You might have noticed very little/no activity here over the past two weeks: partly this is because I'm very busy (preparing for Dinosaurs - A Historical Perspective, among other things), but it's also because I currently have no internet access at home. Sigh. In an effort to add something new, here's the long-planned, third and final part in the series on Europe's cat fauna, adapted from the Big Cats in Britain talk 'The deep time history of Britain's felid fauna'. In a previous article we looked at homotheres, lions and leopards, and in a second one at jaguars, pumas and cheetahs. This time round we deal with the smaller cats: the lynxes, and the members of the domestic cat lineage...
Lynxes are instantly recognisable: they're proportionally long-limbed, they have a very shortened tail, a facial ruff and tufted ears, and they also have a reduced dentition compared to most other cats (Russell et al. 1995). Three lynxes inhabited Europe during the Pleistocene, and two of these are still with us, the large, ungulate-killing Eurasian lynx Lynx lynx and the much smaller Iberian lynx Lynx pardinus [Eurasian lynx image above taken from wikipedia]. The third species was rather different from living lynxes: it's called Lynx issiodorensis, the Issoire lynx, and it had shorter, stockier limbs and a proportionally bigger head than living lynxes. It fact it was proportioned far more like a pantherine cat like a leopard, and its comparatively large size and big cat-like proportions mean that its remains have sometimes been confused with those of European cheetahs and pumas (Hemmer et al. 2004).
The Issoire lynx didn't survive into the Holocene, but of course the Iberian lynx and Eurasian lynx did [adjacent image shows restoration of Issoire lynx by Mauricio Antón, together with lynx lower jaws]. The Iberian lynx previously occurred throughout central Europe (Kurtén 1968, Kurtén & Grandqvist 1987) - in fact it's been described as the 'original' lynx of western Europe (Hemmer 1979) - but it's restricted today to the Iberian Peninsula, and here it's critically endangered, with a fragmented population limited to between 300 and 500 individuals (Bartolomé 2001, Johnson et al. 2004). During the Pleistocene, Eurasian lynxes colonised essentially the whole of Eurasia, but persecution, habitat loss and hunting have meant that they're now locally extinct across much of western Europe [map below, from wikipedia, shows Eurasian lynx's current range].
In Britain, the Eurasian lynx has generally been regarded as a Pleistocene animal that failed to make it into the Holocene. However, we now know that this is completely wrong. Eurasian lynxes were definitely still present in England and southern Ireland as recently as 9000 years ago. The English records, one of which comes from Devon and the other one from Derbyshire, have been carbon dated: both gave ages of 8800-9500 years before present (Coard & Chamberlain 1999, Bronk Ramsey et al. 2002). That's all pretty good: it shows that lynxes were still in Britain during the Holocene. However, in 1997 came the surprising news that Eurasian lynxes had in fact survived in Scotland until as recently as the 3rd century AD.
Radiocarbon dating of a lynx discovered in 1927 in Reindeer Cave in Sutherland showed that the specimen was about 1770 years old (Kitchener & Bonsall 1997). Further radiocarbon dating of lynx bones from North Yorkshire, published in 2006, revealed even younger ages of about 1550 years old, dating these specimens to the 5th or 6th century (Hetherington et al. 2006). In their paper on these modern-era lynxes, David Hetherington and colleagues noted that a word for lynx - llewyn - was used in various 7th century stories and lullabies, all of which originated from north-western England, roughly the same area where the 5th or 6th century lynx specimens came from. And we know from pollen data that this area was well forested at this time, showing that it remained good potential lynx habitat.
So it seems that lynxes persisted in north-western England until the 7th century at least, and of course it's possible and likely that they survived even more recently than this (the few specimens that have been carbon dated are very unlikely to represent the youngest specimens that exist). Quite why and how lynxes then became extinct in Britain - especially when they survived in Scandinavia and elsewhere in Europe - is open to speculation; it's assumed that deforestation, hunting and persecution did the lynx in. Is it conceivable that they survived from the 5th, 6th or 7th century to even more recent times? Various zoologists in the past - including Bernard Heuvelmans and H. G. Hurrell - have suggested exactly this, and of course we have pretty good historical accounts, like William Cobbett's account from the late 1700s, that seem to describe late-surviving British lynxes. Given the massive impact of people on Britain's mammal fauna within the last couple of hundred years, I personally think that the idea that any mid-sized or large mammal might have survived undetected is very unlikely if not impossible. However, if you'd asked an expert 20 years ago as to whether lynxes might have survived to even the 5th century, I think you'd have been laughed at, so perhaps we should be prepared to be surprised again.
