Scottish wildcat history and evolution including miacids, proailurus and Martelli's wildcat

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History and evolution of Scottish wildcats...


The precise history of the felid family is open to plenty of debate, no one knows the exact story although the fossil record and genetic profiling make it possible for us to start piecing together an outline. The following information is intended as a general guide to the evolution of the cats within the history of the Scottish wildcat in particular. Those perhaps looking for the short and sweet answer to where the Scottish wildcat came from would probably prefer to focus on the next paragraph alone...

The short version

9000 years ago, at the end of the last ice age glaciation, the English Channel formed and isolated a group of European wildcats in the British Isles. Over time they evolved unique characteristics in response to the particular habitats and species resident here growing bigger and more heavily camoflaged. As human led deforestation changed the face of Britain and the species was hunted close to extinction across England, Wales and most of Scotland it evolved beyond its forest led behaviour to be able to hunt over the full range of habitats available to it, and developed a greater mistrust of mankind than any other animal. Today, the small number of wildcats that survived deforestation and human persecution are now heavily outnumbered by human introduced feral domestic cats. The two species are closely related and readily mate, causing the wildcat genes to slowly water down and disappear into the huge domestic genepool. Recent conservation efforts have been led by scientists, naturalists, international grants, commercial sponsorships and the interested public, with support from government and statutory agencies proving slow and, to date, ineffective.

That, however, is just the bare facts and only a glimpse at this remarkable cat's full story...

Artist's impression of a miacid, the common ancestor of all carnivore species. by Sally MacLarty (Cats of Africa, Struik) The long version

Miacids (65-25 million years ago)
Pictured right
As the age of dinosaurs came to an end a vacuum formed for a new dominant life form on the planet. The mammals were quickest to respond and species quickly evolved to fill the various ecological niches. One particular group of creatures called miacids were well adapted to forest life with a long tail for balance, strong claws for climbing and teeth specially developed for eating meat; they were one of the earliest steps in the carnivore chain and would eventually evolve into all the modern carnivores; cats, dogs, bears, even seals.

Artist's impression of proailurus, the first true cat, artist unknown Proailurus (30+ million years ago)
Pictured left
Proailurus first appeared somewhere in the Eurasian forests. Larger than a typical miacid it had even more specialised teeth and fully retractable claws enabling it to hunt on the forest floor whilst keeping its claws razor sharp for climbing and killing. It inherited superb climbing skills, a balancing system suited to three dimensional arboreal life and the ability to always land on its feet; it was the perfect form for an animal that could hunt in tree canopy and forest floor, proailurus literally means first cat.

Artist's impression of felis lunensis, Martelli's wildcat, one of the first of the felis species, artist unknown Felis Lunensis (10 million years ago)
Pictured right
Proailurus developed into a range of cats both large and small that spread across Europe, Africa, Asia and the Americas via a land bridge, constantly evolving new species to deal with each new habitat and prey encountered. By 10 million years ago some of the first small cats that we might easily recognise today began to appear, including the first wildcat Felis Lunensis (Martelli's wildcat). Long since extinct its bones have been found across Europe including Britain which would have been part of the mainland at the time.

Felis Silvestris, the modern European wildcat, photographer unknown Felis Silvestris (2 million years ago)
Pictured left
Some evolutionary refinements led to Lunensis making way for Felis Silvestris; the European wildcat, which is still with us today in the remaining patches of European forest (today they are called Felis Silvestris Silvestris to avoid confusion with the other wildcats). By now the cats were perfectly adapted to life in a dense wild forest with excellent camoflage, night vision and hearing honed to perfection over the millennia. A true superpredator capable of tackling anything its own weight, and plenty of things larger, the species was highly successful and spread across the continent.

British Isles during the last ice age glaciation; ice sheets extended to London and lowered sea levels to create a land bridge joining the islands to mainland Europe

The last glaciation (20,000 years ago)
The European wildcats lived through times of great climactic change with ice age glaciations periodically relocating them south where they would eventually evolve into the various Asian and African forms of the wildcat; variously striped and spotted with thinner coats and less bulky bodies better adapted to life near the warm equator. Members of these wildcat bloodlines would also begin developing into the domestic cat years later around 10,000BC. Just before that, around 20,000 years ago, the last great glaciation began and kilometre-thick ice sheets spread as far south as London (pictured right), with the tree line retreating all the way to the Mediterranean. The wildcats were driven south until around 10,000 years ago when the ice began to retreat and the trees and forest animals could return to Europe.

