Housecats and Wildcats (Fluffy and Silvester)

Ever wondered if different kinds of housecat – like a tabby and a Persian, or street cats and purebreds – are different species?

Biologists say that these animals all have the same formal last name (for species or subspecies) – catus.

So it’s all one group despite appearances or lifestyle.

The biologists then politely excuse themselves and get back to work, because the real challenge is telling housecats and wildcats apart, and they’re working under a wildcat conservation deadline.

What’s a wildcat?

Well, it’s not that beautiful animal at the top of this post. That is a housecat – a pedigreed Norwegian forest cat, in fact. It’s a catus.

Here’s a wildcat, photographed in a German national forest:

800px-European_Wildcat_Nationalpark_Bayerischer_Wald_02
It won’t follow you home, no matter how friendly and gentle you are. European wildcats are mostly untameable. (Sunquist and Sunquist, 2002)

The resemblance between these two particular cats is eerie!

Fluffy the Housecat and Silvester the Wildcat don’t always look that much alike. Cat fanciers have worked hard to maintain a wildcat appearance in the Norwegian fancy-breed.

It’s quite an achievement. The breeders are limited to the human time scale, while European wildcats have had hundreds of thousands of years – since roughly the middle of the Pleistocene – to develop their look. (Yamaguchi and others, 2004)

Wildcats are formally known as Felis silvestris. (It’s usually abbrevated as F. s..)

So, even though the cats above are almost mirror images, they belong in different subgroups – catus and silvestris, respectively.

Still, Nordic “twins” aside, housecats and wildcats do have a lot in common.

Flying wildcat
Spot quiz: Domestic cat, wildcat, or a new Marvel character? The bushy ringed tail, with its blunt tip, clues us in that this is a Scottish wildcat. Actually, its nickname – Highland Tiger – would make a good Marvel character name!

Fluffy and Silvester deserve their own post in this series on cat evolution because:

  • Obviously, they’re closely related. But some of the best minds on the planet can’t agree whether domestic cats and wildcats are two different species (Felis catus and Felis silvestris) or just subgroups of the same one, i.e., F. s. catus and F. s. [wildcat subspecies name]. It’s just words to us, but the uncertainty can really mess up legal efforts to protect wildcats. (Macdonald and others)
  • Our own history is part of the story. Cats have associated with human beings for at least ten thousand years. (Vigne and others, 2012) During that time, we’ve carried Fluffy all over the world. And in the last century and a half, we have also dramatically changed its appearance through a system of artificial selection called the cat fancy.
  • Housecats and wildcats interbreed so much that it’s almost impossible to find “pure” wildcats in some parts of Europe. This prompts warnings that wildcats are at risk for genetic extinction. But doesn’t interbreeding show that catus and silvestris are the same species? And how can having kittens make the parents go extinct? The biologists are working on these and other questions.

Let’s start off with the cats closest to us – the few that are fancy-breeds as well as the many unpedigreed domestic cats that share our homes, yards, and streets.

Then we will get a little better acquainted with Silvester and see how some wildcats might have been domesticated.

After a brief outline of Fluffy’s history, we’ll check out the issue of genetic extinction – what it is and what it could mean for people – especially cat owners – as well as wildcats.

Feral cats, fancy breeds, and housecats

You aren’t imagining things – there really are a lot of cats out there, 600 million of them associated with households and another 600 million unowned, roaming and reproducing freely (Driscoll and others, 2011), i.e., feral cats. (Robertson)

“Feral” does mean “wild,” but these aren’t wildcats. Wildcats are, well, wild – feral cats were once domestic or else have very close relatives that are domesticated.

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Some cats haunt the borderland between feral and domestic.

Feral cats exist wherever people live or have lived. It’s a worldwide problem, and one that we humans have caused. (Robertson)

But in this post they are only relevant to the wildcat conservation issue, so we’ll talk about feral cats later.

What about the hundreds of millions of owned cats? Who really owns whom?

In other words, are housecats really domesticated?

Cows, sheep, and most other domesticated animals depend on us for food and shelter, and we control their breeding. (Driscoll and others, 2009)

This isn’t true of housecats. Fluffy can live quite well outdoors on its own, supplementing store-bought cat food with local prey and, given the opportunity, breeding with feral cats and wildcats.

It’s only the pedigreed cats that need people to keep them going. Some breeds also have physical changes – like hairlessness or very flat faces – that make a life in the wild impossible for them.

