Global late Quaternary megafauna extinctions linked to humans,not climate change

The late Quaternary megafauna extinction was a severe global-scale event. Two factors, climate change and modern humans, have received broad support as the primary drivers, but their absolute and relative importance remains controversial. To date, focus has been on the extinction chronology of individual or small groups of species, specific geographical regions or macroscale studies at very coarse geographical and taxonomic resolution, limiting the possibility of adequately testing the proposed hypotheses. We present, to our knowledge, the first global analysis of this extinction based on comprehensive country-level data on the geographical distribution of all large mammal species (more than or equal to 10 kg) that have gone globally or continentally extinct between the beginning of the Last Interglacial at 132 000 years BP and the late Holocene 1000 years BP, testing the relative roles played by glacial–interglacial climate change and humans. We show that the severity of extinction is strongly tied to hominin palaeobiogeography, with at most a weak, Eurasia-specific link to climate change. This first species-level macroscale analysis at relatively high geographical resolution provides strong support for modern humans as the primary driver of the worldwide megafauna losses during the late Quaternary.     

Climate‐driven ecological stability as a globally shared cause of Late Quaternary megafaunal extinctions: the Plaids and Stripes Hypothesis

Controversy persists about why so many large‐bodied mammal species went extinct around the end of the last ice age. Resolving this is important for understanding extinction processes in general, for assessing the ecological roles of humans, and for conserving remaining megafaunal species, many of which are endangered today. Here we explore an integrative hypothesis that asserts that an underlying cause of Late Quaternary megafaunal extinctions was a fundamental shift in the spatio‐temporal fabric of ecosystems worldwide. This shift was triggered by the loss of the millennial‐scale climate fluctuations that were characteristic of the ice age but ceased approximately 11700 years ago on most continents. Under ice‐age conditions, which prevailed for much of the preceding 2.6 Ma, these radical and rapid climate changes prevented many ecosystems from fully equilibrating with their contemporary climates. Instead of today's ‘striped’ world in which species' ranges have equilibrated with gradients of temperature, moisture, and seasonality, the ice‐age world was a disequilibrial ‘plaid’ in which species' ranges shifted rapidly and repeatedly over time and space, rarely catching up with contemporary climate. In the transient ecosystems that resulted, certain physiological, anatomical, and ecological attributes shared by megafaunal species pre‐adapted them for success. These traits included greater metabolic and locomotory efficiency, increased resistance to starvation, longer life spans, greater sensory ranges, and the ability to be nomadic or migratory. When the plaid world of the ice age ended, many of the advantages of being large were either lost or became disadvantages. For instance in a striped world, the low population densities and slow reproductive rates associated with large body size reduced the resiliency of megafaunal species to population bottlenecks. As the ice age ended, the downsides of being large in striped environments lowered the extinction thresholds of megafauna worldwide, which then increased the vulnerability of individual species to a variety of proximate threats they had previously tolerated, such as human predation, competition with other species, and habitat loss. For many megafaunal species, the plaid‐to‐stripes transition may have been near the base of a hierarchy of extinction causes whose relative importances varied geographically, temporally, and taxonomically.     

Accelerating local extinction associated with very recent climate change

Climate change has already caused local extinction in many plants and animals, based on surveys spanning many decades. As climate change accelerates, the pace of these extinctions may also accelerate, potentially leading to large-scale, species-level extinctions. We tested this hypothesis in a montane lizard. We resurveyed 18 mountain ranges in 2021–2022 after only ~7 years. We found rates of local extinction among the fastest ever recorded, which have tripled in the past ~7 years relative to the preceding ~42 years. Further, climate change generated local extinction in ~7 years similar to that seen in other organisms over ~70 years. Yet, contrary to expectations, populations at two of the hottest sites survived. We found that genomic data helped predict which populations survived and which went extinct. Overall, we show the increasing risk to biodiversity posed by accelerating climate change and the opportunity to study its effects over surprisingly brief timescales.     

