Arid Recovery – A comparison of reptile and small mammal populations inside and outside a large rabbit,cat and fox‐proof exclosure in arid South Australia

Australian arid zone mammal species within the Critical Weight Range (CWR) of 35 g–5.5 kg have suffered disproportionately in the global epidemic of contemporary faunal extinctions. CWR extinctions have been attributed largely to the effects of introduced or invasive mammals; however, the impact of these threatening processes on smaller mammals and reptiles is less clear. The change in small mammal and reptile assemblages after the removal of rabbits, cats and foxes was studied over a 6‐year period in a landscape‐scale exclosure in the Australian arid zone. Rodents, particularly Notomys alexis and Pseudomys bolami, increased to 15 times higher inside the feral‐proof Arid Recovery Reserve compared with outside sites, where rabbits, cats and foxes were still present. Predation by cats was thought to exert the greatest influence on rodent numbers owing to the maintenance of the disparity in rodent responses through dry years and the differences in dietary preferences between rabbits and P. bolami. The presence of introduced Mus domesticus or medium‐sized re‐introduced mammal species did not significantly affect resident small mammal or reptile abundance. Abundance of most dasyurids and small lizards did not change significantly after the removal of feral animals although reductions in gecko populations inside the reserve may be attributable to second order trophic interactions or subtle changes in vegetation structure and cover. This study suggests that populations of rodent species in northern South Australia below the CWR may also be significantly affected by introduced cats, foxes and/or rabbits and that a taxa specific model of Australian mammal decline may be more accurate than one based on body weight.     

Holocene mammal extinctions in the Carpathian Basin: a review

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Predicting small‐mammal responses to climatic warming: autecology,geographic range,and the Holocene fossil record

Forecasting how species will respond to climatic change requires knowledge of past community dynamics. Here we use time‐series data from the small‐mammal fossil records of two caves in the Great Basin of the American West to evaluate how contrasting and variable local paleoclimates have shaped small‐mammal abundance dynamics over the last ～7500 years of climatic change. We then predict how species and communities will respond to future scenarios of increased warming and aridity coupled with continued spread of an invasive annual grass (Bromus tectorum). We find that most community‐level responses to climatic change occur in the mammalian abundance structure at both sites; the dominance of the community by individuals from species with a southern geographic affinity increases with climatic warming. This suggests that responses occurred in situ rather than by the immigration of new taxa over this time interval. Despite predictability at the community‐scale, species‐level relationships between abundance and climate are variable and are not necessarily explained by a species' geographic affinity. Species present at both sites, however, exhibit remarkably similar responses to climate at each site, indicating that species autecology (specifically dietary functional group) is important in determining response to climatic warming. Regression‐tree analyses show remarkable concordance between the two cave faunas and highlight the importance of a granivorous dietary strategy in this desert ecosystem. Under projections of increased temperature and decreased precipitation over the next 50 years, our results indicate that granivores should thrive as communities become more dominated by individuals with a southern geographic affinity. Granivores, however, are negatively impacted by the invasion of cheatgrass. The last century of anthropogenic impacts has thus placed granivores at a greater risk of extinction than predicted under climate‐only scenarios.     

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.     

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.     

Threatened mammals become more predatory after small‐scale prescribed fires in a high‐rainfall rocky savanna

Understanding mechanisms underlying fire regime effects on savanna fauna is difficult because of a wide range of possible trophic interactions and feedbacks. Yet, understanding mechanisms underlying fauna dynamics is crucial for conservation management of threatened species. Small savanna mammals in northern Australia are currently undergoing widespread declines and regional extinctions partly attributable to fire regimes. This study investigates mammal trophic and ecosystem responses to fire in order to identify possible mechanisms underlying these declines. Mammal trophic responses to fire were investigated by surveying mammal abundance, mammal diet, vegetation structure and non‐mammal fauna dynamics in savannas six times at eight sites over a period of 3 years. Known site‐specific fire history was used to test for trophic responses to post‐fire interval and fire frequency. Mammal and non‐mammal fauna showed only minor responses of post‐fire interval and no effect of fire frequency. Lack of fauna responses differed from large post‐fire vegetation responses. Dietary analysis showed that two mammal species, Dasyurus hallucatus and Isoodon auratus, increased their intake of large prey groups in recently burnt, compared to longer unburnt vegetation. This suggests a fire‐related change in trophic interactions among predators and their prey, after removal of ground‐layer vegetation. No evidence was found for other changes in food resource uptake by mammals after fire. These data provide support for a fire‐related top‐down ecosystem response among savanna mammals, rather than a bottom‐up resource limitation response. Future studies need to investigate fire responses among other predators, including introduced cats and dingoes, to determine their roles in fire‐related mammal declines in savannas of northern Australia.     

