Golden Hamsters in the wild are diurnal but in captivity are nocturnal. Do we know why?

in a previous article about Golden Hamsters, I wrote

After Rolf Gattermann’s death from cancer in 2006 at the age of 57, his team of collaborators, who had made Halle a centre of small rodent research, continued publishing their work and started new strands, including field work on the Golden Hamster in southern Turkey. One interesting paper, begun while Gattermann was still alive, reported that while in the laboratory male and female hamsters (whether derived from old line or from newly-caught individuals) are nocturnal, in the wild female hamsters are diurnal. They speculated that a balance in the environment whereby nocturnal predators (owls and foxes) are a more potent threat than diurnal ones (birds-of-prey and snakes) may be responsible.

Since then there have been a number of studies attempting to identify the stimuli which turn hamsters from diurnal in the wild to nocturnal in captivity. As far as I can see there has been no success. However, some smaller effects evoked by potential predators have been recorded. For example, hamsters exposed to the odour of domestic ferrets spent less time out of a simulated burrow. In another species, the pattern of activity was affected by the rodent community in which the hamster was living, the time out and about was shortened when there were aggressive gerbils around.

I wonder whether the nature of the housing has an effect, being reminded of the marked changes in thermoregulatory behaviour observed by Roger Avery, then in Bristol, and also by me, in lizards given different sorts of cage furnishings. The deep and dark burrow system in the wild is very different from the accommodation afforded to pet and laboratory hamsters. Sleeping quarters are not necessarily completely dark and the Golden Hamster in the wild is subject to a very different regime of light/dark periods to that in a cage. Some simple experiments could be done.

In the meantime we still do not know if the presence and perception of predators is responsible for the basic activity pattern in the wild of Golden Hamsters.

McPhee ME, Segal A, Johnston RE. 2010. Hamsters use predator odors as indirect cues of predation risk. Ethology 116, 517-523

McPhee ME, Ribbeck AE, Johnston RE. 2009. Male golden hamsters (Mesocricetus auratus) are more reactive than females to a visual predator cue. Journal of Ethology 27, 137-141

Scheibler E, Wollnik F, Brodbeck D, Hummel E, Yuan S, Zhang, F-S, Zhang X-D, Fu H-P, Wu X-D. 2013. Species composition and interspecific behavior affects activity pattern of free-living desert hamsters in the Alashan Desert. Journal of Mammalogy 94, 448-458. (Note—not on the Golden Hamster)

Chinese Giant Salamanders: More information on those we had in the lab in 1960s Hong Kong

In the last post I noted how Edward George Boulenger had, in his description of a new species in 1924, noted: ‘Head…very much depressed’. I have found some photographs and videos of Chinese giant salamanders which show a similar appearance to that described by Boulenger but many which do not. Boulenger’s species (which had been regarded as synonymous with Andrias davidianus), Andrias sligoi, has been resurrected as a result of molecular phylogenetic analysis published this year. One can argue whether the species rank is warranted but there seems no doubt that the classic lineage, davidianus is different from sligoi and that they are geographically isolated. All sorts of questions then spring to mind: Is the ‘very much depressed’ head characteristic of the sligoi lineage? Or is it only in that state at a certain stage of growth? Are there other tissues present beneath the skin and bone that give a non-depressed appearance, changed by nutritional state, for example?

I have also looked at photographs of giant salamander skulls and replicas online, and again there do seem to be that remain flat posterior to the eyes while other show a rise in the region of the junction between the frontal and parietal bones.

I remembered that a very flat head described by Boulenger was not a feature of the Chinese giant salamanders bought in alive for class dissection at the University of Hong Kong in the 1960s. I took several photographs (and previously discussed some of them here) and they are shown below. To me the head seems more bulldog-like with a short ‘muzzle’ and, to continue the canine analogy, a ‘stop’— the rise in the skull beginning at the line of the eyes; in other words, the heads of these animals bought from the human food chain did not resemble Boulenger’s sligoi or some of the photographs of live specimens in circulation.