The not so Scottish wildcat
Finally, we come to the smaller cats. Conventionally it's been thought that, within recent history, the British Isles have only ever been home to a single small cat species, and this is Felis silvestris, the species most usually referred to (in Britain) as the Scottish wildcat. Of course, this name is entirely inappropriate given that the present restriction of the wildcat to Scotland is an artefact of human persecution: until very recently, the wildcat occurred across essentially the whole of Britain (even occuring in Ireland during the Holocene (Sommer & Benecke 2006), contrary to conventional wisdom).
Human persecution and habitat destruction led to the decline of the wildcat: by about 1800, it had been eradicated from much of southern and central England, by about 1850 it was mostly gone from England entirely, and by the late 1800s it was essentially restricted to the Scottish highlands (Langley & Yalden 1977). This historical pattern of decline was worked out by Peter Langley and Derek Yalden in 1977, but there are enough English sightings of wildcats from the 1800s and 1900s to suggest that small populations might have persisted outside of Scotland. Alternatively, wildcats might have successfully recolonised parts of the country, or been re-introduced to southern regions where they had locally become extinct.
Wildcats interbreed with domestic cats wherever the two meet, meaning that pure-bred wildcats are very rare, if not critically endangered, and this is a problem not just restricted to Britain but affecting wildcats in mainland Europe and Africa as well (Pierpaoli et al. 2003, Yamaguchi et al. 2004). Where hybridisation has occurred, wildcat characteristics continue to crop up in the hybrids, and the result is a continuum with domestic cats at one end, true wildcats at the other, and a range of intermediates in the middle (French et al. 1988, Daniels et al. 1998, Kitchener 1998, Reig et al. 2001). Exactly how and where we draw the line between 'intermediates' and 'true' wildcats is a thorny problem, compounded by the fact that some historical type specimens are apparently hybrids.
Exotic small felids in the British record
Until very recently it was thought that the wildcat was the only small felid that ever inhabited the British Isles within geologically recent times. This idea was challenged in 1997 when a fossil Swamp cat Felis chaus (or Jungle cat) was discovered at Aveley in Essex, in sediments dated to the Middle Pleistocene (Schreve 2001a, b). Unfortunately we only have this one specimen at the moment, and we essentially know nothing about the prehistoric distribution of the Swamp cat in Britain, both in term of geography and in terms of how recently it went extinct.
In fact similar comments can be made about the distribution of the Swamp cat in Europe as a whole: several fossils of this species have been reported from the Middle and Upper Pleistocene of Greece, Germany and France, but all are questionable and Hemmer (1979) wrote that they required confirmation. The absence of the Swamp cat from Holocene sediments in Britain as well as from the rest of Europe suggest that it didn't survive until particularly recently, but - as with the lynxes we looked at a moment ago - evidence either way is very thin on the ground and we need more data. The Aveley specimen has yet to be properly described but a full description will appear soon (D. C. Schreve pers. comm. 2008).
Even more poorly known than the presence of swamp cats in Britain is the suggestion that African wildcats once occurred in the British Isles as well: Scharff (1906) described some lower jaws from Ireland and identified them as being particularly close to Felis silvestris libyca, the African wildcat [adjacent image shows Scharff's Irish cat jaws compared with others]. Based on bones discovered in Edenvale and Newhall caves in County Clare, Scharff thought that there were actually two Irish wildcats - a large one and a small one. Scharff's identification has been repeated a couple of times since: the implication being that maybe Ireland was home to an endemic wildcat, perhaps one quite distinct from the wildcats of Scotland, England and Wales - but the problem here is that African wildcats are very similar, if not identical, to domestic cats in skeletal and tooth characters. In fact domestic cats originated from African wildcats rather than from those of Eurasia, and as a consequence it can be almost impossible to distinguish the bones and teeth of the two.