Declining populations in one of the last refuges for threatened mammal species in northern Australia

Australia has contributed a disproportionate number of the world's mammal extinctions over the past 200 years, with the greatest loss of species occurring through the continent's southern and central arid regions. Many taxonomically and ecologically similar species are now undergoing widespread decline across the northern Australian mainland, possibly driven by predation by feral cats and changed fire regimes. Here, we report marked recent declines of native mammal species in one of Australia's few remaining areas that support an intact mammal assemblage, Melville Island, the largest island off the northern Australian coast. We have previously reported a marked decline on Melville Island of the threatened brush‐tailed rabbit‐rat (Conilurus penicillatus) over the period 2000–2015, linked to predation by feral cats. We now report a 62% reduction in small mammal trap‐success and a 36% reduction in site‐level species richness over this period. There was a decrease in trap‐success of 90% for the northern brown bandicoot (Isoodon macrourus), 64% for the brush‐tailed rabbit‐rat and 63% for the black‐footed tree‐rat (Mesembriomys gouldii), but no decline for the common brushtail possum (Trichosurus vulpecula). These results suggest that populations of native mammals on Melville Island are exhibiting similar patterns of decline to those recorded in Kakadu National Park two decades earlier, and across the northern Australian mainland more generally. Without the implementation of effective management actions, these species are likely to be lost from one of their last remaining strongholds, threatening to increase Australia's already disproportionate contribution to global mammal extinctions.     

Responses of two species of heathland rodents to habitat manipulation: Vegetation density thresholds and the habitat accommodation model

The abundance of two native rodent species, Rattus lutreolus and Pseudomys gracilicaudatus, has been shown to correlate with vegetation density in coastal wet heath. Fox's habitat accommodation model relates relative abundances of such small mammal species to heathland vegetation regeneration following disturbance. Implicit in the model is recognition that it is successional changes in vegetation, not time per se, that drives the responses of small mammal species along a regeneration axis. Using a brush‐cutter we deliberately removed approximately 85% of vegetation around trapping stations and recorded significant reductions in the abundance of both P. gracilicaudatus (an earlier‐stage colonizing species) and R. lutreolus (a late seral‐stage species). A significant decrease in the abundance of only the latter had been demonstrated previously when 60–70% of the vegetation had been removed. Following the brush‐cutting both species re‐entered the mammalian secondary succession at different times, first P. gracilicaudatus followed by R. lutreolus after the vegetation cover thresholds of each species had been reached. The impact of this habitat manipulation experiment was to produce a retrogression of the small mammal succession, experimentally demonstrating causality between changes in vegetation density and subsequent small mammal habitat use.     

Passive recovery of small vertebrates following livestock removal in the Australian rangelands

In arid environments, grazing by exotic herbivores, including domestic livestock, can greatly influence native, small vertebrate assemblages. Whether the removal of livestock facilitates passive recovery of these assemblages depends on habitat condition and the species present. We explore changes in small mammal and reptile species richness, abundance, and composition in a degraded chenopod shrubland dominated by Acacia victoriae ssp. and open Acacia aneura (mulga) woodland destocked in 1976 and 1984, respectively. Data were obtained between 1997 and 2007, from two grazed and two ungrazed sites in each community. Species richness increased at a faster rate in ungrazed open A. aneura woodland, but did not differ significantly between ungrazed and grazed degraded chenopod shrubland. Subsequent analyses at a finer‐scale detected disparate responses in richness and abundance for microhabitat. At this scale, a greater number of species‐specific responses were also detected, including increased abundance of generalist species and decreased abundance of species requiring low cover. These results reiterate the potential for species‐specific responses to livestock that are more apparent in particular microhabitats. Furthermore, this investigation provides evidence for the gradual passive recovery of small mammal and reptile assemblages in both communities, which is facilitated by the removal of livestock in open A. aneura woodland in fair condition, but not degraded chenopod shrubland in poor condition.     

