Pleistocene megafaunal extinctions and the functional loss of long‐distance seed‐dispersal services

Pleistocene extinctions affected mainly large‐bodied animals, determining the loss or changes in numerous ecological functions. Evidence points to a central role of many extinct megafauna herbivores as seed dispersers. An important step in understanding the legacy of extinct mutualistic interactions is to evaluate the roles and effectiveness of megafauna herbivores in seed dispersal. Here we use morphological and ecophysiological allometries to estimate both quantitative and qualitative aspects of seed‐dispersal services likely provided by extinct megafauna. We developed a mechanistic model that encompasses four stages of seed dispersal – seed ingestion, gut retention, animal movement, and seed deposition. We estimate seed‐dispersal kernels through simulations to infer the role of Pleistocene megafauna in promoting long‐distance dispersal and examine how seed dispersal was affected by extinctions. Simulations suggest extinct large‐bodied frugivores would frequently disperse large seeds over a thousand meters, whereas smaller‐bodied frugivores are more likely to deposit the seeds over a few hundred meters. Moreover, events of long‐distance seed dispersal by the extinct megafauna would be up to ten times longer than long‐distance dispersal by smaller‐sized extant mammals. By estimating the combined distribution of seed dispersal distances considering all large‐bodied mammalian frugivores in specific South American Pleistocene assemblages we found that long‐distance dispersal contracted by at least two thirds after the megafauna died out. The disruption of long‐distance dispersal is expected to have consequences for recruitment, spatial and genetic structure of plant populations, population persistence and community composition. Promoting long‐distance seed dispersal was one among other salient features of extinct Pleistocene megafauna that reveal their influence on natural ecosystems. Modeling the consequences of megafaunal extinctions can offer quantitative predictions on the consequences of ongoing defaunation to plant populations and ecological communities.     

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.     

The global decline of freshwater megafauna

Freshwater ecosystems are among the most diverse and dynamic ecosystems on Earth. At the same time, they are among the most threatened ecosystems but remain underrepresented in biodiversity research and conservation efforts. The rate of decline of vertebrate populations is much higher in freshwaters than in terrestrial or marine realms. Freshwater megafauna (i.e., freshwater animals that can reach a body mass ≥30 kg) are intrinsically prone to extinction due to their large body size, complex habitat requirements and slow life‐history strategies such as long life span and late maturity. However, population trends and distribution changes of freshwater megafauna, at continental or global scales, remain unclear. In the present study, we compiled population data of 126 freshwater megafauna species globally from the Living Planet Database and available literature, and distribution data of 44 species inhabiting Europe and the United States from literature and databases of the International Union for Conservation of Nature and NatureServe. We quantified changes in population abundance and distribution range of freshwater megafauna species. Globally, freshwater megafauna populations declined by 88% from 1970 to 2012, with the highest declines in the Indomalaya and Palearctic realms (?99% and ?97%, respectively). Among taxonomic groups, mega‐fishes exhibited the greatest global decline (?94%). In addition, freshwater megafauna experienced major range contractions. For example, distribution ranges of 42% of all freshwater megafauna species in Europe contracted by more than 40% of historical areas. We highlight the various sources of uncertainty in tracking changes in populations and distributions of freshwater megafauna, such as the lack of monitoring data and taxonomic and spatial biases. The detected trends emphasize the critical plight of freshwater megafauna globally and highlight the broader need for concerted, targeted and timely conservation of freshwater biodiversity.     

