Frank Wall: Army Medical Officer and Snakeman of India

I first heard of Frank Wall in the 1960s, in John Romer’s checklists of Hong Kong amphibians and reptiles (see my recent post here). I later heard of his publications on Indian snakes but until I read his potted biography in that wonderful series, Contributions to the History of Herpetology, nothing else. I have now found a more material on his life. Herpetology was an area in which his profession coincided with his interest in snakes. Military authorities have long had great concern about venomous snakes and the treatment of snakebite

George Wall—Frank Wall’s father

Frank Wall dedicated his book on the snakes of Ceylon to the memory of his father, George Wall, reproducing in full the obituary in the Ceylon Independent. A great deal of background to the life of Frank can be gained from the roots his father established in England and in Ceylon and to the wide interests of George in matters scientific as well as in commerce, politics, engineering, journalism and the life of the people of the country of what is now known as Sri Lanka but still to stamp collectors and tea drinkers as Ceylon.

George Wall, who was born in Ashton-under-Lyne, now part of Greater Manchester, on 21 December 1820, was the son of a Methodist New Connexion minister*. Employed from the age of 18 in Manchester by Joseph Whitworth, the great engineer and he of the eponymous screw thread, George Wall became ill in the dank of Manchester and sought employment in warmer climes for the sake of his health. He moved to Ceylon in 1846 as manager of a group of estates belonging to the Ceylon Plantation Company around Kandy. Coffee—not tea—was the crop then being grown and his fortunes rose. In 1854 he founded George Wall & Co, dealing in coffee and the management of estates. In 1870 Ceylon exported 45 million kilograms of coffee and Wall’s company made a lot of money. But then came disaster. The firm foundered in 1879 as coffee rust fungus wiped out the plantations. The fungus reached Ceylon in 1875 and by 1889 production was down to 2 million kilograms. In under 20 years most coffee plantations had been destroyed.

George Wall did not spend his entire time in Ceylon during the time his company was active. He returned to Manchester in 1859, becoming a partner in Sir Joseph Whitworth’s company, where he took a large part in the manufacture and testing of the small arms that made the company so famous. He left for Ceylon again in 1863. His obituary for the Royal Astronomical Society noted that he had observatories at his two residences and ‘devoted much time and expense to astronomical studies’ but that his reduced circumstances after his firm crashed in 1879 and failing eyesight prevented continuation of these activities.

Not knowing the detailed political history of colonial Ceylon I find it difficult to estimate his impact on decisions taken there or in London. He had two spells as chairman of the chamber of commerce in Colombo and represented the colony at the parliamentary enquiry into the affairs of Ceylon in London. He was chairman of the planters’ association and a member of Colombo’s municipal council. He clearly found himself, along with other planters and businessmen, at odds with the colonial government in which he had served a term on the legislative council. He was clearly very active in the abolition of a grain tax (for which he was awarded the gold medal of the pro free trade Cobden Club), in distributing rice from his mills during a famine and in such projects as the breakwater for Colombo harbour, all of which were said at the time to have been of great benefit to the local population. He wrote articles and pamphlets under the pseudonym ‘Speculum’ and edited the Ceylon Independent from 1889 until his death, in London where he had travelled for treatment, on 18 December 1894.

Memorial fountain to George Wall in Colombo (Google Earth)

Apart from his other major interests, George Wall was an amateur botanist, with a particular interest in ferns, publishing two works on the ferns of Ceylon in 1873 and 1879. He was a fellow of the Linnean Society. Botanical interest in the Wall family—George had thirteen children by two wives—was continued by one son, Arnold, who served as Professor of English in Christchurch, New Zealand, for decades.

George Wall is the epitome of that great wave of amateur (in the best sense of the world) interest in natural history that swept Victorian Britain and which resulted in lasting work of great scholarship in many parts of the Empire.

Frank Wall

Frank Wall was born in the relative cool of Nuwara Eliya, the hill town still favoured by holidaymakers from Colombo, on 21 April 1868. He appears in a list of boys educated at Harrow School on Wikipedia but I do not know if that list was compiled from school records; I suspect not. He is listed in the 1881 Census as a pupil at Totteridge Park School while the 1891 Census shows him living there as a medical student. In the meantime he began his medical education at the Ceylon Medical College in Colombo. The Times of 9 April 1890 shows he passed part II of the old conjoint diploma of the Royal Colleges of Physicians and Surgeons in anatomy from Mr Cooke’s School of Anatomy and Physiology in London. Cooke’s private school—the activities of which were the cause of frequent complaints by neighbours—was often the recourse of those from the regular medical schools who needed extra tuition in order to pass the anatomy exam. Wall qualified from the Middlesex Hospital medical school, MRCS LRCP, in 1892. In 1893 he was accepted by the Indian Medical Service. whereupon he trained for a year at the Army Medical School at Netley. In 1894 he left for India and was commissioned in the IMS as a Surgeon-Lieutenant on 29 January 1895.

Wall served as a medical officer with a number of regiments across the Indian subcontinent, including Ceylon and Burma. He was officially part of the Madras Presidency IMS establishment. He was promoted Surgeon-Captain in 1898 and Major in 1907. It was as a Captain that he served in the expeditionary force during the Boxer Rebellion, as I described previously in relation to the snakes of Hong Kong; he received the China War Medal for service in 1900.

Frank Wall in 1919
from here

Advanced to Brevet Lieutenant-Colonel in 1914 and to the substantive rank in 1915 he served in France and Belgium from October 1914 until December 1915; he was Mentioned in Despatches twice in that year. Malcolm Smith, fellow medical man and herpetologist, states in an obituary that Wall’s CMG, awarded  in 1915, was for work at Boulogne, the site of the military hospital complex in France. 

Wall then served in Mesopotamia (Indian Expeditionary Force D) from January 1916 to May 1917. He was appointed Assistant Director of Medical Services (i.e. head of medical services for an army division) in January 1917. Promotion to full Colonel came in January 1921 and then, in 1923, he was appointed an Honorary Surgeon to the King, denoted by the postnominal KHS, until his retirement on 5 March 1925.

Captain Frank Wall married Margaret Georgina Cusse, the daughter of a farmer, in Wiltshire on 30 June 1903. A son, Frank William‡, was born in Bangalore, a major army cantonment, on 9 April 1904. Smith noted that he married Mildred Constance Evans in 1924. There are records from around 1911 onwards to suggest that Margaret Georgina was resident in England while Wall was in India. The probate notice for Margaret Georgina, who died in 1933, states that she was the wife of Frank Wall. However, it would appear that Wall and Margaret had divorced since a family history of the Evans family indicates that Frank did indeed marry Mildred Constance in 1924. Mildred, born in 1873, sailed from Liverpool on 1 August 1924 bound for Colombo on board Bibby Line’s SS Yorkshire. The family history on ancestry.com compiled by Brian Kent (Mildred was his paternal great-aunt) suggests Frank and Mildred were married in Colombo or Nuwara Eliya, which explains why I have been unable to find a record of the marriage in UK records. 

Snakes

Malcolm Smith was clearly well acquainted with Frank Wall and his work on snakes. He wrote:

Wall’s interest in the snakes of India began as soon as he reached that country. In 1895 his name appeared in the list of members of the Bombay Natural History Society and it was in connection with that Society and as a regular contributor to its journal that he came to be recognized as the leading authority in India on the snakes of the country. His first article appeared in 1897, his last in 1925. A complete list of his writings will be found in my volume on the snakes of India and Indo-China (1943) in the fauna of British India series. All together they number 88. Most of his writings consisted of short notes or articles on Indian snakes, dealing with their structure and habits and with taxonomy; but Wall never wrote unless he had a good point to make, and he did not waste words in expressing himself. He took infinite pains to verify his facts, and whatever he recorded is accurate. As a writer he had an easy and lucid style. 

His larger works include The Snakes of Ceylon; The Poisonous Terrestrial Snakes of our British Indian Dominions, a small volume that ran into four editions; and A Popular Treatise on the Common Indian Snakes, This last work, which was published in serial form, is beautifully illustrated in color. It is a great pity that this valuable and interesting series of articles was never brought together and issued in book form. 

