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peak, far more than half of the continent became technically arid’. 2 Major extinctions resulted.
    A few carnivores survived. Two were ancestors of the thylacine and the quoll genus, both of them hunters. It may be that specialist scavenging came to be an important niche, with the thylacine in particular ensuring a supply of carrion through its habit of selective feeding. This may be how the devil line arose. The species has no known earlier ancestry, unlike both the thylacine and quoll, which trace back at least 25 million years. The extinct species Glaucodon balabacensis from the Pliocene (around 5 million–2 million years ago) is described as an ‘intermediate form’ between quoll and devil. 3 Although this suggests evolutionary experimentation in response to the increasingly dry environment, speculation based on fragmentary fossil evidence must be treated with care.
    Australia’s Miocene fossil record was considered poor until, in the early 1980s, the rich Riversleigh fossil deposits in northwestern Queensland were properly surveyed. At some 100 sites, huge numbers of limestone-encased fossils are preserved in ancient cave systems and waterways. ‘Almost half of what we know about the evolution of Australian mammals in the last 30 million years comes from bones found at a single site in the Riversleigh fossil beds. Half of that was unearthed in one hour.’ 4
    Riversleigh was granted World Heritage status, together with the much younger limestone fossil sites at Naracoorte Caves, in southeastern South Australia. There the devil is represented in the extraordinarily rich Fossil Chamber, a huge cave into which animals fell over a period of some 300 000 years, creating a gigantic cone of well-preserved bone deposits. Although Naracoorte and Riversleigh contain a wealth of information yet to be tapped, they have enabled a vivid reconstruction of Australia’s relatively recent but mysterious age of marsupial megafauna.
    These giant creatures established themselves as the continent became colder and more arid. They dominated during the most recent Ice Age into the Pleistocene Epoch but were then subject to rapid mass extinction, a process that began about 70 000 years ago and ended when the last of them died away about 20 000 years ago, though these time-spans are as controversial as the reasons put forward to explain the extinctions.
    Sarcophilus laniarius is the devil species found in the Naracoorte Fossil Chamber. It was about 15 per cent larger than a modern devil, making its body mass about 50 per cent greater. But caution is necessary. ‘The relationships between the living Tasmanian Devil and the larger Pleistocene form are in doubt . . . The living animal may either be a dwarfed version of S. laniarius or possibly a different species that coexisted with the latter.’ 5 It was the eminent nineteenth-century palaeontologist Richard Owen (who discovered and classified S. laniarius ) who originally proposed the idea of different coexisting sizes, based on fossils discovered in the Wellington caves of New South Wales in 1877.
    Giant devil bones have also been found in Queensland, Western Australia, New South Wales and Tasmania. The earliest fossil evidence is from the Fishermans Cliff locality in southwestern New South Wales, where the species is described as S. moornaensis . The first appearances of S. laniarius are in a fossil deposit in the eastern Darling Downs of southeastern Queensland, and in the Victoria Cave deposit in South Australia. Dating these sites is difficult, but the species certainly was present between 70 000 and 50 000 years ago. The Mammoth Cave site in Western Australia, where S. laniarius has also been found, may be as old as 70 000 years. The Devil’s Lair cave deposit in Western Australia is dated at 11 000 to 30 000 years old and shows evidence of both devils and Aboriginal inhabitants. More recent deposits from the Holocene Epoch (the past 10 000– 11