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 

Housekeeping: Replacement photographs

Unfortunately, a number of photographs I have added to posts over the years have now been removed from Getty Images. They therefore disappeared from my blog posts. I have now been through all the posts and replaced the blank spaces with other photographs.

I have also amended a few articles and made a note of when this was done at the foot of each post.

History of the Golden Hamster: 6. Golden Hamsters kept in Scotland at the turn of the 20th Century

All the captive Golden Hamsters up to the 1970s were descended from a single litter collected by Israel Aharoni near Aleppo in 1930. That statement is correct but I was surprised when I learnt of a colony of Golden Hamsters being kept in Britain for thirty years until around 1910.

Seven years ago, I was scanning the magazines in W.H. Smith’s and noticed in a now-defunct but then I think new monthly, Small Furry Pets, an article by Dr Karl Shuker on the history of the Golden Hamster.

I was particularly interested in one paragraph:

Nothing more emerged regarding this newly-described species until 1879, when some live examples were brought back to England by James Skene, who had been working in the diplomatic service in Syria. These thrived and bred for 30 years…Then in 1910 the last generation of progeny descended from those original specimens died without breeding, and the golden hamster once again sank back into obscurity.

I was so intrigued that I bought the magazine and kept a copy of the article.

The only reference online to the hamsters brought back by James Skene was to an article written in 1992 by Chris Henwood, a noted keeper and writer on small rodents, for the now-defunct British Hamster Association:

So now if you take the old story as the truth then little was heard of the Golden Hamster for almost exactly 100 years. Wrong! In fact a group of unknown number was brought from Syria to the UK by James Skeene. He had been British Consul to Syria and on his retirement in 1880 returned to Britain. I can sadly find no reference that he wrote anything about the species in general or his animals in particular, and if anyone can help with this area I would be very grateful. It does however appear that they bred in this country at least until 1910 when the last individuals appear to have either died or been destroyed (perhaps on the death of Skeene?).

4th edition of Reynolds's
booklet

Only when I obtained the paper by the late Rolf Gattermann and his colleagues on the collection and ecology of Golden Hamsters in the late 1990s did I get any further information. They listed the collections (dead and alive) over the years, and for 1880 listed Skene, with the information having come from a book published in 1954 by H.W. Reynolds and published by the Zoological Society of London.

I could not find a copy of Reynolds’s book (really a booklet since it is only 32 pages) for sale. But the ZSL Library has two copies and their superb service to Fellows soon had a copy on my desk. This is what Reynolds wrote:

A single preserved specimen, without any particulars recorded, is in the Beirut Museum, and I am indebted to the Rev. Dr. Rosslyn Bruce D.D., F.L.S., for the only other authentic record of specimens being found in the last century. According to Dr Rosslyn Bruce, his grandfather, James Henry Skene, of Rubislaw—who was for many years Consul-General at Aleppo, caught and bred a large number of Golden Hamsters and upon his retirement, in 1880, he brought back to Edinburgh a number of live specimens which were shared among is family and friends. This stock is understood to have bred in captivity for some thirty years, then the strain died out.

In this story we have three people of interest:

James Henry Skene (1812-1886)

Skene was a soldier, writer and diplomat. He served in the 73rd Regiment of Foot (later and eventually incorporated into the Black Watch). He sold his commission to settle in Greece, travel and write books on the region. His wife, Rhalou, whom he married in 1832, was a Fanariot. According to the family history, ‘He eventually became attached to the services of Lord Stratford de Redcliffe, English [sic] Ambassador at Constantinople, and for his services during the Crimean War was appointed Vice-Consul at Constantinople, and afterwards Consul-General at Aleppo, from which office he retired in 1880, and died at Geneva on 3rd October, 1886’.

Skene’s daughter, Jane, was the mother of Rosslyn Bruce*.

Rosslyn Bruce (1871-1956)

Bruce is listed as a clergyman and a naturalist but I suspect that is the wrong order of precedence of his interests. He was in many ways the epitome of the old CofE wealthy parson. He had many interests—including hoaxes—and clearly enjoyed pulling the wool over the eyes of church authorities. He was born in Nottinghamshire, the son of Canon Lloyd Stewart Bruce and Jane Skene, one of eleven children. After divinity at Oxford (he was secretary of the Union but lost an election for the presidency to the evolution-denier Hilaire Belloc) he became a curate in Soho where he campaigned for better living conditions for children whose parents were on the stage and for the education of mothers in the care of babies (still necessary 120 years later).