Wall’s attitude toward the study of ophiology was many-sided. He tackled the problem from every point of view, not only observing his species in the field, but dissecting them in an attempt to correlate structure with function, and keeping them in captivity to learn more about their ways and modes of life. Wherever he went, he collected. By offering small rewards to the natives he induced them to hunt for him, and in that way many thousands of snakes passed through his hands. Every specimen was critically examined. He made careful notes, and it was upon his notes and his excellent memory that he relied for his identifications and for what he pub­lished. His collection of skulls, all prepared by himself, is a very complete one. Every genus and every common species is represented, often by several specimens. After his retirement he presented this collection, together with his voluminous notes, to the British Museum (Natural History). To the Museum also came the types of all new species described by him, as well as many other specimens illustrating characters of particular interest. 

Wall’s work on snakes, although not his interest in them, ended when he left the country. He was not a Museum worker. His one contribution to pure systematics, namely, A Handlist of the Snakes of the Indian Empire, was, he told me, the most boring piece of work that he ever undertook. His interest was in the living creatures, and when these were no longer avail­able to him he turned to other activities to occupy his time. But his work in India will not soon be forgotten. It is due to him more than any other man that our knowledge of the habits and distribution of Indian snakes.

Malcolm Smith described Wall’s donation of specimens to the Natural History Museum in London. An article in a 2016 issue of Hornbill, the magazine of the Bombay Natural History Society, states that Wall also distributed specimens to many museum in India, ‘especially in the Natural History Section of Prince of Wales Museum of Western India,  Mumbai’.

Typical pages from Wall's series on Indian snakes. The plates were by John Green of London

    

The many-sided attitude to Wall’s study of snakes is evident in his book on the snakes of Ceylon published in 1921. Incidentally, the full title is Ophidia Taprobanica or The Snakes of Ceylon, Taprobanica being derived from the ancient Greek name for the island. Thus he wrote of all aspects of each species. For example, for the python, P. molurus, he has sections as follows: names in Tamil and Sinhalese; Synonymy, History, Identification, Coloration, Habits (haunts, disposition, strength, striking posture, nocturnal or diurnal, hibernation, progression, hissing, sloughing); Food;  Thirst; Breeding (the sexes. method of reproduction, season, period of gestation, period of incubation, number of eggs in the clutch, the eggs); Growth (before hatching, early life, maturity, maximum length, longevity); Parasites (ectozoa, entozoa); Lepidosis; Dentition; Distribution. That one species accounts for 31 pages. The only point I would make in addition to those made by Malcolm Smith is that he had correspondents throughout India, Burma and Ceylon who sent him specimens.

I have found litte by contemporary authors on Frank Wall. However, this Wall story, which shows his interest and venomous snakes and the treatment of snakebite, from the blog of Frederic Alois Friedel:

…here is some background: my father, Alois, a German clock maker and “horologist,” who migrated to Asia to set up navigational stations for ships, was imprisoned during WW I, in a British POW camp in Ahmedabad, in the state of Gujarat†. At the time it was a fairly rural place, surrounded by jungles. The camp had a snake problem, and one of the leading herpetologists at the time was called in to conduct research. Colonel Frank Wall was a member of the Bombay Natural History Society and had published a number of scientific articles and books on snakes…Col. Wall needed an assistant, but none of the British guards were willing to touch snakes. Then he heard about a “crazy German POW” who, he was told, was handling reptiles with equanimity.

So Wall recruited Alois to be his assistant. Together they studied the four poisonous snakes of the region and in fact developed the first anti-venoms against their bite (they injected horses with small amounts of snake venom, until they were immune, drew their blood and centrifuged it to get the serum, which was used to produce the antidote).

Smith noted that a complete list of Wall’s publications was included in the former’s book in the Fauna of British India series. A  more complete list was compiled by Simon Campden-Main for the Smithsonian Institution in 1969.

Retirement

One of ‘other activities’ to which Malcolm Smith referred was genealogy. In his will, Wall left typescript copies of a book he had compiled, Wall of County Derby, on the family from 1539 to 1930 and another on Wills of the Wall family in Lichfield, Staffordshire, to the Reference Library in Derby. The librarian noted that neither bore a date but wondered if Wall had left them to the library because he had used it while compiling the information.

Wall and Mildred retired to England, first to Tonbridge in Kent. In 1939 they were living at Hill Rise, Quarry Hill Road; they employed a resident housemaid. By 1944 the Walls had moved to 30 Milton Road, Bournemouth. Frank Wall died, aged 82, on 19 May 1950 in a nursing home. Mildred died on 16 October 1955 at a nursing home in Cornwall.

Mildred Wall; Frank Wall’s sister, Rowena, Duchess of Somerset; Frank Wall. At Bradley House, Maiden Bradley, Wiltshire, the seat of the Duke of Somerset. Late 1920s. Rowena married Edward Hamilton Seymour in 1881; he became 16th Duke of Somerset in 1925 on the death of his distant cousin, the 15th Duke who died without issue. From Brian Kent’s family tree on ancestry.com.

Frank Wall in 1935

*not as stated in Wikipedia quoting the same source (!); confirmed by parish records. His date of birth is as given in the parish records and not a day earlier (20 December) as in his obituary for the Royal Astronomical Society. Nor have I found any evidence that he was educated at Harrow, again as stated by Wikipedia—does that article confuse him with his son who was a Harrow boy.

‡Frank William Wall (1904-1970) married Thelma Geraldine Lightbody, Wall’s second wife Mildred’s niece.

†I suspect the author is mistaken in the location. The internment camp for German civilians was at Ahmednagar (in Maharashtra), not Ahmedabad (in Gujarat).

Anon. 1895. [George Wall obituary notice]. Monthly Notes of the Royal Astronomical Society 55, 202.

Anon. 2014. Wall, Frank (1868-1950). In Contributions to the History of Herpetology (Volume 1, revised and expanded), Edited by Kraig Adler, pp 71-72. Society for the Study of Reptiles and Amphibians.

Anon. 2016. Frank Wall. Pioneer snakeman of India. Hornbill (July-September 2016), 14.

Campden-Main SM. 1969. Bibliography of the herpetological papers of Frank Wall (1868-1950), 1898-1928. Smithsonian Herpetological Information Services: Washington DC.

Crawford DG. 1930. Roll of the Indian Medical Service 1615-1930. London: Thacker.

Smith M. 1951. Frank Wall, 1868-1950. Copeia 1951, 113-114.

Wall F. 1921. Ophidia Taprobanica or The Snakes of Ceylon. HR Cottle Government Printer, Ceylon.

Captain Wall and the Lady’s Hong Kong Snake

The Indian Medical Service had a distinguished history not only in its purpose of providing the medical requirements of the Indian army but also in improving civilian health. The medical officers often had wide research interests in such non-medical fields as natural history, chemistry, physics, languages and religions. One was awarded a Nobel Prize; several became Fellows of the Royal Society.

In 1900, Britain found itself short of troops. The Boer War was in full spate when the Boxer Rebellion in China broke out. In order to take part in the eight-nation alliance’s campaign, the British government sent troops from the Indian Army, including, therefore, the Indian Medical Service, to form the China Expeditionary Force. As a result, Captain Frank Wall found himself spending time in Shanghai and five and a half months in Hong Kong. Wall had a long interest in herpetology. He visited two museums in Shanghai and the City Hall Museum in Hong Kong in order to make a list of the snake species they had. In addition he visited private individuals with preserved snake collections as well collecting snakes himself.

On 15 December 1902 the Zoological Society of London received a paper from Frank Wall with the wonderful title of ‘A prodromus of the snakes hitherto recorded from China, Japan. and the Loo Choo islands; with some notes’; it was published in 1903. His list of snakes from Hong Kong included those he had found ‘in a swamp near the Sanatorium on the Peak, whilst being drained during the campaign against malarial mosquitoes’. His medical duties must have been at the military sanatorium where some of the Indian Army soldiers would have been patients. Long demolished, the site of the sanatorium is known, and there are photographs on the Gwulo website.

Wall was rather critical of the Hong Kong museum collection:

In the City Hall Museum in Hongkong, out of about one hundred specimens from the territory above mentioned, I found many misnamed and others unidentified. I was informed that during a typhoon some years previously a large case, containing specimens, was blown over and the contents wrecked. Out of the debris labels were recovered as far as possible and replaced, but some were evidently incorrectly reattached and others were destroyed. This circumstance may render the accuracy of some of the records open to question. That this collection is far from representative is evidenced by the fact that during five and a half months' residence in this Port I obtained six species which were not to be found in the Hongkong Museum. 