So, technically, only fancy-breeds have been completely domesticated. The rest are still working on it. (Bradshaw and others)

Of course, that’s an extreme definition of “domestication,” though it is scientifically correct. Housecats really are only semidomesticated.

But people have been breeding fancy cats from housecats since Victorian times.

This is a very recent phenomenon compared to dog breeding. Cat breeding probably got such a late start because it’s only done for looks, not for any practical reason. (Lipinski and others)

Egyptian and cat
No one today expects cats to fetch things. Did the ancient Egyptians train them?

Cats in the fancy are still close to their household roots, while even stray dogs, in developed countires, have purebreds in their background. (Kurushima and others)

If pedigreed cats come from ordinary cats, why is there so much fabulous (and occasionally weird) variety on display at a cat show?

A lot of human patience, applied genetics, and luck are certainly involved, but racial differences matter, too.

Cats have races just like we do, and for much the same reason.

There weren’t that many domestic cats at first. They went their separate ways, traveling into various parts of the world with people. Each group then became more or less isolated and had to adapt to different conditions.

This is why, for example, some cats are lithe acrobats while others are ruggedly built powerhouses.

As well, mutations that have given various kitties everything from extra toes to long hair happened in some places but not others. (Lyons, n.d.; Todd)

For instance, a mutation in one group of Asian domestic cats reset the operating temperature for certain pigment genes. As a result, areas on a developing kitten’s fur that are exposed to more warmth before birth get lots of pigment. The result: Siamese cats! (Wastlhuber)

Siamese kitten use
An adorable mutant. Because eye color and coat color are genetically linked, all pointed cats have blue eyes. (Starbuck and Thomas)

Humans played a role in this, too.

Mutations tend to disappear quickly unless some factor allows them to completely replace the usual genes. (Todd)

In that one part of Asia, people liked the dark-pointed look of those cats. They kept and bred more of them, and the mutation became permanent in this group. (Simpson, 1953; Todd)

Geneticists currently recognize at least eight housecat races (Kurushima and others):

  • European
  • Mediterranean
  • Egyptian
  • Iraqi/Iranian
  • Arabian Sea
  • Indian
  • Southeast Asian
  • East Asian

It’s no coincidence that these regions are also human population centers.

Most of the major fancy-breeds come from just four races. Siamese cats, for example, are Southeast Asian. Various Shorthairs come from the European race, Turkish Vans hail from the Mediterranean, and the Sokoke belongs to the Arabian Sea race. (UC-Davis [b])

humanities-1531928_640
I don’t know if this “pied piper” has a pedigree, but there is the Turkish Van’s general coat pattern and its expressive tail.

Some geneticists have actually fine-tuned this to the point where they can say, for instance, that the Norwegian forest cat breed belongs to the Northern European/Nordic subgroup. (Lyons, n.d.)

British shorthairs are part of the same European cat race, but they don’t look like the Norwegian forest cat because they’re in a different subgroup.

Calico brit-480x619
This kitty has a veddy, veddy British look, but the gene mutation for its calico coloring probably originated in Asia Minor or the Indian subcontinent. (Todd; Wastlhuber)

Human intervention has sometimes affected cat races in surprising ways.

Take Persian cats. These luxuriant long-hairs should belong to the Iraqi/Iranian race of cats, and perhaps they did back in the 1500s when traders carried them into Europe. (Wastlhuber)

But when they became very popular, Persian cats were intensively bred for an extreme look, especially in France and England. As a result, they now belong to the European feline race – they have lost their historic genetic signature. (Lipinski and others; Wastlhuber)

As we’ll see later, if something like this ever happens to wildcats because of interbreeding with domestic cats, it will be called genetic extinction.

Things are much simpler with moggies – cats outside the fancy, i.e., most of the cats you see around.

They have the eight genetic races, of course, but apart from us taking cats all over the world, artificial selection hasn’t played as big a role with moggies.

It was the cats with some unusual feature that were usually picked up. The experts call this “novelty selection.” (Todd)

Ship cats in hammock
Besides their striking color pattern, the cat on the left has an extra digit on each forepaw.

As a result, it’s actually possible to trace human movements in the past by mapping some outstanding feature in today’s cats. Usually it’s coat color/pattern, but it can be something else.

For example, the mapping in North America shows clusters of cats with polydactyly (an extra toe) in Halifax and Boston. Geneticists suspect that’s because eighteenth-century Loyalists in Boston took their polydactyl cats with them when they moved to Canada during the American Revolution. Halifax would have attracted them because of its strong commerical ties to Boston. (Todd

As for coat patterns and colors, there’s quite a choice!