Precipitation buffers temperature-driven local extinctions of moths at warm range margins

Species' distributions are moving polewards in response to climate change, and although range expansions of relatively warm-adapted species are widely reported, reports of range retractions in cool-adapted species are less common. Here, we analysed species' distribution shifts for 76 cool-adapted moths in Great Britain using citizen science occurrence records from the National Moth Recording Scheme over a 40-year period. Although we find evidence for trailing edge shifts to higher latitudes, shifts in species' range centroids are oriented towards the north-west, and are more closely correlated with directional changes in total precipitation than average temperature. We also found that species' local extinction risk is higher in areas where temperature is high and precipitation is low, but this risk diminishes as precipitation increases. Adaptation efforts should therefore focus on maintaining or increasing water availability as the climate continues to change.     

Organism activity levels predict marine invertebrate survival during ancient global change extinctions

Multistressor global change, the combined influence of ocean warming, acidification, and deoxygenation, poses a serious threat to marine organisms. Experimental studies imply that organisms with higher levels of activity should be more resilient, but testing this prediction and understanding organism vulnerability at a global scale, over evolutionary timescales, and in natural ecosystems remain challenging. The fossil record, which contains multiple extinctions triggered by multistressor global change, is ideally suited for testing hypotheses at broad geographic, taxonomic, and temporal scales. Here, I assess the importance of activity level for survival of well‐skeletonized benthic marine invertebrates over a 100‐million‐year‐long interval (Permian to Jurassic periods) containing four global change extinctions, including the end‐Permian and end‐Triassic mass extinctions. More active organisms, based on a semiquantitative score incorporating feeding and motility, were significantly more likely to survive during three of the four extinction events (Guadalupian, end‐Permian, and end‐Triassic). In contrast, activity was not an important control on survival during nonextinction intervals. Both the end‐Permian and end‐Triassic mass extinctions also triggered abrupt shifts to increased dominance by more active organisms. Although mean activity gradually returned toward pre‐extinction values, the net result was a permanent ratcheting of ecosystem‐wide activity to higher levels. Selectivity patterns during ancient global change extinctions confirm the hypothesis that higher activity, a proxy for respiratory physiology, is a fundamental control on survival, although the roles of specific physiological traits (such as extracellular pCO₂ or aerobic scope) cannot be distinguished. Modern marine ecosystems are dominated by more active organisms, in part because of selectivity ratcheting during these ancient extinctions, so on average may be less vulnerable to global change stressors than ancient counterparts. However, ancient extinctions demonstrate that even active organisms can suffer major extinction when the intensity of environmental disruption is intense.     

Annual temperature variation as a time machine to understand the effects of long‐term climate change on a poleward range shift

Range shifts due to annual variation in temperature are more tractable than range shifts linked to decadal to century long temperature changes due to climate change, providing natural experiments to determine the mechanisms responsible for driving long‐term distributional shifts. In this study we couple physiologically grounded mechanistic models with biogeographic surveys in 2 years with high levels of annual temperature variation to disentangle the drivers of a historical range shift driven by climate change. The distribution of the barnacle Semibalanus balanoides has shifted 350 km poleward in the past half century along the east coast of the United States. Recruits were present throughout the historical range following the 2015 reproductive season, when temperatures were similar to those in the past century, and absent following the 2016 reproductive season when temperatures were warmer than they have been since 1870, the earliest date for temperature records. Our dispersal dependent mechanistic models of reproductive success were highly accurate and predicted patterns of reproduction success documented in field surveys throughout the historical range in 2015 and 2016. Our mechanistic models of reproductive success not only predicted recruitment dynamics near the range edge but also predicted interior range fragmentation in a number of years between 1870 and 2016. All recruits monitored within the historical range following the 2015 colonization died before 2016 suggesting juvenile survival was likely the primary driver of the historical range retraction. However, if 2016 is indicative of future temperatures mechanisms of range limitation will shift and reproductive failure will lead to further range retraction in the future. Mechanistic models are necessary for accurately predicting the effects of climate change on ranges of species.     