Human influence on distribution and extinctions of the late Pleistocene Eurasian megafauna

Late Pleistocene extinctions are of interest to paleontological and anthropological research. In North America and Australia, human occupation occurred during a short period of time and overexploitation may have led to the extinction of mammalian megafauna. In northern Eurasia megafaunal extinctions are believed to have occurred over a relatively longer period of time, perhaps as a result of changing environmental conditions, but the picture is much less clear. To consider megafaunal extinction in Eurasia, we compare differences in the geographical distribution and commonness of extinct and extant species between paleontological and archaeological localities from the late middle Pleistocene to Holocene. Purely paleontological localities, as well as most extinct species, were distributed north of archaeological sites and of the extant species, suggesting that apart from possible differences in adaptations between humans and other species, humans could also have a detrimental effect on large mammal distribution. However, evidence for human overexploitation applies only to the extinct steppe bison Bison priscus. Other human-preferred species survive into the Holocene, including Rangifer tarandus, Equus ferus, Capreolus capreolus, Cervus elaphus, Equus hemionus, Saiga tatarica, and Sus scrofa. Mammuthus primigenius and Megaloceros giganteus were rare in archaeological sites. Carnivores appear little influenced by human presence, although they become rarer in Holocene archaeological sites. Overall, the data are consistent with the conclusion that humans acted as efficient hunters selecting for the most abundant species. Our study supports the idea that the late Pleistocene extinctions were environmentally driven by climatic changes that triggered habitat fragmentation, species range reduction, and population decrease, after which human interference either by direct hunting or via indirect activities probably became critical.     

Freshwater megafauna diversity: Patterns,status and threats

Aim

Freshwater megafauna remain underrepresented in research and conservation, despite a disproportionately high risk of extinction due to multiple human threats. Therefore, our aims are threefold; (i) identify global patterns of freshwater megafauna richness and endemism, (ii) assess the conservation status of freshwater megafauna and (iii) demonstrate spatial and temporal patterns of human pressure throughout their distribution ranges.

Location

Global.

Methods

We identified 207 extant freshwater megafauna species, based on a 30 kg weight threshold, and mapped their distributions using HydroBASINS subcatchments (level 8). Information on conservation status and population trends for each species was extracted from the IUCN Red List website. We investigated human impacts on freshwater megafauna in space and time by examining spatial congruence between their distributions and human pressures, described by the Incident Biodiversity Threat Index and Temporal Human Pressure Index.

Results

Freshwater megafauna occur in 76% of the world’s main river basins (level 3 HydroBASINS), with species richness peaking in the Amazon, Congo, Orinoco, Mekong and Ganges‐Brahmaputra basins. Freshwater megafauna are more threatened than their smaller counterparts within the specific taxonomic groups (i.e., fishes, mammals, reptiles and amphibians). Out of the 93 freshwater megafauna species with known population trends, 71% are in decline. Meanwhile, IUCN Red List assessments reported insufficient or outdated data for 43% of all freshwater megafauna species. Since the early 1990s, human pressure has increased throughout 63% of their distribution ranges, with particularly intense impacts occurring in the Mekong and Ganges‐Brahmaputra basins.

Main conclusions

Freshwater megafauna species are threatened globally, with intense and increasing human pressures occurring in many of their biodiversity hotspots. We call for research and conservation actions for freshwater megafauna, as they are highly sensitive to present and future pressures including a massive boom in hydropower dam construction in their biodiversity hotspots.

     

Using carbon isotope analysis of the diet of two introduced Australian megaherbivores to understand Pleistocene megafaunal extinctions