Today’s moggies are a mixed lot of long- or short-haired tabbies (look for that “M” marking on Fluffy’s forehead); ticked tabbies (no stripes, but alternating light and dark colors on each hair); solid colored; piebald (black and white); pointed; and black-, gray-, orange-, and/or white-patched cats. (Starbuck and Thomas)

Mapping generally shows the effects of people moving into the city from the countryside. Cats in London and Paris, for instance, have more in common with each other than either group has with rural cats! (Todd)

The maps also indicate that orange cats got established in Asia Minor first and spread westward. Also, black cats (technically, those with”nonagouti” fur coloring) probably traveled west from their original focus in Greece or Phoenicia around 500 BC. (Todd)

The map for black cats (see Todd, page 103) shows them missing from inland Europe, except for the Seine/Rhône waterway. There are also two clusters of black cats, one along the northwestern coast of Africa and another that covers part of the British Isles.

No one has an explanation for this distribution. I wonder if it reflects selective “culling” of black cats in Europe during the “witchcraft” persecution of cats and people during the Middle Ages.

If so, some cats escaped on ships and also, perhaps, found refuge in Scotland, parts of Ireland and England, and northwest Africa.

Scotland was also once considered a refuge for wildcats. As we’ll see later, further research has shifted this view of Scottish wildcats 180 degrees – most of them now have domestic-cat genes! (Rhymer and Simberloff; Yamaguchi and others, 2015)

Wildcats, the species

It’s hard to describe how Silvester and Fluffy are different, apart from their very different tolerances for people and the fact that domestic cats are generally nimbler and come in many more colors than wildcats. (Montague and others)

800px-Felis_silvestris_gordoni
An Arabian wildcat: “Felis lybica gordoni”. The only clue that this is not Fluffy is the black tail tip and rings. If this snoozer was sitting or standing, you would also see the beautiful long legs of a steppe-lineage wildcat. Forest cats are more stocky.

Experts are still debating what specific physical or genetic features define a wildcat. (Yamaguchi and others, 2004)

Starting in the nineteenth century, biologists identified wildcats by location and appearance. That system got so cumbersome – and the basic differences between many species were so few – that in 1951, they simply erected a species name Felis silvestris and then made each of the various wildcat groups a subspecies. (Sunquist and Sunquist, 2002)

What it boils down to is that all wildcats got a third scientific name.

For example, gordoni is the Arabian wildcat’s third name. The European wildcat is F. s. silvestris.

I wonder if European wildcats get named Silvestris twice – Felis silvestris silvestris – because these are the oldest wildcats. They first evolved in Europe some 450,000 years ago. (Kurtén; Yamaguchi and others, 2004)

One of the latest wildcat filing arrangements puts the 21 currently recognized subspecies (Lyons, 2013) into one of these five basic wildcat groups (Yamaguchi and others, 2015):

1. and 2. African wildcats have short fur and the lithe steppe-cat look, with some tabby striping of their legs and body. (Sunquist and Sunquist, 2002)

There are two groups:

  • F. s. cafra is found in southern and southeastern Africa.
  • F. s. lybica lives in northern Africa, coastal Arabia, the Near and Middle East, southwestern Asia, and on most Mediterranean islands. This is gordoni’s group.
Felis_Lybica_Sarda_03
“F. s. lybica”, probably. The wildcat is stalking something, but see those long legs and the ringed tail with a black tip? (Herbst; Pierpaoli and others; but also discussion of Sardinian wildcats in Yamaguchi and others, 2015.)

3.  F. s. silvestris. Europe’s rugged forest cats look massive, but they’re about the same weight as the much slimmer-appearing African wildcats. Their favorite prey is rodents, but they also go after hares, rabbits, and even young deer. (Sunquist and Sunquist, 2002)

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This classic western European wildcat is different from the Nordic wildcat we saw earlier, but both have that powerful, shaggy forest-cat appearance. (Yamaguchi and others, 2015)

4.  F. s. ornata. The dainty Asian wildcat is another steppe wildcat like Lybica and Cafra. Ornata has yellowish fur on its ears and, sometimes, small ear tufts. It’s the only spotted wildcat, with a coat that’s usually short but will vary by season and according to the wildcat’s age. (Sunquist and Sunquist, 2002)

 

5.  F. s. bieti.  This cat has more common names than any other wildcat. This Chinese steppe, mountain, desert, or grass cat is about twice as big as Fluffy and has been sighted only in a small area that includes Sichuan, Shanxi, Gansu, Inner Mongolia, Xingjiang, and Qjinghai. (Sunquist and Sunquist, 2002)

Chinese_Mountain_Cat_(Felis_Bieti)_in_XiNing_Wild_Zoo
“Where’s my lawyer?”

