Salt Glands in Iguanas; Video footage of a South American iguana swimming along a reef

The metaphorical ink had hardly dried on my post (22 March 2019) about salt glands in Galapagos Marine and Land Iguanas and their hybrids when my attention was drawn to a recent video of a common South and Central American iguana, the Green or Linnean Iguana (I. iguana), swimming along a reef off Curaçao.

It's not everyday you spot a land iguana roaming the reef in Curaçao 🦎. 🎥 : IG user turtleandray_photography pic.twitter.com/RzoNE5fZJ3

— PADI (@PADI) 1 April 2019

The question raised, of course, is: was it was feeding on the reef. I am sure that iguanas could swim for hours in sea water without taking in any sodium chloride but eating is a different matter since, like the Galapagos Marine Iguana it is difficult to see how ingestion of sea water could be avoided.

Fundamental questions about the salt gland of a number of iguanid lizards, which appear to be able to swap between a high sodium to a high potassium secretion according to diet or salinity of water ingested, remain unanswered. Some of the questions may be basic, for example, do high sodium and high potassium secretions come from the same gland in the head, or the same cells of one gland? We need some proper physiology and cell biology to find out. And more observations from divers of the type shown in the video.

For more information on the sighting of iguanas in the sea, see Greg Mayer’s post on Jerry Coyne’s WEIT here.

How Chameleons Work: The Tongue. Alexander Sand’s Legacy

Alec Sand, under the name before he changed it by deed poll, Zoond, determined the mechanism by which the chameleon’s tongue is projected at its prey. There has been an additional and very important twist in the story that has come only recently but the essence of where the force is applied and by which muscle was all down to Sand 85 years ago.

There had been several attempts to explain the working of the tongue before the 1930s. Sand began his paper:

The projection of the chameleon's tongue constitutes a unique problem in muscular organisation. It has frequently been recorded that a chameleon can project its tongue to a distance as great, or slightly greater than the length of the animal's body from nose to anus, and that projection takes place with instantaneous rapidity. Various theories have been advanced to explain this remarkable mechanism, yet still it remains imperfectly understood. The variety of the explanations offered is clearly due to the fact that investigation of this problem has been entirely anatomical, and that no attempt has hitherto been made to apply the test of experiment to it Such an experimental investigation forms the subject of this paper. The experiments here described deal with the tongue, the hyoid apparatus and the accessory muscles, and permit of the formulation of a complete account of this highly complicated and highly perfected response. 

Sand was able to dismiss some of the earlier ideas from the 19th Century very easily, as had others before him. Inflation of the tubular tongue by air from the lungs was impossible because there was no pathway for air to reach the lumen of the tongue. The idea that a penile-like engorgement with blood was responsible, suggested in 1828, was shot down in flames by Duvernoy in 1836. Things happened too fast for it to be other than a highly specialised muscular mechanism.

In a series of experiments, Sand determined which muscle was responsible and how it worked. In essence the knob of the tongue throws itself off a long bony projection aimed at the prey. It does so by squeezing that projection (the entoglossal process). As the muscle tightens it spreads so that it starts to slip off the tapered end of the entoglossal process and, still contracting, close down the lumen of the tongue completely. That squeezing force then becomes a force acting against the end of the entoglossal process and the tongue is projected forwards, carrying the corrugated neck of the tongue behind. The sticky tongue plus prey is hauled back into the mouth by the weak muscles in the extended neck of the tongue.

There has sometimes been confusion as to how the basic mechanism works, not helped by inadequate or no explanation in textbooks. For example, Angus Bellairs in his book, Reptiles, from 1957, did not describe the mechanism in the text, despite it being characteristic of chameleons, but just showed a tiny diagram drawn from Sand’s description with a misleading arrow for the direction of force.

