Holocene mammal extinctions in the Carpathian Basin: a review

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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.     

A Peculiar New Pampatheriidae (Mammalia: Xenarthra: Cingulata) from the Pleistocene of Argentina and Comments on Pampatheriidae Diversity

Pampatheriidae are a group of cingulates native to South American that are known from the middle Miocene to the lower Holocene. Two genera have been recognized between the lower Pleistocene and the lower Holocene: Pampatherium Gervais and Ameghino (Ensenadan, Bonaerian and Lujanian, lower Pleistocene–lower Holocene) and Holmesina Simpson (Blancan, Irvingtonian, upper Pliocene–lower Holocene). They have been mainly differentiated by their osteoderm morphology and cranio-dental characters. These taxa had a wide latitudinal distribution, extending from the southern part of South America (Península Valdés, Argentina) to North America (Florida, USA). In this contribution, we describe a new genus and species of Pampatheriidae for the lower and middle Pleistocene of Buenos Aires Province and for the upper Pleistocene of Santa Fe Province (Argentina).The new taxon is represented by disarticulated osteoderms, one skull element, two thoracic vertebrae and a right femur and patella. It has extremely complex osteoderm ornamentations and particular morphological characters of the cranial element and femur that are not found in any other species of the family. This new taxon, recorded in the lower–middle Pleistocene (Ensenadan Stage/Age) and in the upper Pleistocene–early Holocene (Lujanian Stage/Age), is incorporated to the Pleistocene mammal assemblage of South America. Finally, the Pampatheriidae diversity is greater during the Lujanian Stage/Age than the Ensenadan Stage/Age.     

The Impact of 850,000 Years of Climate Changes on the Structure and Dynamics of Mammal Food Webs

Most evidence of climate change impacts on food webs comes from modern studies and little is known about how ancient food webs have responded to climate changes in the past. Here, we integrate fossil evidence from 71 fossil sites, body-size relationships and actualism to reconstruct food webs for six large mammal communities that inhabited the Iberian Peninsula at different times during the Quaternary. We quantify the long-term dynamics of these food webs and study how their structure changed across the Quaternary, a period for which fossil data and climate changes are well known. Extinction, immigration and turnover rates were correlated with climate changes in the last 850 kyr. Yet, we find differences in the dynamics and structural properties of Pleistocene versus Holocene mammal communities that are not associated with glacial-interglacial cycles. Although all Quaternary mammal food webs were highly nested and robust to secondary extinctions, general food web properties changed in the Holocene. These results highlight the ability of communities to re-organize with the arrival of phylogenetically similar species without major structural changes, and the impact of climate change and super-generalist species (humans) on Iberian Holocene mammal communities.     

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.     

The ghosts of mammals past: biological and geographical patterns of global mammalian extinction across the Holocene

Although the recent historical period is usually treated as a temporal base-line for understanding patterns of mammal extinction, mammalian biodiversity loss has also taken place throughout the Late Quaternary. We explore the spatial, taxonomic and phylogenetic patterns of 241 mammal species extinctions known to have occurred during the Holocene up to the present day. To assess whether our understanding of mammalian threat processes has been affected by excluding these taxa, we incorporate extinct species data into analyses of the impact of body mass on extinction risk. We find that Holocene extinctions have been phylogenetically and spatially concentrated in specific taxa and geographical regions, which are often not congruent with those disproportionately at risk today. Large-bodied mammals have also been more extinction-prone in most geographical regions across the Holocene. Our data support the extinction filter hypothesis, whereby regional faunas from which susceptible species have already become extinct now appear less threatened; they may also suggest that different processes are responsible for driving past and present extinctions. We also find overall incompleteness and inter-regional biases in extinction data from the recent fossil record. Although direct use of fossil data in future projections of extinction risk is therefore not straightforward, insights into extinction processes from the Holocene record are still useful in understanding mammalian threat.     

