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.     

Late-Quaternary megafauna extinctions have strongly reduced mammalian vegetation consumption

Aim

How much stronger would the effects of herbivorous mammals be in natural ecosystems if human-linked extinctions and extirpations had not occurred? Many mammal species have experienced range contractions, and numerous species have gone extinct in the late Quaternary, completely or in large part linked to human pressures. Therefore, herbivore consumption rates in seemingly natural ecosystems will deviate from their pre-anthropogenic state. Here, we estimate the size of this deviation.

Location

Terrestrial systems, globally.

Time period

Current.

Major taxa studied

All late-Quaternary terrestrial mammals.

Methods

We estimated and mapped vegetation consumption rate by all late-Quaternary terrestrial mammals. We did this through the estimation of natural densities and dietary needs. We mapped their consumption rate in both current ranges and present-natural ranges, that is estimated ranges in the absence of human-linked range contractions and extinctions. We compared these estimated consumption rates to current net primary productivity (NPP). We summarized the results across ecosystem types everywhere as well as for only the last remaining wilderness areas.

Results

We estimate that wild mammals consume a median of 11% of NPP (at the scale of 96.5 km × 96.5 km grid cells) in current natural areas and that this would have been much higher in the absence of extinctions and extirpations, namely 21%. Looking at the change per grid cell, the mammal losses result in a median 42% reduction in consumption rate. Importantly, we estimate very similar declines in herbivory in what are considered the last remaining wilderness areas.

Main conclusions

Our results suggest that the natural interaction of mammalian herbivores with vegetation in ecosystems across the world is strongly reduced by prehistoric and historic to recent species losses, even in the last remaining wilderness areas, likely with major effects on ecosystem structure and functioning.     

White‐tailed deer as the last megafauna dispersing seeds in Neotropical dry forests: the role of fruit and seed traits

Endozoochory is a prominent form of seed dispersal in tropical dry forests. Most extant megafauna that perform such seed dispersal are ungulates, which can also be seed predators. White‐tailed deer (Odocoileus virginianus) is one of the last extant megafauna of Neotropical dry forests, but whether it serves as a legitimate seed disperser is poorly understood. We studied seed dispersal patterns and germination after white‐tailed deer gut passage in a tropical dry forest in southwest Ecuador. Over 23 mo, we recorded ca 2000 seeds of 11 species in 385 fecal samples. Most seeds belonged to four species of Fabaceae: Chloroleucon mangense, Senna mollissima, Piptadenia flava, and Caesalpinia glabrata. Seeds from eight of the 11 species dispersed by white‐tailed deer germinated under controlled conditions. Ingestion did not affect germination of C. mangense and S. mollissima, whereas C. glabrata showed reduced germination. Nevertheless, the removal of fruit pulp resulting from ingestion by white‐tailed deer could have a deinhibition effect on germination due to seed release. Thus, white‐tailed deer play an important role as legitimate seed dispersers of woody species formerly considered autochorous. Our results suggest that more research is needed to fully understand the ecological and evolutionary effects of the remaining extant megafauna on plant regeneration dynamics in the dry Neotropics.     

Terrestrial effects of moderately nearby supernovae

Recent data indicate one or more moderately nearby supernovae in the Early Pleistocene, with additional events likely in the Miocene. This has motivated more detailed computations, using new information about the nature of supernovae and the distances of these events to describe in more detail the sorts of effects that are indicated at the Earth. This short communication/review is designed to describe some of these effects so that they may possibly be related to changes in the biota around these times.     

Habitats of Pleistocene megaherbivores reconstructed from the frozen fauna remains

The Late Pleistocene landscape in northern Eurasia and North America was inhabited by a specific megafaunal complex, which largely disappeared during the Pleistocene/Holocene transition. Vegetation changes are considered as one of the factors responsible for these extinctions, but the structure and composition of the Pleistocene vegetation are still poorly known. Here we complement previous studies by comparing the taxonomic composition of the plant remains found in the gastrointestinal tracts of the frozen carcasses of Pleistocene megaherbivores with the species composition of the current Siberian vegetation. We compiled a dataset of palaeobotanical records from frozen individuals of Pleistocene megaherbivores found in northern Siberia and Beringia and dated to the period from more than 50 kyr BP to 9 kyr BP. We also compiled a dataset of vegetation plots from several regions in Siberia. We analysed the similarity in taxonomic composition of plants between these two datasets using a novel method that accounts for variable taxonomic resolution in palaeobotanical data. For most megaherbivore individuals, plant remains in their gastrointestinal tracts corresponded to tundra, forest and mire vegetation, while they showed low similarity to steppe. This pattern was relatively constant over time, showing no remarkable differences between the Last Glacial Maximum and the periods before and afterwards. This suggests that during the Upper Pleistocene, a mosaic of mesic and wet vegetation types such as tundra with patches of forests and mires was common in northern Siberia and Beringia. In contrast, the steppe was rare to absent in the landscape or underused by the megaherbivores as a pasture since they found enough food in the widespread mesic and wet habitats with more productive vegetation.     

Microsatellite genotyping reveals end-Pleistocene decline in mammoth autosomal genetic variation

The last glaciation was a dynamic period with strong impact on the demography of many species and populations. In recent years, mitochondrial DNA sequences retrieved from radiocarbon-dated remains have provided novel insights into the history of Late Pleistocene populations. However, genotyping of loci from the nuclear genome may provide enhanced resolution of population-level changes. Here, we use four autosomal microsatellite DNA markers to investigate the demographic history of woolly mammoths (Mammuthus primigenius) in north-eastern Siberia from before 60 000 years ago up until the species' final disappearance c.4000 years ago. We identified two genetic groups, implying a marked temporal genetic differentiation between samples with radiocarbon ages older than 12 thousand radiocarbon years before present (ka) and those younger than 9ka. Simulation-based analysis indicates that this dramatic change in genetic composition, which included a decrease in individual heterozygosity of approximately 30%, was due to a multifold reduction in effective population size. A corresponding reduction in genetic variation was also detected in the mitochondrial DNA, where about 65% of the diversity was lost. We observed no further loss in genetic variation during the Holocene, which suggests a rapid final extinction event.     

Process-explicit models reveal pathway to extinction for woolly mammoth using pattern-oriented validation

Pathways to extinction start long before the death of the last individual. However, causes of early stage population declines and the susceptibility of small residual populations to extirpation are typically studied in isolation. Using validated process-explicit models, we disentangle the ecological mechanisms and threats that were integral in the initial decline and later extinction of the woolly mammoth. We show that reconciling ancient DNA data on woolly mammoth population decline with fossil evidence of location and timing of extinction requires process-explicit models with specific demographic and niche constraints, and a constrained synergy of climatic change and human impacts. Validated models needed humans to hasten climate-driven population declines by many millennia, and to allow woolly mammoths to persist in mainland Arctic refugia until the mid-Holocene. Our results show that the role of humans in the extinction dynamics of woolly mammoth began well before the Holocene, exerting lasting effects on the spatial pattern and timing of its range-wide extinction.     

Using ancient DNA and coalescent-based methods to infer extinction

DNA sequences extracted from preserved remains can add considerable resolution to inference of past population dynamics. For example, coalescent-based methods have been used to correlate declines in some arctic megafauna populations with habitat fragmentation during the last ice age. These methods, however, often fail to detect population declines preceding extinction, most likely owing to a combination of sparse sampling, uninformative genetic markers, and models that cannot account for the increasingly structured nature of populations as habitats decline. As ancient DNA research expands to include full-genome analyses, these data will provide greater resolution of the genomic consequences of environmental change and the genetic signatures of extinction.