Frogs of China. Alice Boring’s Life and Work. 3. Herklots and Hong Kong Amphibians

As I mentioned in Parts 1 and 2 of this series Geoffrey Herklots sent specimens of Hong Kong amphibians to Peking for Alice Boring to identify. In the first of those papers all of which were published in Hong Kong Naturalist she wrote that the material was collected under the supervision of Dr Herklots for a period of a little over one year, from December 1930 to January 1932…Herklots had arrived at the University of Hong Kong in 1928 to teach biology, just like Boring in Peking, to medical students. Further specimens were sent in 1933 and possibly later but of these only one had not been represented previously. The specimens, number unknown, were collected on Hong Kong Island, on Lantau as well as in Kowloon and the New Territories.

Alice Boring identified 16 species (a 17th Herklots realised had been collected elsewhere and I have omitted any reference to it). To date 24 species of amphibians have been recorded in Hong Kong. Of the ones recorded before 1930 the Herklots collection missed only one.

The following is a checklist of Hong Kong amphibians, using the current scientific name. The ones marked § are those identified by Boring.

§Hong Kong Newt Paramesotriton hongkongensis

Leaf-litter Toad Leptolalax laui. First recorded in Hong Kong 1979

§Short-legged Toad Megophrys brachykolos (from tadpole as M. boettgeri and an adult tentatively as M. longipes )

§Asian Common Toad Duttaphrynus melanostictus

Hong Kong Cascade Frog Amolops hongkongensis. First recorded in Hong Kong 1950

South China Cascade Frog Amolops ricketti. First recorded in Hong Kong 2004

§Paddy Frog Fejervarya limnocharis

§Chinese Bullfrog Hoplobatrachus rugulosus

§Günther’s Frog Sylvirana (Hylarana) guentheri

Brown Wood Frog Hylarana latouchii. First recorded in Hong Kong 1984

§Three-striped Grass Frog Hylarana macrodactyla

§Two-striped Grass Frog Hylarana taipehensis

Big-headed Frog Limnonectes fujianensis (not recorded in Hong Kong between 1917 and 1987)

§Rough-skinned Floating Frog Occidozyga lima

§Green Cascade Frog Odorrana chloronata

§Lesser Spiny Frog Quasipaa exilispinosa (as Q. spinosa). Identified as species in 1975 (see below)

Giant Spiny Frog Quasipaa spinosa. Adults first discovered in Hong Kong 1978 (see below)

Romer’s Tree Frog Liuixalus romeri. Discovered in Hong Kong 1952

§Brown Tree Frog Polypedates megacephalus

§Spotted Narrow-mouthed Frog Kalophrynus interlineatus

§Asiatic Painted Frog Kaloula pulchra

Butler’s Pigmy Frog Microhyla butleri. First recorded in Hong Kong 1964

§Ornate Pigmy Frog Microhyla fissipes

§Marbled Pigmy Frog Microhyla pulchra

G.A.C. Herklots

As I read Boring’s papers I became intrigued about the two species of Quasipaa, exilispinosa and spinosa, now known to occur in hill and mountain streams. One, Q. spinosa is much larger than the other, reaching 14 cm—the largest anuran in Hong Kong. Although the drawing used to illustrate Boring’s paper is clearly of Q. exilispinosa (the webbing of the hind limbs does not reach the tip of the toes whereas in Q. spinosa it does) she wrote: ‘My material from Hong Kong includes two very large specimens and 7 medium-sized ones…’ Unfortunately, she provided no quantitative information but there remains the possibility that the collection sent by Herklots did contain the two species of spiny stream frogs rather than just the one.

This collaborative effort between Alice Boring and Geoffrey Herklots was the first organised survey of the amphibians of Hong Kong. It provided the basis on which others have built, starting, of course, with John Romer who began his activities soon after his arrival in the late 1940s.

I do not know if Boring and Herklots ever met. There was one opportunity in Hong Kong in 1937 when Boring, caught away from Yenching during the Japanese advance, made her way back to the university by first travelling south.