The Irish wildcat remains didn't come from a prehistoric fauna - they were discovered in layers where domestic mammals were abundant - and, in an evaluation of Scharff's wildcats, Stelfox (1965) concluded that they were domestic cats, the larger ones being males, the smaller ones females. Rather than accept the idea that African wildcats somehow got into Ireland without leaving a fossil record elsewhere in Europe, it does seem more sensible to conclude that Scharff's Irish wildcats were just domestic cats (Fairley 1975).
Additional small cat species might have been present in Pleistocene Europe - there are some doubtful Middle Pleistocene German fossils and more reliably identified Czech fossils that might represent Pallas's cat Otocolobus manul for example (Kurtén 1968, Hemmer 1979) - but there's presently no evidence that additional species occurred in Britain. Incidentally, there is no fossil or archaeological evidence for rabbit-headed cats, an observation which supports the view that they are teratological rather than a genuine taxonomic entity [adjacent image, from wikipedia, shows Pallas's cat].
And that bring us to the end of our look at Europe's cat fauna. The big sabre-toothed cat Homotherium latidens was still present in northern Europe during the Late Pleistocene, but almost certainly didn't persist to the Holocene. Lions remained in Spain and south-eastern Europe until the early Holocene. Leopards inhabited Britain and the rest of Europe during much of the Pleistocene, but by the Late Pleistocene they were gone from northern Europe. The European jaguar lived alongside the leopard in several locations during the Middle Pleistocene. Pumas and cheetahs occurred across Europe and Asia during the Early and Middle Pleistocene. Pumas were present in Britain but neither pumas nor cheetahs seem to have survived into the Late Pleistocene or Holocene. Lynxes definitely survived in Britain until the 1st century AD, may well have persisted until the 6th or 7th centuries, and were perhaps still present even more recently than this. Jungle cats were present in Britain during the Middle Pleistocene but there is as yet no evidence that they survived into the Holocene. There's no evidence that African wildcats were present in Ireland, despite suggestions from the early 1900s that they were. Finally, European wildcats were here throughout the Pleistocene and Holocene, and have just about managed to survive until today.
Europe's cat fauna - and Britain's cat fauna - was obviously a lot more diverse in the recent past than it is today. In fact, we have a thoroughly impoverished fauna compared to what we 'should' have.
That's all for now. Will be seeing some of you on Wednesday (23rd April) at the Natural History Museum for the CEE's Modern Functional Anatomy Workshop. Will be discussing what happened there here at Tet Zoo, internet connection permitting.
Refs - -
Bartolomé, J. 2001. A creature great and small. BBC Wildlife 19 (8), 22-27.
Bronk Ramsey, C., Higham, T. F. G., Owen, D. C., Pike, A. W. G. & Hedges, R. E. 2002. Radiocarbon dates from the Oxford AMS system: Archaeometry datelist 31. Archaeometry 44 (supplement 1), 1-149.
Coard, R. & Chamberlain, A. T. 1999. The nature and timing of fauna change in the British Isles across the Pleistocene Holocene transition. Holocene 9, 372-376.
Daniels, M. J., Balharry, D., Hirst, D., Kitchener, A. C. & Aspinall, R. J. 1998. Morphological and pelage characteristics of wild living cats in Scotland: implications for defining the 'wildcat'. Journal of Zoology 244, 231-247
Fairley, J. 1975. An Irish Beast Book. The Blackstaff Press, Belfast.
French, D. D., Corbett, L. K. & Easterbee, N. 1988. Morphological discriminants of Scottish wildcats (Felis silvestris), domestic cats (F. catus) and their hybrids. Journal of Zoology 214, 235-259.
Hemmer, H. 1979. Fossil history of living Felidae. Carnivore 2: 58-61.
- ., Kahlke, R.-D. & Vekua, A. K. 2004. The Old World puma - Puma pardoides (Owen, 1846) (Carnivora: Felidae) - in the Lower Villafranchian (Upper Pliocene) of Kvabebi (East Georgia, Transcaucasia) and its evolutionary and biogeographical significance. Neues Jahrbuch fur Geologie und Paläontologie, Abhandlungen 233, 197-233.