PLEISTOCENE EXTINCTIONS OF DEEP-SEA BENTHIC FORAMINIFERA: THE SOUTH ATLANTIC RECORD

Abstract:  Sixty-two species and 19 genera of elongate, cylindrical benthic foraminifera disappeared from the deep-sea in the south-east Atlantic (ODP Sites 1082 and 1083) and the Atlantic sector of the Southern Ocean (ODP Site 1088) during the Early and Middle Pleistocene as part of the global extinction of the families Pleurostomellidae, Stilostomellidae and portions of the Nodosariidae. During the mid-Pleistocene Climate Transition (1·2–0·6 Ma) in the Southern Ocean, these extinct taxa exhibited three pulses of glacial decline in abundance and diversity separated by partial interglacial recoveries. Beneath the high-productivity Benguela Current upwelling region (Sites 1082, 1083), glacial declines in the extinct taxa were suppressed by favourable high organic-carbon flux and consequent low-oxygen bottom conditions. Here two major pulses of diversity loss occurred at c.  1·3–1·2 Ma and 1·0–0·7 Ma. At all three locations, the most dramatic decline in abundance and diversity occurred c.  0·85–0·80 Ma (marine isotope stage 20), and the final disappearance of Extinction Group taxa was completed by 0·67 Ma beneath the Benguela Current and 0·60 Ma in the Southern Ocean. We speculate that this period of enhanced global extinctions was linked to a pulsed decline in glacial temperatures and/or increase in ventilation of deep and intermediate water masses, associated with polar ice cap growth since the late Pliocene.     

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.     

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.     

Contemporary climate change alters the pace and drivers of extinction

Contemporary climate change is expected to affect the distributions of most species, but the nature, tempo, and mechanics of contemporary range shifts are still largely speculative. Here, we use fine‐scale distributional records developed over the past Century, combined with spatially comprehensive microclimatic data, to demonstrate a dramatic shift in the range of a climate‐sensitive mammal and to infer the increasingly important role of climate in local extinctions of this species across a 38.2 million‐ha area. Changes in the distribution of the American pika (Ochotona princeps) throughout the Great Basin ecoregion were characterized using records from 1898–2008, revealing a nearly five‐fold increase in the rate of local extinction and an 11‐fold increase in the rate of upslope range retraction during the last ten years, compared with during the 20th Century. Four of ten local pika extinctions have occurred since 1999, and across this ecoregion the low‐elevation range boundary for this species is now moving upslope at an average rate of about 145 m per decade. The rapid, ecoregional range shift of this small, talus‐dwelling species stands in remarkable contrast with the dynamics and determinants of endangerment previously observed for most terrestrial species, and to earlier extinction determinants for O. princeps in this region. Further investigation of widely distributed species will clarify rates at which biotic response to environmental change is occurring, and how factors driving that change are evolving in different portions of the earth.     

Ecosystem dynamics through the past 2000 years as revealed by fossil mammals from Lamar cave in Yellowstone National Park,USA

Lamar Cave, a late Holocene paleontological site in Yellowstone National Park, USA, has yielded 36 fossil mammal species from 10 stratigraphic units representing about 2000 years. The fossil fauna is similar to the mammals in the park today and affords a unique opportunity to investigate paleocommunity and ecosystem dynamics through time.

