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A Brief History of the Fox

WE ARE REDESIGNING THE FOX: its diet, terri­tory size, social interactions, and its longevity and causes of death have all been changed by us. Even their body fats are impregnated with our lifestyle, carrying residues as diverse as fire retardants and nuclear radiation. Their days are filled with human-made noises, human-made landscapes, and human-made risks.

But foxes have not spent their evolutionary history sunbathing on greenhouse roofs or evading aggressive pet cats, let alone treading on broken glass or eating left­over pizza. Wild nature has been twisted out of joint in Brit­ain; except for the lonely saltmarshes of the north Nor­folk coast, very little has not been reshaped by our finger­prints. But to understand the fox among us, we must first consider the world as it was before.

I AM FOLLOWING a wild wolf with a hind foot as wide as my hand. The paw print is written in the soft clay of a path flanked by wrinkled old trees that take circuitous routes to the sky, their branches bending under a red squirrel’s leaps, their bark festooned with furry moss. Looking past their trunks, I see more trees, and yet more: shadows of green upon green, and woodpeckers laugh from within. There is not a sound nor sight save of natural things – just tree frogs purring at dusk, and pure, sweet, forest air. Where the canopy has been opened by a giant’s fall, new saplings race to the light over the carpet of wild garlic flowers. Deer consume many such infant trees and are themselves taken by wolves; the leftover bones fall to foxes.

This is Białowieża, lowland Europe’s closest match to a truly wild forest. It mantles the border of Poland and Belarus – a living palace of oak and hornbeam, and a mortuary of naturally dead trees that sport brilliantly coloured fungi, nourishing new life where they crumble. Old growth or primeval forests are relatively widespread in North America, although their fate is the subject of bitter battles between loggers and environmentalists. On the European Plain, we only have Białowieża, and its history has been uneven, from bison-hunting Russian Tsars to the Nazis who murdered Polish patriots under gently shadow­ing leaves.

Yet today, the rails laid by Germans to export timber in World War I are overgrown by wild pansies and chick­weed. Disputes over logging in the buffer zone aside, the forest remains largely in control of itself, as it has been for most of the last 8,000 years. Human tragedies, triumphs and the entire Roman Empire have risen and fallen, and, all the while, Białowieża has quietly evolved along its own lines, its vast compendium of living things predating, competing, and joining into symbiotic relationships with each other. It is the last benchmark: a reference point that explains how European foxes are when the human touch is light.

That path where I found sign of the wolf’s wander­ing is also a datasheet of information. The damp ground reveals more tracks – of wild boar, roe deer, red deer, bison – and here, not three metres from the wolf’s trail, a fox foot­print.

I could probably fit all four of its little paws into the track of its distant relation.

It is living here as a pure wild animal, sustaining itself on other life supported by the forest, and when it dies, its flesh will grow more trees.

What is a fox in such a place? I’ve caught glimpses on my camera traps: visually indistinguishable from foxes in gritty south London boroughs, they trot confidently down trails trampled by gargantuan bison bulls. They balance deftly on fallen tree trunks beside rivers that flood when they please. They lope across tracks at night, under crystal­line stars untainted by light pollution.

Białowieża

There are questions that all wildlife biologists ask of their study species, and most of them would take a lifetime of research to answer. Where do they live, and how are they distributed within those areas? Do the regions with the highest activity have characteristics in common? What about their social interactions with their own kind – do they defend territories? When do they breed, and how far do the young disperse? In a forest with uncountable forms of life, with which of the plants, mammals, birds, reptiles, and invertebrates do they interact as predator, as prey, as disperser of seeds, as bringer of change through some-­ thing as subtle as digging the mud to create a den?

Unravelling these mysteries requires deciding a re­search hypothesis, collecting data to test that hypoth­esis, and exploring those data with statistics or maps. I have no recourse for such ventures during my holidays in Białowieża. But I do spy some clues: fox scat, for example, the ubiquitous silent witness to their diet.

It is full of the bristling black hairs of wild boar.

Boar are as far beyond a fox’s hunting prowess as is an elephant – fourteen times heavier, and armed with tusks and teeth. They are quite capable of fending off leopards. Yet here in a near-pristine forest, they comprise a startling portion of vulpine diets.

The link is the wolf. About 30 per cent of the fox’s winter diet in Białowieża is carrion, specifically red deer and wild boar. Some may have starved, but many were killed by wolves and lynx. Despite the popular image of wolves as marauding Vikings, they inadvertently assist many other creatures. Ravens, martens and eagles are among those which feast on the wolf’s leftovers.

The beauty of this forest is its completeness; nothing is wasted, and ecological relationships almost forgotten in Britain’s radically altered landscapes shine clear.

Perhaps the message from Białowieża, then, is that the wild fox is an interactive creature. They are part of a living web that cycles energy from plants to herbivores to predators to scavengers, a natural dodgems in which spe­cies knock into each other. Sometimes that brings death to one and food to another; sometimes there is mutual gain. Wherever it is, whatever the backdrop, a fox can never be understood outside of its relationship with the rest of the natural world.

But what is a fox in the first place?

A FOX’S SKULL, like a wolf’s footprint, fits neatly in my hand. In its way, it is a birth certificate, a genealogical sign­post that reveals the fox’s relationship to other mam­mals, living and extinct. Not for nothing did Victorian scientists spend so many hours measuring bones; patterns emerge from meticulous study, and the bewildering array of ver­tebrate life can be organised into logical groups. In recent decades, genetic analysis has refined our understanding.

