Human Overkill Research Articles

Overkill, glacial history, and the extinction of North America’s Ice Age megafauna

The end of the Pleistocene in North America saw the extinction of 38 genera of mostly large mammals. As their disappearance seemingly coincided with the arrival of people in the Americas, their extinction is often attributed to human overkill, notwithstanding a dearth of archaeological evidence of human predation. Moreover, this period saw the extinction of other species, along with significant changes in many surviving taxa, suggesting a broader cause, notably, the ecological upheaval that occurred as Earth shifted from a glacial to an interglacial climate. But, overkill advocates ask, if extinctions were due to climate changes, why did these large mammals survive previous glacial−interglacial transitions, only to vanish at the one when human hunters were present? This question rests on two assumptions: that previous glacial−interglacial transitions were similar to the end of the Pleistocene, and that the large mammal genera survived unchanged over multiple such cycles. Neither is demonstrably correct. Resolving the cause of large mammal extinctions requires greater knowledge of individual species’ histories and their adaptive tolerances, a fuller understanding of how past climatic and ecological changes impacted those animals and their biotic communities, and what changes occurred at the Pleistocene−Holocene boundary that might have led to those genera going extinct at that time. Then we will be able to ascertain whether the sole ecologically significant difference between previous glacial−interglacial transitions and the very last one was a human presence.

Sea otters, kelp forests, and the extinction of Steller’s sea cow

The late Pleistocene extinction of so many large-bodied vertebrates has been variously attributed to two general causes: rapid climate change and the effects of humans as they spread from the Old World to previously uninhabited continents and islands. Many large-bodied vertebrates, especially large apex predators, maintain their associated ecosystems through top-down forcing processes, especially trophic cascades, and megaherbivores also exert an array of strong indirect effects on their communities. Thus, a third possibility for at least some of the Pleistocene extinctions is that they occurred through habitat changes resulting from the loss of these other keystone species. Here we explore the plausibility of this mechanism, using information on sea otters, kelp forests, and the recent extinction of Steller's sea cows from the Commander Islands. Large numbers of sea cows occurred in the Commander Islands at the time of their discovery by Europeans in 1741. Although extinction of these last remaining sea cows during early years of the Pacific maritime fur trade is widely thought to be a consequence of direct human overkill, we show that it is also a probable consequence of the loss of sea otters and the co-occurring loss of kelp, even if not a single sea cow had been killed directly by humans. This example supports the hypothesis that the directly caused extinctions of a few large vertebrates in the late Pleistocene may have resulted in the coextinction of numerous other species.

Revisiting Paleoindian exploitation of extinct North American mammals

In 2002, we assessed all sites known to us that had been suggested to provide evidence for the association of Clovis-era archaeological material with the remains of extinct Pleistocene mammals in North America. We concluded that, of the 76 sites we assessed, 14 provided compelling evidence for human involvement in the death and/or dismemberment of such mammals. Of these sites, 12 involved mammoth (Mammuthus), the remaining two mastodon (Mammut). Here, we update that assessment. We examine Clovis-era, and earlier, sites reported since 2002, as well as sites examined previously but for which additional information has become available. Our assessment leads us to exclude Hebior (Wisconsin) from the list of accepted sites, and to add El Fin del Mundo (Sonora) and Wally's Beach (Alberta). There are now 15 sites on our list, providing what we find to be compelling evidence for human involvement in the death and/or dismemberment of five genera of now-extinct late Pleistocene mammals: Equus, Camelops, Cuvieronius, Mammut, and Mammuthus. As in 2002, however, we note this is a small fraction of the 37 genera that disappeared at the end of the Pleistocene, and for this and other reasons we remain highly skeptical that human overkill was responsible for their extinction.

Local extinction and unintentional rewilding of bighorn sheep (Ovis canadensis) on a desert island.