Paddy Frog (Fejervarya limnocharis)

Boring AM. 1934. The amphibia of Hong Kong Part I. Hong Kong Naturalist 5, 8-22

Boring AM. 1934. The amphibia of Hong Kong Part II. Hong Kong Naturalist 5, 95-107

Boring AM. 1936. The amphibia of Hong Kong Part III. Hong Kong Naturalist 7, 11-14

Karsen SJ, Lau M W-N, Bogadek A. 1998. Hong Kong Amphibians and Reptiles. Second Edition. Hong Kong: Provisional Urban Council

Frogs of China. Alice Boring’s Life and Work. 2. Legacy: Liu Cheng Chao

Alice Boring made good use of her time on long leave in 1928-29. From correspondence with the leading American herpetologists of the day and by working at the American Museum of Natural History in New York she had, by the time she sailed back to China, compiled a checklist of amphibians from all parts of China. The checklist was published with Nathaniel Gist Gee (1876-1937)*. According to her biography she was instilled with what was then the American way of doing taxonomy: collecting a wide range of specimens over a wide area in order to determine the degree of variation within and between species. She established a network of people who sent her specimens as well as sending students and collectors on collecting trips to various parts of China. Because museums depend on access to specimens she sent and exchanged amphibians for study

Geoffrey Herklots at the University of Hong Kong, was, to go right back to the opening paragraph of Part 1, part of that network that existed throughout China and to the U.S.A. He sent collections of frogs to Peking for her to identify. In turn she published the findings in Hong Kong Naturalist.

Boring was a founder member of the Peking Natural History Society and it was in the the Society’s bulletin and a handbook that she published most of her work. However, it was not all taxonomy as some biographers have implied. One paper was on Bidder’s organ while another was on seasonal changes in the reproductive organs of frogs and toads, both continuing the sort of work she was doing in Maine before she left for China.

Liu Cheng Chou
from Contributions to the History
of Herpetology

Boring’s major legacy was a student Liu Cheng Chao (1900-1976). Not officially her student, Alice Boring provided advice and encouragement for Liu’s interest in amphibians at Yenching. After graduation, Liu lost his herpetological papers and books in 1931 when forced to leave Northeastern University at Mukden when the Japanese moved in. Boring took great trouble in  arranging for him to study for a Ph.D. in the U.S.A. funded by the Rockefeller Foundation. He went to Cornell to work with Albert Hazen Wright (1879-1970). Back in China, Liu was first at Suzhou University but the Japanese advance took him to Chengdu in Sichuan and the West China Union University. It was from there that he made 11 long field trips to the hills and mountains on the edge of the Tibetan Plateau. On one he was seriously ill with typhus fever. In September 1946 Liu returned to the U.S.A. and was based at what was then called the Chicago Natural History Museum, now the Field Museum of Natural History.

Liu was with some big names of classical herpetology in Chicago: Karl P. Schmidt, Clifford H. Pope and Robert F. Inger. Pope, of course, had travelled extensively in China and written a book on the reptiles. During his year there Liu wrote his major work Amphibians of Western China. Much more than a catalogue of amphibians it describes the field trips, the peoples he encountered and the geography. He discusses the adaptations of adults and tadpoles in relation to habitat as well as the history and cultural significance of the various species. It is not surprising that after this tour-de-force Liu was described as China’s most prominent herpetologist.

By the time Amphibians of Western China was published in 1950 (Liu’s Preface was dated 1 November 1948) he had returned to Yenching University as head of department. In 1951 he moved to Chengdu as President of Sichuan Medical College. He became a member of the Chinese Academy of Sciences. He and his wife Hu Shu Chin (Shu-qin) (1914-1992) later published Chinese Tailless Amphibians in 1961.

Liu acknowledge the part Boring had played in his career in Amphibians of Western China:

My former teacher at Yenching University, Dr. Alice M. Boring, has my grateful remembrance for her continued encouragement and help during the war years. 

In 1945 Liu named a spectacular amphibian, Vibrissaphora (now Leptobrachium) boringii in Alice Boring’s honour. This species is known as the Emei Moustache Toad because the males in the breeding season have sharp tubercles around the upper lip. These spines are used in defence of their nests. A frog full of vim and vigour named for a ‘dame full of vim and vigor’. And, surely, it should have the common name, Boring’s Toad?

A plate from Liu's book showing the species named for Alice Boring
I have added a red arrow to show the sharp tubercles

Another plate from Amphibians of Western China

A modern photograph of Boring's Toad
from Hudson & Fu 2013

*Yenching University had, incidentally, originally lined Gee up for Boring’s job when he was free to take it. That is why she was only offered a two-year appointment.