His family controlled the living at Clifton in Nottinghamshire and Bruce was installed as rector there. (This story has many of the elements of an Anthony Trollope novel.) He broke his leg while fox hunting, an episode recounted in detail in the Nottingham Evening Post of 22 October 1907. A year later he married Rachel Gurney, a member of the banking family immortalised by W.S. Gilbert in his line from the judge (‘and a good judge too’) in Trial by Jury on how he had become a wealthy lawyer ‘by falling in love with a rich attorney’s elderly ugly daughter‘: ‘at length I became as rich as the Gurneys…’

From Clifton (later the site of the largest council housing estate in Europe) he moved to Edgbaston, Birmingham in 1912. During the First World War he went on Noel Baker’s humanitarian mission to the Balkans and served as chaplain to British troops in France.

In 1923 he became rector at Hustmonceux in East Sussex—and stayed there. The obituarist for The Times (21 January 1956) clearly thought he should have risen higher in the church hierarchy given he background and education. However, he clearly upset the upper reaches as newspaper explained:

He did much to beautify the ancient church of Herstmonceux, and fancifully explained how supernatural agencies had introduced new embellishments, for which the necessary faculty had not first been obtained from higher ecclesiastical authority. Such unorthodoxy of behaviour stood as a barrier to preferment that his abilities and devotion undoubtedly deserved.

The obituary went on to praise the gaiety of his company and his ‘deep love of all fellow creatures’.

Bruce’s interests emerge in the penultimate paragraph:

Rosslyn Bruce’s lifelong pastime was the care and breeding of animals. Horses, monkeys, birds, mice, rats, iguanas, snakes, dogs, ferrets, even bears and an elephant all passed through his cherishing hands. He was known as the greatest expert on smooth fox terriers, and bred several champions. He judged at dog shows far  and wide. One of Queen Victoria’s pets was a Skye terrier from his kennels.

Local press reports show that he gave talks about his animals kept in Nottinghamshire, Birmingham and Hurstmonceux and other animals can be added to the list: canaries, lovebirds, pigeons and a tortoise in his study; kangaroo, porcupines, racoons, chameleons, eagles and fleas. He told the Sussex Agricultural Express that his ambition as a boy was ‘to become the proprietor of a menagerie’.

His winding-up the church hierarchy extended to the news media. He told a reporter he had bred green mice—and the story, in modern terminology, went viral.

Rosslyn Bruce (centre) in 1916

Bruce appeared in the newspapers for another very sad reason that still resonates from 1912. Bruce’s sister, Kathleen, married Robert Falcon Scott—Scott of the Antarctic—and it was Rosslyn who informed the world in February 1913 that Kathleen, en route to New Zealand to meet the returning Terra Nova Expedition, had been informed of her husband’s death on the trek back from the South Pole.

There can be no doubt that the Reverend Doctor Bruce knew what he was talking about when he told Reynolds about the hamsters brought back to Scotland by Skene in 1880. I am still curious about the family and friends who kept the hamsters and how Bruce knew they had died out around 1910. Certainly, a number of the Skene family lived in Edinburgh (and helped bring up the eleven siblings of the  Bruce family after their mother, Jane, died in 1880, and their father in 1886).

Skene had many relatives and descendants. Do any of them know anything more or of anything written about his hamsters? And did Bruce himself have any of the offspring of Skene’s hamsters?

Harold William Reynolds (1907-1966)

UPDATE: Comments below have been used to update the text below.

1st Edition

I have not been able to find out much about Reynolds, the author of the hamster book who was told of Skene’s hamsters by Rosslyn Bruce. Local newspapers show that he was secretary of a fanciers’ hamster club for the midlands and north of England and in 1949 was living at 7 Milford Road, Walton-on-the-Hill, Stafford. He and other fanciers imported ‘panda’ hamsters—a black and white colour form—which had first been bred in the U.S.A. He always appeared in print, as was usual then, as H.W. My impression, from the emphasis on the suitability of hamsters for keeping in schools, is that he was a schoolteacher. However, I found on his marriage certificate that he was a civil servant.

I have found no trace of him after 1950. The fourth edition of his booklet, Golden Hamsters, was published by the Zoological Society of London. The first edition, though, he published himself in 1948 as ‘Belvedere Booklet No. 1’. The website of a hamster breeder in Cambridge states:

Mr Reynolds was a member of the earliest hamster clubs in the UK and held the show prefix ‘Belvedere’ for his animals—which was also the name of his house.

His book obviously sold well to run to four editions. But why was the 4th and last, in 1954, published by the Zoological Society of London and not by Reynolds himself? My guess is that the booklet was useful to recommend to those seeking advice from the Zoo on hamsters during the hamster craze. Why Reynolds revised but did not publish the last edition remains unknown.