As you can see, Bieti doesn’t resemble the other wildcats very much. There is ongoing scientific debate whether it really is a wildcat or a separate Felis species. (Driscoll and others, 2007; Werdelin and others, 2010; Yamaguchi and others, 2015)

That’s Silvester as experts view it today (and maybe also Felis bieti). Now, which one turned into Fluffy?

Given the strong resemblance, biologists always have felt that it had to be wildcats that got domesticated, not some other small-cat species.

But was it just one of these wildcat groups or several?

The domestication of cats

It isn’t easy to pin down the exact moment when a few wildcats threw in with us – there’s no hard evidence of the event.

Wildcats and domestic cats do have very different color vision, and the European wildcat’s gut is shorter than Fluffy’s for some reason (Kitchener and others, 2010), but only bones and teeth can last during the millennia that have passed since cats were domesticated.

And, as paleontologists who specialize in cats know and lament, feline skeletons – from tigers on down – are almost identical, except for size. (Sunquist and Sunquist, 2002; Turner and Antón)

Wildcats and housecats are about the same size. It’s impossible to tell their skeletons apart. (Sunquist and Sunquist, 2002)

So when archaeologists find ancient cat and human bones together without any other clue of a connection – like paintings or some other sign of human affection for the cat – it’s difficult to prove beyond any doubt that this little feline was a pet. (Driscoll and others, 2007)

One of the oldest such finds ia a 9,500-year-old grave in Cyprus. It contains both human and cat remains, but there is no clear evidence that the cat was a pet. Common sense says it’s likely enough, since cats aren’t native to the island. (Vigne and others, 2012)

After all, who would want to take a wildcat sailing?

The earliest unquestionable proof of domestic cats goes back, of course, to Egyptian tomb paintings, some four thousand years ago. (Lipinski and others; Sunquist and Sunquist, 2002)

Cat stealing egg
The artists also caught a feline egg thief in the act.

 

Both Cyprus and Egypt are in Lybica’s range, and Ornata (the Asian wildcat) isn’t far away. However, for a long time scientists couldn’t find a definite link between any wildcat and housecats.

The big breakthrough came in 2007, when researchers reported that genetic markers show a very close relationship between Lybica and Fluffy. (Driscoll and others, 2007, 2009)

They were even able to demonstrate that the Lybica-group wildcats living in remote parts of Israel, the United Arab Emirates, Bahrain, and Saudi Arabia could be the housecat’s ancestral group! (Driscoll and others, 2007)

The team also found some intriguing genetic clues about how cats were domesticated.

According to these experts, it might have gone like this:

As the world warmed up after the end of the last ice age, some 10,000 to 11,000 years ago, Neolithic hunter-gatherer people settled down on what were then good farmlands in the Fertile Crescent region.

Fertile Crescent
Note that Egypt isn’t part of the Fertile Crescent.

 

As farms spread and villages were established, the first house mice showed up. Then whatever local wildcats could tolerate humans began to exploit this new and very reliable source of prey.

The cats soon became part of the human household, although they kept one paw set firmly on wild ground, so to speak. Competition from other wildcats in the neighborhood would limit how tame they could become. (Herbst)

Now, there is no field evidence whatsoever to back up this story. Geneticists do say that it explains the otherwise mysterious presence and timing of five feline “Eves” that left their traces in Fluffy’s DNA. (Driscoll and others, 2009)

And the events certainly are plausible.

Neolithic people were so good at domesticating animals and plants that scientists call them “the first geneticists” (Driscoll and others, 2009), but even they wouldn’t have been able to catch and tame wildcats for pest control purposes.

Instead, evolution worked it out through natural selection. (That’s a little ironic, considering what a strong role artificial selection plays today in the cat fancy!)

First there was probably a mutation that made some African wildcats tolerant of people. (Driscoll and others, 2009)

It wouldn’t have any effect until people settled down in the wildcat’s range. At that point, cats with this “domestic” mutation would start using human infrastructure as hunting grounds.