This one had lunged at my wife's hand with mouth open, and is ready to take on all comers

The bulldog-like shape of the head is particularly evident here

In the last post I noted the distance between the nostrils in Chinese and Japanese giant salamanders. I did a rough calculation from the photograph above (it had to be rough given the perspective). The distance between the nostrils was approximately 24% of head width, close to 21.5, the mean of five specimens in U.S. museums, detailed by Liu in his book published in 1950.

The pattern of tubercles on the head of our lab specimens is similar to that shown in a drawing in Liu’s book for A. davidianus but not that of sligoi shown in a drawing by or for Boulenger published on the ZSL website.

Is it, therefore, likely that the 1960s Hong Kong specimens were not from the sligoi genetic lineage but one of the other lineages from the mountain ranges of north-western China?

The question is interesting because it would suggest the main source for the trade in live salamanders going through Guangzhow (Canton), said to be the centre of the trade, was not the closer Nanling mountains but the mountains on the edge of the Tibetan plateau to the north and west.

In the old zoology store in the now-demolished Northcote Science Building there were some preserved specimens. My recollection is that there were several giant salamanders of modest size in jars of spirit rather than formalin. Did they survive the grand clear out when the department moved out and then moved on again to the Kadoorie Biological Sciences building? I was told a lot of stuff had been thrown out when I was external examiner in the department in the late 1990s but would somebody please have a look. If they still exist they could add something to the molecular phylogeny research and the pattern of trade for the human food market.

I have been fascinated by giant salamanders ever since seeing my first, the large Japanese one that lived in the vestibule of the aquarium at London Zoo. I never saw it move. The dozen or so for class use in the vertebrate course in Hong Kong were fascinating during the short time they were there. They were decidedly feisty; my wife had to move quickly as salamander wrangler when they made a lunge for her fingers during the photoshoot on the roof of the Northcote Science Building. I know they had to be ordered, with notice when they were needed. Food/Chinese medicinal items often came into Kong Hong from Chinese provinces like Yunnan and Sichuan far to the west in the1960s. There was a shelf of bear paws in the Chinese Merchandise Emporium on Queen’s Road.

We went through giant salamander country in Sichuan two years ago, with roads descending alongside the rocky streams and the slacker water in flatter valleys. Unfortunately, because many of us would like to have seen the set-up, the only salamander farm we saw was deserted, with overgrown or empty outdoor tanks.

Reading about what is known about giant salamanders I am struck by the need for more basic biology, particularly their reproductive biology. A major research facility, aimed at multigenerational captive breeding, could, alongside further studies in the wild, well be justified. If the Giant Panda deserved such a major effort, surely the Giant Salamander does as well. Given the massive increase in internal tourism in China, a well-designed centre with stunning exhibits could be a major attraction and a useful source of income.

I can only hope that the conservation measures being supported by ZSL and partners in China are successful and the survival of the giant salamanders in the wild and throughout their range can be assured.

References are in previous post.

The Giant Salamander found in Hong Kong but not of Hong Kong in 1920

Edward George Boulenger

In my last post I described how Edward George Boulenger (1888-1946), Curator of Reptiles and then of the Aquarium at London Zoo, described what he thought was a new species of giant salamander. He sent his account to the Society on 2 January 1924 where it was read on 4 March:

The Zoological Society early last year received a notification from the Marquess of Sligo, who was then in Peking, that whilst recently visiting Hong Kong he had come across a giant salamander which was living in captivity in the Botanical Gardens there, and that he had prevailed upon Sir Reginald Stubbs, the Governor of the Island*, to present it to the Society. 

and

I have therefore no hesitation in describing this salamander as new, and I name it Megalobatrachus sligoi, after the Marquess of Sligo, who was responsible for securing the animal for the Zoological Society. 

After describing the specimen he concluded with:

The Marquess of Sligo has kindly provided me with the following notes on the history of this interesting batrachian :-  

‘‘In April 1920 there was an unusually violent storm of wind and rain in Hong Kong which did much damage to the Botanical Gardens. Among other damage, it caused an 18-inch drain to be choked, causing in its turn the pipe to burst. The result was the scouring out of a long length of drain snd much ground round about it. The giant salamander was found on the scour, having evidently been washed down from somewhere, and thrown out at the end. Since its capture it lived in a circular basin 4 feet in diameter with 6 inches depth of water. It was fed once a day on live tadpoles. At times when the supply of tadpoles ran short, small quantities of raw beef were substituted.”  

