Thursday, 15 February 2018

What is the Chinese Mountain Cat aka Steppe Cat, Desert Cat, Felis bieti?

The Chinese Mountain Cats that we saw by spotlight on the Tibetan Plateau in Sichuan have found themselves to be in a state of taxonomic confusion and disagreement. This is not surprising  or unusual for a small cat since there are relatively few specimens in museums and the division of small cats into robust taxons is well known to be difficult. Add to that the differences in species concepts adopted and vigorously defended by different workers with the addition with several possible interpretations of DNA data, it is easy to see why recent papers have pulled the animal in two directions: first it was lumped into the local Wildcat (itself the subject of controversy); then it was restored as a proper species.

In 1892 Alphonse Milne-Edwards described a new species of cat; he named it Felis bieti for Felix Biet (1838–1904) a French missionary. Neither a type specimen nor the locality was noted in the original description but Glover Morrill Allen (1879-1942) in his book explained that Prince Henri d’Orléans (1867-1901) brought back the two specimens to Paris from Tongolo and Tatsienlu in Sichuan. Henri d’Orléans, noted duellist and anglophobe, led an expedition, at the insistence of, and paid for by, his father from Siberia to Indo-China.

Shortly after Milne-Edwards, Eugen Alexander Büchner (1861-1913) of St Petersburg described what turned out to be the same species as Felis pallida, a name actually appropriate to its appearance and one which Allen used for the common name, Pale Desert Cat in his book. A number of other common names have been used for F. bieti: Chinese Mountain Cat; Chinese Steppe Cat; Chinese Desert Cat.

It seems that nobody has found a modern location of Tongola but Tatsienlu is now known as Kangding on what, historically, was the border between China and Tibet (and, incidentally, about 60 km west of where we were watching Red Pandas).

Although my video as well as several other videos available on YouTube were taken at night by spotlight, it can be seen that the appearance of the cats is the same as a still photograph taken by day by Caroline Simpson two months before our visit:

Chinese Mountain Cat aka Chinese Desert Cat
Chinese Mountain Cat on the Tibetan Plateau, September 2017
Photograph by Caroline Simpson (from Flickr)

It is quite clear that all—except, and possibly importantly, one—of the still photographs and videos taken of ‘mountain cats’ on the Tibetan Plateau near Ruoergai in Sichuan conform with the description of the F. bieti. And this is where I went back into the classical descriptive zoology of the early 20th Century.

Reginald Innes Pocock FRS (1863-1947) wrote his Catalogue of the Genus Felis but it had to be edited and published posthumously by the Natural History Museum in 1951. There is a perfect description of ‘our’ cat under the subspecies F. b. bieti (which Pocock himself had proposed earlier to distinguish it from another subspecies that I will return to later and in a subsequent post). I will not repeat the description here other than to point out that he did not specifically mention the tail although he did describe it for his other subspecies; the tails were clearly similar.

I am therefore entirely content with the conclusion that ‘our’ Chinese Mountain Cats on the Tibetan Plateau in Sichuan were what had been originally described as Felis bieti.

As I mentioned earlier the molecular phylogeneticists have not left the difficult problem of small cat taxonomy untackled. Thus we had a paper in Science in 2007 which, in considering the origins of the domestic cat, lumped, amongst other lumpings, Felis bieti into the Wildcat as the subspecies, Felis silvestris bieti. It is this classification that one finds on many current websites. However, I did not find the authors’ argument persuasive (see below for a possible interpretation) since models drawn from mitochondrial DNA (i.e. maternal line) differed from those built from microsatellite (nuclear DNA) data.

One of the authors of the paper described, Andrew Kitchener, of what is now ungrammatically known as National Museums Scotland, then took the opposite view in 2009. After modelling the geographical distribution from 18,000 years ago until the present, he, with E.E. Rees of the University of Alberta at Edmonston, wrote:
The palaeo-DDMs [Deduced Distribution Model] provide no clear support for the redesignation of the Chinese steppe cat Felis bieti as a subspecies of F. silvestris [Wildcat] based on Driscoll et al.’s (2007) phylogeographical analysis, except that the effective loss of the central Asian distribution during the LGM [Last Glacial Maximum] may have resulted in introgression between F. bieti and F. s. ornata, in small isolated habitat fragments. Distinctive morphology and possible sympatry with Asian wildcats suggest that the Chinese steppe cat is a distinct species whether using the BSC or PSC.

The latter argument—that Felis bieti—is a ‘good’ species appears to be holding sway since the species has been retained in the IUCN’s Cat Specialist Group’s Cat Classification Task Force report, Revised Taxonomy of the Felidae published very recently. 

