Pleistocene megafaunal extinctions – were humans to blame?

Australia and the Americas had an extremely diverse fauna of large beasts (giant wombats and kangeroos in Australia; elephants, bears, big cats, camelids, ground sloths etc in the Americas) until the last glaciation and the warming period that led into the Holocene interglacial. The majority of these megafauna species vanished suddenly during that recent period. To a lesser extent something similar happened in Eurasia, but nothing significant in Africa. Because the last glacial cycle also saw migration of efficient human hunter-gatherers to every other continent except Antarctica, many ecologists, palaeontologists and anthropologists saw a direct link between human predation and the mass extinction (see Earth-Pages of April 2012. Earlier humans had indeed spread far and wide in Eurasia before, and the crude hypothesis that the last arrivals in Australasia and the Americas devoured all the meatiest prey in three continents had some traction as a result: predation in Eurasia and Africa by earlier hominids would have made surviving prey congenitally wary of bipeds with spears. In Australia and the Americas the megafauna species would have been naive and confident in their sheer bulk, numbers, speed and, in some cases, ferocity. Other possibilities emerged, such as the introduction of viruses to which faunas had no immunity or as a result of climate change, but none of the three possibilities has gained incontrovertible proof. But the most popular, human connection has had severe knocks in the last couple of years. A fourth, that the extinctions stemmed from a comet impact proved to have little traction.

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Megafauna in a late-Pleistocene landscape including woolly mammoths and rhinoceroses, horses, and cave lions with a carcass. (credit: Wikipedia)

Since the amazing success of analysing the bulk DNA debris in sea water – environmental DNA or eDNA – to look at the local diversity of marine animals, the analytical and computing techniques that made it possible have been turned to ancient terrestrial materials: soils, permafrost and glacial ice. One of the first attempts revealed mammoth and pre-Columbian horse DNA surviving in Alaskan permafrost, thanks to the herds’ copious urination and dung spreading. Several articles in the 24 July 2015 issue of Science review ancient DNA advances, including eDNA from soils that chart changes in both fauna and flora over the last glacial cycle (Pennisi, E. 2015. Lost worlds found. Science, v. 349, p. 367-369). Combined with a variety of means of dating the material that yield the ancient eDNA, an interesting picture is emerging. The soil and permafrost samples potentially express ancient ecosystems in far more detail than would fossil animals or pollens, many of which are too similar to look at the species level and in any case are dominated by the most abundant plants rather than showing those critical in the food chain.

Nunavut tundra
Plants of the Arctic tundra in Nunavut, Canada (Photo credit: Wikipedia)

The first major success in palaeoecology of this kind came with a 50-author paper using eDNA ‘bar-coding’ of permafrost from 242 sites in Siberia and Alaska IWillerslev, E. and 49 others 2014. Fifty thousand years of Arctic vegetation and megafaunal diet. Nature, v. 506, p. 47-51. doi:10.1038/nature12921). Dividing the samples into 3 time spans – 50-25, 25-15 (last glacial maximum) and younger than 15 ka – the team found these major stages in the last glacial cycle mapped an ecological change from a dry tundra dominated by abundant herbaceous plants (forbs including abundant anemones and forget-me-not), to a markedly depleted Arctic steppe ecosystem then moist tundra with woody plants and grasses dominating. They also analysed the eDNA of dung and gut contents from ice-age megafauna, such as mammoths, bison and woolly rhinos, where these were found, which showed that forbs were the mainstay of their diet. Using bones of large mammals 6 member of the team also established the timing of extinctions in the last 56 ka (Cooper, A. et al. 2015. Abrupt warming events drove Late Pleistocene Holarctic megafaunal turnover. Science, DOI: 10.1126/science.aac4315), showing 31 regional extinction pulses linked to the rapid ups and downs of climate during Dansgaard-Oeschger cycles in the run-up to the last glacial maximum. By the end of the last glacial maximum, the megafauna were highly stressed by purely climatic and ecological factors. Human predation probably finished them off.