More recent research, as I said above, has added to that story because high-speed photography couple with physiological knowledge has demonstrated that the muscle, aptly called the accelerator muscle, just cannot do the amount of work needed in the time it takes for the tongue to reach its prey. There is an additional, elastic, process that accounts for the projection of the tongue. The accelerator muscle squeezing inwards act to compress layers of connective tissue, the intralingual sheaths, that lie between the muscle and the entoglossal process. The collagen fibres within those sheaths are arranged such that when compressed by the muscle, strain within the fibres increases. Thus the intralingual sheaths build up a store of energy that is released suddenly as the muscle and sheaths themselves slide off the end of the entoglossal process. It also seems that the thickened tip of the entoglossal process provides a passive block allowing the accelerator muscle to generate considerable pressure on the intralingual sheaths before the forward edges of the sheath and overlying muscle can move forward over the tip to initiate the sudden release of energy from the sheaths and the continued contraction of the muscle. Although other muscles help in the process, their role is minor compared to that of the accelerator muscle.

The presence of the intralingual sheaths, which are in layers and extend like a telescope, Star Wars Light Sabres or the tubes of a photographic tripod, were of course described by the 19th Century anatomists. Any function, though, except for providing lubrication or mechanical protection, was not considered.

Diagram modified from Martin & Wolfe (Chameleons, London: Blandford, 1992) to show the direction of contraction by the accelerator or ring muscle on the intralingual sheaths and the bony entoglossal process

Elasticity to generate rapid motion is known from other cases in animals. There is still though nothing quite like seeing the speed of the ballistic mechanism of the chameleon’s tongue. It is not surprising that it has excited such interest in the world of biomechanics. Experiments have been done on it, mathematical models have been built of it and manipulators designed and constructed on the principles attributed to it. But while Sand may not have foreseen the importance of the elastic component, the whole process is driven, as he said, by that ‘massive sphincter-like ring muscle’ of the tongue knob.

A great advantage to a ballistic mechanism (with the stored energy built up from muscle power) compared with one relying on muscle contraction acting in real time, is not only the velocity of the movement but its relative resistance to a lowering of temperature. Since first keeping chameleons in the early 1960s I have had an interest (plus an unsuccessful grant application in 1968 over which I sometimes occasionally rant) in what happens to reptiles as their temperature changes during the day. Yes, they can by behavioural thermoregulation achieve a preferred body temperature which is optimal for their metabolism (chameleons are often desperate to flatten their bodies in the first rays of the morning sun and absorb heat) but may be operating sub-optimally for much of the time. In that respect it is particular interest that ballistic tongue projection works at high performance over a 20°C range. By comparing, projection (elastic) with retraction (simply muscle-powered) of the tongued in the Veiled Chameleon (Chamaeleo calyptratus) at different environmental temperatures, Christopher Anderson and Stephen Deban found that peak velocity and power declined by 10-19% with a 10°C drop in temperature for projection of the tongue compared with a greater than 42% decline for retraction. That finding means that chameleons may be slower in the cold of early morning but that they can feed and take advantage of prey whose movements may be affected by the cold.

Chameleons come in different sizes, from Brookesia micra at 2.9 cm long to what is thought to be the largest, Parson’s Chameleon (Calumma parsonii)  at 68 cm. By comparing the performance of the tongue of 20 species over a five-fold difference in length Christopher Anderson found that small species ‘project their tongues proportionately further than large species, achieving projection distances of 2.5 body lengths’. The small chameleons achieved the highest accelerations and outputs of power ever recorded in any movement of a reptile, bird or mammal. The whole arrangement of the jaws and tongue is such that small chameleons can capture eat relatively large prey items compared with large chameleons.

Finally, Sand’s observations on the chameleons in his laboratory discovered the trick that has been used to get chameleons in captivity to feed when the supply of live insects has dried up.