The Task Force was chaired by Andrew Kitchener and comprised a Core Group, an Expert Group and a Review Group. Given the wide membership, backgrounds and adherence to different species concepts, it is not at all surprising that disagreements remain. To complicate matters a little further, the Wildcat of Eurasia has been split/resplit into two species, Felis silvestris, the European Wildcat and Felis lybica, the African and Asian wildcats, so the section on Felis bieti reads:
There has been no recent taxonomic study of this species since Groves* (1980), although Driscoll et al. (2007) showed that Felis bieti was basal to Felis silvestris (sensu lato) according to mtDNA, but a sister taxon to Felis lybica ornata from microsatellites, which suggests that Felis bieti may have an ancient hybrid origin, possibly during the last glaciation when the distribution of F. l. ornata was apparently restricted to a very small area in Central Asia (Kitchener & Rees 2009). F. bieti is morphologically distinct and is supposedly sympatric with F. l. ornata, which would also preclude its recognition as a subspecies of F. silvestris/lybica. However, C. Driscoll (pers. obs.) maintains this species as a subspecies within F. silvestris (sensu lato).
The stated distribution of Felis bieti has tightened over the years, not as a result of habitat loss, but as a result of excluding misidentified specimens. Li He, Rosa García-Perea, Ming Li and Fuwen Wei began their paper of a study made in 2000 and 2001:
One of the problems is the difficulty of identification. This cat is almost unknown to Chinese scientists, and the few specimens kept at Chinese institutions are mostly misidentified. A similar confusion applies to local hunters, who do not know what species they are dealing with when they capture the animal (pers. obs.). Fur traders are probably more familiar with the identification of the species, but they do not have an interest in the geographical origin of any pelts that they receive. Therefore it is common to find incorrect information in the local literature, with the species often confused with manul Otocolobus manul, Asiatic wildcats of the Felis silvestris [lybica] ornata group, or lynx Lynx lynx (e.g. Wang, 1990, 1991). Thus any information on the distribution and occurrence of the species requires careful evaluation of the source.
The authors looked at 45 specimens and living individuals, 168 reports from the literature and 21 records from local hunters. However, I found the criteria for inclusion as F. bieti to be unclear and while some of the records were obviously wrong, I have the suspicion that further studies will reveal a wider distribution, particularly to the west. As a result of this study though, the distribution map below is the one that is currently accepted.

modified from Sanderson et al. 2010

We found it difficult at night to estimate the size of the Chinese Mountain Cats we saw. The Guide to the Mammals of China says about twice the size of a domestic cat but the data given by Pocock do not support such a large difference. The head-body length of skins in museums tend to be longer than in real life and so absolute size is not very informative. Some of the skins are from the fur trade and could well have been stretched. Pocock’s data showed a range of 16-23 inches for the skins of feral domestic cats from various parts of the world compared with 28-33.5 inches for specimens collected as, or assigned to, Felis bieti bieti. Taking the medians of 19.5 inches for the feral cats and 30.3 for F. bieti, the latter are about 1.5 times the length of a feral domestic cat. But given that feral cats tend to be smaller than those kept in the house, about a third larger seems a fair estimate. Indeed, Allen’s description begins, 'About the size of a house cat…'

Ours, and other sightings of Chinese Mountain Cats on the Tibetan Plateau in recent years, have been made possible by the efforts of that indefatigable guide to, and font of knowledge of, the birds and mammals of Sichuan, Sid Francis. There is, though, a fairly recent photograph from one of his trips to the Tibetan Plateau in 2015 of a  cat which does not appear to conform to the others. That cat was seen during and photographed by Yann Muzika; the photograph was shown in Small Wild Cat Conservation News in May 2015:

from Small Wild Cat Conservation News, May 2015

To my untrained eye it looks more like a Wildcat, i.e. Felis silvestris/lybica, even accounting for any possible seasonal shift in pelage. If so, the presence of this species together with F. bieti would demonstrate sympatry (i.e. both occur in the same area) and provide conclusive evidence that Felis bieti is indeed a ‘good’ species. In that respect it is interesting that the distribution map for the Wildcat in A Guide to the Mammals of China (which also has a good painting of F. bieti, if a little angular) has a series of question marks for that area of Sichuan.

I do have one main concern left though. There are photographs of animals, clearly identified as F. bieti that do not really conform with the photographs and videos shown and referred to in this article. I will deal with that question, with some delving into the classical literature and much inadequate knowledge in a subsequent post.

There is very much more to be discovered about the Chinese Mountain Cat and indeed many of the small cats of the world. We just counted ourselves lucky and privileged to have seen Chinese Mountains Cats and Pallas’s Cats on the grasslands of the eastern Tibetan Plateau.

*Colin Groves died on 30 November 2017.

The following will also provide references to other research on the Chinese Mountain Cat

Allen GM. 1938. The mammals of China and Mongolia. New York: American Museum of Natural History.

Driscoll CA, Menotti-Raymond M, Roca AL, Hupe K, Johnson WE, Geffen E, Harley EH, Delibes M, Pontier D, Kitchener AC, Yamaguchi N, O’Brien SJ, Macdonald DW. 2007. The near eastern origin of cat domestication. Science 317, 519-523. doi:10.1126/science.1139518

He L, Garcia-Perea R, Li M, Wei F. 2004. Distribution and conservation status of the endemic Chinese Mountain Cat Felis bieti. Oryx 38, 55-61.

Kitchener AC, Rees E. 2009. Modelling the dynamic biogeography of the wildcat: implications for taxonomy and conservation. Journal of Zoology 279, 144-155.

Kitchener AC, Breitenmoser-Würsten C, Eizirik E, Gentry A, Werdelin L, Wilting A, Yamaguchi N, Abramov AV, Christiansen P, Driscoll CA, Duckworth JW, Johnson WE, Luo SJ, Meijaard E, O’Donoghue P, Sanderson J, Seymour K, Bruford MW, Groves C, Hoffmann M, Nowell K, Timmons Z, Tobe SS. 2017. A revised taxonomy of the Felidae. The final report of the Cat Classification Task Force of the IUCN/SSC Cat Specialist Group. Cat News Special Issue (11). ISSN 1027-2992

Pocock RI. 1951. Catalogue of the genus Felis. London: British Museum

Sanderson J, Yufeng Y, Naktsang D. 2010. Of the only endemic cat species of China.

Wozencraft WC. 2008 In, Smith AT, Xie Y. editors, A Guide to the Mammals of China. Princeton: Princeton University Press.

1 comment:

  1. In Uzbekistan recently, we were shown a photo of a Steppe Cat, native to their country. It looked like the first of your photos, definitely not the like the second photo, as it had stripes only on its legs. We were told it was half as big again as a domestic cat.