It was not the specimens in the old museum, nor those he collected himself, that have echoed down the years for all those interested in the fauna of Hong Kong. After describing the known distribution of Dendrophis pictus, he wrote: 

------------------------------

One specimen of this snake in spirit was given me by a lady, who assured me she had obtained it on Hongkong Island.

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Who, incidentally, was the unidentified lady who kept a snake preserved in alcohol? There cannot be too many candidates amongst the residents of 1901. 

In describing that one specimen from Hong Kong he started a debate which continued for 80-odd years. Was it a genuine record? The question was asked because that species had not been seen in the wild nor had any other preserved specimens turned up from any part of Hong Kong—Hong Kong island, the Kowloon peninsula, the mainland New Territories nor the offshore islands? Had the lady acquired it from a relict population wiped out by building work? Could it have been a stowaway in a ship’s cargo—a route of entry of a number of reptiles found in Hong Kong?

Checklists of Hong Kong snakes did, of course, continue to include what is now known as Dendrelaphis pictus, the Painted Bronze Back or Bronzeback. The first edition of the book on Hong Kong’s amphibians and reptiles published in 1986 included the species (with a photograph from elsewhere) in a full page entry, the same as other species. In a review I queried whether, in what was essentially a book aimed at popular readership, it might give the reader the impression that it might be have been or could be spotted in the wild. In the second edition, published in 1998, it was again included because there was positive news: another specimen of Dendrelaphis had turned up in unusual circumstances and in an unusual place.

Dendrelaphis pictus - photograph not from Hong Kong
from Karsen SJ, Lau M W-N, Bogadek A. 1998

Passengers on the present fast boats from Hong Kong to Macau—and those in the old days on the slow boats—pass a number of islands in Hong Kong waters south of the large island of Lantau. One of these, on the port side heading for Macau, is where the specimen was found. Shek Kwu Chau, known to the old sailors as Coffin Island, was uninhabited until 1962. Then a drug rehabilitation centre was established on the island and it became, and remains, closed to visits by the public. The government medical officer at the centre from 1971 until 1984 was Dr James Barrie Hollinrake. During that time he accumulated a collection of pickled snakes, as James D. ‘Skip’ Lazell, who made long herpetological research visits to the islands, including Shek Kwu Chau, of Hong Kong in the 1980s, explained:

Dr Barrie Hollinrake, who runs the place, pickled anything that bit anyone on the island between 1971 and 1984. A medical doctor, he reasoned, that should the victim develop severe symptoms, treatment would depend on correct identification of the biter. Exactly so!

Amazingly, no one ever did develop symptoms, and Dr. Hollinrake was about to dispose of his collection when I heard about it. The snakes and spiders went to the Smithsonian (USNM), but jars and jars and dozens and dozens of vipers, kraits, and all those others—including the amazing, probably novel rarities, are now safely at the Museum of Comparative Zoology [Harvard].

In a paper published with his wife, Wenhua Lu, Lazell went on the write that among Hollinrake’s specimens was a ‘beautiful female Dendrelaphis pictus’. However, he was unable to compare this specimen with the one given to Frank Wall because what happened to the latter is not known. Wall gave all of his later material from India, Burma and Ceylon to the Natural History Museum in London when he retired. Lazell found the Hong Kong specimen was not there, nor, unsurprisingly, in Harvard. Could it still exist in some dusty museum collection? Did Wall take the pickled snake back with him to India where he was part of the IMS establishment in the Madras Presidency? Could it have been or, indeed, still be at the Chennai Museum, in operation since the 1840s? A further clue is contained in an article from 2016 in Hornbill, the magazine of the Bombay Natural History Society. That article states that he distributed specimens to many museum in India, ‘especially in the Natural History Section of Prince of Wales Museum of Western India,  Mumbai’.

Scale counts are the long-established method of distinguishing one species of snake from another. The Lazells compared the scale counts of the Shek Kwu Chau specimen with comparable published data for D. pictus from the range of the species in China. They noted that it had 184 ventrals compared with 186-193 for female D. pictus. There was obviously a misprint in the paper because there is a range (rather than a single value) in the number of subcaudal scales and that range is identical to the published data for D. pictus. Because of the differences, and possibly others—I have not seen the original paper—Lazell described the specimen from Shek Kwu Chau as a new species, Dendrelaphis hollinrakei in a paper published as a Memoir of the Hong Kong Natural History Society in 2002.

What has happened in the past 20 years? The latest reliable information I can find on the status of the Dendrelaphis specimen from Shek Kwu Chau is that written in 2012 for the entry in the IUCN Red List by Michael Lau, one of the authors of the 1986 and 1998 editions of the book on Hong Kong amphibians and reptiles and who had been on Shek Kwu Chau with Lazell. Key points are:

• There have been surveys in the island and no specimens have been found.
• Research is needed to determine if this is a valid species.
• The known specimen of this snake was referred to Dendrelaphis pictus by Karsen et al. (1998), although Lazell and Lu (1990) had previously recognized its distinctiveness. It was formally described as a new species by Lazell (2002). It is similar to D. subocularis, and Lazell (2002) hypothesizes that this may be a relict species that diverged from the common ancestor of this species and D. pictus. Wall (1903) reported D. pictus from Hong Kong; as this specimen has now been lost (Lazell and Lu 1990) it is unclear whether this record is referable to D. hollinrakei.

There have been a number of attempts to sort out the confused status of Dendrelaphis pictus (as far as I can see all based on classical quantitative morphological differences) which in its original sense ranged widely from India to Indonesia. There  have been numerous taxonomic ‘splits’. Perhaps the question marks hanging over the whole Dendrelaphis genus explain why the Agriculture Fisheries and Conservation Department checklist of snakes simply lists Dendrelaphis sp. But even if the taxonomy of the Shek Kwu Chau specimen is re-examined as part of a wider assessment of the Dendrelaphis genus the problem remains of  explaining why individuals have survived, or survived until comparatively recently, in seemingly very low numbers on one island, Shek Kwu Chau, and until over 100 years ago, possibly on another—Hong Kong Island—on the edge of the Pearl River Delta.

I find it odd that just as with Wall’s presumed specimen from Hong Kong Island no other sightings or specimens have turned up either on Shek Kwu Chau or on Lantau. Dendrelaphis species are active snakes, mainly arboreal and often out and about during daytime.

But that isn’t the only serpentine gem from Shek Kwu Chau. In Dr Hollinrake’s spirit jars were two Jade Vine Snakes or Oriental Whip Snakes, Ahaetulla prasina, a species never recorded dead or alive in Hong Kong, but known in South China. And then there’s the lizard, Bogadek’s Burrowing Lizard, first discovered in 1987 on Hei Ling Chau and then on the nearby Sunshine Island but also on Shek Kwu Chau…a story and species for another day.

I found I had Shek Kwu Chau in the background of a family photograph
taken in 2017 on Lantau. The island is about 1.4 x 1.1 km

There is a variety of views on old or relatively old records that seem ‘problematical’, ranging from the collector or reporter being mistaken, their discovery a fraud, a hoax or the result of intentional or accidental introduction, to the acceptance of the record as genuine for which a biological explanation must be sought. Is it possible that the Bronzeback and Oriental Whip Snake were present in Hong Kong and adjacent regions but brought to local extinction by events unknown, leaving relict populations on the offshore islands where they also slowly died out or been extirpated, being arboreal and easy to see, by an expanding or new human population?

Adam Francis has left the Bronze back out of his recent field guide for the obvious reason that the casual herper is highly unlikely to see one. However, If you find yourself wandering nonchalantly amongst trees and shrubs anywhere in the Hong Kong countryside and you spot a slender snake gliding amongst the twigs and branches, you will be entitled to shout Eureka! But carry the thought that most people hiking in Hong Kong never see a snake and the vast majority would not spot or recognise a Bronzeback. There is I suppose the possibility that some could have been hiding in plain sight since the unknown lady gave a pickled specimen to Captain Wall a hundred and twenty years ago.