They were kept indoors on a small privet bush planted in a tub, and most of the original stock have now survived in this situation for over three months. Every day a batch of house-flies is released in the room, and the tree is sprayed with water, of which the chameleons require a regular supply, as Gadow has pointed out It is generally stated that the chameleons will only take live food. I have found that they have extraordinarily poor discrimination. Not only will they take dead, even mouldy flies, but also such miscellaneous objects as a dried up piece of chameleon embryo, raw or cooked liver, a small piece of twig, the head of a burnt match and a piece of putty have been repeatedly taken. The three latter were never swallowed, but were always rejected soon after being drawn into the mouth. But the object, whatever it be, must be kept in motion, which can be done by presenting it on the end of a dissecting needle. These observations are of interest in that they show that neither auditory nor olfactory stimuli play any part in attracting a chameleon to its food. The stimulus is purely visual, and apparently any oscillating object of appropriate size, indefinite colour, and irregular shape will induce a chameleon to shoot out its tongue and seize it.

Middle Dwarf Chameleon (Bradypodion thamnobates)
Photographed in my office in the late 1980s when I had a breeding group.

Below I have just shown a few of the key papers. Those interested can find references to other and earlier work there.

Anderson CV. 2016 Off like a shot: scaling of ballistic tongue projection reveals extremely high performance in small chameleons. Scientific Reports Reports 6, 18625 doi:10.1038/srep18625

Anderson CV, Deban SM. 2010 Ballistic tongue projection in chameleons maintains high performance at low temperature. Proceedings of the National Academy of Science of the USA 107, 5495–5499. doi:10.1073/pnas.0910778107

Anderson CV, Deban SM. 2012. Scaling of the ballistic tongue apparatus in chameleons. Journal of Morphology 273,1214–1226 doi: 10.1002/jmor.20053 

Debray A. 2011 Manipulators inspired by the tongue of the chameleon. Bioinspiration & Biomimetics 6, 1–15. doi:10.1088/1748-3182/6/2/026002

de Groot JH, van Leeuwen JL. 2004 Evidence for an elastic projection mechanism in the chameleon tongue. Proceedings of the Royal Society B 271, 761–770. doi:10.1098/rspb.2003.2637 

Moulton DE, Lessinnes T, O’Keeffe S, Dorfmann L, Goriely A. 2016 The elastic secrets of the chameleon tongue. Proceedings of the Royal Society A 472, 20160030. http://dx.doi.org/10.1098/rspa.2016.0030 

Zoond, A. 1933. The mechanism of projection of the chameleon’s tongue. Journal of Experimental Biology 10, 174-185.

How Chameleons Work: Great discoveries in the 1930s by Alexander Zoond aka Alexander or Alec Sand (1901-1945)

SPLAT. KERPOW. The words of a schoolboy comic hardly do justice to the chameleon launching its tongue to capture its prey. That whole process, from moving the eyes into stereoscopic mode, preparing the tongue for launch, the explosion of the tongue from the mouth, to the prey being dragged into the mouth and chewed must surely rank as one of the great sights in the living world. Equally the colour-changing abilities of these lizards has to be seen in real time to be appreciated.

Sadly, and the story even ends on a very sad note, the name of the man who made the most important discoveries in how the chameleon tongue works and of how colour changes are controlled has been largely forgotten. Partly, this is because he changed his name and partly it is because he died young and in tragic circumstances.

The name of the author that appears in list of references to the work on chameleons in the early 1930s is A. Zoond. The only reason I knew that he changed his name is that ETB Francis told me so; he did that while we were talking about his work on chameleons I had supplied him with. Zoond, he said, changed his name to Sand. In fact both names appeared on the extensive reading list ETBF handed out to honours zoology students for his course on vertebrates, a list so extensive a modern student would have an apoplexy, a. from shock over the quantity, and b., from having to work out whole fields of research for themselves.

Alexander Sand
In the tropical, white, uniform of a
Lieutenant, R.N.V.R.
Photograph used in his Royal Society
Obituary Notice

Alexander Zoond or Alexander Sand and, sometimes, Alec Sand is one of my scientific heroes from the then new world of experimental biology. He is sometimes labelled as a comparative physiologists but what he did was excellent physiology; there is nothing to be gained by adding ‘comparative’ or demeaning its fundamental importance because it was not done on mammals. Only fairly recently have I got to know more about him and only last week discovered his tragic end.