Anon. 2014. Boring, Alice M. In Contributions to the History of Herpetology (Volume 1, revised and expanded), Edited by Kraig Adler, pp 107-108. Society for the Study of Reptiles and Amphibians.

Anon. 2007. Hu, Shu-qin (1914-1992). In Contributions to the History of Herpetology (Volume 2,), Edited by Kraig Adler, pp 207-208. Society for the Study of Reptiles and Amphibians.

Anon. 2014. Liu, Cheng-chao (1900-1976). In Contributions to the History of Herpetology (Volume 1, revised and expanded), Edited by Kraig Adler, pp 123-124. Society for the Study of Reptiles and Amphibians.

Fu L. 2017. Nathaniel Gist Gee’s contribution to biology in modern China. Protein & Cell 8, 237-239 DOI 10.1007/s13238-016-0318-x 

Hudson, CM, Fu, J. 2013. Male-biased sexual size dimorphism, resource defense polygyny, and multiple paternity in the Emei Moustache Toad (Leptobrachium boringii). PLoS ONE 8(6): e67502. doi:10.1371/journal.pone.0067502 

Liu C-C. 1950. Amphibians of Western China. Fieldiana: Zoology Memoirs Volume 2. Chicago: Chicago Natural History Museum

Ogilvie MB, Choquette CJ, 1999. A Dame Full of Vim and Vigor: a Biography of Alice Middleton Boring: biologist in China. Amsterdam: Harwood.

Zheng. 2015. Alice M. Boring: a pioneer in the study of Chinese amphibians and reptiles. Protein & Cell 6, 625-627 DOI 10.1007/s13238-015-0165-1 

Frogs of China. Alice Boring’s Life and Work. 1. Biography

As we explored the Hong Kong University Compound after our arrival in November 1965 we came across the lily pond. Seeing the occasional plop as a frog jumped into the water, we decided to continue the exploration in the evening. As well as collecting two nightwatchmen as spectators, several frogs were caught. We had no idea what they were so the next day tried to identify them. Fortunately, there was a tiny collection of offprints in the old zoology department’s office. Some were the checklists given by John Romer but others were from the Hong Kong Naturalist series of the 1930s. How those offprints had survived or had been obtained later we had no idea since the then newly-opened science building had been stripped of everything during the Japanese Occupation. However, they were there and as well as identifying the frogs as Günther's frog, Sylvirana (formerly Rana and Hylarana) guentheri, we became acquainted with the name of the author of the key paper, Alice M. Boring.

Alice Middleton Boring*

Only a couple of years ago when I read her short biography in Contributions to the History of Herpetology did I realise that she was the doyenne of amphibians in China. I also found a book on her life had been published in 1999, A Dame Full of Vim and Vigor. That provides a highly detailed account of her entire life together with background on political and military events unfolding in China during the 20th century. After reading it I see why the authors chose that title.

Alice Boring, as well as living in China in interesting times—very interesting times—was, in the early decades of the 20th century, part of the cutting edge of biological research in the U.S.A. Then, in China, she realised that it would be impossible to pursue such studies and turned to describing and cataloguing the amphibians of China. It is for these latter works of scholarship that she is still remembered today, rather than for the original research on cytogenetics with world leaders in their fields of her earlier years.

From a school in Philadelphia and part of a largely Quaker family, Alice Middleton Boring, gained admission to the Quaker-founded women’s college Bryn Mawr in Pennsylvania. Teaching at the college were Nettie Stevens (1861-1912) who amongst some confusion about credit at the time appears to have discovered sex chromosomes. There was also Thomas Hunt Morgan (1866-1945) who went on to win a Nobel Prize for his work on genetics. After a Ph.D. under Stevens she worked in Würzburg (with Stevens) where they annoyed Theodor Boveri as “the purest blood suckers” (i.e. visiting scientists who visit for a very short time, take up the host’s time and resources and emerge with only superficial findings) and then in Naples.

Returning to the U.S.A. in 1910, Boring got her first job in the agricultural college of the University of Maine where her boss was Raymond Pearl (1879-1940). It would take a book to describe Pearl’s life and career, his many wrong-headed ideas as well as some better ones. Suffice it to say that Boring collaborated with Pearl on reproduction and the plumage pattern of domestic chickens. She advanced rapidly through the ranks to be Associate Professor by 1913. Then, in 1918 at the age of 35, she upped sticks for China on a two-year appointment.