Francis A, 2021. A Field Guide to the Snakes of Hong Kong. Hong Kong: Adam Francis with Robert Ferguson. ISBN 978-988-74586-2-3. 196 pages.

Karsen SJ, Lau M W-N, Bogadek A. 1986. Hong Kong Amphibians and Reptiles. First Edition. Hong Kong: Urban Council.

Karsen SJ, Lau M W-N, Bogadek A. 1998. Hong Kong Amphibians and Reptiles. Second Edition. Hong Kong: Provisional Urban Council. ISBN 962-7849-05-7

Lazell JD. 1993. Snakes of Shek Kwu Chau. Porcupine, January-February 1993 (4), 7.

Lazell JD. 2002. The herpetofauna of Shek Kwu Chau, South China Sea, with descriptions of two new colubrid snakes. Memoirs of the Hong Kong Natural History Society 25, 1-82.

Lazell JD, Lua W. 1990. Four remarkable reptiles from South China Sea Islands, Hong Kong Territory. Asiatic Herpetological Research 3, 64-66.

Peaker M. 1987. [Review of] Hong Kong Amphibians and Reptiles. Herptile 12, 36-37.

Wall F. 1903. A prodromus of the snakes hitherto recorded from China, Japan. and the Loo Choo islands; with some notes. Proceedings of the Zoological Society of London 1903, 84-102.

UPDATED 3 September 2021

A Field Guide to the Snakes of Hong Kong. A new book

Thirty four years ago I reviewed a book on the reptiles and amphibians of Hong Kong. Because of the efforts of three individuals, inspired by the pioneering work that John Romer did in his spare time in the decades after World War II, more species had been discovered in Hong Kong and more information was accruing on their distribution and life histories. That book, published in 1986, and the second edition of 1998, which again incorporated another phase of new knowledge, have been the standard popular works on the topic. A number of snake species hitherto undiscovered in Hong Kong were found by the simple expedient of searching the deep concrete water catchments after heavy rain for those individuals which had fallen or had been washed in. Since then a newer generation has been observing and photographing snakes in the wild, particularly by exploring after dark with powerful torches the paths and rocky streams.

The results of looking for snakes, chelonians and amphibians after dark have appeared regularly on the YouTube channel SnakeID TV, a must watch for anybody interested in Hong Kong wildlife, and its associated website, hongkongsnakeID.com. The new book, sent to me by AP from Hong Kong, A Field Guide to the Snakes of Hong Kong, is the third result of the triad. It was both written and published by Adam Francis with the help during production of Robert Ferguson.

As we have watched the various episodes on SnakeID TV we have wondered if snakes and chelonians are now more common than they were in the 1960s or if more people are active in looking for snakes. My guess is both. Despite the devastation of the New Territories by the construction of new towns in the late 20th century there have been major beneficial changes: the remarkable recovery of habitat after the removal of trees and shrubs for firewood during the Japanese occupation together with a decrease in the numbers collected for food. Hunters—if that’s the right term for snake and terrapin catchers—scoured all the wild areas to stock the snake shops and ‘wet’ markets.

As with most books printed in Hong Kong this new field guide is beautifully produced. It is a true pocket book size with a true pocket book cover. It is lavishly illustrated and packed with information on 43 species (the sea snakes are not included), as well as general notes on snakes and other sources of information. It is a true field guide because it draws attention to the salient features of identification and how to distinguish between the different species from ones of a similar appearance. Each description is headed with the practical classification of ‘Venomous-Dangerous’ (9 species), ‘Venomous-Not Dangerous’ and ‘Non-Venomous’.

The only species omitted from an overall checklist of Hong Kong snakes are those, four in all, which have only been recorded a few times of which one or two may not be native.

The extent of the information and photographs showing variation are evident from the following two scans of pages showing the Golden Kukri Snake, Oligodon cinereus, an uncommon snake we saw in 1968 just off Route Twisk on the shoulder of the mountain, Tai Mo Shan.

The reader will find that the scientific names of a number of the species have changed both from the earlier book and the checklists shown by the Agriculture Fisheries and Conservation Department and the School of Biological Sciences of the University of Hong Kong. An additional page showing the various synonyms would have been useful and would not have detracted from the book’s popular appeal.

The book does not seem to be available on Amazon. Various sellers of books on reptiles have had copies available via Facebook. Orders can be sent to Hong Kong at the website. My copy was bought in Hong Kong and with airmail suspended had to come by surface mail to UK. I suspect the blockage of the Suez Canal was responsible for its arrival after around 9 weeks. Felt like the 1960s again.

In short: highly recommended! The author and those who helped him should be commended.

Francis A, 2021. A Field Guide to the Snakes of Hong Kong. Hong Kong: Adam Francis with Robert Ferguson. ISBN 978-988-74586-2-3. 196 pages.

See also:

Karsen SJ, Lau M W-N, Bogadek A. 1998. Hong Kong Amphibians and Reptiles. Second Edition. Hong Kong: Provisional Urban Council. ISBN 962-7849-05-7

An uncommon Hong Kong butterfly

Our Hong Kong correspondent sent this photograph of an uncommon butterfly seen this week in a park in Kowloon Tong. It is a swallowtail, Papilio xuthus. Hong Kong is at the southern edge of its range in Asia.

A Zoologist and Forgotten Pioneer of Endocrinology in Britain: Who was Mrs Bisbee?

She published no papers on endocrinology and yet she was the mentor of pioneering endocrinologists who, in turn, disseminated research and teaching in this then burgeoning field in many parts of the world. She also almost fell victim to a partially successful attempt by a British university to remove and disbar married women from employment. But who was this zoologist who, for a while, was a U.S. citizen?

I first heard of the retired lecturer at the University of Liverpool from John Guest Phillips (1933-1987; FRS 1981) and Alan Wright (both Liverpool graduates) in Hong Kong in 1966. John had raised money from the Nuffield Foundation for a research unit in comparative endocrinology in the zoology department of which he was head. I remember the line, ‘…she started everybody off’. Only recently have I uncovered just who some of the ‘everybody’ were, the circumstances of her near-miss dismissal by the university and something of her sad and hidden private life.

Readers of Lord Solly Zuckerman’s (1904-1993; FRS 1943) autobiography From Apes to Warlords 1904-46 (Hamish Hamilton, 1978) cannot fail to have noticed his praise of Mrs Bisbee:

My last year [1924] at Cape Town University [as a preclinical medical student] was a busy one. Mrs. Ruth Bisbee, an exchange lecturer in zoology from Liverpool University, had come that year to Cape Town, and for the first time in my life I learnt what it was to be lectured to in an inspired way, and what a fascinating field of study zoology is. She encouraged me in my ambitious idea of carrying out a comprehensive enquiry into the baboon, and my dissections and other research went on apace.

Zuckerman’s work on baboons led directly to his work with Sir Alan Parkes on the menstrual cycle and its endocrine events, and in turn, to the launching and financing of a new journal in 1939 (which Zuckerman edited for many years), and then a new society, the Society for Endocrinology in 1946.

But Mrs Bisbee’s support of Zuckerman did not end in Cape Town. He wrote of his overspending:

My conscience suffered no twinges when I borrowed from friends, even from Mrs. Bisbee, whom I occasionally saw on her visits to London. She knew only of the scientific work I was doing, not the  way I spent the rest of my time (I am happy to say that in the end I returned to her what I had borrowed)…

In addition to cash, Mrs B was also involved in the ‘hard editing’ of his papers and book that is so essential for a young scientist:

As each chapter [of his book on the social life of monkeys and apes] was finished I sent it for comment to Mrs. Bisbee at Liverpool University. She is the R.C.B. to whom the book is dedicated…I had already sent her the manuscript of my paper on the skull of the chimpanzee. She was tireless and ruthless in her criticism. Draft after draft of the chapters would come back scored through, marked with alterations, and often with sarcastic comments. On one occasion I could bear it no longer, and literally started to weep with frustration. But that was the way to learn…

At Liverpool, Mrs Bisbee supervised students in research in endocrinology. Horace ‘Harry’ Waring (1910-1980), who went on to become the doyen of environmental physiology and endocrinology in Western Australia, did seminal work on the origins of X-zone of the mouse adrenal for an MSc thesis. He noted that she originally suggested the work while helping and criticising throughout. Similarly, James ‘Jimmie’ Munro Dodd’s (1915-1986; FRS 1975) biographical memoir for the Royal Society records:

It was also while at the University of Liverpool that James Dodd first became interested in comparative endocrinology, through the teaching of a remarkable lecturer on the staff called Ruth C. Bisbee, whom he described as quite the best lecturer he ever encountered. It is noteworthy that a majority of the group of scientists that made Britain the centre of development of comparative endocrinology were contemporaries, and many received their undergraduate training at Liverpool in this period.