Aim Australia lost a diverse assemblage of large marsupial herbivores in the late Pleistocene, with suggestions that the extinctions were biased towards browsers. In modern times two bovines, the Asian water buffalo (Bubalus bubalis) and banteng (Bos javanicus), have established feral populations in the Northern Territory, Australia. Buffalo have aggressively expanded throughout the savanna landscape, yet banteng remain near their point of introduction on the Cobourg Peninsula. We hypothesized that this difference is related to feeding ecology, possibly reflecting a legacy of the Pleistocene extinctions. Location Western Arnhem Land, Northern Territory, Australia. Methods Analysing a previously published dataset of body mass and feeding ecology of extinct and extant marsupial herbivores, we evaluated whether browsers were at greater risk of extinction than grazers. We compared the carbon isotope composition and nitrogen content of banteng and buffalo dung in order to evaluate the hypotheses that the differences in invasion success are related to feeding ecology, and that seasonal variation in browse consumption is linked to changing nutritional quality of grass. Results Controlling for body mass, the Pleistocene extinctions were clearly biased towards browsers. Introduced banteng appear to be primarily browsers, with their diets comprising 40% grass in the wet season and 15% in the late dry season. Buffalo have a more variable diet, with an increasing proportion of browse from the wet (30%) to the late dry season (75%), and can therefore be described as switching from grazer to browser. The decline of grass in the diet of both species appears to reflect the decline in the nutritional value of grass through the dry season, an inference supported by the negative relationship between δ¹³C values and the nitrogen content of dung. Main conclusions Banteng and buffalo are much larger than extant native herbivores, of which browsers are restricted to isolated rocky habitats. This suggests that banteng and buffalo have filled niches made vacant following the Pleistocene extinctions. The success of buffalo appears to be related to their greater dietary breadth, which enables them to graze and browse in eucalypt savannas, whilst the browsing banteng remain tethered to a mosaic of rain forest patches. The restriction of browsers may be a long‐range consequence of habitat transformations associated with Aboriginal landscape burning.     

Large mammals and trees in eastern monsoonal China: anthropogenic losses since the Late Pleistocene and restoration prospects in the Anthropocene

Massive human-induced declines of large-sized animals and trees (megabiota) from the Late Pleistocene to the Anthropocene have resulted in downsized ecosystems across the globe, in which components and functions have been greatly simplified. In response, active restoration projects of extant large-sized species or functional substitutes are needed at large scales to promote ecological processes that are important for ecosystem self-regulation and biodiversity maintenance. Despite the desired global scope of such projects, they have received little attention in East Asia. Here, we synthesise the biogeographical and ecological knowledge of megabiota in ancient and modern China, with relevant data mostly located in eastern monsoonal China (EMC), aiming to assess its potential for restoring functionally intact ecosystems modulated by megabiota. We found that during the Late Pleistocene, 12 mammalian megafaunal (carnivores ≥15 kg and herbivores ≥500 kg) species disappeared from EMC: one carnivore Crocuta ultima (East Asian spotted hyena) and 11 herbivores including six megaherbivores (≥1000 kg). The relative importance of climate change and humans in driving these losses remains debated, despite accumulating evidence in favour of the latter. Later massive depletion of megafauna and large-sized (45–500 kg) herbivores has been closely associated with agricultural expansion and societal development, especially during the late Holocene. While forests rich in large timber trees (33 taxa in written records) were common in the region 2000–3000 years ago, millennial-long logging has resulted in considerable range contractions and at least 39 threatened species. The wide distribution of C. ultima, which likely favoured open or semi-open habitats (like extant spotted hyenas), suggests the existence of mosaic open and closed vegetation in the Late Pleistocene across EMC, in line with a few pollen-based vegetation reconstructions and potentially, or at least partially, reflecting herbivory by herbivorous megafauna. The widespread loss of megaherbivores may have strongly compromised seed dispersal for both megafruit (fleshy fruits with widths ≥40 mm) and non-megafruit plant species in EMC, especially in terms of extra-long-distance (>10 km) dispersal, which is critical for plant species that rely on effective biotic agents to track rapid climate change. The former occurrence of large mammals and trees have translated into rich material and non-material heritages passed down across generations. Several reintroduction projects have been implemented or are under consideration, with the case of Elaphurus davidianus a notable success in recovering wild populations in the middle reaches of the Yangtze River, although trophic interactions with native carnivorous megafauna have not yet been restored. Lessons of dealing with human–wildlife conflicts are key to public support for maintaining landscapes shared with megafauna and large herbivores in the human-dominated Anthropocene. Meanwhile, potential human–wildlife conflicts, e.g. public health risks, need to be scientifically informed and effectively reduced. The Chinese government's strong commitment to improved policies of ecological protection and restoration (e.g. ecological redlines and national parks) provides a solid foundation for a scaling-up contribution to the global scope needed for solving the crisis of biotic downsizing and ecosystem degradation.     