U.S. Version

I have just found on eBay’s U.S. site, an American edition of the book with the same text but more photographs. It was published by a Milo G. Denlinger in 1950 and printed in the U.S.A.

Reynolds published a second booklet, ‘Belvedere No. 2’, also in 1948. It was entitled Bishop the Hamster and Other Tales.

Edward Hindle provided an introduction to booklet No. 1 and provided a short review for the Autumn 1948 edition of Zoo Life. His introduction, written when he was Scientific Director at the Zoo, takes us back to the reason I began writing this series in the first place:

I received two pairs of these animals from Dr. S. Adler in the following year, and built up a stock maintained first at Hampstead and subsequently in my department at the University of Glasgow From the outset I had no difficulty in breeding them and was able to distribute specimens to any one interested. The London Zoological Gardens received a few pairs shortly before the war, which bred so prolifically that they were put on sale, and helped to make the Golden Hamster known to a much wider public.
Their rapid increase in general popularity dates from the end of the war and now they seem to have become thoroughly established as a domestic pet. 

UPDATED 23 October 2025

*Francis Rosslyn Courteney Bruce at birth; Rosslyn Francis Courteney at death.

Reynolds HW. 1948. Golden Hamsters. 1st edition. Stafford

Reynolds HW. 1954. Golden Hamsters. 4th edition. London: Zoological Society of London

Reynolds HW. 1950. Golden Hamsters. USA: Milo G Denlinger

Skene WF (editor). 1887. Memorials of the Family of Skene of Skene. Aberdeen: New Spalding Club

History of the Golden Hamster: 5. The rejected hamster

Domestication of the Golden Hamster was a contingent event: contingent on Israel Aharoni being asked to collect hamsters to replace the Chinese hamster; contingent on their capture, rearing and breeding; contingent on their suitability as a host for leishmaniasis, and contingent on their attractiveness and suitability as a domestic pet. Had they not been suitable for research on a nasty human disease or unable to be bred in the laboratory they would soon have been discarded by Saul Adler. Indeed that is exactly what happened to another species of hamster collected by Aharoni in Syria for Adler.

The standard animal for leishmaniasis research was the Chinese Hamster. As I wrote previously Adler imported them from the Far East since his animal house was unable to breed them. There has been some discussion of whether Adler knew there were two species, including the Golden Hamster, in the Middle East. However, it is quite clear from from Adler’s account written in 1948 that it was Cricetulus phaeus that he asked Aharoni to bring back alive, a species closely related to the Chinese Hamster (then Cricetulus griseus) and, therefore, perhaps likely to be suitable as a host for Leishmania. This is what Adler wrote:

Prof. Aharoni undertook to bring specimens of Cricetulus phaeus for our work. In addition to getting specimens of Cricetulus phaeus (which were unsatisfactory because they did not breed in captivity), Prof. Aharoni brought back a litter of eight golden hamsters collected near Aleppo…

So, because the Golden Hamster was found easy to breed and to be a suitable host for Leishmania, that species was retained and maintained in Jerusalem. Mesocricetus (then Cricetus) auratus was in: the smaller Cricetulus phaeus was out.

Cricetulus phaeus has now been lumped into Cricetulus migratorius and is known as the Grey (or Gray in the U.S.A.) Dwarf Hamster. It has a wide distribution from Eastern Europe to Mongolia. The Chinese Hamster is now the Striped Dwarf Hamster, Cricetulus barabensis, formerly griseus, found from Russia to Korea and as far south as Shanghai.

Both species have been bred regularly in captivity. Indeed Chinese Hamsters are fairly standard laboratory animals and sometimes seen in pet shops. Why Adler’s animal house was unable to breed either is not known, since the methods of introducing males and females used for Golden Hamsters also work for these much smaller species.

Adler S. 1948. Origin of the Golden Hamster Cricetus auratus as a laboratory animal. Nature 162, 256-257.

Kryštufek B. 2017. Family Cricetidae (part). Pp 282-283 in: Wilson DE, Lacher TE, Mittermeier RA (editors). Handbook of Mammals of the World, Volume 7, Rodents II. Barcelona: Lynx Edicions.

Haslauer R. 2017. Family Cricetidae (part). Pp 283-285 in: Wilson DE, Lacher TE, Mittermeier RA (editors). Handbook of Mammals of the World, Volume 7, Rodents II. Barcelona: Lynx Edicions.

Hobbs KR. 1987. Hamsters. In, The UFAW Handbook on the Care and Management of Laboratory Animals, edited by TB Poole, pp 377-392. Harlow: Longman.