turbinal bones that magnify smells, the nuchal ligament that strengthens their necks to allow them to run for extended periods with nose to the ground – the physical hallmarks are unambiguous. The fox is a dog.
Or very nearly.
THERE WAS ONCE another forest, more remote to our lives and fainter than Białowieża’s leathery trunks: an ancient jungle that no human saw, in a global climate hotter than we have ever known. Fifty million years ago, the so-called Eocene hothouse featured Earth at its balmiest, basking in temperatures averaging 14 °C (57 °F) higher than present. Ice vanished from even the poles, and lush forests extended across the globe. Palm trees grew in what is now the London basin; extinct primates foraged under them, and the peculiar horse-like ungulate Hyracotherium browsed in their shadows – but there were no foxes. Canids had not yet left their family cradle.
That birthplace was North America; perhaps dogs are the United States’ most successful export. Canids have spent the majority of their evolutionary history restricted to this one continent, and the distant ancestors of foxes are dated to the Eocene jungles of Texas. Prohesperocyon – the first known canid – was a small, omnivorous creature in a forest of astonishing giants, a natural neighbour to the likes of the fearsome Nimravidae, sabre-toothed carnivores resembling great cats. As Earth gradually cooled and dried and the dense jungles were replaced by extensive grasslands, the elongated legs and long strides of the canid body shape clearly provided an advantage in long distance chases. Canidae thrived.
Most of the fox’s extended family is extinct, and known to us only through palaeontology, but the glimpses defy imagination. Among them is the subfamily Borophaginae, containing species that might frighten Cerberus – they were the bone-cracking dogs, with massive teeth capable of extracting marrow from giant carcasses. Epicyon haydeni, the largest of all, is estimated to have weighed four times as much as an average grey wolf, and a pack on the hunt must have been a formidable sight. Yet the dentition of many fossil canids, not least the little Leptocyon – the direct ancestor of the fox – strongly suggests a varied, plant-inclusive diet.
Plate tectonics and the reduction in sea levels during ice ages eventually connected North America to other continents, and its wildlife migrated into South America and Eurasia. By seven million years ago, at least one canid was present in Spain, and Vulpes riffautae – the earliest known fox outside North America – lived in what is now the N’Djamena desert in northwest Chad. Fast forward another few million years, and Vulpes galaticus is part of the Turkish fauna. Vulpes vulpes, the red fox – the only species that we have in modern Britain – is recorded first in Hungary, perhaps 3.4 million years ago, when humans in Africa were just beginning to use stone tools. Recent genetic analysis has provided further hints; it appears that all living red foxes are descended from individuals who lived in the ancient Middle East. From there, they spread across the entire northern hemisphere.
Canids travel. Their long legs and unfussy diets enable them to colonise new habitats with ease. Nature is not a fixed condition, and Britain has experienced many waves of colonisation and extinction over its geological history. But from the perspective of a wild animal, one of the most significant qualities of our island is that it lies rather far to the north. Much as we complain about the weather, it is remarkably mild for a country on the same latitude as Moscow. Take away the Gulf Stream, and it would be time to buy some serious winter clothing. Add all the geographical and solar phenomena which regularly cause the world to have ice ages, and the Thames Valley becomes cold, hard tundra.
I have tried to breathe normally in temperatures of -35°C (-31°F), on a day in Alberta when humans seemed disinclined to be outside. Gulping such air is like swallowing swords; your lungs baulk at the freezing blast, yet it is nothing compared to the frigid temperatures reached in the wind-chill from a continent-wide ice sheet several kilometres thick. The Pleistocene Epoch played a long game of catch-and-release with Britain, repeatedly coating the northern half of the country with dense ice and then releasing it in warm interludes called interglacials. We live in an interglacial now called the Holocene. It has lasted nearly 12,000 years, but it probably won’t continue forever.
© Red fox skulls and jaw and jaws from Ightham Fissure, near Maidstone. Plate III in Sidney H. Reynolds, A Monograph of the British Pleistocene Mammalia. Vol. II, Part III: The Canidae, London, 1909 (Wikimedia Commons)
A study of red fox bones. This one lived during the Pleistocene in what is now Kent.
Our wildlife has been dictated by ice. The fossil record suggests that red foxes appear in the interglacials, historically alongside a wealth of other creatures that would monopolise attention if glimpsed on safari in Africa. Our first British foxes perhaps scavenged on the carcass of a straight-tusked elephant predated by cave lions, and certainly would have heard the whooping laughs of spotted hyena. The next time you wonder why a fox sits and watches you rather than bolting in panic, remember they have had to judge the risk from very dangerous predators for thousands of millennia, and their evolved strategy is waiting at a safe distance with access to a known safe spot, such as a den or – these days – a gap in a fence. If they had run further than required from each Pleistocene sabre-toothed cat, European jaguar and cave bear, the energy wastage would have crippled them.
Meanwhile, Arctic foxes – along with woolly mammoths, wild horses and reindeer – were present during the colder times. Red foxes disappeared from Britain entirely, surviving in the relatively mild refugia in Spain and the Balkans. Sometimes, while walking in my native Surrey Hills, I try to imagine what those glacial millennia must have been like. The glaciers never extended this far south, but the bite from the wind must have been excruciatingly bitter, and the landscape would have been an austere mixture of bare rock and frozen snow. Fed by the ice sheet were huge rivers of cyan-blue glacial meltwater.
And foxes must have drunk from them. But were they the Arctic or red species?
Over in Somerset, one cave dated to 12,000 years ago contains fossils of both, but red foxes have always returned north and displaced their smaller Arctic cousins in times of mild climate, and they continue to do so on the modern thawing-line of Sweden. In any case, while lions and hyenas did not migrate back to Britain after the ice retreated, foxes rapidly did, even as tundra budded with crowberry bushes and mugwort, and finally grew trees once more.
SO THE FOX trotting across a Clapham street is directly descended from individuals that encountered species wondrous beyond our most outlandish fairy stories, survived extremes of climate that we have never known, and crossed land bridges long lost beneath the sea. Human culture is such a late entry into the story of the fox that it would seem disingenuous to mention it – except, of course, we have a strong bias towards it.
No one will ever know where the first Homo sapiens laid eyes upon a living fox, or how the two species perceived each other. As pre-history continues, our fossils and theirs begin to overlap in palaeontological sites, a silent testimony to forest meetings that have passed into the veil of unwritten time. But 16,000 years ago, when Palaeolithic painters were drawing steppe bison in the Spanish cave of Altamira, a woman of unknown name died in what is now Jordan, in a site called ‘Uyun al-Hammam. Her body was laid among flint and ground stone, and a red fox was carefully placed beside her ribs, resting with her for eternity on a bed of ochre.
We cannot perceive the meaning. Was this a pet, or an animal kept for its ceremonial significance? The care in the joint burial is believed to suggest some emotional link between human and fox, beyond that shown to wildlife perceived as food or clothing. It has been speculated that these pre-Natufian people coexisted with foxes that were at least half domesticated. Perhaps they scavenged rubbish on the edge of camps, along with the earliest dogs. Perhaps the behaviour so often complained about in London is more ancient than we think.
In any case, it is clear that foxes held a strong cultural significance for the later peoples of the Levant. They are commonly found in human graves in Kfar Hahoresh (modern Israel), dated to around 8,600 years ago, while stone carvings of foxes with thick brushes adorn the pillars of Göbekli Tepe in Turkey, believed to be the world’s oldest temple. In Mesolithic Britain, humans who hunted deer by the