Ecological and evolutionary legacy of megafauna extinctions

For hundreds of millions of years, large vertebrates (megafauna) have inhabited most of the ecosystems on our planet. During the late Quaternary, notably during the Late Pleistocene and the early Holocene, Earth experienced a rapid extinction of large, terrestrial vertebrates. While much attention has been paid to understanding the causes of this massive megafauna extinction, less attention has been given to understanding the impacts of loss of megafauna on other organisms with whom they interacted. In this review, we discuss how the loss of megafauna disrupted and reshaped ecological interactions, and explore the ecological consequences of the ongoing decline of large vertebrates. Numerous late Quaternary extinct species of predators, parasites, commensals and mutualistic partners were associated with megafauna and were probably lost due to their strict dependence upon them (co‐extinctions). Moreover, many extant species have megafauna‐adapted traits that provided evolutionary benefits under past megafauna‐rich conditions, but are now of no or limited use (anachronisms). Morphological evolution and behavioural changes allowed some of these species partially to overcome the absence of megafauna. Although the extinction of megafauna led to a number of co‐extinction events, several species that likely co‐evolved with megafauna established new interactions with humans and their domestic animals. Species that were highly specialized in interactions with megafauna, such as large predators, specialized parasites, and large commensalists (e.g. scavengers, dung beetles), and could not adapt to new hosts or prey were more likely to die out. Partners that were less megafauna dependent persisted because of behavioural plasticity or by shifting their dependency to humans via domestication, facilitation or pathogen spill‐over, or through interactions with domestic megafauna. We argue that the ongoing extinction of the extant megafauna in the Anthropocene will catalyse another wave of co‐extinctions due to the enormous diversity of key ecological interactions and functional roles provided by the megafauna.     

The impact of the megafauna extinctions on savanna woody cover in South America

Has land surface cover in South America been impacted by the loss of most large herbivores following the severe Pleistocene and Early Holocene megafauna extinctions on this continent? Here, we estimate how mean savanna woody biomass may have changed in the Americas following these extinctions by creating an empirical model to understand how large herbivores impact savanna woody biomass. To create this empirical model, we combine a large recently published dataset of savanna woody cover from Lehmann et al. (2014) (n = 2154 plots) with estimates of mammals ranges and weights from the IUCN database. We evaluate how variables such as number of megaherbivores (mammal species ≥ 1000 kg), log10 sum species weights, and total number of mammal species predict changes to woody cover by using both ordinary least squares regression analysis (OLS) and simultaneous auto‐regressive (SAR) analysis to control for spatial autocorrelation. Both number of megaherbivores and log10 sum species weights, which both disproportionately weight for megaherbivores, significantly explained much (～ 5–13%) variance in woody cover, but the third variable weighting all animals equally, did not. We then combined these biotic variables with abiotic variables such as temperature, precipitation, and fire frequency to create a model predicting 36% of the variance of savanna woody cover. We used this model combined with estimated range maps of extinct South American megafauna to estimate that had those South American megafauna not gone extinct, total savanna woody cover in South America could possibly have decreased by ～ 29% and that savannas would likely have been more open like current African savannas.     

Contrasts in the large herbivore faunas of the southern continents in the late Pleistocene and the ecological implications for human origins

Aim

Africa is renowned for the current abundance and diversity of its large mammals. The aim of this study was to assess distinctions evident in the functional composition of continental large herbivore faunas during the late Pleistocene before extinctions depleted the megafauna outside Africa.

Location

The African large herbivore fauna was compared with that formerly inhabiting South America, Australia, North America, Eurasia and tropical Asia during the late Pleistocene.

Methods

Pleistocene faunas were reconstructed from the literature in terms of their relative body size composition, grazer/browser contributions and taxonomic representations, omitting forest and island species.