This is when another evolutionary factor called “phenotype” – what the cat looks like – would kick in.

If the mutation had happened to a big cat, people would just kill it or drive it away. But Lybica was small and cute, as well as an excellent mouser.

The farmers and villagers not only let it move in, they probably also helped domestication along by offering food treats and selecting the friendliest and most controllable cats. (Montague and others)

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“You’re welcome. Now, how about some worship?” The wildcats probably did some controlling, too.

 

Natural selection is all about having more kittens than the other wildcats. (Simpson, 1953; Turner and Antón)

That’s what Lybica did on the new “Easy Street.” Its numbers grew, the “domestic” mutation became fixed in this population, and Silvester turned into Fluffy.

Perhaps this mutation has happened in other cat lineages, too.

A little over five thousand years ago, Neolithic people in northwestern China lived with small cats for several centuries.

This is surprising, since Lybica’s home is on the other side of that huge continent. Too, housecats in China today – like those all over the world – have the Lybica genetic markers. (Hu and others)

Nevertheless, those Neolithic cats apparently were pets. One lived to an old age and another seems to have eaten a lot of grain – an unusual diet for a hypercarnivore LINK unless it’s scrounging table scraps. (Hu and others)

Researchers eventually established that these people had domesticated leopard cats, not wildcats! (Vigne and others, 2016)

Leopard_cat_India
Leopard cat: “Prionailurus bengalensis.” Remember this little fellow from last time?

 

In light of that fact, it’s interesting that leopard cats and Felis were the last two cat lineages to evolve (O’Brien and Johnson), and that some analyses even make these two sister groups. (Nyakatura and Bininda-Emonds)

Could they have some sort of a genetic connection that would make them both more prone to a “domestic” mutation relatively soon after the lineage appeared?

Or did it happen to other lineages, which are millions of years older than these two, and then fade away because people weren’t around yet?

Pet leopard cats would be sweet. Unfortunately, domestication didn’t take with them the way it did with Lybica.

After a few hundred years the Neolithic small cats vanished from the record. No other domesticated cats have been found in China between then and about two thousand years ago, when traders from the west brought in Fluffy. (Hu and others; Vigne and others, 2016)

Chinese cat sellers
A Western traveler sketched this late eighteenth-century Chinese cat market.

 

An outline of housecat history

It’s not 100% proven yet, but let’s assume that Lybica was domesticated in the Fertile Crescent about ten thousand years ago, while human beings there were developing agriculture and towns.

If Neolithic people brought cats to Cyprus, they probably also carried them to other Mediterranean islands and coastal lands. (Randi and Ragni)

However, nothing definite is known about Fluffy until roughly 3,600 years ago, when housecats first showed up in Egypt’s tomb paintings. (Sunquist and Sunquist, 2002)

They may have reached Egypt by way of Ethiopia and Nubia. (Wastlhuber)

A thousand years after housecats first had their portraits painted, they were honored members of many Egyptian families. It was a capital crime to kill one. Cats were mummified by the hundreds of thousands – each buried with some mummified mice and little milk containers to help it along in the afterworld. (Sunquist and Sunquist, 2002)

Bastet
Bastet, the ultimate “cat lady.”

 

The cult of Bastet, the cat-headed goddess, was also in full swing. Some 700,000 pilgrims visited her cat-filled temple at Bubastis each year for what sounds like several days of partying – a happy, alcohol-fueled celebration of love and fertility. (Sunquist and Sunquist, 2002; Wastlhuber)

Egypt made it illegal to export cats, but of course it’s impossible to keep such a good thing secret for long.

Domestic cats first appeared in Grecian art around 500 BC. Phoenician traders also smuggled cats into Europe. They’re probably the ones who introduced Fluffy to the British Isles, although after the Romans took over Egypt, housecats traveled throughout the empire with the legions. (Sunquist and Sunquist, 2002; Wastlhuber)

Cats went eastward, too, with monks and trade caravans.

Genetic studies confirm that Fluffy traveled to Asia early. Once there, it was isolated from western cats for a long time. In addition, various regional groups of Asian housecats were kept apart and developed on their own. (Lipinski and others)

India was the first Asian country to include cats in a religion. (Wastlhuber)

Buddhist monks brought cats to Japan around the sixth century A.D. Two cats were kept in every temple to protect religious manuscripts from rodents. (Wastlhuber)

This guardian function eventually evolved into maneki-neko: the little cat statue at the door, one paw raised to invite visitors in.