Remains of an extralimital species in Yellowstone today, Microtus ochrogaster are found only in the oldest cave deposits, dated at 1695 ± 60 yr B.P. Disappearance of this species by about 1550 years ago marks evolution into the ecosystem that has persisted into the present and is recognizable by a fossil mammal assemblage that is virtually identical to the modern one. Relative abundance fluctuations in the mammal populations imply community resilience through the past 2000 years. Trends in relative abundances of small mammals, particularly the inverse relationship between Microtus and Spermophilus are interpreted as evidence of declining grass cover near Lamar Cave between approximately 1500 and 1000 years ago. The initiation of this environmental change may well have stimulated the ecosystem change marked by Microtus ochrogaster extirpation. Support for this hypothesis is found in a pollen record from a nearby lake.     

Fire‐mediated niche‐separation between two sympatric small mammal species

Fire is a key ecological process influencing the population dynamics of small mammals. Whilst shifting competitive advantage amongst small mammal species following a single fire event is well‐documented, there has been little investigation of the potential influence of fire frequency on small mammal interspecific interactions. In this study, we investigated the effect of fire frequency on the abundance of two small dasyurid mammals, Antechinus stuartii and A. flavipes, which occur sympatrically in some parts of their range. The two antechinus species are known to have different habitat preferences, so it is possible that fire regimes may promote their coexistence in areas of sympatry by altering vegetation structure. To investigate this possibility, we estimated the abundance of both species using replicate sites which differed in the number of times burnt (1–4) during the last four decades, but with identical time‐since‐fire. Proportionally, we captured greater numbers of A. stuartii in less frequently burnt sites and greater numbers of A. flavipes in more‐frequently burnt sites. Hence, fire may mediate niche‐separation between these two species. To clarify further this pattern of response to fire frequency, we investigated which structural habitat variables differed between fire frequencies, and compared antechinus abundances with structural vegetation characteristics. We found a trend for lower ground cover density under higher fire frequencies. This offers one potential explanation of the patterns of abundance that we observed. Our study provided insights into the complexities of small mammal responses to fire, and strongly suggests that fire could mediate competitive interactions between species.     

Body size changes in bivalves of the family Limidae in the aftermath of the end‐Triassic mass extinction: the Brobdingnag effect

Reduction in body size of organisms following mass extinctions is well‐known and often ascribed to the Lilliput effect. This phenomenon is expressed as a temporary body size reduction within surviving species. Despite its wide usage the term is often loosely applied to any small post‐extinction taxa. Here we assess the size of bivalves of the family Limidae (Rafineque) prior to, and in the aftermath of, the end‐Triassic mass extinction event. Of the species studied only one occurs prior to the extinction event, though is too scarce to test for the Lilliput effect. Instead, newly evolved species originate at small body sizes and undergo a within‐species size increase, most dramatically demonstrated by Plagiostoma giganteum (Sowerby) which, over two million years, increases in size by 179%. This trend is seen in both field and museum collections. We term this within‐species size increase of newly originated species in the aftermath of mass extinction, the Brobdingnag effect, after the giants that were contemporary with the Lilliputians in Swift's Gulliver's Travels. The size increase results from greater longevity and faster growth rates. The cause of the effect is unclear, although it probably relates to improved environmental conditions. Oxygen‐poor conditions in the Early Jurassic are associated with populations of smaller body size caused by elevated juvenile mortality but these are local/regional effects that do not alter the long‐term, size increase. Although temperature‐size relationships exist for many organisms (Temperature‐Size Rule and Bergmann's Rule), the importance of this is unclear here because of a poorly known Early Jurassic temperature record.     

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.     