Results

Although the three southern continents were closely similar in the overall species richness of large herbivores that they supported during the late Pleistocene, South America had a predominance of very large herbivores, while most of Australia's mammalian herbivores were relatively small and those of Africa were intermediate. Africa had many more grazers, especially in the size range 100–1000 kg, than other continents. The South American pattern resembled that in North America and Eurasia, while Africa and Australia diverged in different ways.

Main conclusions

Neither the total extent of savannas in each continent nor the morphological features enabling bovid radiation seemed adequate on their own to explain the greater richness of macrograzers in Africa. Africa is characterized by the widespread occurrence of arid/eutrophic savannas, which are unrepresented in other continents. The prevalence of savanna is partly attributable to the high elevation of interior eastern and southern Africa, associated with relatively low rainfall, and to the comparatively high soil fertility, related to volcanic influences. This promoted an abundance and diversity of medium‐sized grazing ruminants unrivalled elsewhere. Indigenous grasses in South America and Australia are less well adapted to withstand severe grazing than the African grasses introduced to support livestock. The locally high abundance of African ungulates presented conditions that facilitated the adaptive transition by early hominins from plant‐gatherers to meat‐scavengers.     

Historical stocking data and 19th century DNA reveal human‐induced changes to native diversity and distribution of cutthroat trout

Many species are threatened with extinction and efforts are underway worldwide to restore imperilled species to their native ranges. Restoration requires knowledge of species' historical diversity and distribution. For some species, many populations were extirpated or individuals moved beyond their native range before native diversity and distribution were documented, resulting in a lack of accurate information for establishing restoration goals. Moreover, traditional taxonomic assessments often failed to accurately capture phylogenetic diversity. We illustrate a general approach for estimating regional native diversity and distribution for cutthroat trout in the Southern Rocky Mountains. We assembled a large archive of historical records documenting human‐mediated change in the distribution of cutthroat trout (Oncorhynchus clarkii) and combined these data with phylogenetic analysis of 19th century samples from museums collected prior to trout stocking activities and contemporary DNA samples. Our study of the trout in the Southern Rocky Mountains uncovered six divergent lineages, two of which went extinct, probably in the early 20th century. A third lineage, previously declared extinct, was discovered surviving in a single stream outside of its native range. Comparison of the historical and modern distributions with stocking records revealed that the current distribution of trout largely reflects intensive stocking early in the late 19th and early 20th century from two phylogenetically and geographically distinct sources. Our documentation of recent extinctions, undescribed lineages, errors in taxonomy and dramatic range changes induced by human movement of fish underscores the importance of the historical record when developing and implementing conservation plans for threatened and endangered species.     

Quaternary record of aridity and mean annual precipitation based on δ15N in ratite and dromornithid eggshells from Lake Eyre, Australia

The cause(s) of the late Pleistocene megafauna extinction on the Australian continent remains largely unresolved. Unraveling climatic forcing mechanisms from direct or indirect human agents of ecosystem alteration has proven to be extremely difficult in Australia due to the lack of (1) well-dated vertebrate fossils and (2) paleo-environmental and -ecological records spanning the past approximately 100 ka when regional climatic conditions are known to have significantly varied. We have examined the nitrogen isotope composition (δ(15)N) of modern emu (Dromaius novaehollandiae) eggshells collected along a precipitation gradient in Australia, along with modern climatological data and dietary δ(15)N values. We then used modern patterns to interpret an approximately 130-ka record of δ(15)N values in extant Dromaius and extinct Genyornis newtoni eggshells from Lake Eyre to obtain a novel mean annual precipitation (MAP) record for central Australia spanning the extinction interval. Our data also provide the first detailed information on the trophic ecology and environmental preferences of two closely related taxa, one extant and one extinct. Dromaius eggshell δ(15)N values show a significant shift to higher values during the Last Glacial Maximum and Holocene, which we interpret to indicate more frequent arid conditions (<200 mm MAP), relative to δ(15)N from samples just prior to the megafauna extinction. Genyornis eggshells had δ(15)N values reflecting wetter nesting conditions overall relative to those of coeval Dromaius, perhaps indicating that Genyornis was more reliant on mesic conditions. Lastly, the Dromaius eggshell record shows a significant decrease in δ(13)C values prior to the extinction, whereas the Genyornis record does not. Neither species showed a concomitant change in δ(15)N prior to the extinction, which suggests that a significant change in vegetation surrounding Lake Eyre occurred prior to an increase in local aridity.     