Plant controls on Late Quaternary whole ecosystem structure and function

Plants and animals influence biomass production and nutrient cycling in terrestrial ecosystems; however, their relative importance remains unclear. We assessed the extent to which mega‐herbivore species controlled plant community composition and nutrient cycling, relative to other factors during and after the Late Quaternary extinction event in Britain and Ireland, when two‐thirds of the region's mega‐herbivore species went extinct. Warmer temperatures, plant–soil and plant–plant interactions, and reduced burning contributed to the expansion of woody plants and declining nitrogen availability in our five study ecosystems. Shrub biomass was consistently one of the strongest predictors of ecosystem change, equalling or exceeding the effects of other biotic and abiotic factors. In contrast, there was relatively little evidence for mega‐herbivore control on plant community composition and nitrogen availability. The ability of plants to determine the fate of terrestrial ecosystems during periods of global environmental change may therefore be greater than previously thought.     

Climate change impact on Balearic shearwater through a trophic cascade

A recent study showed that a critically endangered migratory predator species, the Balearic shearwater Puffinus mauretanicus, rapidly expanded northwards in northeast Atlantic waters after the mid-1990s. As a significant positive correlation was found between the long-term changes in the abundance of this seabird and sea temperature around the British Isles, it was hypothesized that the link between the biogeographic shift and temperature occurred through the food web. Here, we test this conjecture and reveal concomitant changes in a regional index of sea temperature, plankton (total calanoid copepod), fish prey (anchovy and sardine) and the Balearic shearwater for the period 1980-2003. All three trophic levels exhibit a significant shift detected between 1994 and 1996. Our findings therefore support the assertion of both a direct and an indirect effect of climate change on the spatial distribution of post-breeding Balearic shearwater through a trophic cascade.     

A review of Quaternary range shifts in European aquatic Coleoptera

Aim To undertake a quantitative review of the Quaternary fossil record of European water beetles to evaluate their geographical and temporal coverage, and to characterize the extent and typology of the shifts in their geographical ranges. Location Europe. Methods We compiled Quaternary water beetle records from public databases and published references. We included in the analyses species of 10 families of aquatic Coleoptera, and recorded range shifts through the comparison of the location of fossil remains with the current distribution of the species. We explored the ecological representativeness of the fossil record, as well as the relationship between range shifts and the habitat type of the species. Results Our final data set included over 9000 records for 259 water beetle species. Fossil remains of aquatic beetles have been documented exclusively north of 42° N, with most of the records from the British Isles and virtually none from southern Europe or the Mediterranean Basin. Over 80% of the records were from the Late Glacial and the Holocene periods (the last 15 kyr), and overall approximately 20% of the species have been recorded outside their present range (23% excluding Holocene records). Most range shifts were southern or western extensions of currently widespread, northern species, with 10 species displaying major range shifts through the Palaearctic. Lentic species were significantly more likely to have experienced major range shifts, even accounting for the general ecological bias of the fossil record towards lentic habitats. Main conclusions Our results show that the Quaternary record of aquatic Coleoptera is geographically, temporally and ecologically skewed, necessitating caution when extrapolating general conclusions about range changes and ecological stability to other areas or periods on the basis of such scattered evidence. Most central and northern European species for which there are fossil records seem to have conserved their ranges through the Late Pleistocene, with geographical shifts mostly restricted to species with current widespread north Palaearctic or Holarctic distributions. Major range shifts through the Palaearctic are taxonomically uneven, suggesting either an idiosyncratic behaviour of taxa depending on ecological or phylogenetic factors, or a sampling artefact produced by the limited availability of taxonomic expertise.     