Late quaternary turnover of mammals in Borneo: the zooarchaeological record

The Quaternary has been a period of repeated, oscillating patterns of climate change. Global fluctuations in sea level affected the island status of Borneo, which was probably joined to continental Asia for more than half of the last 250,000 years. Alternating connection and isolation, coupled with the ecological barrier of a savanna corridor running from the Malay Peninsula to Java during periods of marine recession, are reflected in the present mammal fauna of Borneo. 38% of mammal species (excluding bats) are endemic, and some distinctive species or subspecies are confined to the north of the island. No known sites in Borneo match the Early and Middle Pleistocene regional sources in eastern Java. However, caves at Niah, Sireh and Jambusan, Sarawak, and Madai, Sabah, provide a zooarchaeological record covering the past 50,000 years. The Late Pleistocene mammals of Borneo included ten species also present among a Javan Middle Pleistocene savanna-adapted assemblage. Of these, four are categorised as ‘megafuana’: a giant pangolin, Javan rhinoceros, Malay tapir and tiger; the Sumatran rhinoceros can be added. In addition, there are less secure Pleistocene records of Asian elephant from Sarawak and Brunei. Holocene canid remains from Madai could either be the dhole or an early domestic dog. Palynological data combined with the mammal fauna confirm that around 45,000 years ago the vicinity of Niah was vegetated by closed forest. The continuous presence of a suite of arboreal specialists, including large primates, indicates that forest cover persisted through the terminal Pleistocene. Among local extinctions, the giant pangolin apparently disappeared early in this period, but tiger, Javan rhinoceros and tapir probably survived into the last millennium. Human predation of juveniles may account for the loss of the large ungulates, but the disappearance of tiger needs another explanation. Despite hunting pressure throughout the terminal Pleistocene and Holocene, a population of orangutan survived at Niah until perhaps the last millennium. Size diminution observed among large, medium and small mammal species is interpreted as the selective impact of environmental change. Once more is known about their ecology, changes in the bat fauna of Niah cave may provide indicators of environmental impacts affecting the wider mammal community during the later Holocene. In conclusion, it is recommended that the three nations, Brunei Darussalam, Malaysia and Indonesia, should support the WWF sponsored ‘Heart of Borneo’ as the most hopeful project to provide sustainable management of the rare and threatened forest-adapted wild mammals of the island.     

Ancient DNA suggests modern wolves trace their origin to a Late Pleistocene expansion from Beringia

Grey wolves (Canis lupus) are one of the few large terrestrial carnivores that have maintained a wide geographical distribution across the Northern Hemisphere throughout the Pleistocene and Holocene. Recent genetic studies have suggested that, despite this continuous presence, major demographic changes occurred in wolf populations between the Late Pleistocene and early Holocene, and that extant wolves trace their ancestry to a single Late Pleistocene population. Both the geographical origin of this ancestral population and how it became widespread remain unknown. Here, we used a spatially and temporally explicit modelling framework to analyse a data set of 90 modern and 45 ancient mitochondrial wolf genomes from across the Northern Hemisphere, spanning the last 50,000 years. Our results suggest that contemporary wolf populations trace their ancestry to an expansion from Beringia at the end of the Last Glacial Maximum, and that this process was most likely driven by Late Pleistocene ecological fluctuations that occurred across the Northern Hemisphere. This study provides direct ancient genetic evidence that long‐range migration has played an important role in the population history of a large carnivore, and provides insight into how wolves survived the wave of megafaunal extinctions at the end of the last glaciation. Moreover, because Late Pleistocene grey wolves were the likely source from which all modern dogs trace their origins, the demographic history described in this study has fundamental implications for understanding the geographical origin of the dog.     