The British Isles (9,000 years ago)
As the ice retreated and the European wildcats returned home, this time joined by stone age man, sea level rise was also happening as a result of all the melting ice. The English Channel formed cutting off Britain and part of the wildcat population forever. Densely forested at the time the wildcat flourished on the island and, isolated from the European genepool, took on its own specialised form; growing big (one skeleton discovered measured 4 feet from nose to tail and record keepers related stories of them reaching 5 feet) with a thick coat and tail to protect it from the ever changing weather.

The badge of the Clan Chattan

The Roman invasion (2,000 years ago)
Under the Romans cities grew larger, people more numerous, and hunting became a sport as well as a necessity. The domestic cat was introduced in larger numbers whilst wild carnivores like the British lynx were pushed to the abyss of extinction and fell over not long after the Romans left. Wildcats were also prized for their wonderful fur though somehow evaded extinction, learning quickly never to trust mankind.

Scotland at the time was ruled by the Picts, an artistic but fearsome race that scared the Romans so much they constructed the 80 mile long Hadrian's Wall and the 40 mile long Antonine Wall to keep them out of the Empire. Picts seem to have worshipped animals and spirits and in the area now called Caithness ("Land of the Cats") lived the Catti tribe; legend states that on first landing in Scotland they were attacked by wildcats which they barely managed to overcome and were so impressed that the wildcat became their symbol, for a time the whole of the West Highlands was called simply "Cat".

Thousands of years later the Celts arrived in the form of Clan Sutherland, equally impressed by the wildcats that they boasted ensured you couldn't find a rat anywhere in the region (rabbits only arrived after 1066) they also still have a wildcat as their leader's crest. The Sutherland's weren't the only Celts to find inspiration in the Scottish wildcat either, around Strathspey and the Cairngorms you can find a confederation of clans called Clan Chattan ("Clan Cat") which includes clans such as Mackintosh, Macpherson, Macbean and MacGillivray all proudly displaying wildcats in their heraldry (Clan Chattan's badge shown above).

Scottish wildcat fights pack of terriers, hand coloured engraving from 1860 The Middle Ages (500 years ago)
Chroniclers recorded a still extensive woodland well populated with wildcats as recently as the Middle Ages but changes were on their way; the agricultural and industrial revolutions ate up the forests and began polluting Britain's natural resources whilst numbers of people and the size of towns and cities exploded. In Scotland even the Highlands was no longer safe from modernisation after the failed Jacobite rebellions; forest-devastating industries such as shipbuilding rapidly expanded. Wildcats were still hunted for their fur and also became classed as vermin accused of annihilating stocks of lamb, game birds and chickens as part of an anti-wildcat propaganda campaign that still lurks today. Modern research has strongly suggested that pure wildcats had little to do with it and most of the conflict emerged from hybrids with their reduced fear of mankind inherited from domestic cat genes. The engraving (above) from 1800 perpetuates the wildcat's almost mythological fierceness; terriers lining up to be swiftly dispatched, leaving it up to the mad axeman to apply the coup de grace. Throughout this time land development, persecution and hybridisation were having a terrible toll and the wildcat gradually began to disappear from England, Wales, Ireland and southern Scotland.

Victorian Britain (100 years ago)
Persecution reached a peak in the Victorian era with the advent of sporting estates where the natural landscape was altered by mass deforestation to create heathland habitats ideal for raising game birds and farming sheep. And whilst the same modern research suggests most conflict came via hybrids, wildcats were firmly blamed with eating into the profits of estates and farms. Gamekeepers were paid a bounty for every wildcat killed, and the total amount killed in a year was used as a yardstick to gauge their ability to do their job. Even 50 years ago keepers would hang wildcat carcasses out on a gibbet to show the landowner that they were doing their job, all the time propagating the myth of the cats destroying stocks of precious game birds.

A fearsome looking Scottish wildcat with black grouse, by A. Thorburn, 1902 Museums of the time paid keepers a bounty for bringing them in the largest specimens of dead wildcat to be stuffed and put on display, invariably in wild snarling poses with oversized sets of fake teeth to terrify polite society with. These same skins finally had some use in modern research into hybridisation and wildcat genetics revealing a very low proportion of killed cats to be true wildcats even 150 years ago; they were either very good at dodging bullets or rare visitors to the human altered landscape. Whatever impact persecution had, hybridisation was undoubtedly pushing the wildcat hard; it can be seen taking a stronger and stronger hold in the coat markings of these skins. Around this time though, some relief came, unexpectedly in the shape of the First World War. With all able bodied men called up to fight, the human persecution of wildcats dropped off significantly, and after the war the UK found itself in a very different economic climate; sporting estates began to break up and increasing attention turned to replanting the forests.