Warning times for species extinctions due to climate change

Climate change is likely to become an increasingly major obstacle to slowing the rate of species extinctions. Several new assessment approaches have been proposed for identifying climate‐vulnerable species, based on the assumption that established systems such as the IUCN Red List need revising or replacing because they were not developed to explicitly consider climate change. However, no assessment approach has been tested to determine its ability to provide advanced warning time for conservation action for species that might go extinct due to climate change. To test the performance of the Red List system in this capacity, we used linked niche‐demographic models with habitat dynamics driven by a ‘business‐as‐usual’ climate change scenario. We generated replicate 100‐year trajectories for range‐restricted reptiles and amphibians endemic to the United States. For each replicate, we categorized the simulated species according to IUCN Red List criteria at annual, 5‐year, and 10‐year intervals (the latter representing current practice). For replicates that went extinct, we calculated warning time as the number of years the simulated species was continuously listed in a threatened category prior to extinction. To simulate data limitations, we repeated the analysis using a single criterion at a time (disregarding other listing criteria). Results show that when all criteria can be used, the Red List system would provide several decades of warning time (median = 62 years; >20 years for 99% of replicates), but suggest that conservation actions should begin as soon as a species is listed as Vulnerable, because 50% of replicates went extinct within 20 years of becoming uplisted to Critically Endangered. When only one criterion was used, warning times were substantially shorter, but more frequent assessments increased the warning time by about a decade. Overall, we found that the Red List criteria reliably provide a sensitive and precautionary way to assess extinction risk under climate change.     

Ecological niches as stable distributional constraints on mammal species, with implications for Pleistocene extinctions and climate change projections for biodiversity

Aim Theoretical work suggests that species’ ecological niches should remain relatively constant over long‐term ecological time periods, but empirical tests are few. We present longitudinal studies of 23 extant mammal species, modelling ecological niches and predicting geographical distributions reciprocally between the Last Glacial Maximum and present to test this evolutionary conservatism. Location This study covered distributional shifts in mammal species across the lower 48 states of the United States. Methods We used a machine‐learning tool for modelling species’ ecological niches, based on known occurrences and electronic maps summarizing ecological dimensions, to assess the ability of ecological niches as modelled in one time period to predict the geographical distribution of the species in another period, and vice versa. Results High intertemporal predictivity between niche models and species’ occurrences indicate that niche conservatism is widespread among the taxa studied, particularly when statistical power is considered as a reason for failure of reciprocal predictions. Niche projections to the present for 8 mammal taxa that became extinct at the end of the Pleistocene generally increased in area, and thus do not support the hypothesis of niche collapse as a major driving force in their extinction. Main conclusions Ecological niches represent long‐term stable constraints on the distributional potential of species; indeed, this study suggests that mammal species have tracked consistent climate profiles throughout the drastic climate change events that marked the end of the Pleistocene glaciations. Many current modelling efforts focusing on anticipating climate change effects on species’ potential geographical distributions will be bolstered by this result — in essence, the first longitudinal demonstration of niche conservatism.     

Eco‐evolution on the edge during climate change

We urgently need to predict species responses to climate change to minimize future biodiversity loss and ensure we do not waste limited resources on ineffective conservation strategies. Currently, most predictions of species responses to climate change ignore the potential for evolution. However, evolution can alter species ecological responses, and different aspects of evolution and ecology can interact to produce complex eco‐evolutionary dynamics under climate change. Here we review how evolution could alter ecological responses to climate change on species warm and cool range margins, where evolution could be especially important. We discuss different aspects of evolution in isolation, and then synthesize results to consider how multiple evolutionary processes might interact and affect conservation strategies. On species cool range margins, the evolution of dispersal could increase range expansion rates and allow species to adapt to novel conditions in their new range. However, low genetic variation and genetic drift in small range‐front populations could also slow or halt range expansions. Together, these eco‐evolutionary effects could cause a three‐step, stop‐and‐go expansion pattern for many species. On warm range margins, isolation among populations could maintain high genetic variation that facilitates evolution to novel climates and allows species to persist longer than expected without evolution. This ‘evolutionary extinction debt’ could then prevent other species from shifting their ranges. However, as climate change increases isolation among populations, increasing dispersal mortality could select for decreased dispersal and cause rapid range contractions. Some of these eco‐evolutionary dynamics could explain why many species are not responding to climate change as predicted. We conclude by suggesting that resurveying historical studies that measured trait frequencies, the strength of selection, or heritabilities could be an efficient way to increase our eco‐evolutionary knowledge in climate change biology.     