Sir Reginald Stubbs informs me that it is practically certain that the new salamander is not a Hong-Kong-born creature. It appears that specimens which have from time to time been brought over from the mainland and been placed in the fountain in the Botanical Gardens have escaped. It is therefore highly probable that the salamander under discussion is one of these animals. 

The 6th Marquess and
Marchioness of Sligo

The arrival of the salamander at the Zoo in early June was announced in The Times on 14 July 1923. However, there was no detail of how it had been transported from Hong Kong to London. It did though survive for many years. In 1945 the preserved corpse was given a catalogue number by Natural History Museum where it was sent as the holotype since it was used to describe a new species. It was a major exhibit in the Fresh-water Hall at the opening of the new (and now recently closed) aquarium in April 1924.

Not only was the location of its find surprising but also its size—99 cm in length. How long had it been in the fountain, or living in a drain in or above the gardens? What size was it when released? Had it moved into a drain to avoid the heat (giant salamanders are animals of montane streams)?

Sir Reginald Stubbs

Sir Reginald Stubbs (1876-1947) was a-less-than-successful governor of Hong Hong from 1919 until 1925. The Marquess of Sligo was the 6th Marquess, George Ulrick Browne (1856-1935) who was a fellow of the Zoological Society.

In the Zoological Society’s library there was found watercolour and a pen-and-ink drawing of the specimen. Both were shown on the ZSL website in a post of 3rd April this year written by Ann Sylph, the Society’s librarian. Whether prepared by or for Boulenger is not known but they could well have been used to illustrate his paper that was read on 4 March 1924 as some sort of poster, perhaps for attendees to peruse. However they were not published as part of his paper. The watercolour was included in the recent paper on molecular phylogeny by Turvey and his co-workers from ZSL, Natural History Museum, Toronto, Kunming and Chengdu.

Watercolour in the Zoological Society of London's Library illustrating
Megalobatrachus maximus and M. sligoi

Pen-and-ink drawing from the Zoological Society of London's library

Why did Boulenger think he had a new species?

Although a giant Chinese Salamander had been described by Blanchard under the name of Sieboldia davidiana [i.e. Andrias davidianus], by most authorities the giant salamander of China was regarded as identical with Megalobatrachus maximus (now Andrias japonicus] of Japan. I have myself examined a number of Chinese specimens in the British Museum collection, and can find no distinction between them and the previously described Japanese species. We naturally assumed that the Hong Kong specimen would prove to be the well-known and only Old World species which occurs in the mountain streams of both Japan and China. On arrival in our gardens early in June, the salamander, a large specimen measuring 99 cm. in length, appeared to me to differ in various respects from any salamander previously recorded. Investigation showed that not only was the head longer and flatter than in Megalobatrachus maximus, and without the characteristic tubercles of that species, but that the nostrils were much more widely separated from one another. As a result of there being no prominent tubercles on the head, the eyes, which in M. maximus are scarcely discernible with the naked eye, are in this Hong Kong specimen quite prominent. I have therefore no hesitation in describing this salamander as new… 

He showed the differences as follows:

As I said in the previous post, Boulenger’s form sligoi was regarded as a synonym of davidianus by later workers in the U.S.A. and the name disappeared until the recent paper on molecular phylogeny resurrected it.

The interpretation of Boulenger’s limited data and description was that it was confounded by his using the Japanese form as comparator. He stated that he examined Chinese and Japanese material in the Natural History Museum and found no difference between them. The Natural History Museum had 12 specimens of giant salamander in the 1920s (the present total of 13 includes Boulenger’s specimen). Eight were from Japan; 5 from China. All incidentally are listed today as Andrias japonicus. Why I do not know, perhaps it is some arcane rite of those in the museum world not to update scientific names.

Others did find a difference between the Chinese and Japanese specimens, so did Boulenger actually make more than a cursory examination of the 12 specimens available to him? Perhaps he thought the nostrils in his specimen were more widely spaced than in any of them. I have not, of course, looked at the specimens in London that were available to Boulenger; it would interesting and informative to do so.