Seed dispersal kernel of the largest surviving megaherbivore—the African savanna elephant

Owing to the late Pleistocene extinctions, the megafauna of Europe, Australia and the Americas disappeared, and with them the dispersal service they offered megafaunal fruit. The African savanna elephant, the largest remaining megaherbivore, offers valuable insights into the seed dispersal services provided by extinct megafauna in prehistoric times. Elephant seed dispersal studies have for the most part concentrated on African and Asian forest elephants. African savanna elephants are morphologically distinct from their forest counterparts. Like the forest elephants they consume large quantities of fruit from a large number of tree species. Despite this little is known of the savanna trees that rely on elephants for their dispersal or the spatial scale at which these seeds are dispersed. We combined information from feeding trials conducted on four park elephants with field telemetry data from 38 collared elephants collected over an 8‐year period in APNR/Kruger National Park to assess the seed dispersal service provided by savanna elephants. This study provides the first detailed account of the spatial scale at which African savanna elephants disperse seeds. Our mechanistic model predicts that 50 percent of seeds are carried over 2.5 km, and distances up to 65 km are achievable in maximum gut passage time. These findings suggest the savanna elephant as the longest distance terrestrial vertebrate disperser yet investigated. Maintaining their ecological role as a seed disperser may prove a significant factor in the conservation of large‐fruited tree diversity within the savannas. These results suggest that extinct megafauna offered a functionally unique dispersal service to megafaunal fruit.     

Do human activities raise species richness? Contrasting patterns in United States plants and fishes

In a given area, human activities usually cause the extinction of native species and the establishment of non‐native species. A key conservation issue is whether non‐native establishment tends to outpace native species extinction to produce a net gain in species richness. To determine this, empirical data must be accumulated at various scales. I show that, within the United States, the number of established non‐native plant species per state does tend to outpace the number of extinct and threatened species per state. The net gain in plant species is strongly and positively correlated with human population density. Continuation of this trend predicts substantial gains in net plant species richness for all states in the United States as human population grows. This contrasts with freshwater fishes, where most states show a net loss of species diversity as extinct and threatened species exceed established non‐native species. Changes in fish diversity do not correlate strongly with human population or non‐native species but are largely driven by the decline of native fish species.     

Megafaunal extinctions and the disappearance of a specialized wolf ecomorph

The gray wolf (Canis lupus) is one of the few large predators to survive the Late Pleistocene megafaunal extinctions [1]. Nevertheless, wolves disappeared from northern North America in the Late Pleistocene, suggesting they were affected by factors that eliminated other species. Using skeletal material collected from Pleistocene permafrost deposits of eastern Beringia, we present a comprehensive analysis of an extinct vertebrate by exploring genetic (mtDNA), morphologic, and isotopic (delta(13)C, delta(15)N) data to reveal the evolutionary relationships, as well as diet and feeding behavior, of ancient wolves. Remarkably, the Late Pleistocene wolves are genetically unique and morphologically distinct. None of the 16 mtDNA haplotypes recovered from a sample of 20 Pleistocene eastern-Beringian wolves was shared with any modern wolf, and instead they appear most closely related to Late Pleistocene wolves of Eurasia. Moreover, skull shape, tooth wear, and isotopic data suggest that eastern-Beringian wolves were specialized hunters and scavengers of extinct megafauna. Thus, a previously unrecognized, uniquely adapted, and genetically distinct wolf ecomorph suffered extinction in the Late Pleistocene, along with other megafauna. Consequently, the survival of the species in North America depended on the presence of more generalized forms elsewhere.     

