Extant Megafauna Research Articles

Worldwide Late Pleistocene and Early Holocene population declines in extant megafauna are associated with Homo sapiens expansion rather than climate change

The worldwide extinction of megafauna during the Late Pleistocene and Early Holocene is evident from the fossil record, with dominant theories suggesting a climate, human or combined impact cause. Consequently, two disparate scenarios are possible for the surviving megafauna during this time period - they could have declined due to similar pressures, or increased in population size due to reductions in competition or other biotic pressures. We therefore infer population histories of 139 extant megafauna species using genomic data which reveal population declines in 91% of species throughout the Quaternary period, with larger species experiencing the strongest decreases. Declines become ubiquitous 32–76 kya across all landmasses, a pattern better explained by worldwide Homo sapiens expansion than by changes in climate. We estimate that, in consequence, total megafauna abundance, biomass, and energy turnover decreased by 92–95% over the past 50,000 years, implying major human-driven ecosystem restructuring at a global scale.

A wide megafauna gap undermines China's expanding coastal ecosystem conservation.

To fulfill sustainable development goals, many countries are expanding efforts to conserve ecologically and societally critical coastal ecosystems. Although megafauna profoundly affect the functioning of ecosystems, they are neglected as a key component in the conservation scheme for coastal ecosystems in many geographic contexts. We reveal a rich diversity of extant megafauna associated with all major types of coastal ecosystems in China, including 218 species of mammals, birds, reptiles, cephalopods, and fish across terrestrial and marine environments. However, 44% of these species are globally threatened, and 78% have not yet been assessed in China for extinction risk. More worrisome, 73% of these megafauna have not been designated as nationally protected species, and <10% of their most important habitats are protected. Filling this wide "megafauna gap" in China and globally would be a leading step as humanity strives to thrive with coastal ecosystems.

Africa as an evolutionary arena for large fruits.

Strong paleoclimatic change and few Late Quaternary megafauna extinctions make mainland Africa unique among continents. Here, we hypothesize that, compared with elsewhere, these conditions created the ecological opportunity for the macroevolution and geographic distribution of large fruits. We assembled global phylogenetic, distribution and fruit size data for palms (Arecaceae), a pantropical, vertebrate-dispersed family with > 2600 species, and integrated these with data on extinction-driven body size reduction in mammalian frugivore assemblages since the Late Quaternary. We applied evolutionary trait, linear and null models to identify the selective pressures that have shaped fruit sizes. We show that African palm lineages have evolved towards larger fruit sizes and exhibited faster trait evolutionary rates than lineages elsewhere. Furthermore, the global distribution of the largest palm fruits across species assemblages was explained by occurrence in Africa, especially under low canopies, and extant megafauna, but not by mammalian downsizing. These patterns strongly deviated from expectations under a null model of stochastic (Brownian motion) evolution. Our results suggest that Africa provided a distinct evolutionary arena for palm fruit size evolution. We argue that megafaunal abundance and the expansion of savanna habitat since the Miocene provided selective advantages for the persistence of African plants with large fruits.

Multiproxy analysis of permafrost preserved faeces provides an unprecedented insight into the diets and habitats of extinct and extant megafauna

The study of faecal samples to reconstruct the diets and habitats of extinct megafauna has traditionally relied on pollen and macrofossil analysis. DNA metabarcoding has emerged as a valuable tool to complement and refine these proxies. While published studies have compared the results of these three proxies for sediments, this comparison is currently lacking for permafrost preserved mammal faeces. Moreover, most metabarcoding studies have focused on a single plant-specific DNA marker region. In this study, we target both the commonly used chloroplast trnL P6 loop as well as nuclear ribosomal ITS (nrITS). The latter can increase taxonomic resolution of plant identifications but requires DNA to be relatively well preserved because of the target length (∼300–500 bp). We compare DNA results to pollen and macrofossil analyses from permafrost and ice-preserved faeces of Pleistocene and Holocene megafauna. Samples include woolly mammoth, horse, steppe bison as well as Holocene and extant caribou. Most plant identifications were found using DNA, likely because the studied faeces contained many vegetative remains that could not be identified using macrofossils or pollen. Several taxa were, however, identified to lower taxonomic levels uniquely with macrofossil and pollen analysis. The nrITS marker provides species level taxonomic resolution for commonly encountered plant families that are hard to distinguish using the other proxies (e.g. Asteraceae, Cyperaceae and Poaceae). Integrating the results from all proxies, we are able to accurately reconstruct known diets and habitats of the extant caribou. Applying this approach to the extinct mammals, we find that the Holocene horse and steppe bison were not strict grazers but mixed feeders living in a marshy wetland environment. The mammoths showed highly varying diets from different non-analogous habitats. This confirms the presence of a mosaic of habitats in the Pleistocene ‘mammoth steppe’ that mammoths could fully exploit due to their flexibility in food choice.

Are we eating the world's megafauna to extinction?