Bighorn sheep (Ovis canadensis) were not known to live on Tiburón Island, the largest island in the Gulf of California and Mexico, prior to the surprisingly successful introduction of 20 individuals as a conservation measure in 1975. Today, a stable island population of ∼500 sheep supports limited big game hunting and restocking of depleted areas on the Mexican mainland. We discovered fossil dung morphologically similar to that of bighorn sheep in a dung mat deposit from Mojet Cave, in the mountains of Tiburón Island. To determine the origin of this cave deposit we compared pellet shape to fecal pellets of other large mammals, and extracted DNA to sequence mitochondrial DNA fragments at the 12S ribosomal RNA and control regions. The fossil dung was 14C-dated to 1476–1632 calendar years before present and was confirmed as bighorn sheep by morphological and ancient DNA (aDNA) analysis. 12S sequences closely or exactly matched known bighorn sheep sequences; control region sequences exactly matched a haplotype described in desert bighorn sheep populations in southwest Arizona and southern California and showed subtle differentiation from the extant Tiburón population. Native desert bighorn sheep previously colonized this land-bridge island, most likely during the Pleistocene, when lower sea levels connected Tiburón to the mainland. They were extirpated sometime in the last ∼1500 years, probably due to inherent dynamics of isolated populations, prolonged drought, and (or) human overkill. The reintroduced population is vulnerable to similar extinction risks. The discovery presented here refutes conventional wisdom that bighorn sheep are not native to Tiburón Island, and establishes its recent introduction as an example of unintentional rewilding, defined here as the introduction of a species without knowledge that it was once native and has since gone locally extinct.

Reply to Brook et al: No empirical evidence for human overkill of megafauna in Sahul

We welcome comments by Brook et al. (1), supporters of human-driven models, on our review of the role of climate in Pleistocene faunal extinctions in Sahul (Pleistocene Australia–New Guinea) (2). In response, we begin on a point of agreement: the fossil fauna record on which our respective arguments are based is sparse, although our understanding of Pleistocene environmental conditions is improving (3⇓–5). However, we also flag a basic point of difference. Unlike Brook et al., who focus on the ∼50 extinct Australian species (an artificial distinction because Australia was part of the larger landmass Sahul) and the 50 ka since human arrival, we consider the bigger picture, …

Flightless ducks, giant mice and pygmy mammoths: Late Quaternary extinctions on California's Channel Islands

Explanations for the extinction of Late Quaternary megafauna are heavily debated, ranging from human overkill to climate change, disease and extraterrestrial impacts. Synthesis and analysis of Late Quaternary animal extinctions on California's Channel Islands suggest that, despite supporting Native American populations for some 13,000 years, few mammal, bird or other species are known to have gone extinct during the prehistoric human era, and most of these coexisted with humans for several millennia. Our analysis provides insight into the nature and variability of Quaternary extinctions on islands and a broader context for understanding ancient extinctions in North America.

Overdone overkill – the archaeological perspective on Tasmanian megafaunal extinctions

The reasons for megafaunal extinction in Australia have been hotly debated for over 30 years without any clear resolution. The proposed causes include human overkill, climate, anthropogenic induced habitat change or a combination of these. Most protagonists of the human overkill model suggest the impact was so swift, occurring within a few thousand years of human occupation of the continent, that archaeological evidence should be rare or non-existent. In Tasmania the presence of extinct megafauna has been known since the early twentieth century ( Noetling, 1912; Scott, 1911, 1915) with earlier claims of human overlap being rejected because of poor chronology and equivocal stratigraphic associations. More recent archaeological research has not identified any megafauna from the earliest, exceptionally well-preserved late Pleistocene cultural sites. In 2008 however an argument for human induced megafaunal extinctions was proposed using the direct dates from a small sample of surface bone from two Tasmanian non-human caves and a museum sediment sample from an unknown location in a cave, since destroyed by quarrying ( Turney et al., 2008). Turney et al. (2008) supplemented their data with published dates from other Tasmanian caves and open sites to argue for the survival of at least seven megafauna species from the last interglacial to the subsequent glacial stage. To investigate the timing of extinctions in Tasmania and examine the latest claims, new excavations and systematic surveys of limestone caves in south central Tasmania were undertaken. Our project failed to show any clear archaeological overlap of humans and megafauna but demonstrated that vigilance is needed when claiming survival of megafauna species based on old or suspect chronologies. The results of our six-years of fieldwork and dating form the first part of the present paper while, in the second part we assess the data advanced by Turney et al. (2008) for the late survival of seven megafauna species. A model of human prey selection and the reasons for the demise of a range of marsupials, now extinct, are discussed in the third part of the paper.