Synchronous genetic turnovers across Western Eurasia in Late Pleistocene collared lemmings

Recent palaeogenetic studies indicate a highly dynamic history in collared lemmings (Dicrostonyx spp.), with several demographical changes linked to climatic fluctuations that took place during the last glaciation. At the western range margin of D. torquatus, these changes were characterized by a series of local extinctions and recolonizations. However, it is unclear whether this pattern represents a local phenomenon, possibly driven by ecological edge effects, or a global phenomenon that took place across large geographical scales. To address this, we explored the palaeogenetic history of the collared lemming using a next‐generation sequencing approach for pooled mitochondrial DNA amplicons. Sequences were obtained from over 300 fossil remains sampled across Eurasia and two sites in North America. We identified five mitochondrial lineages of D. torquatus that succeeded each other through time across Europe and western Russia, indicating a history of repeated population extinctions and recolonizations, most likely from eastern Russia, during the last 50 000 years. The observation of repeated extinctions across such a vast geographical range indicates large‐scale changes in the steppe‐tundra environment in western Eurasia during the last glaciation. All Holocene samples, from across the species' entire range, belonged to only one of the five mitochondrial lineages. Thus, extant D. torquatus populations only harbour a small fraction of the total genetic diversity that existed across different stages of the Late Pleistocene. In North American samples, haplotypes belonging to both D. groenlandicus and D. richardsoni were recovered from a Late Pleistocene site in south‐western Canada. This suggests that D. groenlandicus had a more southern and D. richardsoni a more northern glacial distribution than previously thought. This study provides significant insights into the population dynamics of a small mammal at a large geographical scale and reveals a rather complex demographical history, which could have had bottom‐up effects in the Late Pleistocene steppe‐tundra ecosystem.     

Unraveling the consequences of the terminal Pleistocene megafauna extinction on mammal community assembly

Recent studies connecting the decline of large predators and consumers with the disintegration of ecosystems often overlook that this natural experiment already occurred. As recently as 14 ka, tens of millions of large‐bodied mammals were widespread across the American continents. Within 1000 yr of the arrival of humans, ～ 80% were extinct including all > 600 kg. While the cause of the late Pleistocene (LP) extinction remains contentious, largely overlooked are the ecological consequences of the loss of millions of large‐bodied animals. Here, we examine the influence of the LP extinction on a local mammal community. Our study site is Hall's Cave in the Great Plains of Texas, which has unparalleled fine‐grained temporal resolution over the past 20 ka, allowing characterization of the community before and after the extinction. In step with continental patterns, this community lost 80% of large‐bodied herbivores and 20% of apex predators at the LP extinction. Using tightly constrained temporal windows spanning full glacial to modern time periods and comprehensive faunal lists, we reconstruct mammal associations and body size distributions over time. We find changes in alpha and beta diversity, and in the statistical moments associated with periods of climate change as well as with the LP extinction event. Additionally, there is a fundamental change in the composition of herbivores, with grazers being replaced by frugivores/granivores starting about 15 ka; the only large‐bodied grazer remaining today is the bison Bison bison. Moreover, the null model program PAIRS reveals interesting temporal patterns in the disassociation or co‐occurrence of species through the terminal Pleistocene and Holocene. Extinct species formed more significant associations than modern ones, and formed more aggregated pairs than do modern species. Further, negative species associations were about three times stronger than positive ones, suggesting that competitive interactions or environmental filtering are a strong force in community structure.     

Body mass‐related changes in mammal community assembly patterns during the late Quaternary of North America

The late Quaternary of North America was marked by prominent ecological changes, including the end‐Pleistocene megafaunal extinction, the spread of human settlements and the rise of agriculture. Here we examine the mechanistic reasons for temporal changes in mammal species association and body size during this time period. Building upon the co‐occurrence results from Lyons et al. (2016) – wherein each species pair was classified as spatially aggregated, segregated or random – we examined body mass differences (BMD) between each species pair for each association type and time period (Late Pleistocene: 40 000 ¹⁴C–11 700 ¹⁴C ybp, Holocene: 11 700 ¹⁴C–50 ybp and Modern: 50–0 yr). In the Late Pleistocene and Holocene, the BMD of both aggregated and segregated species pairs was significantly smaller than the BMD of random pairs. These results are consistent with environmental filtering and competition as important drivers of community structure in both time periods. Modern assemblages showed a breakdown between BMD and co‐occurrence patterns: the average BMD of aggregated, segregated and random species pairs did not differ from each other. Collectively, these results indicate that the late Quaternary mammalian extinctions not only eliminated many large‐bodied species but were followed by a re‐organization of communities that altered patterns of species coexistence and associated differences in body size.     