Mike Tomkies pictured in 2005 from the film Last of the Scottish Wildcats

Late 1900s
By now confined to Scotland the cats were known as Felis Sylvestris Grampia or the Scottish wildcat. Defined as a subspecies of the European wildcat due to its larger size, stronger markings and its ability to make use of all the habitats available to it, the classification has provoked a great deal of debate as to whether it is a true sub-species or an isolated island population of the European wildcat, currently the official classification is island population, though many argue in favour of switching back to a true sub-species.

Myths of man-killing wildcats still persevered, in his 1950's book 'British Wild Animals' naturalist H. Mortimer Batten clearly disliked the animal, citing a Scottish crofter and a shepherd being killed by female wildcats in separate incidents, and describing the best way to trap and kill them, much in line with natural history books in the 1800s talking of wildcats killing armed gamekeepers. Wildcats were even suspected of hanging from branches by a hook at the end of their tails and dropping onto unsuspecting passers by!

In the 70's the author Mike Tomkies (pictured above), who spent much of his life living in the Scottish wilderness, came to share his life with a group of wildcats and hybrids over nearly a decade and wrote several books of his observations telling of a fearless, honest, intelligent and cautious cat, spectacular in its survival skills, terribly threatened by the actions of man and more than worthy of our respect and protection. Mike was also the first to carry out research on those museum skins, trying to get a handle on hybridisation and noting the gradual impact in coat markings and body shape the hybridisation was having, without question he inspired the modern Scottish wildcat conservation movement that would emerge in later decades.

Wildcats were finally offered legal protection in 1988, though debate was to rage as to whether the specific legislation was fit for purpose, what exactly threatened the wildcat, how many of them there were and how best to save them, all covered in more detail on the Scottish wildcat conservation page.

Wildcat Haven logo

Post 2000
Official conservation efforts via Scottish Natural Heritage and their various partners can be largely summarised as taking a hands-off approach that preferred to look at public awareness and PR activities rather than direct fieldwork, whilst many other scientists and conservationists tried to push for more direct action. The frustration led to independent efforts to rebuild a captive breeding program from 2000 and establishment of the Scottish Wildcat Association in 2007. The charity quickly secured funding from the US and China to begin efforts in the field in what would become the Wildcat Haven conservation project, named after the Tomkies book and based not far away from where he had lived.

Various scientists identified hybridisation as the primary threat and estimates of the Scottish wildcat population were made as being around 400 individuals by Oxford University in 2004 and just 35 by the SWA in 2012. Official SNH efforts were still claiming there were as many as 4,000 in 2012, in spite of spending half a million pounds of public money trying to prove it without managing to find a single pure wildcat, or for that matter managing to remove a single feral cat from the wild or complete any piece of research that didn't have to be redone; we are currently on the fourth population survey which has already had its methodology questioned in peer review, and claims that genetics research is being carried out have been made since 1998, finally bearing fruition in independent hands at the University of Chester after just two years of research.

Today, there continue to be two stories to wildcat conservation which are often muddied as many experts attempt to remain impartial and see the best way forward to support both sets of opinion. Government backed efforts continue to focus on public awareness activities and the recent SNH Scottish Wildcat Conservation Action Plan has gone as far as re-classifying wildcats with a relaxed definition that allows for significant hybridisation. Whilst such a methodology effectively condemns the pure wildcat to extinction, it also means that government efforts cannot fail to 'save' the 'wildcat', so long as the majority of the general public fail to appreciate that not every brown tabby is a wildcat. The independent efforts continue to circle around the Wildcat Haven project which is focused almost entirely on fieldwork, recognises that there are probably less than 100 pure wildcats left, and aims to save those pure wildcats and end hybridisation entirely.

Few people outside of the SNH PR department or those organisations that benefit from associating closely with SNH believe that the true Scottish wildcat has much time left, or that it is likely to exist outside of the West Highlands region. The clock is ticking down on estimates by numerous conservationists over the last decade that the Scottish wildcat could be gone forever around 2016, just as the wider public is beginning to appreciate what it may be about to lose...

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