Ian Chester Jones (1916-1986) (Dale Medallist of the Society for Endocrinology in 1976) was also encouraged by Mrs Bisbee initially in genetics (see later) and then in endocrinology to work, as with Waring, on the mouse adrenal. Chester Jones built up his own research group in Liverpool and then moved to Sheffield, taking John Phillips with him†. 

The importance of Mrs Bisbee in fostering endocrinology in its early days is clear. But who was the ‘remarkable lecturer’? With the help of genealogical search sites, old newspapers and material held in the archives of the University of Liverpool I have found something about her as well as the tribulations of being a female university scientist in Britain in the first half of the 20th century.

Ruth Culshaw Bamber was born on 30 November 1889, the elder of two daughters of a gamekeeper at Lathom, near Ormskirk, Lancashire. Records show she was educated at Burscough Bridge Methodist School from the age of five. She lived in Burscough, also near Ormskirk, for most of her life. The next record is of her graduating from the University of Liverpool, 17 mlles from home, in 1912 with an ordinary degree. The honours degree was an extra year and she graduated with a 1st in 1913. That was followed by an MSc in 1914. The PhD had not then been introduced in British universities and even when it was there was considerable opposition; in Liverpool it was called the ‘phantom doctorate’. Miss Bamber was appointed lecturer in 1915. Until 1919 zoology in Liverpool was called ‘natural history’ and in that year (see below) she was (re)appointed as ‘lecturer in zoology’.

In those early years, Miss Bamber worked closely with Sir William Herdman FRS (1858-1924) the marine biologist and oceanographer, and with his daughter, who worked in the department as a volunteer. She worked on occasion at the marine laboratory at Port Erin on the Isle of Man, an activity restricted by access to shipping in the First World War; she was Edward Forbes Exhibitioner there in August 1915, working on tubularians. An early task in her career was working on fish collected from the Red Sea in an exercise of classical zoology. In doing so she described two new species, the Ornate or Winged Pipefish, Halicampus macrorhynchus, and the Small-Fin Worm Eel, Neenchelys microtretus, for which she erected a new genus and a new family, the Neenchelidae. There is no clue in these early years that she would lead her students—and herself—into pursuits of burgeoning and more important biological topics.

Halicampus macroehynchus Bamber 1915
Nhobgood Nick Hobgood, CC BY-SA 3.0
<https://creativecommons.org/licenses/by-sa/3.0>
via Wikimedia Commons

In 1919 her life changed but perhaps not in the way she expected. 

In April 1918 George Allen Bisbee, aged 34, applied to the US Government for a passport; he intended to visit France and Britain for the YMCA, starting in June. On 24 September when he had to register for the draft, he was secretary of the American YMCA in Liverpool; he requested exemption from serving in the US army because he was a ‘regular pastor in charge of a church’. It was in Liverpool that he met Ruth Bamber and the next record is of her sailing for New York from Southampton on 3 September 1919 as a first-class passenger on board the White Star Liner Adriatic. Tellingly, the records show she intended the USA to be her permanent residence. She married George Bisbee on 13 September in Manhattan, the day the ship docked in New York. However, something clearly went amiss because Ruth Bamber (travelling under a British passport) was back in Liverpool on 2 October; she had travelled alone, listed as a ‘housewife’, on Cunard’s chartered ship Orduña from New York on 23 September—two weeks after her marriage. A distant cousin - see comment below - heard local stories that indicated why she left George Bisbee so quickly. Although I have found no evidence that George was already married, the term 'wives' might have been used in a wide sense.

It is highly unlikely, judging from the passenger lists and shipping records, that she ever met George Bisbee again. He was a school teacher and then qualified as a lawyer; he appears not to have remarried and died in California in 1963.

The reason for her appointment twice as a lecturer, in 1915 and then in 1919 is unclear. It is possible that the first was a temporary appointment for the length of Herdman’s tenure of the chair. Or she could have resigned as was required of a woman at marriage but, wishing to remain on the staff, re-applied for her own job. However, given her initial intention to marry and remain in the USA, her application to have her old job back may have come as she returned to Liverpool from her 10-day stay in the USA. Whatever the reason and intention, she was appointed lecturer in zoology in 1919. 

Back in Liverpool at the age of 30, being married to, but separated from, an American and being known as Mrs Bisbee brought two problems: nationality and eventually continued employment. In 1919 a woman marrying an American became a US citizen and the former Miss Bamber clearly believed she had renounced her British nationality. By 1924 the marriage had been annulled (the actual date has not come to light) and she travelled to Canada for the British Association meeting in Toronto as an American citizen. In 1925 she applied for and was granted renaturalisation as a British subject.

Despite what was clearly a rapid separation and eventual annulment, Ruth Bamber remained known as Mrs Bisbee. It is difficult to appreciate a hundred years later how separation, divorce and, perish the thought, annulment were perceived in 1920s Britain. Scandalous, shameful and unmentionable just about sums it up. In the Methodist world she inhabited (she became superintendent of the Burscough Methodist Sunday school) any revelation of her private life would have been taboo. Indeed Mrs Bisbee did not report to the university the fact that after her marriage was annulled she was no longer a married woman. As a result she came to be included in an infamous incident of the 1930s.

In 1933 the vice-chancellor persuaded Liverpool university’s council that on marriage a woman’s tenure would be ended automatically. Glossed over in the official history was the decree that married women already on the staff would be obliged to resign and reapply for a post which might not exist or be at a lower grade. The national outrage that followed is outwith the scope of this article. However, records in the university archives show that Mrs Bisbee did resign as instructed in 1932 but then withdrew her resignation when it was revealed that she was not actually married. Others who were married were treated shabbily; one woman was given a temporary contract for a year at a lower grade. The whole affair became a cause célèbre and the university was forced to reverse its policy in 1934. Even then one downgraded woman was told that if she applied for her old job she would not get it. The vice-chancellor responsible sailed on to become, as Sir Hector Hetherington, the principal of Glasgow University where a building is named after him*.

After 1919 and the retirement of Herdman, Mrs Bisbee’s research activity changed. Leonard Doncaster FRS was appointed to the chair of zoology. He was a geneticist working on some of the unexplained phenomena of inheritance including the tortoiseshell coloration in cats. With coat colour determined by one autosomal gene and one co-dominant gene with two alleles on the X chromosome, the tortoiseshell phenotype should not occur in male cats. However, sometimes (around 1 in 3000) a male tortoiseshell appears. Nearly all of those are sterile. However, even more rarely a fertile tortoiseshell male is born. Doncaster was suggesting the sort of mechanism that might be at play to account for male tortoiseshells when he died only a year after moving to Liverpool from Cambridge. Mrs Bisbee took up and continued Doncaster’s research, working with Emma Catherine Herdman (1899-1953) who, like Ruth Bamber, had taken a master’s degree in her father’s old department. Catherine Herdman was a lady of independent means** and worked as a volunteer in the zoology department and then, from 1930, as honorary lecturer until her early death from scleroderma.

Catherine Hardman and Bamber, who always published with her name as, ‘Ruth C. Bamber (Mrs Bisbee)’, the style, used in university staff lists well into the 1980s—which reduced the chances of a correspondent committing the solecism of addressing a Mrs as Miss or a Miss as Mrs—published their work on cats (kept in a cattery at the university) in a series of papers published between 1922 and 1933. During their studies they acquired from a breeder a fertile male tortoiseshell called ‘Lucifer’. Although their research did not finally establish the genetical and developmental bases of tortoiseshell males, they did examine and eliminate many possible explanations and one, the suggestion of an extra bit of X chromosome, came very close to the state of present knowledge of an additional X chromosome. The discovery of the XXY condition in some human males came 25 years after Bisbee and Herdman. XXY accounts for the occurrence of the rare tortoiseshell male cats that are sterile. The likes of the fertile ‘Lucifer’ are now attributed to the very rare events of chimaerism and mosaicism. The frustration of not being able to provide at the time a definitive explanation for tortoiseshell tom cats or for such anomalies as the appearance of a few white hairs in black cats, where black is recessive, is evident in their papers, in comments made by Mrs Bisbee to newspapers and in her talk to the British Association meeting in South Africa in 1929 in which she questioned a key tenet of Mendelian genetics. 