Millennial-scale faunal record reveals differential resilience of European large mammals to human impacts across the Holocene

The use of short-term indicators for understanding patterns and processes of biodiversity loss can mask longer-term faunal responses to human pressures. We use an extensive database of approximately 18 700 mammalian zooarchaeological records for the last 11 700 years across Europe to reconstruct spatio-temporal dynamics of Holocene range change for 15 large-bodied mammal species. European mammals experienced protracted, non-congruent range losses, with significant declines starting in some species approximately 3000 years ago and continuing to the present, and with the timing, duration and magnitude of declines varying individually between species. Some European mammals became globally extinct during the Holocene, whereas others experienced limited or no significant range change. These findings demonstrate the relatively early onset of prehistoric human impacts on postglacial biodiversity, and mirror species-specific patterns of mammalian extinction during the Late Pleistocene. Herbivores experienced significantly greater declines than carnivores, revealing an important historical extinction filter that informs our understanding of relative resilience and vulnerability to human pressures for different taxa. We highlight the importance of large-scale, long-term datasets for understanding complex protracted extinction processes, although the dynamic pattern of progressive faunal depletion of European mammal assemblages across the Holocene challenges easy identification of ‘static’ past baselines to inform current-day environmental management and restoration.     

Phylogeography and demographic history of Shaw's Jird (Meriones shawii complex) in North Africa

Palaeoenvironmental data and climatic reconstructions show that the Mediterranean ecoregion of North Africa underwent drastic ecological changes during the Pleistocene. Given its rich palaeontological record, North Africa is a pertinent region for documenting the role of climate change and human mediated‐habitat changes on the demography and genetic structure of faunal species. In the present study, we collected data from this species in Morocco, Algeria, and Tunisia, and we combined molecular (mitochondrial and nuclear DNA sequences, microsatellites), fossil, palaeoenvironmental, and human context data to propose an explanation for the fluctuations of populations belonging to the Meriones shawii complex in the past. Genetic and fossil data both indicate a strong bottleneck in Moroccan populations at the Middle Holocene (last interglacial optimum) compared to the Late Pleistocene. Our mitochondrial DNA data suggest a diversification event within Morocco corresponding to the 130–125 kya interglacial optimum. Given that (1) major demographic changes in the M. shawii complex coincide with the interglacial optimums, and (2) the impact of human activities on the landscape and faunal communities was moderate during the Middle Holocene (beginnings of the Neolithic culture), our results demonstrate that climate, rather than anthropogenic influences, likely explains the M. shawii complex population decline in the Holocene.     

The Liang Bua faunal remains: a 95 k.yr. sequence from Flores, East Indonesia

Excavations at Liang Bua, a limestone cave on the island of Flores, East Indonesia, have yielded a well-dated archaeological and faunal sequence spanning the last 95 k.yr., major climatic fluctuations, and two human species - H. floresiensis from 95 to 17 k.yr.¹, and modern humans from 11 k.yr. to the present. The faunal assemblage comprises well-preserved mammal, bird, reptile and mollusc remains, including examples of island gigantism in small mammals and the dwarfing of large taxa. Together with evidence from Early-Middle Pleistocene sites in the Soa Basin, it confirms the long-term isolation, impoverishment, and phylogenetic continuity of the Flores faunal community. The accumulation of Stegodon and Komodo dragon remains at the site in the Pleistocene is attributed to Homo floresiensis, while predatory birds, including an extinct species of owl, were largely responsible for the accumulation of the small vertebrates. The disappearance from the sequence of the two large-bodied, endemic mammals, Stegodon florensis insularis and Homo floresiensis, was associated with a volcanic eruption at 17 ka and precedes the earliest evidence for modern humans, who initiated use of mollusc and shell working, and began to introduce a range of exotic animals to the island. Faunal introductions during the Holocene included the Sulawesi warty pig (Sus celebensis) at about 7 ka, followed by the Eurasian pig (Sus scrofa), Long-tailed macaque, Javanese porcupine, and Masked palm civet at about 4 ka, and cattle, deer, and horse - possibly by the Portuguese within historic times. The Holocene sequence at the site also documents local faunal extinctions - a result of accelerating human population growth, habitat loss, and over-exploitation.     