640px-Maneki_neko_at_Asasuka

 

At first, cats were an indoor luxury in Japan, but in 1602, when pests threatened the silkworm industry, the government ordered all cats released. (Wastlhuber)

Meanwhile, back at the Mediterranean, the old cult of Bastet was dead by 100 B. C. (Wastlhuber), but it left Fluffy carrying some heavy baggage in medieval Europe.

Cats may have been very good at keeping down the rats, but European Christians saw Fluffy’s Egyptian background in terms of heathens and idols . . . and witchcraft. (Wastlhuber)

Some of today’s housecat myths began during this period. Witches could turn into a cat a total of nine times. A black cat that crossed your path was marking Satan’s way. (Sunquist and Sunquist, 2002)

But there was nothing funny about it back then. People believed that stuff. Such excuses made cats fair game for torture and killing throughout Europe. It was horrible.

Then, in the seventeenth century, France’s Cardinal Richelieu decided that he was a cat person and kept several of them as pets.

And that was the end of widespread cat persecution in Europe! (Sunquist and Sunquist, 2002)

Richelieu at La Rochelle
Richelieu at the siege of La Rochelle, a (very) few factual details of which are given in Dumas’ “Three Musketeers.” When this man told Europe that cats were cool, Europe listened.

 

When the Age of Sail arrived, sailors brought cats on board not only for pest control but also for luck and a supposed feline ability to foresee storms. (Wastlhuber)

It also helped that cats take up very little room and are very entertaining, especially after weeks to months of nothing else to look at but the sky and the sea.

This phase of Fluffy’s history probably accounts for a lot of the variety seen in moggies today.

Remember “novelty selection”? Given the enormous number of cats in any major port, that individual cat might stand out in a crowd, but it wouldn’t matter much to the gene pool. Take it somewhere else, especially to a place where housecats aren’t well established yet, and its “novelty” genes can get established fairly easily. (Todd)

Churchill use
Winston Churchill didn’t give Blackie, his ship’s cat, a chance to meet any felines on the American ship during a World War II naval conference in 1941.

 

So on top of the eight races of cats was added an amazing assortment of looks, making housecats the most varied group in the modern cat family.

Yet all this diversity is only skin deep. Each and every one of those moggies and fancy-breeds has the same last name: catus.

And as we saw above, it’s the first name that gives biologists and conservationists so much trouble.

Wildcats and genetic extinction

Evolution and extinction go together. Every species has an origin, and it also eventually comes to an end.

What happens in between those two points is so complex that evolutionists must use mathematics and graphs to study it.

Just as meteorologists use H’s and L’s on maps to talk about the weather, evolutionary researchers describe things like an “adaptive peak” that sails majestically across the graphic landscape.

Fortunately for the rest of us, none of that is needed to get a basic picture of the origin of wildcats.

It’s not a very complete picture. The wildcat’s genome hasn’t been sequenced yet (Oliveira and others), and there are very few wildcat fossils around to study.

Still, paleontologists are pretty sure that wildcats first appeared during the era of sabertooths and ice ages, millions of years after the other modern cat lineages developed.

  • The oldest known”Felis” lived in what’s now Kenya over four million years ago. (Werdelin and others)
  • About two million years ago, there was a Felis cat – F. lunensis, or Martelli’s cat – in southwestern Europe (Werdelin and others; Yamaguchi and others, 2004) and probably elsewhere around the Mediterranean. It was a little bigger than a wildcat, with some cranial differences, but Martelli’s cat resembled Silvester more than any other early small cat did. (Kurtén)
  • Around 350,000 to 450,000 years ago, the first known wildcats showed up in the geologic record as fossils in what are now French and German caves. (Kurtén; Yamaguchi and others, 2004)
  • Those early wildcats were rugged and probably shaggy forest cats. The lithe steppe-cat line began at some point between 130,000 and 20,000 years ago – probably at around the coldest part of the last ice age (Pierpaoli and others) – when a few wildcats left Europe’s arctic landscape for the warm and dry Middle East. Fossils of these rather large animals have been found in Palestine. The ancestors of Lybica and Cafra then headed south into Africa. Ornata’s forebears went eastward into central Asia. (Kurtén; Yamaguchi and others, 2004)
  • Around 10,000 years ago, as we have seen, some Lybica wildcats hooked up with humans in the Fertile Crescent. (Vigne and others, 2012)
  • The domestic-cat wave began happening to wildcats some 3,000 years ago (Randi and Ragni), when Neolithic humans, then Phoenicians, and eventually the Romans carried Fluffy into Britain and Europe, where it met Silvester for the first time. These two feline lineages have been enthusiastically sharing genes ever since.