The job she went to was a new one. Rockefeller money had bought out the old Union Medical College in Peking from the joint control of British and American missionary societies and renamed it the Peking Union Medical College. There was though pretty open warfare between the missionaries and Rockefeller employees even though in the end the board that ran the college had input from both. Boring arrived at the time of expansion into a new pre-clinical school. She was fiercely anti-missionary, taught there for two years and at the end of her contract returned to the U.S.A. as professor of zoology at Wellesley College.

Boring was apparently frustrated by Wellesley. Zoology was treated as a poor relation. She entered into negotiations to return to China. It was intended that a new university organised on American lines, Yenching, formed by merging various colleges in Peking (I will stick to the Wade-Giles transliteration instead of the Pinyin ‘Beijing’) should act as a pre-clinical feeder to the Peking Union Medical College, and should take over that function entirely from the College. The university itself was eager not to give Boring a long-term position since they had somebody else lined up for the permanent job. Eventually, she was reassured about her many concerns by the head of the university, John Leighton Stuart (1876-1962) and after making sure she would not be on a missionary’s salary, and that the position at Wellesley would be kept open for her return, she was all set to go: “It would seem like a bit of real contribution to civilization as opposed to teaching in America”. She did not return to Wellesley, preferring Chinese students eager to learn to American young ladies keener to acquire a Mrs as a pre-nominal than a bachelor’s degree after it.

Alice Boring arrived at Yenching University in 1923. She  landed at a time of considerable political and military turmoil which continued until and after her final departure 27 years later. Like her boss, Stuart, Boring took an anti-Western stance. Anti-missionary initially, her position became more nuanced when she argued that the christian groups were also opposed to imperialism and capitalism. She—and Stuart who was born in China, the son of American missionaries—supported the political aspirations of the students, as anti-Western and anti-Japanese rioting flared up and as battles raged between warlords.

Yenching University ca 1937*

Boring was remembered as an austere and tough but kindly task-mistress in preparing students for admission to the medical school. Reading the accounts of her student laboratory sessions with regular tests, my impression is one of overteaching and more like a school class than what would have been expected at the time in an English university. She certainly demanded high standards but also ensured the students were looked after and became proficient in the couth and culture required to move in Western circles and to spend time in the U.S.A. for postgraduate studies. Students who succeeded clearly thought highly of her, both for her knowledge and as a mentor even though she was known by students of both sexes to favour the men. That she was prim, proper, fierce and opinionated there is no doubt. But beneath the prim exterior was something very kind and a little more wild. On a camping trip in Mongolia soon after her arrival in China she wrote: ‘…we take shower baths in the morning by rolling naked in the long wet grass’. Her brother wrote of her as enthusiasm as a ‘menace to society’.

Teaching clearly occupied a great deal of her time (she was in her green lab coat and spectacles nicknamed the ‘Green Grasshopper’ by her students) in Peking. Research was a different matter. She decided that it would be impossible to continue her interest in what was then seen as modern biology. Instead she turned towards the description and taxonomy of the amphibians and reptiles of China. During a long leave at the University of Pennsylvania in 1928-1929 she was in contact with the leading herpetologists of the day to learn all she could about the field and what she needed to do in order to become proficient. At the same time she avoided most of her old acquaintances in cytology and genetics who would undoubtedly have looked down their noses at a move to the world of dead frogs and museums. Pearl though, with whom she was in contact, was supportive. She had already met and helped Clifford H. Pope (1899-1974) on one of his numerous expeditions to China. Further assistance and support was obtained from G.K. Noble (1894-1940) and Albert Hazen Wright (1879-1970).