AbstractMany of the world's vertebrates have experienced large population and geographic range declines due to anthropogenic threats that put them at risk of extinction. The largest vertebrates, defined as megafauna, are especially vulnerable. We analyzed how human activities are impacting the conservation status of megafauna within six classes: mammals, ray‐finned fish, cartilaginous fish, amphibians, birds, and reptiles. We identified a total of 362 extant megafauna species. We found that 70% of megafauna species with sufficient information are decreasing and 59% are threatened with extinction. Surprisingly, direct harvesting of megafauna for human consumption of meat or body parts is the largest individual threat to each of the classes examined, and a threat for 98% (159/162) of threatened species with threat data available. Therefore, minimizing the direct killing of the world's largest vertebrates is a priority conservation strategy that might save many of these iconic species and the functions and services they provide.

Trophic rewilding as a climate change mitigation strategy?

The loss of megafauna at the terminal Pleistocene has been linked to a wide range of Earth-system-level changes, such as altered greenhouse gas budgets, fire regimes and biome-level vegetation changes. Given these influences and feedbacks, might part of the solution for mitigating anthropogenic climate change lie in the restoration of extant megafauna to ecosystems? Here, we explore the potential role of trophic rewilding on Earth's climate system. We first provide a novel synthesis of the various ways that megafauna interact with the major drivers of anthropogenic climate change, including greenhouse gas storage and emission, aerosols and albedo. We then explore the role of rewilding as a mitigation tool at two scales: (i) current and near-future opportunities for national or regional climate change mitigation portfolios, and (ii) more radical opportunities at the global scale. Finally, we identify major knowledge gaps that complicate the complete characterization of rewilding as a climate change mitigation strategy. Our perspective is urgent since we are losing the Earth's last remaining megafauna, and with it a potential option to address climate change.This article is part of the theme issue 'Trophic rewilding: consequences for ecosystems under global change'.

The Casa del Diablo cave (Puno, Peru) and the late Pleistocene demise of megafauna in the Andean Altiplano

During the Late Quaternary Extinction event South America lost ∼83% of all its late Pleistocene megafaunal genera. As in other regions of the world, the debate about the possible drivers behind these extinctions revolves around the role of humans arriving into the continent and on the effect of the climatic changes characteristic of the Pleistocene-Holocene transition. The availability of precise chronological information, in order to estimate the timing of extinction of the different taxa affected, is critical for solving such debate. Here we present an updated study of the late Pleistocene mammalian deposits from Casa del Diablo Cave (CdD) in the Altiplano of Peru. The study includes an updated list of the mammalian faunas found in the cave and 11 taxon-specific XAD radiocarbon dates from extinct and extant megafauna bones. We compare this new chronology to the timing of major environmental changes and human arrival in the area, as well as with other megafaunal discoveries from the high Andes. The radiocarbon dates from CdD fall in the time window between 23 and 12.8 cal kyr BP. Compared to other records of extinct megafauna in the high Andes, the one from CdD presents in general younger occurrences. No temporal overlap between humans and extinct megafauna emerges from comparing first dates of appearance of humans in the Altiplano, and last appearance dates of extinct megafauna from CdD. However, the possibility of temporal overlap among the records becomes evident when we compare confidence intervals calculated to estimate true times of human arrival and megafaunal local extinctions.

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

AbstractEndozoochory 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.

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.

Saving the World's Terrestrial Megafauna.

From the late Pleistocene to the Holocene, and now the so called Anthropocene, humans have been driving an ongoing series of species declines and extinctions (Dirzo et al. 2014). Large-bodied mammals are typically at a higher risk of extinction than smaller ones (Cardillo et al. 2005). However, in some circumstances terrestrial megafauna populations have been able to recover some of their lost numbers due to strong conservation and political commitment, and human cultural changes (Chapron et al. 2014). Indeed many would be in considerably worse predicaments in the absence of conservation action (Hoffmann et al. 2015). Nevertheless, most mammalian megafauna face dramatic range contractions and population declines. In fact, 59% of the world’s largest carnivores (≥ 15 kg, n = 27) and 60% of the world’s largest herbivores (≥ 100 kg, n = 74) are classified as threatened with extinction on the International Union for the Conservation of Nature (IUCN) Red List (supplemental table S1 and S2). This situation is particularly dire in sub-Saharan Africa and Southeast Asia, home to the greatest diversity of extant megafauna (figure 1). Species at risk of extinction include some of the world’s most iconic animals—such as gorillas, rhinos, and big cats (figure 2 top row)—and, unfortunately, they are vanishing just as science is discovering their essential ecological roles (Estes et al. 2011). Here, our objectives are to raise awareness of how these megafauna are imperiled (species in supplemental table S1 and S2) and to stimulate broad interest in developing specific recommendations and concerted action to conserve them.