A quick perusal of the photographs of giant salamanders available online shows the nostrils more widely separated in davidianus than in japonicus, a reflection of the pointed snout of the latter. His comparison (as evident in the description and diagrams) show that his maximus was indeed japonicus, both with respect to the pattern of tubercles on the head and the distance between the nostrils.

Having used what is clearly japonicus to compare with his specimen from Hong Kong, it is not surprising that he found the latter different. Later herpetologists had no difficulty in shooting his new species down in flames. Liu collected morphometric data from Chinese and Japanese specimens in U.S. museums and presented the raw data for 5 from China and 10 from Japan. Expressed as a percentage of head width (to allow for the difference in size of the specimens), the mean distance between the nostrils was 21.5% in davidianus and 17.6% in japonicus. Using Boulenger’s drawing to take the same measurements (with all sorts of caveats about doing so) the value for ‘maximus’ is 16.7%, i.e. very close to the Liu values for japonicus, while for sligoi it is 29%, a value markedly higher than Liu’s specimens of davidianus. However, taking measurements from the photograph of Boulenger’s preserved specimen shown in the 2019 paper, and estimating from the shape of the snout the likely position of the nostrils (which I cannot discern) the figure I come up with is 22%—no different from the mean of Liu’s 5 specimens of davidianus.

Boulenger's specimen preserved in the Natural History Museum
Even given the shrinkage that occurs in spirit was not Boulenger's claim of a length of 99 cm
when received just a little ambitious?
Photograph from Turvey et al. 1991

The other key point of Boulenger’s description is: 'Head…very much depressed'. The watercolour and drawing make this statement clear to understand although the photograph of the preserved specimen is less convincing. Such a very flat head was not though, I recall, a feature of the Chinese giant salamanders bought in for class dissection at the University of Hong Kong in the 1960s. I will deal with these in my final article in this series, along with the photographs I took.

It is also possible to see from Boulenger’s drawing (which would of course not have been seen by later authors since it was not in the published paper) that the pattern of tubercles on the head is not like that shown for davidianus in Liu’s book or that which can be discerned from my photograph of the specimens for class dissection in the 1960s. Again, this point could be checked on museum specimens.

The evidence from Boulenger’s paper and the illustrations discovered later suggests to me that that there could be difference in the shape of the head and possibly in the pattern of tubercles on the head in the sligo form. Changes in shape during growth of the animal cannot be ruled out, of course. The whole question of whether there are consistent morphological differences between the genetic lineages such as davidianus (in a narrow sense) and sligoi could be settled by modern morphometric analysis on a larger number of specimens.

So, definitive molecular genetics and very tentative (or even tenuous) morphology indicate that the allopatric forms davidianus and sligoi are distinct. The discussion of their status as biological species continues. In the meantime would it not be worth treating them as subspecies, Andrias davidianus davidianus and A. d. sligoi, along with the un-named genetic line from the Huangshan mountains?

Edward George, son of the great George Albert Boulenger FRS (1858-1937) who had retired from the Natural History Museum (then still existing under the awful name of British Museum (Natural History)) in 1920, does seem to have been right about his specimen—there was something different but not necessarily for the reasons that he stated. Boulenger senior who was one of the authorities who lumped the Chinese and Japanese forms into one species.

It would be fascinating to know the sources of supply to the main centre of the trade in giant salamanders, Guangzhou, or Canton to old China hands, and whether ones from nearer (sligoi) or further (davidianus) away were more common as access to remote areas and political changes occurred over the decades of the last century. Was the one washed out of a drain in Hong Kong in 1920 typical of those coming in to the food markets at the time?

There have been claims that a specimen of giant salamander recorded by Arthur Sowerby in 1923 was the largest ever recorded—1.75 metres long. Since it was captured near Guiyang in Guizhou province, it falls within the range of the sligoi lineage and, therefore, the latter may achieve a greater size than davidianus from further north and west, and hence claim the title of world’s largest amphibian.