Boring returned to Yenching to enjoy the new laboratory she had designed and the formal opening of the university. However all was not sweetness and light. First there was the need to raise cash after the Wall Street Crash of 1929; Boring helped but reluctantly. Second there was Stuart, a widower since 1926, who attracted a coterie of female fans. Boring emerged as top dog after what can only be described as a cat-fight with her former friend and house-mate Grace Boynton who taught English. While Boynton was on leave, Boring began to have Sunday dinner with Stuart. When Boynton returned she joined them. Boring asked her to leave. The resultant mutual loathing lasted for many years even though in the end they came to some form of accommodation if not renewed friendship. Students saw Boring and Stuart walking hand in hand over the bridge on campus and tongues began to wag. One staff member noted that Boring became, in effect, social secretary and hostess to Leighton Stuart. She rode with him and travelled with him to make up a foursome on a camping trip. Members of the English department wrote a poem about her ‘something about the power behind the power behind the power behind the throne’. Whatever happened between them though, Stuart avoided permanent commitment. Indeed, during the Japanese occupation he brought one Alice Gregg to the campus. Gregg described herself as “just Stuart’s Shanghai Sweetie”. Grace Boynton described Boring as “frantic” but that Stuart “never seemed aware that AMB was green with jealousy”.

Leighton Stuart's house, Yenching University*

External political forces continued to have major effects on the university campus and student body. After Chiang Kai-shek became established as top warlord in 1930 to head a Nationalist government, Japanese encroachment and occupation of Manchuria was not met by firm action. Student protests demanding action against Japan erupted. Some faculty members backed the students; others Chiang Kai-shek. Boring met Madame Chiang (a former Wellesley student) and was impressed. Time after time, some incident sparked off protests and strikes. Eager to ensure the students did not waste all their time on patriotic protests, Boring once found herself on the wrong side of them when she attempted to hold a lecture during a demonstration.

1937 saw a major shift. Japanese forces pushed into China and occupied Peking. At the time Boring and other staff members of Yenching were on local leave 70 miles south of Peking. They made their way back via Canton and Hong Hong and the university carried on operating with the Japanese in occupation while to the south were the Nationalists and the Communists. Students from the south crossed the lines to and from Yenching, eager to get the best education in Yenching while becoming open to accusations of being unpatriotic. There was a degree of rapprochement between Boring and Boynton with the latter becoming the conduit for information from Yenching to Boring’s family in the U.S.A. since she chose to be evacuated (until returning to unoccupied China) while Boring was insistent on staying on.

After the attacks on Pearl Harbour and on Hong Kong, the foreigners in Yenching were interned, first on campus and then into houses left empty by American diplomats. Finally on 25 March 1943 they were moved to a camp—foul it goes without saying—in Shandong. However, she was not there that long because Alice Boring was down for repatriation in exchange for Japanese internees. She sailed with other Yenching staff on 24 August on board Teia Maru (crowded, dirty and short of food and water) for Goa, then belonging to neutral Portugal. A neutral ship, the Swedish M.S. Gripsholm then carried the 1,440 Americans and Canadians to Jersey City.

Alice Boring clearly felt it hard to be back in the U.S.A at the age of 60. Nobody, least of all her family, was really interested in her life in China and like so many long-term expatriates she remained wrapped up in Chinese affairs. She was a supporter of Chiang Kai-shek, even while recognising the shortcomings of his regime. A couple of temporary teaching jobs occupied some of her time. It is not surprising that she headed back to Yenching as soon as it was possible to do so.

Returning to Yenching in September 1946, exactly one year after VJ-Day, she set about gathering equipment for her course. But the civil war was close. Stuart was made U.S. Ambassador to the Chinese government—an episode that did not end well. The end of 1948 brought Nationalist forces retreating through the campus followed by very polite Communist soldiers who got on well with both staff and students. The initial soft approach of the new regime to Western interests hardened. Political events and family tragedies beckoned Boring home. She left Yenching in August 1950 and died on 18 September 1955, aged 72. Grace Boynton, Alice Boring’s rival for Leighton Stuart’s attention, was given the job best described as ‘historical executor’, sorting her letters and papers on her life at Yenching for archiving.

In Part 2 I will cover Alice Boring’s contributions to herpetology and her lasting legacy in China.

*Photographs from International Mission Photography Archive, University of Southern California

Ogilvie MB, Choquette CJ, 1999. A Dame Full of Vim and Vigor: a Biography of Alice Middleton Boring: biologist in China. Amsterdam: Harwood.

Anon. 2014. Boring, Alice M. In Contributions to the History of Herpetology (Volume 1, revised and expanded), Edite by Kraig Adler, pp 107-108. Society for the Study of Reptiles and Amphibians.

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