity for ecological research to take advantage of opportunities arising from major natural disturbances. We argue that the increased anticipatory capability resulting from these lessons is critical given that ecological surprises may become more prevalent because of climate change and multiple and interacting environmental stressors.

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