Some megafaunas of the recent past

Harvey was an imaginary, 2 m tall rabbit which befriended Elwood P. Dowd in Mary Chase’s 1944 comedy of errors named after the said rabbit, filmed in 1950 and starring James Stewart as the affable though deranged Dowd. Though not so tall, a giant fossil rabbit (relative to modern rabbits) weighing it at 12 kg has emerged from the prolific Late Neogene cave deposits of Minorca (Quintana, J. Et al. 2011. Nuralagus rex, gen. et sp. nov., an endemic insular giant rabbit from the Neogene of Minorca (Balearic Islands, Spain). Journal of Vertebrate Paleontology, v. 31, p. 231-240). At about 3 times heavier than Barrington my lagomorphophagic (rabbit-eating to the uninitiated) cat, this would have been, to him, a beast best avoided, as the name N. rex might suggest. So unexpected was a gigantic rabbit that, interestingly, it was first mistaken for a fossil tortoise, albeit one lacking a carapace.

Island faunas have long been recognized as havens for peculiar trends in evolutionary successions, either towards dwarfism as in the case of the tiny elephants on which H. floresiensis preyed until quite recently on the Indonesian island of Flores or gigantism as in this remarkable case. As the authors infer, on account of the creature’s ‘…(short manus and pes with splayed phalanges, short and stiff vertebral column with reduced extension/flexion capabilities), and the relatively small size of sense-related areas of the skull (tympanic bullae, orbits, braincase, and choanae)…’ this was a rabbit which sadly could not hop. This un-rabbit-like locomotion may well have been a result of it not having needed to hop, being so large as to challenge seriously the largest Neogene predators on the island – lizards – and thereby saving energy. For much the same evolutionary logic, neither did N. rex have long ears, having less need to detect a stealthy nemesis.

The demise of Late Neogene megafaunas in general has often been ascribed to human intervention. Though N. rex became extinct at around 3 Ma and avoided human predation, later giants did not fare so well. A case in point is the celebrated wooly mammoth, the last of the steppe mammoths, that first appeared in the fossil record of Siberia around 750 ka ago (Nicholls H. 2011. Last days of the mammoth. New Scientist, v. 209 (26 March 2011), p. 54-57). DNA evidence from hairs preserved in permafrost suggests that ancestors of the steppe mammoth line diverged with that of Asian elephants from African elephant ancestors around 5 Ma. Interestingly, ancestral steppe mammoths – without shaggy coats but having the archetypical curved tusks – roamed Africa until 3 Ma when they disappear to reappear in Europe and Asia, yet without adaptation to cold until the onset of northern glaciations around 2.5 Ma. At that point the true steppe mammoths evolved increased tooth enamel needed for a diet of mainly silica-rich grasses to resist wear. The family spread to North America when sea-level fell to expose the sea floor of the Bering Straits. The woolly mammoth is the star partly because specimens periodically turn up almost perfectly preserved in permafrost. This has allowed almost half of a full DNA sequence to be restored. Preserved haemoglobin from a woolly mammoth shares with that from modern musk oxen an ability to release oxygen at temperatures well below zero so that they could function even if their extremities became chilled.

The Woolly Mammoth at the Royal BC Museum, Vic...
Reconstructed woolly mammoth at the Royal BC Museum, Victoria, British Columbia (Image via Wikipedia)

Astonishingly, all elephants urinate so copiously that they soak their range lands in DNA, though it only lingers in ultra cold climes. This bizarre fact encouraged a large team of palaeobiologists to comb frozen soils in an alluvium section in Arctic Alaska for mammoth DNA (Haile, J and 17 others, 2009. Ancient DNA reveals late survival of mammoth and horse in interior Alaska.  Proceedings of the National Academy of Sciences of the USA, v. 106, p. 22352–22357). Mammoth DNA turned up in soils as young as 10.5 ka. Moreover mammoth overlapped with human occupation for several millennia, casting doubt on theories that mammoth extinction resulted either from human predation or the introduction of epidemic disease that might have felled mammoths quickly: they declined gradually. Yet the empirical fact that steppe mammoths in general and the woolly mammoth in particular survived through at least 8 major glacial-interglacial transitions only to become extinct at the start of the current Holocene interglacial period at the same time as humans recolonised the frigid desert of Arctic latitudes, where woolly mammoths could survive except at the last glacial maximum surely points to some influence that arose from human activity.