Of course, we eventually carried Fluffy all over the world, too, but in this section we’re focusing on the housecat-wildcat connection.

That has been most thoroughly studied in Europe, though it may also be an issue in northern Africa. (Yamaguchi and others, 2015)

The problem is that no one knows how much of that 450,000-year genetic heritage of European wildcats remains after thousands of years of interbreeding wth domestic cats.

It’s not unusual that housecats and wildcats can mate and produce fertile hybrids, even though they are usually put into different groups – catus and silvestris.

About ten percent of all animal species do this. Hybridization is most common when both groups are closely related and evolved relatively recently. (Mallet)

That’s certainly true of wildcats and domestic cats.

Sometimes the genes of one side invade the other’s genome after hybridization. This is called introgression, and it happens regularly, too. (Mallett; Shurtliff)

Hybridization and introgression can be harmful or helpful. Each case is different.

It’s at this point that many scientists bring out the graphs, adaptive peaks, etc. (Mallet; Rhymer and Simberloff; Simpson, 1953)  Moving right along, experts still have much to learn about wildcat hybridization and introgression. (Oliveira and others)

Most of the domestic cats involved are feral. In Britain there are about 1.2 million feral cats, compared with 1,000 to 4,000 wildcats, all of whom live in the Scottish Highlands. (Macdonald and others; Yamaguchi and others, 2015)

It’s easy to see why there is concern about the wildcat genome being “swamped.” There may be as few as four hundred Scottish wildcats that are still relatively free of domestic-cat genes. (Kitchener and others, 2005)

Hungary is another introgression hotspot, but wildcats in Italy, northeastern France, and the Iberian peninsula are only minimally affected. (Oliveira and others)

Some wildcats and domestic cats in Italy may even have achieved reproductive isolation! (Randi)

That inability to interbreed used to be the only way to define a species until late in the twentieth century, when newly developed molecular techniques showed researchers just how complicated the speciation process really is.

The first laws to protect rare wildlife used this “reproductive isolation” definition, too. That’s why hybrids aren’t legally protected. Domestic cats aren’t considered a threatened species, of course, but wildcats are.

It’s these well-intentioned human laws that make the first part of Fluffy’s scientific name so important.

The basic legal assumption at present is that Fluffy and Silvester are two separate species – F. catus and F. silvestris.

The laws, therefore, are written to protect F. silvestris.

But quite a good argument can be made – above and beyond the whole interbreeding thing – that Fluffy and Silvester belong to the same overall species, in different subgroups.

If the courts ever accept that view, legal protection of wildcats will be difficult, to say the least.

Officially recognizing domestic cats as a wildcat subgroup – F. s. catus – would red-list practically every housecat on Earth, which is absurd.

To go the other route and put Silvester into Fluffy’s F. catus group would delist wildcats. Those that haven’t been “swamped” by housecat genes yet would then quickly disappear via genetic extinction or be hunted into extinction for the fur trade or exterminated as pests.

I can’t do justice to the legal quagmire this F. versus F. s. issue threatens to lead us all into. See Macdonald and others, “…an enigma wrapped in a riddle inside a conundrum,” for more details.

In the meantime, out in the field, conservationists are doing what they can to preserve as many wildcats as possible.

In some places, like Switzerland’s Jura Mountains, relatively simple steps are possible, like monitoring the introgression level in local wildcats. Efforts can also be made to protect the relatively “pure” wildcats in Italy, Germany, and the Iberian peninsula. (Nussberger and others)

Conservationists need to be much more proactive in hotspots. The Scottish Natural Heritage agency, for example, has come up with an action plan that includes a captive breeding program, neutering and vaccinating feral cats (due to public concerns, feral cats are trapped and released in this program rather than killed, which happens in many other countries), strict land management guidelines, and identifying at least five zones where wildcats can be fully protected.

mother and kitten
Mother and kitten Scottish wildcats in a breeding program at the British Wildlife Centre.

 

The plan also calls for public education on the importance of neutering pets and keeping them indoors. Those living in wildcat protection zones would be discouraged from keeping cats as pets.

As a cat owner, would you go along with that? It is awfully tempting to say, ‘they’re all cats – what’s all this fuss about?’