Temperature change effects on marine fish range shifts: A meta-analysis of ecological and methodological predictors

The current effects of global warming on marine ecosystems are predicted to increase, with species responding by changing their spatial distributions. Marine ectotherms such as fish experience elevated distribution shifts, as temperature plays a key role in physiological functions and delineating population ranges through thermal constraints. Distributional response predictions necessary for population management have been complicated by high heterogeneity in magnitude and direction of movements, which may be explained by both biological as well as methodological study differences. To date, however, there has been no comprehensive synthesis of the interacting ecological factors influencing fish distributions in response to climate change and the confounding methodological factors that can affect their estimation. In this study we analyzed published studies meeting criteria of reporting range shift responses to global warming in 115 taxa spanning all major oceanic regions, totaling 595 three-dimensional population responses (latitudinal, longitudinal, and depth), with temperature identified as a significant driver. We found that latitudinal shifts were the fastest in non-exploited, tropical populations, and inversely correlated with depth shifts which, in turn, dominated at the trailing edges of population ranges. While poleward responses increased with rate of temperature change and latitude, niche was a key factor in predicting both depth (18% of variation) and latitudinal responses (13%), with methodological predictors explaining between 10% and 28% of the observed variance in marine fish responses to temperature change. Finally, we found strong geographical publication bias and limited taxonomical scope, highlighting the need for more representative and standardized research in order to address heterogeneity in distribution responses and improve predictions in face of changing climate.     

Late Quaternary climate legacies in contemporary plant functional composition

The functional composition of plant communities is commonly thought to be determined by contemporary climate. However, if rates of climate‐driven immigration and/or exclusion of species are slow, then contemporary functional composition may be explained by paleoclimate as well as by contemporary climate. We tested this idea by coupling contemporary maps of plant functional trait composition across North and South America to paleoclimate means and temporal variation in temperature and precipitation from the Last Interglacial (120 ka) to the present. Paleoclimate predictors strongly improved prediction of contemporary functional composition compared to contemporary climate predictors, with a stronger influence of temperature in North America (especially during periods of ice melting) and of precipitation in South America (across all times). Thus, climate from tens of thousands of years ago influences contemporary functional composition via slow assemblage dynamics.     

用组合模型综合比较的方法分析气候变化对朱鹮潜在生境的影响

气候变化的不确定性和物种与环境关系的不确定性使气候变化生物学的研究充满变数。为了降低不确定性,人们开始用组合模型综合比较的方法研究物种对气候变化的响应。以朱鹮(Nipponia nippon)为研究对象,介绍组合模型综合比较方法的特点。朱鹮曾经高度濒危,目前种群大小在迅速恢复中;然而其分布区依旧狭小,气候变化可能是朱鹮面临的新威胁。应用BIOMOD模型中的9种模型,选择了每年的最低温和最高温、温度的季节性变异、每年的总降水量和降水的季节性变异共5个气候因子,依据WorldClim气候数据的CGCM2气候模型的A2a排放情形,计算了朱鹮当前(1950—2000年)的适宜生境和2020年、2050年、2080年3个阶段的潜在生境范围。结果表明朱鹮潜在生境将逐渐北移,生境中心脱离现在的保护区。因此,制定朱鹮的长期保护策略是必要的。9个模型在预测结果上、变量权重上和拟合优度的指标上都有差异,反映了模型本身的不确定性。气候变化的生物学效应比较复杂,应用多个模型进行综合比较,可以尽可能地减少模型所导致的误差。     