Frugivory and seed dispersal by tapirs: an insight on their ecological role

Tapirs are one of the last extant megafauna species that survived the Pleistocene extinctions. Given their size and digestive system characteristics, tapirs might be the last potential seed disperser of plant species that were previously dispersed by other large mammal species that are now extinct. We compiled evidence from 39 published scientific studies showing that tapirs have a key role as seed dispersers and seed predators. Tapirs play an important role either through seed predation or by facilitating the recruitment of seeds over long distances, therefore influencing the diversity of plant species in the ecosystem. Neotropical tapirs might have a unique role as long-distance seed dispersers of large seeds (<20 mm) because they are capable of depositing viable large seeds in favorable places for germination that even large-bodied primates cannot disperse. Given the high diversity of seed species found in tapir diet, more information is needed on the identification of seed traits that allow the survival of seeds in the tapir's gut. Moreover, further studies are necessary on the role of tapirs as seed dispersers and predators; in particular considering spatial patterns of dispersed seeds, seed viability, effect of dung, and seed density in tapir latrines, and the effect of deposition sites on germination and seedling survival. Because all tapir species are highly threatened, it is paramount to identify gaps in our knowledge on the ecological role of tapirs and, in particular, on critical and endangered plant-tapir interactions to avoid possible trophic cascading effects on ecosystem function.

Late Quaternary Radiocarbon Chronology of Extinct and Extant Megafauna in Northern Eurasia and North-west North America

Late Quaternary Radiocarbon Chronology of Extinct and Extant Megafauna in Northern Eurasia and North-west North America

BIODIVERSITY LETTER: The North American Pleistocene overkill hypothesis and the re‐wilding debate

AbstractThe conservation agenda to re‐wild North America may or may not be realistic in terms of political ecology. However, it represents a real conservation recommendation to re‐wild North America with the extant megafauna most closely related to those that became extinct at the end of the Pleistocene. The recommendation is based on the presumption that society bears an ethical responsibility to re‐wild because humans caused the extinctions. However, the extent to which Pleistocene megafaunal extinctions were the result of overkill is hotly debated. As a result, the ethical imperative for North American re‐wilding should be questioned. It will not be questioned unless members of the conservation community read the extensive archaeological and geological literature concerning the North American Pleistocene extinctions. Overlooking the assumptions underlying this particular recommendation is costly to conservation science and to archaeology because it represents an over‐simplified, unwieldy and troubling fusion of the two.

Seed Dispersal and Establishment of Endangered Plants on Oceanic Islands: The Janzen-Connell Model, and the Use of Ecological Analogues

BackgroundThe Janzen-Connell model states that plant-specific natural enemies may have a disproportionately large negative effect on progeny close to maternal trees. The majority of experimental and theoretical studies addressing the Janzen-Connell model have explored how it can explain existing patterns of species diversity in tropical mainland areas. Very few studies have investigated how the model's predictions apply to isolated oceanic islands, or to the conservation management of endangered plants. Here, we provide the first experimental investigation of the predictions of the Janzen-Connell model on an oceanic island, in a conservation context. In addition, we experimentally evaluate the use of ecological analogue animals to resurrect the functional component of extinct frugivores that could have dispersed seeds away from maternal trees.Methodology/Principal FindingsIn Mauritius, we investigated seed germination and seedling survival patterns of the critically endangered endemic plant Syzygium mamillatum (Myrtaceae) in relation to proximity to maternal trees. We found strong negative effects of proximity to maternal trees on growth and survival of seedlings. We successfully used giant Aldabran tortoises as ecological analogues for extinct Mauritian frugivores. Effects of gut-passage were negative at the seed germination stage, but seedlings from gut-passed seeds grew taller, had more leaves, and suffered less damage from natural enemies than any of the other seedlings.Conclusions/SignificanceWe provide the first experimental evidence of a distance-dependent Janzen-Connell effect on an oceanic island. Our results potentially have serious implications for the conservation management of rare plant species on oceanic islands, which harbour a disproportionately large fraction of the world's endemic and endangered plants. Furthermore, in contrast to recent controversy about the use of non-indigenous extant megafauna for re-wilding projects in North America and elsewhere, we argue that Mauritius and other oceanic islands are ideal study systems in which to empirically explore the use of ecological analogue species in restoration ecology.

Frugivory and seed dispersal by Asian elephants, Elephas maximus, in a moist evergreen forest of Thailand

The Asian elephant Elephas maximus is the only living species of the genus Elephas (Elephantidae) that evolved in Africa c. 5–6 million y ago and migrated into Eurasia (Sukumar 2003). The Asian elephant is one of the few remaining extant megafauna (Owen-Smith 1988) and has disappeared from c. 95% of its historical range (Sukumar 2006). Asian elephants eat fruit when it is available, defecating intact seeds, of which some later germinate in the dung (Lekagul &amp; McNeely 1977, Ridley 1930). However, to date there has been no detailed study of frugivory and seed dispersal by Asian elephants (Corlett 1998). The only common feature of the fruit reported to be eaten by Asian elephants is their relatively large size, but there is no evidence that they are exclusive dispersers of any plant species (Corlett 1998), in contrast to the more frugivorous African forest elephants, Loxodonta africana cyclotis (Babweteera et al. 2007, Chapman et al. 1992, Cochrane 2003, Feer 1995).