New carbon dates link climatic change with human colonization and Pleistocene extinctions

Drastic ecological restructuring, species redistribution and extinctions mark the Pleistocene-Holocene transition, but an insufficiency of numbers of well-dated large mammal fossils from this transition have impeded progress in understanding the various causative links. Here I add many new radiocarbon dates to those already published on late Pleistocene fossils from Alaska and the Yukon Territory (AK-YT) and show previously unrecognized patterns. Species that survived the Pleistocene, for example, bison (Bison priscus, which evolved into Bison bison), wapiti (Cervus canadensis) and, to a smaller degree, moose (Alces alces), began to increase in numbers and continued to do so before and during human colonization and before the regional extinction of horse (Equus ferus) and mammoth (Mammuthus primigenius). These patterns allow us to reject, at least in AK-YT, some hypotheses of late Pleistocene extinction: 'Blitzkrieg' version of simultaneous human overkill, 'keystone' removal, and 'palaeo-disease'. Hypotheses of a subtler human impact and/or ecological replacement or displacement are more consistent with the data. The new patterns of dates indicate a radical ecological sorting during a uniquely forage-rich transitional period, affecting all large mammals, including humans.

Megafauna extinction: A paleoeconomic theory of human overkill in the pleistocene

After centuries of debate, paleontologists are converging towards the conclusion that human overkill caused the massive extinction of large mammals in the late Pleistocene. This paper revisits the question of megafauna extinction by incorporating economic behavior into the debate. We allow for endogenous human population growth, and labor allocation decisions involving activities such as wildlife harvesting and (proto) agriculture. We find that the role of agriculture in deciding the fate of megafauna was small. In contrast, the presence of ordinary small animals that have been overlooked in previous non-economic extinction models is likely to have been much more important.

Rapid body size decline in Alaskan Pleistocene horses before extinction.

About 70% of North American large mammal species were lost at the end of the Pleistocene epoch. The causes of this extinction--the role of humans versus that of climate--have been the focus of much controversy. Horses have figured centrally in that debate, because equid species dominated North American late Pleistocene faunas in terms of abundance, geographical distribution, and species variety, yet none survived into the Holocene epoch. The timing of these equid regional extinctions and accompanying evolutionary changes are poorly known. In an attempt to document better the decline and demise of two Alaskan Pleistocene equids, I selected a large number of fossils from the latest Pleistocene for radiocarbon dating. Here I show that horses underwent a rapid decline in body size before extinction, and I propose that the size decline and subsequent regional extinction at 12,500 radiocarbon years before present are best attributed to a coincident climatic/vegetational shift. The present data do not support human overkill and several other proposed extinction causes, and also show that large mammal species responded somewhat individualistically to climate changes at the end of the Pleistocene.

Mitigating Human Effects on European Biodiversity through Traditional Animal Husbandry

Abstract:Livestock grazing (without modern fertilizers), mowing, and other traditional methods of animal husbandry are used in Europe for managing human‐made habitats such as seminatural grasslands. From a review of essential literature, I hypothesize that traditional animal husbandry partially compensates for the loss of natural processes that have been suppressed by humans. There is indirect evidence that livestock grazing and mowing have made possible the continued existence of many species threatened by the human overkill of megaherbivores and other large herbivores. Many species that were dependent on natural fires and floods may have benefited from grazing and mowing, which also may be effective tools for mitigating the negative effects of eutrophication. As partial surrogates, traditional grazing and mowing have obscured the importance of natural disturbances to European biodiversity. Thus, the end of traditional animal husbandry, together with the suppression of natural disturbances, may cause even more adverse effects to biodiversity than is generally recognized.