Mrs Bisbee and Catherine Herdman regularly attended British Association meetings, including those held in Toronto (1924) and Cape Town (1929). Their interest in cat genetics—not just in the tortoiseshell—attracted publicity and the attention of cat owners and breeders. Newspaper reports show they also worked on the Manx breed where the length of the tail varies upwards from and including zero. Newspaper reports and other records also show that Mrs Bisbee was an active member of the Liverpool Biological Society, the Liverpool Naturalists’ Field Club; she was also a Fellow of the Linnean Society.

Doncaster’s successor in Liverpool, William John Dakin (1883-1950), although remembered for marine biology, also did some genetics research, on abnormalities in domestic chickens. Mrs Bisbee also became involved, investigating a form of inherited dwarfism; the results appeared as a letter in Nature.

Interest in genetics extended to the determination of coat colour in other mammals. Before Ian Chester Jones turned to endocrinology under Mrs Bisbee’s guidance she had him working on the roan coloration of dairy shorthorn cattle. She was also acknowledged for advice in research on congenital tail deformity in pigs.

Mts Bisbee did have her detractors. The wife of a former colleague, herself an undergraduate and postgraduate student in zoology at Liverpool, told me that women had a much harder time than the men. The latter, she assured me, had ‘only to smile nicely’ for preferment; she had one by-then well-known endocrinologist in mind as an exemplar of Mrs Bisbee’s perceived bias.

Because knowledge of her role in promoting endocrinology is now limited to finding acknowledgement of her instigation and interest in published papers and because only those available online show in searches, it is difficult to be certain of when her interest in endocrinology blossomed to the point where she could point postgraduate students to research in that subject. With work on the adrenal published by Robert Roaf (1913-2007) showing experiments that began in early 1932, it would seem that endocrinology replaced genetics as her main interest in the early 1930s. There also seems little doubt that the topic that excited her interest was the X-zone of the mouse adrenal, discovered in the 1920s; both Waring and Roaf were launched into studying its origin in the mouse and to search for evidence of its existence in the rabbit, respectively. Roaf, incidentally, was the son of Catherine Herdman’s sister and the professor of physiology at Liverpool; he became an orthopaedic surgeon and later returned to Liverpool to the chair in that subject.

During the Second World War, Mrs Bisbee’s interests in endocrinology and genetics extended into cancer research. A PhD student, Vera E Jones, worked on the formation and disposal of haemosiderin in the uterine wall of mice after the placenta breaks away during parturition. She found that in a strain of mice highly prone to mammary cancer the haemosiderin, which is confined to phagocytic macrophages, is formed more slowly and disposed of much more slowly. Vera Jones was awarded her PhD in Liverpool in 1944. She went on to work at the National Institute for Research in Dairying at Shinfield. In 1945 she joined the Gerontological Research Unit in  Oxford working on endocrinology.

Mrs Bisbee retired from the university in 1955 as a lecturer, the same grade as she was appointed in 1915. The last reference to her work and the only one concerned with endocrinology I can find was in the Cheshire Observer of 23 January 1954. She gave a ‘talk on glands’ to a local Women’s Institute.

Ruth Culshaw Bamber (Mrs Bisbee) died on 7 June 1970, aged 80. She was then living at an isolated house, occupied by her sister’s family, near Kendal in Westmorland, now Cumbria.

Mrs Bisbee’s scientific legatees fostered the study of endocrinology in many parts of the world, from Hong Kong in the east to California in the west. They have served the Society for Endocrinology as officers and editors. Perhaps we should do more to commemorate an important figure from a time when it was not the done thing to be listed as a co-author of a student’s paper even when the topic and direction was not the student’s. If anybody deserves to be remembered as the foster mother of the science of endocrinology in Britain, that accolade should surely be awarded to Ruth Culshaw Bamber (Mrs Bisbee).

†Which makes Mrs Bisbee my ‘scientific grandmother’ in a succession of research students:

Mrs Bamber ▶︎ Ian Chester Jones ► John Guest Phillips ► MP

*Sir Hector Hetherington, as principal of Glasgow University, was chairman of the board of my research institute from 1936 until 1961. Delving into the archives one day in the early 1980s I came across minutes and correspondence on employment and accommodation matters. I remember being shocked, even allowing for employment practice at the time, about his attitude to a female member of staff and his enquiries into the domestic abilities of a wife before agreeing to the housing of an employee on site. Had, I known what I know now I would have realised that he had previous form.

**Catherine Herdman adopted three children (two boys and a girl), all of whom took the surname Herdman.

UPDATED 29 APRIL 2025

Kelly T. 1981. For Advancement of Learning. The University of Liverpool 1881-1981. Liverpool: Liverpool University Press.

Sumpter JP, Follett BK. 1991. James Munro Dodd. 26 May 1915-15 December 1986. Biographical Memoirs of Fellows of the Royal Society 37, 128-147.

Acknowledgement of Mrs Bisbee in published papers

Brooksbank NH. 1958. Congenital deformity of the tail in pigs. British Veterinary Journal 114, 50-52.

Chester Jones. I. 1947. The inheritance of red, roan and white coat colour in dairy shorthorn cattle. Journal of Genetics 48, 155-163.

Jones VE, 1947. A comparative histological study of haemosiderin in the uteri of mice of cancerous and non-cancerous strains. Quarterly Journal of Microscopical Science 88, 479-490.

Roaf R. 1935. A study of the adrenal cortex of the rabbit. Journal of Anatomy 70, 126-135.

Waring H. 1935. The development of the adrenal gland of the mouse. Quarterly Journal of Microscopical Science 78, 329–366.

Papers published by Mrs Bisbee:

Bamber RC. 1915. Reports on the marine biology of the Sudanese Red Sea, from collections made by Cyril Crossland, M.A., D.Sc., F.L.S. XXII. The fishes. Journal of the Linnean Society, Zoology 31, 477–485.

Bamber RC. 1922. The male tortoiseshell cat. Journal of Genetics 12, 209-216

Bamber RC. 1927. Genetics of domestic cats. Bibliographia Genetica 3, 5-52.

Bamber RC, Herdman EC. 1927. The inheritance of black, yellow and tortoiseshell coat-colour in cats. Journal of  Genetics 18, 87-97. 

Bamber RC, Herdman EC. 1927. Dominant black in cats with its bearing on the question of the tortoiseshell males—a criticism. Journal of Genetics. 18, 219.

Bamber RC, Herdman EC. 1928. The problem of the tortoiseshell male cat. Zeitschrift für induktive Abstammungs- und Vererbungslehre. Supplement 1, 387-390.

Bamber RC (Mrs Bisbee). 1930. Segregation. Report of the 97th Meeting of the British Association for the Advancement of Science, South Africa—1929, pp 333-334.

Bamber RC (Mrs Bisbee), Herdman EC. 1931. Two new colour-types in cats. Nature 127, 558.

Bamber RC, Herdman EC. 1931. The incidence of sterility amongst tortoiseshell male cats. Journal of Genetics 24, 355-357.

Bamber RC, Herdman EC. 1932. A report on the progeny of a tortoiseshell male cat, together with a discussion of his gametic constitution. Journal of Genetics 26, 115-128.

Bamber RC (Mrs Bisbee). 1932. An inherited abnormality in Rhode Island Red poultry. Nature 129,  762.

Bamber RC (Mrs Bisbee). 1933. Correlation between white coat colour, blue eyes and deafness in cats. Journal of Genetics 27, 407-413.

Is the intestinal spiral valve of cartilaginous fish a Tesla valve?