Faunal relationships and zoogeographical affinities of mammals in north-west Africa

The nonmarine mammal fauna of the Maghreb region of north‐west Africa is related to that from three potential source areas: the northern Palaearctic (Europe and south‐west Asia; here referred to as the European fauna), subsaharan Africa (the African fauna) and the arid Palaearctic (Sahara Desert: the desert fauna). On the basis of geographical distribution patterns, this fauna divisible into two groups: the bats, whose affinities are most closely related to southern Europe and south‐west Asia, and nonflying species, most closely related to subsaharan Africa but with an appreciable northern Palaearctic element. These affinities are even more pronounced if desert fauna are removed from the analysis. The nonflying European fauna probably colonized via south‐west Asia and north Africa, rather than direct from western Europe. The results demonstrate that terrestrial habitat barriers are less of an impediment to dispersal, for all mammals except bats, than even narrow stretches of water. The fauna of the Maghreb may be undergoing faunal relaxation, following immigration from tropical Africa and south‐west Asia during mesic phases in the Late Pleistocene and early Holocene.     

Holocene vegetation change and the mammal faunas of South America and Africa

Aim Although sharing many similarities in their vegetation types, South America and Africa harbour very dissimilar recent mammal faunas, not only taxonomically but also in terms of several faunistic patterns. However late Pleistocene and mid‐Holocene faunas, albeit taxonomically distinct, presented many convergent attributes. Here we propose that the effects of the Holocene climatic change on vegetation physiognomy has played a crucial role in shaping the extant mammalian faunistic patterns. Location South America and Africa from the late Pleistocene to the present. Methods Data presented here have been compiled from many distinct sources, including palaeontological and neontological mammalian studies, palaeoclimatology, palynology, and publications on vegetation ecology. Data on Pleistocene, Holocene and extant mammal faunas of South America and Africa allowed us to establish a number of similar and dissimilar faunistic patterns between the two continents across time. We then considered what changes in vegetation physiognomy would have occurred under the late Pleistocene last glacial maximum (LGM) and the Holocene climatic optimum (HCO) climatic regimes. We have ordained these proposed vegetation changes along rough physiognomic seral stages according to assumptions based on current botanical research. Finally, we have associated our hypothesized vegetation changes in South America and Africa with mammalian faunistic patterns, establishing a putative causal relationship between them. Results The extant mammal faunas of South America and Africa differ widely in taxonomical composition; the number of medium and large species they possess; behavioural and ecological characteristics related to herbivore herding, migration and predation; and biogeographical patterns. All such distinctions are mostly related to the open formation faunas, and have been completely established around the mid‐Holocene. Considering that the mid‐Holocene was a time of greater humidity than the late Pleistocene, vegetation cover in South America and Africa would have been dominated by forest or closed vegetation landscapes, at least for most of their lower altitude tropical regions. We attribute the loss of larger‐sized mammal lineages in South America to the decrease of open vegetation area, and their survival in Africa to the existence of vast savannas in formerly steppic or desertic areas in subtropical Africa, north and south of the equator. Alternative explanations, mostly dealing with the disappearance of South American megamammals, are then reviewed and criticized. Main conclusions The reduction of open formation areas during the HCO in South America and Africa explains most of the present distinct faunistic patterns between the two continents. While South America would have lost most of its open formations within the 30° latitudinal belt, Africa would have kept large areas suitable to the open formation mammalian fauna in areas presently occupied by desert and semi‐arid vegetation. Thus, the same general climatic events that affected South America in the late Pleistocene and Holocene also affected Africa, leading to our present day faunistic dissimilarities by maintaining the African mammalian communities almost unchanged while dramatically altering those of South America.     