Pleistocene extinctions: the pivotal role of megaherbivores

Two alternative hypotheses have been advanced to explain the demise of about half of the mammalian genera exceeding 5 kg in body mass in the later Pleistocene. One hypothesis invokes climatic change and resulting habitat transformations. This fails to predict the increased likelihood of extinctions with increasing body size, greater severity in both North and South America than in Eurasia or Australia, lack of simultaneous extinctions in Africa and tropical Asia, and the absence of extinctions at the end of previous glacial periods. The other hypothesis invokes human predation as the primary cause. This fails to explain the simultaneous extinctions of a number of mammalian and avian species not obviously vulnerable to human overkill. I propose a “keystone herbivore” hypothesis, based on the ecology of extant African species of megaherbivore, (i.e., animals exceeding 1,000 kg in body mass). Due to their invulnerability to non-human predation on adults, these species attain saturation densities at which they may radically transform vegetation structure and composition. African elephant can change closed woodland or thicket into open grassy savanna, and create open gaps colonized by rapidly-regenerating trees in forests. Grazing white rhinoceros and hippopotamus transform tall grasslands into lawns of more nutritious short grasses. The elimination of megaherbivores elsewhere in the world by human hunters at the end of the Pleistocene would have promoted reverse changes in vegetation. The conversion of the open parklike woodlands and mosaic grasslands typical of much of North America during the Pleistocene to the more uniform forests and prairie grasslands we find today could be a consequence. Such habitat changes would have been detrimental to the distribution and abundance of smaller herbivores dependent upon the nutrient-rich and spatially diverse vegetation created by megaherbivore impact. At the same time these species would have become more vulnerable to human predation. The elimination of megaherbivore influence is the major factor differentiating habitat changes at the end of the terminal Pleistocene glaciation from those occurring at previous glacial-interglacial transitions.

Pleistocene overkill and the associational critique

The “associational critique”, the claim there are insufficient associations between extinct Pleistocene mammalian genera and cultural materials to support a model of human overkill, has been challenged by Grayson (1984). He argues that it is based on a faulty assumption regarding the sample of terminal Pleistocene megafauna, and that analysis using the radiocarbon record of extinct fauna demonstrates that there are actually more extinct genera found in archaeological association than would be expected by chance. He concludes that the associational critique is weakly based. However, his analysis warrants careful examination for, by including only radiocarbon-dated sites, and utilizing dates of questionable reliability, it is subject to sample bias. Re-analysis suggests that a rejection of the associational critique is premature. There are far too few sites showing evidence of human predation to support a model of human overkill.

Production of sulphur in the soil of Paris

In 2002, we assessed all sites known to us that had been suggested to provide evidence for the association of Clovis-era archaeological material with the remains of extinct Pleistocene mammals in North America. We concluded that, of the 76 sites we assessed, 14 provided compelling evidence for human involvement in the death and/or dismemberment of such mammals. Of these sites, 12 involved mammoth (Mammuthus), the remaining two mastodon (Mammut). Here, we update that assessment. We examine Clovis-era, and earlier, sites reported since 2002, as well as sites examined previously but for which additional information has become available. Our assessment leads us to exclude Hebior (Wisconsin) from the list of accepted sites, and to add El Fin del Mundo (Sonora) and Wally's Beach (Alberta). There are now 15 sites on our list, providing what we find to be compelling evidence for human involvement in the death and/or dismemberment of five genera of now-extinct late Pleistocene mammals: Equus, Camelops, Cuvieronius, Mammut, and Mammuthus. As in 2002, however, we note this is a small fraction of the 37 genera that disappeared at the end of the Pleistocene, and for this and other reasons we remain highly skeptical that human overkill was responsible for their extinction.