The fountain in the Botanic Gardens, Hong Kong 1966. I was trying out my new Exakta 35 mm camera.
The configuration of the fountain was changed to this format in 1932 but the size and location were
unchanged from the 1920s.

    

*Boulenger did not know his geography. Hong Kong comprised, and still does, part of mainland China as well as Hong Kong and other islands.

Adler K. (Editor). 2012. Boulenger, Edward G. (1888-1946). In, Contributions to the History of Herpetology, Volume 3, p 208. Society for the Study of Amphibians and Reptiles

Boulenger EG.. 1924. On a new giant salamander, living in the Society’s gardens. Proceedings of the Zoological Society of London, 1924, 173–174 

Liu C. 1950. Amphibians of Western China. Chicago, Chicago: Chicago Natural History Museum.

Turvey ST, Marr MM, Barnes I, Brace S, Tapley B, Murphy RW, Zhao E, Cunnigham AA. 2019. Historical museum collections clarify the evolutionary history of cryptic species radiation in the world’s largest amphibians. Ecology and Evolution 2019;00:1–15. https://doi.org/10.1002/ece3.5257

Chinese Giant Salamanders: How many species are there?

Two well-publicised papers, published in 2018 and 2019, based on analysis of mitochondrial and nuclear genes, propagate the view that there is more than one species of giant salamander in China, that these species are ‘cryptic’ in that they do not differ in appearance, and that some are in imminent danger of extinction through ignorance of their existence. The authors certainly established that there are a number of distinct genetic lineages of giant salamander in China, enough to convince adherents to the phylogenetic species concept that they constitute different species. However, whether they are ‘good’ biological species is a matter that can be debated endlessly.

Giant salamanders in China in the wild are endangered because of over-collecting for the human food trade and from loss of habitat. Those working on the origins, evolution and conservation of these animals face major difficulties. The escape or release of wild-caught and farmed animals transported alive in large numbers throughout China in recent decades means that the geographical origin of newly-collected specimens cannot be guaranteed. By using specimens in museums collected before the vast increase in trade, the authors of the 2019 paper have overcome this problem but at a cost; the number of available specimens was small—just 17 individuals—since the DNA of many early specimens had been degraded by preservation in formalin.

The lineages are associated with the mountain ranges formed during the great geological upheavals that formed the Tibetan plateau starting about 3.3 million years ago. The authors of the 2019 paper realised that two of the lineages could be attributed to species that had been named in the past.

One lineage was be attributed to the classical domain of the Chinese giant salamander—in the northern Yangtze/Sichuan region—and conforms with Andrias davidianus. A second lineage from the Pearl River tributaries in the Nanling Mountains of south-eastern China appears to be that described by Edward George Boulenger in 1924 as a separate species, Andrias sligoi, that was later lumped into A. davidianus. A third lineage from the Huangshan mountains of north-eastern China has never been described as a separate form previously and could be a new, as yet un-named, species.

Boulenger noted morphological differences between what he named Megalobatrachus (now Andrias) sligoi and M. maximus. He would, wouldn’t he, since that is how new species were described then. But just as at the present with molecular genetic data, different people in the trade regarded some differences as too small to justify separation into a new species; others regarded the tiniest difference sufficient.

At that time Boulenger took his father's (George Albert Boulenger) view that any observable differences between Chinese and Japanese giant salamanders were too small to consider them as two species and continued to lump them into one, M. maximus. While he could find no consistent difference between the Chinese and Japanese forms, others claimed they could and the species maximus was split again into the Chinese, davidianus, and the Japanese, japonicus.

Ignoring the arrangement of tubercles on the head (which appear to differ somewhat in the Chinese and Japanese species) Boulenger also differentiated sligoi by its longer, flatter, and smoother head, and by the shorter distance between the eye and lip.

How Boulenger distinguished his new species, M. sligoi

Boulenger’s new species did not survive as such. Later authors argued it was no different or insufficiently different from davidianus and into the latter it was lumped. Liu in his Amphibians of Western China, published in 1950, condemned Boulenger’s erection of sligoi as a separate species:

Neither the length of the head (which is difficult to define, and which was not defined by Boulenger) nor the relative distance of the eye from the labial border are valid characters.

