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