So imagine an Ark with all the world’s animals entering two by two: birds, reptiles, amphibians, and other, more exotic creatures. A fox would enter the deck reserved for mammals, for it has fur and suckles its young, along with a diaphragm and a neocortex. But a fox is clearly differ­ent from its mammalian relatives. It is certainly not a bat, giraffe or horse. To divide mammals logically, Noah must look at their jaws. Teeth are not only clues to diet, but also indicate who is related to whom.

Foxes have 42 teeth inside their narrow muzzles, among them the trademark of the order Carnivora. Their last upper pre-molar and first lower molar are called carnas­sials, modified with a tall shearing edge for slicing flesh. All 300 or so Carnivorans possess carnassials. Your pet cat has them – in fact they are exceptionally well developed in felids, which are the most meat-specialised of the entire order. Pandas, on the other hand, have flattened carnas­sials because they are largely vegetarian.

What about foxes? Their carnassials are specialised for nothing. Berries, mice, insect larvae – they can eat it all. That adaptability is the main reason why the wild dog family has been such a runaway evolutionary success.

Foxes are of the canine kind. Many fox-watchers, including myself, enjoy the cat-like grace of the species: the delicate pounce and careful footsteps. But beneath that rich orange fur, foxes are undeniably a canid – that is, a member of Canidae, the dog family. Turning the skull over, the base of the ear sockets is fused into a bony casing called a tympanic bulla, protecting the fragile inner-ear bones. In canids it is uniquely divided by a septum: a thin, bony wall. It is believed that this extra echo-chamber enhances their ability to hear low frequencies.

Five toes with claws that are not fully retractile, a long muzzle filled with delicate 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 shore of extinct Lake Flixton – in the North Yorkshire archaeological site of Star Carr – must have been aware of their small red neighbours. Bones from two foxes have been found at this ancient settlement, along with those of Britain’s first known domestic dogs, but there is no indication of what role, if any, canids played in their culture.

© Prince Hanzoku Terrorised by a Nine-Tailed Fox, Utagawa Kuniyoshi (1798–1861) (Wikimedia Commons)

A nine-tailed kitsune in nineteenth-century Japanese art.

Later, as humanity discovered the joy of story-telling, foxes joined the cast. The oral literature of native Ameri­cans occasionally opts for a fox as a trickster, albeit a potentially handy one; according to one Apache legend, it was Fox who stole fire from the fireflies and introduced it to Earth. It is across the Pacific in Japan, however, that fox folklore reaches its most astounding heights. Kitsune – the revered fox of Japanese myth, poetry and traditional belief – has existed in human thoughts for many centuries. It even makes an appearance in what may be the world’s oldest novel: the eleventh-century epic The Tale of Genji, where a human character debates whether the figure by a tree is a woman or a shapeshifting vulpine. Kitsune delight, deceive and confuse in countless other legends; while the theme of pretending to be an attractive woman is frequent, other tales relive how they mislead travellers by light­ing ghost fires at night, assume the form of cedar trees, or even become the guardian angels of samurai. Today, anime writers continue the kitsune tradition.

BACK IN EUROPE, by Roman times the uneasy relation­ship between foxes and agriculture had woven itself into religious rituals – in the festival of Cerealia, for example, live foxes were released into the Circus Maximus with burning torches tied to their tails. Seven hundred years later, Aesop’s tales also provide a nod to fox interactions with farmers, and – to a lesser extent – with their neigh­bouring wildlife. My favourite Aesop fable features a wolf taking a fox to court for theft; given the vast quantity of wolf-killed carrion that real foxes consume, it seems vaguely reasonable.

Old English literature picks up similar themes. The Fox and the Wolf, a rhyming poem from the thirteenth century, stars a fox who helps himself to some chickens and then tricks a wolf into taking the blame:

A fox went out of the wood

Hungered so that to him was woe

He ne was never in no way

Hungered before half so greatly.

He ne held neither way nor street

For to him (it) was loathsome men to meet

To him (it) were more pleasing meet one hen

Than half a hundred women.

He went quickly all the way

Until he saw a wall.

Within the wall was a house.

The fox was thither very eager (to go)

For he intended his hunger quench

Either with food or with drink

And so it continues, with the hungry fox trapping him­self in a well before deceiving a wolf named Sigrim into taking his place. Ironically, this poem was written about the same time that the wolf’s howl was finally falling silent in southern Britain.

Did the fox notice the disappearance of its distant rela­tive? Perhaps, unconsciously. As shown in Białowieża and elsewhere, the wolf was a provider as well as rival, a power­ful force in the wildwood whose absence has changed these islands as much as a spoke missing from a wheel. Some species have sharply increased, and others have probably declined.

Yet civilisation has done more than simply rip out culturally troublesome natives while boosting deer and grouse for hunting. We have released millions upon mil­lions of non-native animals into the countryside: rabbits from Spain, fallow deer from Persia, sheep from Mesopo­tamia, hens from south-east Asia, cats from Africa. Our trading ships accidentally added black rats from India and house mice from the Middle East, while American grey squirrels, Japanese sika deer and even Australian red-necked wallabies joined our countryside from zoos. We have persuaded ourselves that the six million sheep of Scotland are part of the ‘natural’ scene, but the Highland ecosystem evolved with none. Even the Scottish red deer population of 300,000 is far higher than in the time of the wolf. These changing grazing pressures affect the rodents and berries that foxes eat, and near-total deforestation has altered their territory sizes and feeding habits.

In a flash of geological time, we have rewritten the fox’s wildwood, in ways both graphic and subtle. We have added, taken away, replanted and concreted.

And the fox that once played its natural dodgems with the rest of the natural web will inevitably interact with the components of the new Britain that we have designed without ecological aforethought.

The fox is not an intruder into our world.

We have simply laid our modern ambitions over the landscape it already knew.