Stand by, English ‘A’ level pupils of a certain age. You will not be able to read this article without a nostalgic whiff of formaldehyde tickling the nostrils. You will remember the first dissection in the series of vertebrate types, a dogfish, and how quickly your attention was drawn by the dissection guide to the structure in the intestine labelled the ‘spiral valve’.

For those reading this overseas I should start by saying that the dogfish sold for dissection by biological suppliers was the Lesser Spotted Dogfish, also known as the Rough Hound. The accepted name for the beast among the fishy cognoscenti is Small-spotted Catshark. The scientific name is Scyliorhinus canicula but it was long known as Scyllium canicula, both names coined by Linnaeus. Why anything named for a dog should then get ‘cat’ as a common name I do not know.

If you wanted to know the function of the spiral valve this is what was to be found in Parker & Haswell*: ‘Absorption takes place largely in the colonic region, which is equipped with an anterior spiral valve. This begins at the end of the duodenum; it is a richly glandular fold of tissue which runs spirally around the intestine and both retards the too rapid passage of food and affords a more extensive area for absorption of digested substances’. The later editions of J.Z. Young’s Life of Vertebrates describes the spiral valve as ‘a tightly wound flap or ridge’.

It is highly pertinent to quote from the text book co-authored by Thomas Jeffery Parker (1850-1897; FRS 1888) since in 1880 he published a seminal contribution to the form and function of spiral valve, as well as devising a fixation method for studying its shape in situ. At that time Parker was lecturer at Bedford College in London and demonstrator in Thomas Henry Huxley’s department at the Royal School of Mines. Shortly after he gave the paper at a meeting of the Zoological Society of London on 16 December 1879 he moved to the University of Otago in New Zealand. He was diabetic and died aged 47.

By studying a number of species of elasmobranch particularly skates and rays, Parker showed there was a considerable amount of variation in the form of the spiral valve; he also calculated the increase in surface area as a result of its presence, in an early example of a mathematical treatment of anatomical features, and argued that passage of digesta would be relatively slow through the ‘valve’.

A recent paper in Proceedings of the Royal Society takes Parker’s work of 140 years earlier on the spiral valve further both in terms of form and function. The 3D diagrams the authors constructed from CT scans are essentially similar to the 2D ones drawn by Parker using his method of distending and hardening the intestine with dilute chromic acid and then cutting holes through the wall to examine the retained 3D structure inside.

The authors confirmed the four types of spiral valve found in different species (and adapted Parker’s drawings to illustrate them): (a) Column, (b) Scroll, (c) Funnels pointed posteriorly, (d) Funnels pointed anteriorly. They then measured the flow-rate of liquids of different viscosities in both directions through fixed specimens of spiral valves of the different types. Flow through the spiral valve was lower in the posterior to anterior direction than in the anterior to posterior direction; statistically significant differences were found for the two types of funnel structure but not for the column or scroll types even though these was a trend in the same direction. I have some concerns about the method used to determine flow rate and the seemingly high degree of variation observed with what was dead, fixed tissue. Those looking at the paper might be mystified by the unexplained notation used to denote statistical significance; I was and still am.

From Leigh et al. 2021

Note: The Order of types of spiral valve is not the same as in the first diagram

The authors suggest that the spiral valve in those elasmobranch fish which have the funnel types can act as a partially effective Tesla valve, favouring movement of digesta through the intestine compared with the reverse direction. In their own words:

The spiral intestine…would allow segmental contractions to better mix digesta in the spiral intestine without the risk of much back-flow. Little is known about intestinal motility in sharks. Typically, overall evacuation rate is used to estimate the length of time that digesta remains in the gastrointestinal tract of sharks. However, by understanding contractile capabilities of the different segments (proximal, spiral and distal intestines) of the shark digestive system separately, we can begin to establish transit rates at specific points throughout the gut. We have begun to do this by measuring the average number of contractions per minute and the average amount of time necessary to move material of a known viscosity through the spiral intestine of S. suckleyi. 

And end with:

In conclusion, our flow rate data suggest that the spiral intestine is acting as a flapper-less Tesla valve, which would promote unidirectional flow without any parts that are susceptible to blockage. We have established, quantitatively, that flow rate is slowed in the spiral intestine. Additionally, the flow rate was slowed significantly more when the two funnel-shaped spiral intestines (anterior and posterior funnels) were subjected to flow in the posterior to anterior direction. This indicates that at least funnel-shaped spiral intestines are capable of producing unidirectional flow, although the spiral and scroll do to a lesser extent. This could explain why digesta transit rates vary among species and among different spiral intestine structures. Further investigation of these unique intestinal structures as a component of the digestive success of sharks is necessary to understanding their function and evolution. The new techniques produced by this project lay the groundwork for future investigations involving the spiral intestine, and for understanding the functional role of the digestive tract in sharks, fishes and vertebrates in general. 

Much has been made of the claimed Tesla valve-like properties of the spiral valve in the popular scientific press, as did the authors in the introduction in a ‘now ask of the beasts, and they shall teach thee’ theme in which biological mechanisms can lead to applications in engineering. Nikola Tesla, incidentally, patented the no-moving parts valve in 1920. I remain somewhat sceptical. While the authors have shown that the various forms of spiral valve CAN act as a sort of Tesla valve there is still way to go before it is finally established that in real life, spiral valves DO act as a Tesla valve, since that function could be an epiphenomenon of a mechanism that evolved, as Parker suggested, to ensure a high efficiency of absorption by means of a slow transit time and a large surface area. Whether or not a Tesla type effect is physiologically relevant would depend on such factors as the the timing and magnitude of peristaltic waves, the muscular tone of the gut at rest and the host of other possible factors, including the thick sludge-liked digesta, that determine transit through the gut. Some good gut physiology seems called for.

The authors also had me mystified on another aspect of their work. They mapped the type of spiral valve onto a cladogram of shark families. The family Scyliorhindae was denoted as having the ‘Scroll’ type spiral valve since two species from that family CT-scanned showed such a structure. However, Parker showed clearly that our old friend the Lesser Spotted Dogfish, Scyliorhinus canicula (closely related, according to the source of the cladogram, to one of the species CT scanned), has the ‘Funnels pointed anteriorly’ type of spiral valve.

Parker's drawing of
dogfish spiral valve

This is what Parker wrote:

In a large specimen of Scyllium canicula [a synonym of Scyliorhinus canicula] I find an especially interesting type of valve…This intestine is shown in fig. 5, Pl. XI.: there are twelve turns to the valve , all but the last of which are strongly deflected forwards, producing a structure which must offer an immense amount of resistance to the passage of intestinal contents, and, of course, making a decidedly greater proportional increase of surface than in any of the cases recorded of the Ray. The difficulty of cleaning out the intestine afforded a good criterion of the forms of these points; the finely divided contents stuck so tightly between the successive “cones,” that a stream of water was often quite insufficient to dislodge them. In fact the chyle (if one may apply the term to what rather resembled fine mud) completely filled up the whole available space in the intestine, so that, although the animal was preserved entire in spirit, the gut and its valve were in as good a condition for examination as if the former had been carefully emptied and distended with spirit while fresh. The pyloric valve was very perfect, having the form of a short conical tube projecting into the bursa entiana, with a very small aperture at its apex. This, of course, brought about the result referred to, that only finely divided matter could find its way into the intestine. 

Another point I may mention about this specimen is the great thickness of its walls at the posterior end ; the thickness was actually greater than the diameter of the lumen at that part. This may have been a mere individual abnormality; but it seems not impossible that this increase of muscular substance had relation to the great force necessary to drive on the contents in a gut with so peculiar a spiral valve. 

In a smaller specimen of the same species there were eight turns to the valve, of which the first five had a forward, the last three a backward direction…

Given that description I can see why ‘valve’ is the right description for the spiral structure.

Parker raised the intriguing possibility that the form of the spiral valve may differ considerably either between individuals of the same species and/or during growth and development of an individual.

A ‘why’ question to be asked, of course, is why have different types of spiral valve evolved?

To conclude, although I found the authors of the new paper had failed to convince me on a number of points, they should, if nothing else, have excited more interest in the functions of the spiral valve and of such old (the spiral valve appears to have been first described in 1671) but important topics in how fish work.

…And, in case you are asking, no, I cannot remember any detail whatsoever of the spiral valve in the three dogfish I have dissected, other than the fleshy folds and their semi-solid contents.