Faunal histories from Holocene ancient DNA

Recent studies using ancient DNA have been instrumental in advancing understanding of the impact of Holocene climate change on biodiversity. Ancient DNA has been used to track demography, migration and diversity, and is providing new insights into the long-term dynamics of species and population distributions. The Holocene is key to understanding how the past has impacted on the present, as it bridges the gap between contemporary phylogeographic studies and those with inference on Pleistocene patterns, based on ancient DNA studies. Here, we examine the major patterns of Holocene faunal population dynamics and connectivity; highlighting the dynamic nature of species and population responses to Holocene climatic change, thereby providing an 'analogue' for understanding potential impacts of future change.     

Seedling dynamics of <Emphasis Type="Italic">Acer mono</Emphasis> and <Emphasis Type="Italic">Fagus crenata</Emphasis>: an environmental filter limiting their adult distributions

Spatial pattern of a biotic population may change over time as its component individuals grow or die out, but whether this is the case for desert annual plants is largely unknown. Here we examined seasonal changes in spatial patterns of two annuals, Eriogonum abertianum and Haplopappus gracilis, in initial (winter) and final (summer) densities. The density was measured as the number of individuals from 384 permanent quadrats (each 0.5 m × 0.5 m) in the Chihuahuan Desert near Portal, Arizona, USA. We used three probability distributions (binomial, Poisson, and negative binomial or NB) that represent three basic spatial patterns (regular, random, and clumped) to fit the observed frequency distributions of densities of the two annuals. Both species showed clear clumped patterns as characterized by the NB and had similar inverse J-shaped frequency distribution curves in two density categories. Also, both species displayed a reduced degree of aggregation from winter to summer after the spring drought (massive die-off), as indicated by the increased k-parameter of the NB and decreased values of another NB parameter p, variance/mean ratio, Lloyd’s Index of Patchiness, and David and Moore’s Index of Clumping. Further, we hypothesized that while the NB (i.e., Poisson-logarithmic) well fits the distribution of individuals per quadrat, its components, the Poisson and logarithmic, may describe the distributions of clumps per quadrat and of individuals per clump, respectively. We thus obtained the means and variances for (1) individuals per quadrat, (2) clumps per quadrat, and (3) individuals per clump. The results showed that the decrease of the density from winter to summer for each plant resulted from the decrease of individuals per clump, rather than from the decrease of clumps per quadrat. The great similarities between the two annuals indicate that our observed temporal changes in spatial patterns may be common among desert annual plants.     

Mass extinctions do not explain skew in interspecific body size distributions

In several higher animal taxa, such as mammals and birds, the distribution of species body sizes is heavily skewed towards small size. Previous studies have suggested that small‐bodied organisms are less prone to extinction than large‐bodied species. If small body size is favourable during mass extinction events, a post mass extinction excess of small‐bodied species may proliferate and maintain skewed body size distributions sometime after. Here, we modelled mass extinctions and found that even unrealistically strong body mass selection has little effect on the skew of interspecific body size distributions. Moreover, selection against large body size may, counter intuitively, skew size distributions towards large body size. In any case, subsequent evolutionary diversification rapidly erases these rather small effects mass extinctions may have on size distributions. Next, we used body masses of extant species and phylogenetic methods to investigate possible changes in body size distributions across the Cretaceous–Paleogene (K‐Pg) mass extinction. Body size distributions of extant clades that originated during the Cretaceous are on average more skewed than their subclades that originated during the Paleogene, but the difference is only minor in mammals, and in birds, it can be explained by a positive relationship between species richness and skewness that is also present in clades that originated after the transition. Hence, we cannot infer from extant species whether the K‐Pg mass extinctions were size‐selective, but they are not the reason why most extant bird and mammal species are small‐bodied.     

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.