Learning from the past to prepare for the future: felids face continued threat from declining prey

Many contemporary species of large‐felids (≥ 15 kg) feed upon prey that are endangered, raising concern that prey population declines (defaunation) will further threaten felids. We assess the threat that defaunation presents by investigating a late Quaternary (LQ), ‘present‐natural’ counterfactual scenario. Our present‐natural counterfactual is based on predicted ranges of mammals today in the absence of any impacts of modern humans Homo sapiens through time. Data from our present‐natural counterfactual are used to understand firstly how megafauna extinction has impacted felid communities to date and secondly to quantify the threat to large‐felid communities posed by further declines in prey richness in the future. Our purpose is to identify imminent risks to biodiversity conservation and their cascading consequences and, specifically, to indicate the importance of preserving prey diversity. We pursue two lines of enquiry; first, we test whether the loss of prey species richness is a potential cause of large‐felid extinction and range loss. Second, we explore what can be learnt from the large‐scale large‐mammal LQ losses, particularly in the Americas and Europe, to assess the threat any further decline in prey species presents to large‐felids today, particularly in Africa and Asia. Large‐felid species richness was considerably greater under our present‐natural counterfactual scenario compared to the current reality. In total, 86% of cells recorded at least one additional felid species in our present‐natural counterfactual, and up to 4–5 more large‐felid species in 10% of the cells. A significant positive correlation was recorded between the number of prey species lost and the number of large‐felid species lost from a cell. Extant felids most at risk include lion and Sunda clouded leopard, as well as leopard and cheetah in parts of their range. Our results draw attention to the continuation of a trend of megafauna decline that began with the emergence of hominins in the Pleistocene.     

Megafauna extinction,tree species range reduction,and carbon storage in Amazonian forests

During the Late Pleistocene and early Holocene 59 species of South American megafauna went extinct. Their extinction potentially triggered population declines of large‐seeded tree species dispersed by the large‐bodied frugivores with which they co‐evolved, a theory first proposed by Janzen and Martin (1982). We tested this hypothesis using species range maps for 257 South American tree species, comparing 63 species thought to be primarily distributed by megafauna with 194 distributed by other animals. We found a highly significant (p < 0.001) decreased mean range size of 26% for the megafauna dispersed fruit (n = 63 species) versus fruit dispersed by other animals (n = 194), results which support the hypothesis. We then developed a mathematical model of seed dispersal to estimate the theoretical impact of megafauna extinction on tree species range and found the estimated dispersal capacity (Φ_seed) of a 2 g seed decreases by > 95% following disperser extinction. A numerical gap dynamic simulations suggests that over a 10 000 yr period following the disperser extinctions, the average convex hull range size of large‐seeded tree species decreased by ～ 31%, while the estimated decrease in population size was ～ 54%, indicating a likely greater decrease in species population size than indicated by the empirical range patterns. Finally, we found a positive correlation between seed size and wood density of animal‐dispersed tree species implying that the Late Pleistocene and early Holocene megafaunal extinctions reduced carbon content in the Amazon by ～ 1.5 ± 0.7%. In conclusion, we 1) provide some empirical evidence that megafauna distributed fruit species have a smaller mean range size than wind, water or other animal‐dispersed species, 2) demonstrate mathematically that such range reductions are expected from megafauna extinctions ca 12 000 yr ago, and 3) illustrate that these extinctions may have reduced the Amazon's carbon storage capacity.     