But Liu did not say why the characters are not valid!

We therefore seem to have agreement between morphology (whether declared ‘valid characters’ or not) and molecular genetics published 95 years later. However, I too have doubts about Boulenger's comparison (see later article). But Boulenger may have been right all along, and the name given recently, South China Giant Salamander, Andrias sligoi, an appropriate one. However, that is only if you agree that species should be defined in that way and not by the biological species concept.

Before accepting the increased number of proposed giant salamander species, perhaps it is worth considering whether the results of the recent artificial mixing up of lineages in both China and Japan (A. davidianus has been released into Japan) have not already invalidated that proposal.

‘Good’ biological species do not or only rarely interbreed in the wild. The classic difficulty with deciding whether species whose ranges do not overlap (i.e. they are allopatric) are ‘good; biological species, is that they do not meet in the wild, and the question of if, say, a geographical barrier were to be removed, would they interbreed naturally, cannot be answered. However, if lineages interbreed freely after translocation, then the case for their being biological species is severely dented if not holed beneath the waterline. Indeed, in the 2018 paper, the authors, do show that hybridisation between lineages has occurred in farms. Similarly, hybridisation between Chinese and Japanese forms has occurred in Japan where the former has been introduced.

On the evidence of hybridisation, then, the case for a number of ‘good’ species of giant salamander, according to the biological species concept, is weak.

The authors of the two papers (many of the authors are common to both) are concerned, rightly, with the conservation of giant salamanders and with the measures that should be taken to ensure their survival in the wild. However, the type of evidence used—that different lineages equal different species—is clearly not acceptable to adherents of the biological species concept. Indeed, if it that premise were correct, two of my daughters-in-law would be of a different species to me and my grandchildren interspecific hybrids. Thus, Jerry Coyne, a world authority on speciation from the University of Chicago, in his blog, Why Evolution is True, wrote:

…virtually every paper I’ve seen on the process of speciation—that is, on the ways that new species come into being—deals not with the accumulation of genetic distance per se, but on the development of reproductive barriers that eventually prevent populations from exchanging genes.

Greg Mayer of the University of Wisconsin, making the same general point from a different case on Coyne’s website:

…At the time, this bothered me, as I saw it as an application of the old morphological species concept, extended to genetic data: if you can tell them apart, they are different species. This is also what Jerry argued against… an arbitrary amount of morphological or genetic difference, or inferred time of separation based on the amount of genetic difference, is not a sound basis for a species concept.

Here, sadly, there may be incompatibility between the politics of conservation and science. Conservationists find it easier to provoke politicians into action with measures to protect species rather than populations and habitats—thus the more species, the better the argument and the greater geographical spread of habitat protection.  However, Jerry Coyne has argued cogently:

…the ‘splitting’ of species [in this case discussing the Giraffe, ‘split’ into different ‘species’ by virtue of differences in genetic lineage] is a conservationist motivation, not an attempt to partition out nature in biologically and evolutionary meaningful ways’.

The research on the giant salamanders reminds us of some fundamental biological questions, like the perpetual problem of how we define a species, as well as some urgent practical conservation concerns. Indeed the research highlights the urgent need to protect, manage and re-introduce populations of giant salamanders in the wild.

However we regard Boulenger’s Megalobatrachus (now Andrias) sligoi, the discovery of his type specimen is an interesting story that I will cover in a further article…and can we determine where those giant salamanders used for class dissection in Hong Kong in the 1960s came from?

Boulenger EG.. 1924. On a new giant salamander, living in the Society’s gardens. Proceedings of the Zoological Society of London, 1924, 173–174 

Turvey ST, Marr MM, Barnes I, Brace S, Tapley B, Murphy RW, Zhao E, Cunnigham AA. 2019. Historical museum collections clarify the evolutionary history of cryptic species radiation in the world’s largest amphibians. Ecology and Evolution 2019;00:1–15. https://doi.org/10.1002/ece3.5257

Yan F, Lü,J, Zhang B, Yuan Z, Zhao H, Huang S, Wei G, Mi X, Zou D, Xu W, Chen S, Wang J, Xie F, Wu M, Xiao H, Liang Z, Jin J, Wu S, Xue C, Tapley B, Turvey ST, Papenfuss TJ, Cunningham AA, Murphy RW, Zhang Y, Che J. 2018. The Chinese giant salamander exemplifies the hidden extinction of cryptic species. Current Biology, 28, R590–R592. https://doi. org/10.1016/j.cub.2018.04.004 

Who connects giant salamanders, my father’s win on a horse and my grandmother’s sewing machine?