Hetherington, D. A., Lord, T. C. & Jacobi, R. M. 2006. New evidence for the occurrence of Eurasian lynx (Lynx lynx) in medieval Britain. Journal of Quaternary Science 21, 3-8.
Johnson, W. E., Godoy, J. A., Palomares, F., Delibes, M., Fernandes, M., Revilla, E. & O'Brien, S. J. 2004. Phylogenetic and phylogeographic analysis of Iberian lynx populations. Journal of Heredity 95, 19-28.
Kitchener, A. C. 1998. The Scottish wildcat - a cat with an identity crisis? British Wildlife 9, 232-242.
- . & Bonsall, C. 1997. AMS radiocarbon dates for some extinct Scottish mammals. Quaternary Newsletter 83, 1-11.
Kurtén, B. 1968. Pleistocene Mammals of Europe. Aldine Press, Chicago.
- . & Grandqvist, E. 1987. Fossil pardel lynx (Lynx pardina spelaea Boule) from a cave in southern France. Ann Zool Fennici 24, 39-43.
Langley, P. J. W. & Yalden, D. W. 1977. The decline of the rarer carnivores in Great Britain during the nineteenth century. Mammal Review 7, 95-116.
Pierpaoli, M., Birò, Z. S., Herrman, M., Hupe, K., Fernandes, M., Ragni, B., Szemethy, L. & Randi, E. 2003. Genetic distinction of wildcat (Felis silvestris) populations in Europe, and hybridisation with domestic cats in Hungary. Molecular Ecology 12, 2585-2598.
Reig, S., Daniels, M. J. & Macdonald, D. W. 2001. Craniometric differentiation within wild-living cats in Scotland using 3D morphometrics. Journal of Zoology 253, 121-132.
Russell, A. P., Bryant, H. N., Powell, G. L. & Laroiya, R. 1995. Scaling relationships within the maxillary tooth row of Felidae, and the absence of the second uper premolar in Lynx. Journal of Zoology 236, 161-182.
Scharff, R. F. 1906. On the former occurrence of the African wild cat (Felis ochreata Gmel.) in Ireland. Proceedings of the Royal Irish Academy 26, 1-12.
Schreve, D. C. 2001a. Mammalian evidence from Middle Pleistocene fluvial sequences for complex environmental change at the oxygen isotope substage level. Quaternary International 79, 65-74.
- . 2001b. Differentiation of the British late Middle Pleistocene interglacials: the evidence from mammalian biostratigraphy. Quaternary Science Reviews 20, 1693-1705.
Sommer, R. S. & Benecke, N. 2006. Late Pleistocene and Holocene development of the felid fauna (Felidae) of Europe: a review. Journal of Zoology 269, 7-19.
Stelfox, A. W. 1965. Notes on Irish "wild cat". The Irish Naturalists' Journal 15, 57-60.
Yamaguchi, N., Kitchener, A. C., Driscoll, C. A., Ward, J. M. & Macdonald, D. W. 2004. Craniological differentiation amongst wild-living cats in Britain and southern Africa: natural variation of the effects of hybridisation? Animal Conservation 7, 339-351.
What language is that? Cumbrish?
Felis silvestris lybica
F. l. libyca. Libya, not Lybia.
A very interesting article, Ive always been a fan of lynxs and I hope the guy in Scotland suceeds in his quest to reintroduce them (along with wolves, moose and other mega fauna).
I was in fact photographing a wildcat (captive obviously!) and finally managed to get one better than the one you've featured. I also got a rather nice photo of a lynx that day too, but unfortunately no where near as good as the one from Wikipedia!
"Llewyn" is a Welsh word, pronounced (as far as I can reproduce the sounds on an English keyboard) as "thloo-in", with a distinct hiss on the "thl" sound. "Llew" on its own means "lion", which might give "llewyn" an approximate meaning of "little lion".