Resolving the relationships of Paleocene placental mammals

The ‘Age of Mammals’ began in the Paleocene epoch, the 10 million year interval immediately following the Cretaceous–Palaeogene mass extinction. The apparently rapid shift in mammalian ecomorphs from small, largely insectivorous forms to many small‐to‐large‐bodied, diverse taxa has driven a hypothesis that the end‐Cretaceous heralded an adaptive radiation in placental mammal evolution. However, the affinities of most Paleocene mammals have remained unresolved, despite significant advances in understanding the relationships of the extant orders, hindering efforts to reconstruct robustly the origin and early evolution of placental mammals. Here we present the largest cladistic analysis of Paleocene placentals to date, from a data matrix including 177 taxa (130 of which are Palaeogene) and 680 morphological characters. We improve the resolution of the relationships of several enigmatic Paleocene clades, including families of ‘condylarths’. Protungulatum is resolved as a stem eutherian, meaning that no crown‐placental mammal unambiguously pre‐dates the Cretaceous–Palaeogene boundary. Our results support an Atlantogenata–Boreoeutheria split at the root of crown Placentalia, the presence of phenacodontids as closest relatives of Perissodactyla, the validity of Euungulata, and the placement of Arctocyonidae close to Carnivora. Periptychidae and Pantodonta are resolved as sister taxa, Leptictida and Cimolestidae are found to be stem eutherians, and Hyopsodontidae is highly polyphyletic. The inclusion of Paleocene taxa in a placental phylogeny alters interpretations of relationships and key events in mammalian evolutionary history. Paleocene mammals are an essential source of data for understanding fully the biotic dynamics associated with the end‐Cretaceous mass extinction. The relationships presented here mark a critical first step towards accurate reconstruction of this important interval in the evolution of the modern fauna.     

Quantifying the Extent of North American Mammal Extinction Relative to the Pre-Anthropogenic Baseline

Earth has experienced five major extinction events in the past 450 million years. Many scientists suggest we are now witnessing a sixth, driven by human impacts. However, it has been difficult to quantify the real extent of the current extinction episode, either for a given taxonomic group at the continental scale or for the worldwide biota, largely because comparisons of pre-anthropogenic and anthropogenic biodiversity baselines have been unavailable. Here, we compute those baselines for mammals of temperate North America, using a sampling-standardized rich fossil record to reconstruct species-area relationships for a series of time slices ranging from 30 million to 500 years ago. We show that shortly after humans first arrived in North America, mammalian diversity dropped to become at least 15%–42% too low compared to the “normal” diversity baseline that had existed for millions of years. While the Holocene reduction in North American mammal diversity has long been recognized qualitatively, our results provide a quantitative measure that clarifies how significant the diversity reduction actually was. If mass extinctions are defined as loss of at least 75% of species on a global scale, our data suggest that North American mammals had already progressed one-fifth to more than halfway (depending on biogeographic province) towards that benchmark, even before industrialized society began to affect them. Data currently are not available to make similar quantitative estimates for other continents, but qualitative declines in Holocene mammal diversity are also widely recognized in South America, Eurasia, and Australia. Extending our methodology to mammals in these areas, as well as to other taxa where possible, would provide a reasonable way to assess the magnitude of global extinction, the biodiversity impact of extinctions of currently threatened species, and the efficacy of conservation efforts into the future.     

Large‐scale patterns of seed removal by small mammals differ between areas of low‐ versus high‐wolf occupancy

Because most tree species recruit from seeds, seed predation by small‐mammal granivores may be important for determining plant distribution and regeneration in forests. Despite the importance of seed predation, large‐scale patterns of small‐mammal granivory are often highly variable and thus difficult to predict. We hypothesize distributions of apex predators can create large‐scale variation in the distribution and abundance of mesopredators that consume small mammals, creating predictable areas of high and low granivory. For example, because gray wolf (Canis lupus) territories are characterized by relatively less use by coyotes (C. latrans) and greater use by foxes (Vulpes vulpes, Urocyon cinereoargentus) that consume a greater proportion of small mammals, wolf territories may be areas of reduced small‐mammal granivory. Using large‐scale, multiyear field trials at 22 sites with high‐ and low‐wolf occupancy in northern Wisconsin, we evaluated whether removal of seeds of four tree species was lower in wolf territories. Consistent with the hypothesized consequences of wolf occupancy, seed removal of three species was more than 25% lower in high‐wolf‐occupancy areas across 2 years and small‐mammal abundance was more than 40% lower in high‐wolf areas during one of two study years. These significant results, in conjunction with evidence of seed consumption in situ and the absence of significant habitat differences between high‐ and low‐wolf areas, suggest that top‐down effects of wolves on small‐mammal granivory and seed survival may occur. Understanding how interactions among carnivores create spatial patterns in interactions among lower trophic levels may allow for more accurate predictions of large‐scale patterns in seed survival and forest composition.