Environmental correlates of the Late Quaternary regional extinctions of large and small Palaearctic mammals

Most studies of mammal extinctions during the Pleistocene–Holocene transition explore the relative effects of climate change vs human impacts on these extinctions, but the relative importance of the different environmental factors involved remains poorly understood. Moreover, these studies are strongly biased towards megafauna, which may have been more influenced by human hunting than species of small body size. We examined the potential environmental causes of Pleistocene–Holocene mammal extinctions by linking regional environmental characteristics with the regional extinction rates of large and small mammals in 14 Palaearctic regions. We found that regional extinction rates were larger for megafauna, but extinction patterns across regions were similar for both size groups, emphasizing the importance of environmental change as an extinction factor as opposed to hunting. Still, the bias towards megafauna extinctions was larger in southern Europe and smaller in central Eurasia. The loss of suitable habitats, low macroclimatic heterogeneity within regions and an increase in precipitation were identified as the strongest predictors of regional extinction rates. Suitable habitats for many species of the Last Glacial fauna were grassland and desert, but not tundra or forest. The low‐extinction regions identified in central Eurasia are characterized by the continuous presence of grasslands and deserts until the present. In contrast, forest expansion associated with an increase in precipitation and temperature was likely the main factor causing habitat loss in the high‐extinction regions. The shift of grassland into tundra also contributed to the loss of suitable habitats in northern Eurasia. Habitat loss was more strongly related to the extinctions of megafauna than of small mammals. Ungulate species with low tolerance to deep snow were more likely to go regionally extinct. Thus, the increase in precipitation at the Pleistocene–Holocene transition may have also directly contributed to the extinctions by creating deep snow cover which decreases forage availability in winter.     

Conservation genetics of the endangered Iberian steppe plant Ferula loscosii (Apiaceae)

Ferula loscosii (Lange) Willk (Apiaceae) is a threatened endemic species native to the Iberian Peninsula. The plant has a narrow and disjunct distribution in three regions, NE, C and SE Spain. Genetic variability within and among 11 populations from its natural distribution was assessed using allozymes. Intermediate levels of genetic diversity were detected in F. loscosii (P(99%) = 36.83; H(E) = 0.125; H(T) = 0.152). However, the highest genetic diversity (58%) corresponded to the threatened populations from SE and C Spain (H(T) = 0.169) rather than the more abundant and larger populations from NE Spain (Ebro valley) (H(T) = 0.122). Low to moderate levels of genetic structure were found among regional ranges (G(ST) = 0.134), and several statistical spatial correlation analyses corroborated substantial genetic differentiation among the three main regional ranges. However, no significant genetic differentiation was found among the NE Spain populations, except for a northernmost population that is geographically isolated. Outcrossing mating and other biological traits of the species could account for the maintenance of the present values of genetic diversity within populations. The existence of an ancestral late Tertiary wider distribution of the species in SE and C Spain, followed by the maintenance of different Quaternary refugia in these warmer areas, together with a more recent and rapid post-glacial expansion towards NE Spain, are arguments that could explain the low genetic variability and structure found in the Ebro valley and the higher levels of diversity in the southern Iberian populations.     

Ecologically heterogeneous populations of the invasive ant Wasmannia auropunctata within its native and introduced ranges

Abstract 1. The biology of most invasive species in their native geographical areas remains largely unknown. Such studies are, however, crucial in shedding light on the ecological and evolutionary processes underlying biological invasions. 2. The present study focuses on the little fire ant Wasmannia auropunctata, a species native to Central and South America that has been widely introduced and which has become invasive throughout the tropics. We characterise and compare several ecological traits of native populations in French Guiana with those in one of its introduced ranges, New Caledonia. 3. We found ecologically heterogeneous populations of W. auropunctata coexisting in the species’ native geographical area. First, we found populations restricted to naturally perturbed areas (particularly floodplains) within the primary forest, and absent from the surrounding forest areas. These populations were characterised by low nest and worker densities. Second, we found dominant populations in recent anthropogenic areas (e.g. secondary forest or forest edge along road) characterised by high nest and worker densities, and associated with low ant species richness. The local dominance of W. auropunctata in such areas can be due to the displacement of other species (cause) or the filling‐up of empty habitats unsuitable to other ants (effect). With respect to their demographic features and ant species richness, the populations of native anthropogenic habitats were to a large extent similar to the invasive populations introduced into remote areas. 4. The results point to the need for greater research efforts to better understand the ecological and demographic features of invasive species within their native ranges.