*This is what the 7th edition of 1962 says. As I recall that is no different from the 6th edition, published in 1940.

The new paper:

Leigh SC, Summers AP, Hoffmann SL, German DP. 2021 Shark spiral intestines may operate as Tesla valves. Proceedings of the Royal Society B 288: 20211359. https://doi.org/10.1098/rspb.2021.1359 

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Parker TJ. 1880. On the intestinal spiral valve in the genus Raia. Transactions of the Zoological Society of London 11, 49-61 plus 2 plates.

Parker TJ, Haswell WA. 1962. A Text-book of Zoology (7th edition revised and largely rewritten by AJ Marshall). London: Macmillan.

Vélez-Zuazo X, Agnarsson I. 2011. Shark tales: A molecular species-level phylogeny of sharks (Selachimorpha, Chondrichthyes). Molecular Phylogenetics and Evolution 58, 207–217 doi:10.1016/j.ympev.2010.11.018 

Young JZ. 1981. The Life of Vertebrates (3rd edition). Oxford: Clarendon Press.

‘On Deposit’ at London Zoo: A new essay on zoo history published

I read with great interest but with some disagreement a recent paper in Archives of Natural History. The Patron’s Review is written by invitation of the Society for the History of Natural History to an outstanding young scholar.  The author, Eleanor Larsson, has written on the acquisition of animals by London Zoo between 1870 and 1910 and in particular the practice of animals being ‘on deposit’.

Over the period in question the death rate of animals at the Zoo was extremely high; on average around 30% died each year (but see below). Therefore at least 1,000 animals were needed every year to replace those which had died. These were obtained by donation, purchase, breeding or exchange. In addition animals belonging to other people were placed in the zoo ‘on deposit’ for short or long periods.

Data extracted from the zoo records by Eleanor Larsson (2021)

Animals ‘on deposit’ could range from those presented to members of the royal family by foreign potentates to those from exotic pet keepers who could not quite bear to give their monkey away even though it had grown up and started to bite people. It is on two categories of depositor that Larsson has concentrated her attention: commercial animal dealers and Walter Rothschild who operated on such a scale that he becomes a category as a single individual.

Animal dealers, including the famous Charles Jamrach, based in the East End of London near the docks, William Cross of Liverpool and even Carl Hagenbeck in Hamburg frequently deposited animals at London Zoo while trying to find a customer. The Zoo benefitted because it could use the opportunity to see the health and condition of the animal at close hand before making an offer to buy. The Zoo also benefitted because it kept the place well stocked and with a turnover and degree of novelty that the Great British Public demanded—and still does demand—without having to shell out for an animal. The dealers benefitted because the Zoo provided a show-case for their wares (sometimes fellows the the society who had private menageries), free accommodation for large animals unsuited to their premises and, with Cross and Hagenbeck, for example, as a holding place for their London purchases while they waited shipment to their final owner.

Rothschild, by contrast, had no commercial interest but was responsible for the peak in numbers of animals on deposit between 1903 and 1907. For his museum and for his own study collectors throughout the world sent specimens. He also bought live animals from the London dealers. Many of his live animals were kept on the family estate at Tring. However, after a cassowary—birds of particular interest to Rothschild—attacked his father, he had to reduce the size of his living collection, became a fellow of the zoological society and transferred animals to London Zoo on deposit. It is obvious that over the years he came to regard the Zoo as part of his personal fiefdom, issuing instructions directly to junior staff even after his particular plan had been blocked by those more senior in the hierarchy. However, the Zoo again benefitted because Rothschild provided financial support for the care of his deposits, as well as donating some of the animals in which he had no further interest. One example Larsson quotes is his funding the construction of a house for 50 Galapagos tortoises he acquired from an expedition in 1897-98. In 1900, Larsson calculated, a quarter of the animals at London Zoo were on deposit from Rothschild. On average Rothschild deposited an average of 700 animals per year. That all came to an end in 1907 as financial problems supervened. The story of his being blackmailed by a peeress over many years is now well known.

For the private depositor of animals at the Zoo, the animal remained his or her property—in death as in life. For many such owners the Zoo was an intermediate stage in getting the animal stuffed and mounted for exhibition in their houses. Rothschild had animals sent directly from the Zoo to his preferred taxonomist for the type of animal in question. Sometimes, he did not wait for the animals to die. In order to work on his monograph on cassowaries he had one ‘killed by order’ and sent to a taxidermist in Cambridge.

Did the policy of holding animals on deposit backfire? Larsson argues that it did. Firstly, the Zoo had no control over when the depositor might remove the animal. Secondly, ‘the Society was often compelled by obligation to put the needs, wants and desire of the depositors above its own’. Thirdly, because the depositors, but not the dealers in this case, had their dead animal sent to a taxidermist, the Society missed the opportunity of determining the cause of death by autopsy and therefore ‘prevented the Society from expanding its knowledge about animal disease and by extension, the care that might be provided to animals in life to reduce the occurrence of disease and death’.

It is on this last point—the main thrust of Larsson’s article—that I disagree with the author. The main rôle of the Society’s Prosector in the decades at the end of the 19th century was the study of comparative anatomy but not pathology. Efforts were made to determine the cause of death, especially earlier in that century by the undiplomatic and irascible first Prosector, James Murie (1832-1925) who held the post for around 5 years until 1870, but a pathologist in addition to the prosector was not appointed until 1903. Therefore, if any of the Society’s activities suffered as a result of not having deposited animals available for dissection it was comparative anatomy. Given the overall death rate though there was plenty of material to keep the Prosector occupied.

I also question whether the failure to determine the cause of death of animals on deposit had any material effect on efforts to decrease the annual mortality and morbidity. It is one thing to determine the cause of death, with the attendant difficulties of distinguishing ‘dying of’ from ‘dying with’, and another to be able to do anything about it. Tuberculosis—a common cause of death—could not be cured. Common infections were often the result of poor nutrition, stress and inappropriate housing but very little could be done about those because the knowledge to do something simply did not exist. Vitamins, for example, had not been thought of, let alone characterised.

In essence I found Larsson’s essay a very useful addition to the history of the Zoological Society of London at the end of the 19th and the beginning of the 20th centuries but I cannot agree with her main point that the policy of accepting animals for exhibition ‘on deposit’ inhibited attempts to decrease the death rate at London Zoo.

Chalmers Mitchell, in his Centenary History, and who became Secretary of the society in 1903, does not mention any particular problems associated with the ‘on deposit’ policy although he clearly had a few problems with Rothschild’s demands in his early years in office. Mitchell never missed the opportunity of having a pop at the regime of this predecessor, Philip Lutley Sclater, and so I am reassured that as far as he was concerned the problem of high rates of sickness and death of the animals was a more general problem, not helped until 1903 by a lack of pathological expertise, and not materially affected by the ‘on deposit’ scheme.

The death rate of around 30%, incidentally, does not accord with the figures obtained by Eric Ashton and Gwynne Vevers in their survey published in 1959. Average annual mortality between 1883 and 1904 was 42%. After the First World War, there was some improvement but even in 1957 it was 28%. 

Although not in the period of Larsson’s essay, holding animals ‘on deposit’ was continued for many decades beyond 1910 but not on the scale of the peak caused by Rothschild’s activities. In the 1950s and 1960s—and perhaps beyond—labels on cages would bear the inscription ‘deposited by’ with the owner’s name. Animal collectors—not just those employed by the Society—lodged their animals at London Zoo while buyers were found or stock sorted. For example, the bird collector Wilfred Frost, stored his animals there as late as 1957.

Larsson E. 2021. “On Deposit”: animal acquisition at the Zoological Society of London, 1870–1910 (Patron's review). Archives of Natural History. 48, 1-21 doi.org/10.3366/anh.2021.0685

Ashton EH, Vevers HG. 1959. The numbers of exhibits, births and deaths in the menagerie at Regent’s Park: 1835-1957, and in Whipsnade Park 1931-1957. Proceedings of the Zoological Society of London 1959, 489-514.

Mitchell PC. 1929. Centenary History of the Zoological Society of London. London: Zoological Society of London

Ellis M. 2010. W.J.C. (Wilfred) Frost: An incomplete biography. Avicultural Magazine 116, 158-183.