Taxonomy and palaeobiology of the largest-ever marsupial, Diprotodon Owen, 1838 (Diprotodontidae, Marsupialia)

To determine accurately the rates of late Pleistocene megafaunal loss, it is fundamentally important to have accurate taxonomic information for every species. In Australia, accurate taxonomic information is lacking for several Pleistocene groups, including the largest marsupial ever to live, Diprotodon Owen, 1838. Diprotodon taxonomy has been complicated by early nomenclatural problems and by the occurrence of two distinct size classes of individuals that do not reflect an ontogenetic series. Traditionally, the two size classes have been regarded as separate species. However, a taxonomic investigation of large samples (> 1000 teeth) of Diprotodon material from several different fossil localities in Queensland, New South Wales, South Australia and Victoria suggests that there is little evidence for the discrimination of more than one morphospecies. Thus, Diprotodon is here considered a monotypic genus and the single morphospecies, D. optatum Owen, 1838 is considered to have been highly sexually dimorphic. By drawing analogy with extant sexually dimorphic megaherbivores and marsupials, the large form was probably male, and the small form was probably female. Diprotodon optatum probably moved in small, gender-segregated herds, and exhibited a polygynous breeding strategy. As a single morphospecies, D. optatum had a near-continental geographical distribution, similar to that of extant megaherbivores, possibly indicating its niche as a habitat generalist.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 153 , 389–417.     

Extinctions of herbivorous mammals in the late Pleistocene of Australia in relation to their feeding ecology: No evidence for environmental change as cause of extinction

Abstract There has been debate over the cause of the extinction of ‘megafauna’ species during the late Pleistocene of Australia. One view is that environmental change, either natural or human‐induced, was the main factor in the extinctions. Some support for this comes from the observation that, among herbivores, most of the species that went extinct were apparently browsers rather than grazers. Browsers would presumably have been more dependent on shrubland and woodland habitats than grazers, and it has been argued that such habitats might have contracted in response to aridity or changed fire regimes in the late Pleistocene. Here, we test this idea by comparing extinction rates of browsers and grazers in the late Pleistocene, controlling for body mass in both groups. We show that in both browsers and grazers the probability of extinction was very strongly related to body mass, and the body mass at which extinction became likely was similar in the two groups. It is true that more browsers than grazers went extinct, but this is largely because most very large herbivores in the late Pleistocene were browsers, not because large browsers were more likely to go extinct than similarly sized grazers. This result provides evidence against some forms of environmental change as a cause of the extinctions.     

Prediction uncertainty of environmental change effects on temperate European biodiversity

Observed patterns of species richness at landscape scale (gamma diversity) cannot always be attributed to a specific set of explanatory variables, but rather different alternative explanatory statistical models of similar quality may exist. Therefore predictions of the effects of environmental change (such as in climate or land cover) on biodiversity may differ considerably, depending on the chosen set of explanatory variables. Here we use multimodel prediction to evaluate effects of climate, land-use intensity and landscape structure on species richness in each of seven groups of organisms (plants, birds, spiders, wild bees, ground beetles, true bugs and hoverflies) in temperate Europe. We contrast this approach with traditional best-model predictions, which we show, using cross-validation, to have inferior prediction accuracy. Multimodel inference changed the importance of some environmental variables in comparison with the best model, and accordingly gave deviating predictions for environmental change effects. Overall, prediction uncertainty for the multimodel approach was only slightly higher than that of the best model, and absolute changes in predicted species richness were also comparable. Richness predictions varied generally more for the impact of climate change than for land-use change at the coarse scale of our study. Overall, our study indicates that the uncertainty introduced to environmental change predictions through uncertainty in model selection both qualitatively and quantitatively affects species richness projections.     

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 Pleistocene. 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 interactions 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 abundance after the LGM. Moreover, the structural organization of Beringian and European communities strongly differed: compared with Europe, species interactions 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.