Interestingly, the online translator from the University of Wales at Lampeter (www. geiriadur.net) gives no result for "llewyn" and translates "lynx" as "lyncs" - obviously just a straight transliteration of the English word. If "llewyn" really did mean "lynx", then it looks like the animal has been extinct in Wales long enough for its original Welsh name to have been forgotten. This contrasts with the wolf, whose Welsh name "blaidd" can still be found in placenames here and there.
Just forgot to add, the use of a Welsh word "llewyn" for lynx in north-west England isn't as contradictory as it sounds. Brittonic Celtic - the forerunner of modern Welsh -was spoken across much of this region, and the Scottish lowlands too, until probably the 9th or 10th century AD. Hence the modern name for the English Lake District - "Cumbria" - which has the same derivation as the Welsh word for Wales - "Cymru" (pronounced "Cum-ree").
Here endeth the History lesson. Now back to Zoology!
I think there is a bit of confusion about the correct spelling for the african subspecies. Both have been used in the literature- libyca/lybica.
Sunquist&Sunquist have it as "lybica"
David, further to Dave's comments the quote is from the Gododdin. See
"Or sawl yt gyrhaedei dy dat ty ae gicwein
O wythwch a llewyn a llwyuein
Nyt anghei oll ny uei oradein"
Llewyn is often translated as lynx. There some uncertainty about this, and about the age of the poem.
... and I should have looked in my "Geiriadur Mawr" [H. Meurig Evans and W.O. Thomas 1958] which helpfully includes lots of words described as obsolete. It doesn't have "llewyn" but it does have "llewynawg" which it translates as a fox. Against that "llwyuein" could be read as "llwynein", like the modern "llwynog" which also means fox and indeed that's how it's usually translated. Would the poet have put two foxes in one line? Well maybe, if the words fitted.
Does Hetherington's paper offer any further explanation?
For curiosity, how many caudal vertebrae does a lynx have? Fewer than a long-tailed cat, or the same number but the vertebrae themselves reduced in size? I ask because I recently noticed (when visiting some U.S. museum that had a keletal mount) that Smilodon -- always portrayed in reconstructions with a lynx-like stub of a tail -- seems to have had as many caudal vertebrae as a "normal" cat, but tiny ones: the caudal skeleton looked as if it belonged on a much smaller cat.
As to the pronunciation of the Welsh ll ... I recall a story about an 18th century clergyman who had been appointed to a bishopric in Wales and was learning Welsh so he could preach to his new flock: his Welsh tutor explained it "Thy lordship must clap thy lordhip's tongue to the roof of his mouth and hiss like a goose."
Something I find really amazing is the sheer size a lynx can reach. There is a speciemen in our local zoological collection, and it is really enormous, not much smaller than a wolf, and some of them grow even larger. If you think about this animal, you often don´t realize that they can grow to the size of a small leopard or puma, and as a result of their long legs, they are even taller. Keeping this in mind, we could perhaps count the lynx as the very last european "big" cat.
Thats just fine, Sordes; a titchy Indian wolf or a huge Timber wolf??
It was a european wolf, and a bit bigger than a german sheperd (But don´t think eurpean wolves are small in general, some of them were really huge). This wolf was stockier than the lynx, and I suppose the lynx weighed around 25kg or so when it was alive. The heaviest male lynxes reach weights close to 40kg, what would be again much bigger.
I wouldn't be surprised to discover that a small population of lynxes had persisted in some restricted region of the British Isles right up to the present day, going quietly about their business and surviving by being extremely nocturnal. Occasional lynx sightings could be folded into local "black dog" legends. British lynxes would have received a new lease on life when assorted "pets" or menagerie specimens escaped or were released in the late 20th century.
Some of your readers might be interested in Dies the Fire and its sequels by S.M. Stirling. The larger story is what happens when persons or forces unknown make modern technology unworkable on the surface of the earth and the survivors find themselves in a weirdly old new world, but one of the major subplots is what happens to the zoo/menagerie/"pet"/circus animals, many of which are kept in enclosures that they could escape if they really wanted to. The new ecosystems formed by all these escapees are fascinating (not to mention dangerous; firearms no longer function). Big cats feature heavily in the newly wild heart of Britain.