Arthur de C. Sowerby

The answer is Arthur de Carle Sowerby (1885-1954) who I wrote about in my series on the Duke of Bedford’s Exploration of East Asia.

Sowerby was referred to in a well-publicised paper on giant salamanders which appeared earlier this year. I have not read Sowerby's original publications but he is shown as having thought there might be more than one species of giant salamander in China and also as describing the pattern of trade in live animals for human consumption.

Sowerby published those papers in 1925. Suffering from severe arthritis, he had just had to give up exploring and working as a collector in China for Robert Sterling Clark (1877-1956) an heir to the Singer sewing machine fortune (my grandmother owned a much-used Singer). Clark first recruited Sowerby as naturalist for his eponymous expedition of 1908-09. As well as his interests in art, geography and natural history, Clark was a well-known race-horse owner. His famous horse, Never Say Die, won the Derby at Epsom in 1954. My father drew Never Say Die, a 33/1 outsider, in a sweepstake.

The cover of the book
Sowerby travelled well armed

Since writing the initial account of Sowerby, I have bought the book on his life written by a distant relative. The travels in China for Clark, in the days of warlords and banditti, were extremely dangerous, and he was lucky to escape with his life. By 1925, though, he was ensconced in Shanghai and publishing The China Journal. It was successful and he, with his second wife, seemed for the first time to be free from a shortage of money. He collected pottery, porcelain and books, all of which he donated to the Heude Museum when he left China in 1946.

Robert Sterling Clark

The Japanese brought this relatively affluent part of Sowerby’s life to an end. An initially successful business venture to develop Manchuria folded in 1931 when the Japanese marched in. For a time Sowerby and his wife, who were both ill, escaped internment when the Japanese took the foreign concessions of Shanghai in December 1941. When his wife died he was interned for the last eight months of the war until the Japanese surrender. After returning to Britain (where he met the then Duke of Bedford, the son of the duke who had funded the Exploration and rescued Père David’s Deer), visiting the U.S.A. and returning briefly to Shanghai, then South Africa and South America, he and his third wife obtained residency visas for the U.S.A. They settled in Washington DC for the last six years of his life.

The link with Clark though was not broken. Clark hired private detectives to investigate the state of Sowerby’s finances during his last years in Washington. As a result Clark sent money through these investigators to assist Sowerby and his wife. The Robert Sterling Clark Foundation in New York has the correspondence between Clark and Sowerby including those thanking Clark for his help. There is, however, no evidence that Sowerby and Clark ever met again or communicated other than through the private investigators.

There is more information on the Clark Expedition here, details of an exhibition held the the Explorers Club, New York, organised by the Sterling and Francine Clark Art Institute in 2012.

The title page of the book on the Clark Expedition

The following photographs and artwork are from the book on the Clark Expedition:

The jerboa named after Sowerby by Oldfield Thomas
now lumped into Dipus sagitta (Northern Three-toed Jerboa)

Clark (left) and Sowerby after a Christmas Day shoot for the pot

Sowerby's colour plate of the snake, Coluber dione, now Elaphe dione

                           

UPDATED 1 December 2019

Clark RS, Sowerby A deC. 1912. Through Shên-kan. London: T Fisher Unwin

Sowerby RR. 1956. Sowerby of China : Arthur de Carle Sowerby F.R.G.S., F.Z.S. Kendal: Titus Wilson.

Turvey ST, Marr MM, Barnes I, Brace S, Tapley B, Murphy RW, Zhao E, Cunnigham AA. 2019. Historical museum collections clarify the evolutionary history of cryptic species radiation in the world’s largest amphibians. Ecology and Evolution 2019;00:1–15. https://doi.org/10.1002/ece3.5257