Here’s the basic question. Is wildcat conservation in Europe worth all this effort, expense, and sacrifice?

Yes and no.

  • Yes. There are still wildcats out there with almost all of their 450,000-year-old heritage intact. We know very little about wildcat genetics and the effects of hybridization and introgression. It would be a shame if those wildcats disappeared before we learned more about them. And it would also be a shame for our children and grandchildren to grow up thinking of wildcats only in the past tense.
  • No. As we briefly mentioned last time, hybridization and introgression are also happening with spotted cats like the oncilla and Geoffroy’s cat in South America, and no group seems to be suffering from it. Of course, this hasn’t been studied very much yet. (O’Brien and Koepfli; Trigo and others) But hybridization seems to be one of the usual patterns of evolution, especially after an ice age. (Barton and Hewitt; Hewitt; Nussberger; Shurtliff) Why not just let the cats evolve as they will?

I can’t go any farther with this. It’s a very complicated issue, and there is something to be said for all points of view. Sorting it out and reaching the best outcome for everybody requires a lot of discussion among scientists, the public, government representatives, and the private sector.

But you now see why I couldn’t include Fluffy and Silvester in the other post about the world’s cats. These two definitely require separate coverage.

I had intended the next post in this series to be the last one, about issues and future research directions. However, before doing that I might attempt an experimental post.

Earth’s climate has evolved dramatically in the last 60 million years or so, and cats/cat-like creatures have evolved along with it over the last 35 million years. That is the epic nature of the story of feline evolution that really grabbed me at first.

I’m not sure it’s possible to cover this, at least for the book, because scientists, as far as I can tell, haven’t reached a consensus on important details of climate evolution.

Too, it may not be possible to cover the feline evolution over such a span of time without resorting to math and graphs.

I’ll think it over. We’ll certainly have at least one more post in this series.

The book is very tentatively scheduled to come out at the end of October 2017, just before Halloween.

Thank you for your interest!


IMAGES
Featured image: Norwegian forest cat. Trish Hamme. CC BY 2.0.

European wildcat in open-air enclosure in Bavarian Forest National Park: Aconcagua (talk). CC BY-SA 3.0.

Flying Scottish wildcat, Cormack by name, at the British Wildlife Centre: Peter Trimming. CC BY 2.0.

Cute barn cats: Alexandre Delbos. CC BY 2.0.

Egyptian fowling with a feline retriever: Illustration from The Cat of Bubastes, 1896. British Library HMNTS 012621.I.5.  Public domain.

Cat show video https://youtu.be/cMXxAkiaAr4 .  IdeateTV.

Seal point Siamese kitten: William Winters. Public Domain.

Turkish Van and child on bike: Candoyi.  Public domain.

Calico British Shorthair: Squeaky Knees. CC BY 2.0.

Ship cats in hammock:  George Silk, Australian War Memorial Flickr collection.  Public domain.

Arabian wildcat: Michal Mañas. CC BY 2.5.

Felis silvestris lybica (Sardinian wildcat): Gurtuju. CC BY-SA 3.0.

European wildcat: Luc Viatour/www.Lucnix.be. CC BY-SA 3.0.

Asian wildcat video https://www.youtube.com/watch?v=Qsy69yIFSWI:  Wild India.

Bieti: 西宁野生动物园.  CC BY-SA 3.0.

Egyptian painting of cat stealing eggs, 1422-1411 BC: The Yorck Project.  Public domain.

Fertile Crescent: Atanas Kostovski. CC 1.0 Universal PD dedication.

Wildcat in tree: Rute Martins of Leoa’s Photography (www.leoa.co.za).  CC BY-SA 3.0.

Leopard cat in Indian Sundarbans. Shan2797. CC BY-SA 4.0.

Chinese cat market. Thomas Allom, 1843, from a sketch by William Alexander in 1793. Public domain.

Statue of Bastet. Walters Art Museum, acquired by Henry Walters in 1913. CC BY-SA 3.0.

Maneki-neko at Asasuka:  Kondo Atsushi.  CC BY-SA 2.0.

Cardinal Richelieu at the siege of La Rochelle: Henri Mott, 1881, at Musée d’Orbigny Bernon. Public domain.

Churchill and Blackie: Captain Horton, War Office official photographer, August 1941, photograph H12756. Imperial War Museums. Public domain.

Scottish wildcat mother and kitten:  Peter Trimming.  CC BY-SA 2.0.


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