Oh, and have you heard of the proposition that feral cats in some port cities have a percentage of F. chaus genes because sailors brought them home from Southeast Asia? Apparently they are semidomesticated pest catchers there.
Occasional lynx sightings could be folded into local "black dog" legends.
Lynxes are never black, though.
I was thinking the two-glowing-eyeballs-and-a-shape-in-the-gloaming variety of Black Dog legend. In the period just after sunset, anything that isn't brightly colored tends to look black.
That makes sense.
But many black dog encounters involve the person SEEING the dog.
I dont know where the black dog legends come from (there are 50 in my country of wiltshire alone) some say it was based upon encounters with rabid animals.
But people in the past were well aware of the symptoms and consequences of a rabid animal, there was nothing mythical there.
Thanks to all for neat comments. Very briefly: while it has been suggested that 'black dogs' might be old sightings of British big cats (e.g., by Di Francis in Cat Country), it's a non-starter, as black dog sightings were never vague, but always very specific in describing all the salient features that allow us to easily distinguish dogs from cats. But black dogs are almost certainly nothing to do with sightings of real animals anyway: in the accounts, the 'dogs' are satanic monsters with eyes that glow red like hot coals, they belch fire or smoke, they sometimes have a phosphorescent glow, and they make the noise of chains or the clip-clop of cloven hooves as they walk. They serve as omens or potents of disaster for people who drink too much, or stay out too late (uh-oh), or are otherwise naughty. When they touch people, the people die. They leave scorch marks when they leap through church doors. And so on..
As for melanistic lynxes, melanism has been recorded (and photographed) in Bobcat Lynx rufus, but I can't recall hearing of it in Eurasian lynx. Google-image 'melanistic bobcat': the individuals that have been photographed look very freaky!
Just a small point about lions surviving into the Holocene in Iberia and southeastern Europe. There is at least one early Holocene record from Italy (Tuscany) too, so "southern Europe" in general would be more appropriate.
There was an iron age tribe from The Caithness/Sutherland region called the Lugi. This is commonly reported as meaning 'Raven People' but this seems to be based on one doubtful source as to the application of the word to the raven (based on association of the bird with the god Lugus.)
The usual Celtic word for Raven is Bran.
A more likely meaning of Lugi is 'Lynx People'. Could it be that this tribe was named after the Lynx and when that animal became extinct they became associated with the Wildcat? The dark age province Cataibh means the land of the Cat or Cat Tribe (Caithness = promontory of Cathaibh.)
An even more speculative possibilty is that these people were the ancestors of the Clan Chattan (Clan of the Cats) the great confederation of the eastern Highlands.
If the derivation of Lugi is correct this is good evidence of the Lynx surviving to a period close to that of the Roman invasion of these islands
[from Darren: sorry, delayed by spam-filter]
With regard to llewyn, in Irish leomhin (pronounce 'mh' like English 'w') was used to mean a small lion. However, an early Irish word "lug" actually meant a lynx. Received wisdom is that it was brought to Ireland from the Celts of the Continent, but it somehow survived in the insular language. This could have been due to contact with Scotland, but it could also mean that the odd lynx was still to be found in Ireland, stravaging around in blissful ignorance of the fact it was supposed to be extinct.
Jesus Armorgaden Western Australia on Sep.16th 2009 night. Polar Bear, giraffe, snakes were all died. Chinese, muslim and Indian are all died in Western Australia.
Jesus also Armorgaden Europe (ex British), Middle East, Africa and Latin America on May 23rd 2009.
Polar bears are all died at North Pole.
GOD didn't create all things such as giraffe, panda, Polar bears and pests. But GOD created Salmon fish for Northern American Teddy Bears.
In China Teddy farms and zoos, Devil Chinese lock up and chop Brown Teddy hands off.
GENESIS GOD'S Daughter
That was odder than usual. Is it some sort of Gnostic thing?
That was odder than usual. Is it some sort of Gnostic thing?
No idea, but if you perform a google search with the terms "crab white" and "genesis" you'll find that he/she/it has left similarly deranged comments on other internet sites (example here). Basically, it's just spam.