Information on him comes mainly from his obituary notice for the Royal Society and a family tree (without much detail) on ancestry.com. F. S. Russell FRS (later Sir Frederick) wrote that Sand was ‘of Jewish-Russian extraction’ and was born in Warsaw on 28 December 1901. The unrest in Poland (his father was a Menshevik) saw the arrival of the family in London when Alec was six. Before Zoond the family name had been Zundelevich or, in South Africa, Zoondevitch.

His schooling in London was entirely on the arts side. After a year at Reading university college came a strange and unexplained move. Reading then was the place for degrees in agriculture but Zoond left for the University of British Columbia in Canada to study dairy bacteriology with the intention of a farming career. After graduating with an agriculture degree, he tried a job in farming in the U.S.A. for a year but found it unrewarding. In 1924 he headed back to Canada, this time to McGill University, for an MSc in bacteriology (his thesis from October 1925, The influence of green manures upon the growth and physiological efficiency of Azotobacter chroococcum, can be found in full online). Appointed to a Demonstratorship in 1926 he had a chance meeting with Lancelot Hogben FRS who was then Assistant Professor of Medical Zoology. Hogben was only at McGill for 18 months and when he was offered the chair of zoology at the University of Cape Town, he took it. With him went Zoond, having turned down a fellowship at Yale, as lecturer.

Lancelot Hogben
Photograph used in his Royal Society
Biographical Memoir

Zoond, in other words, changed field entirely, an attitude entirely in keeping with Hogben’s iconoclasm and loathing of classical zoology and classical zoologists. Hogben wrote:

…I must here explain that the sacred cow of first-year biology in those days was a whimsy of Thomas Henry Huxley known to posterity as the “type system”. This involved the detailed anatomical study of about a dozen species chosen from different groups of the Animal Kingdom, less because they were typical of their phyla or classes than because it was at one time relatively easy to place orders for them with pet shop dealers in London. When I entered my department on the day after arrival, I found the shelves lined with dozens of bottles of Huxleyan types imported in formalin from the seat of the empire. On enquiry, I learned that no living creatures had ever desecrated these hallowed precincts. The same day, I therefore instructed my laboratory steward to empty all the contents of the pickle jars in the yard behind the laboratory ready for the incinerator when the sun had dried them. Henceforth, the motto was to be only the living pass these portals…

Zoond clearly thrived in Cape Town. He published with Hogben on the pH of surface water, with David Slome¹ on the effect of electrolytes on cardiac rhythm and with Enid Charles, (the first wife of Hogben) on respiration in crabs.

In his memoir Hogben did not mention Zoond by name. However, it is clear who is being referred to in the following passage:

When I was not dealing face to face with the student body, Boer-backlashed mentality could make matters disagreeable. The lecturer whom I imported from McGill to replace the relict of my predecessor formed an attachment to one of the only two coloured medical graduates then practising in the Dominion, the other one being her brother-in-law, a leading light in the Moslem Community. Both held the Glasgow degree. When our young colleague confided in us that he was bringing his lady-friend to the University annual dance, both Enid and I scented embarrassment for both. Accordingly, we asked the pair to dine with us so that the lady could come as our own guest. My Jewish students clustered round and saw to it that she was never without a partner. Next day, the more rabid nationalist students held an open air campus meeting of protest.

It was after Hogben, with apartheid in the offing, had left Cape Town in 1930 that Zoond, who had stayed on, completed his work on chameleons, that on the tongue with Joyce Eyre², that on pigmentation with Eyre and with Naomi Adeline Helen Bokenham (later Millard) (1914-1997). The papers were published between 1933 and 1935, as he was leaving or after he had left South Africa.

Cape Dwarf Chameleon (Bradypodion pumilum, in the 1930s known as
Lophosaura pumila and then Microsaura pumila).
All of Zoond's (Sand's) research was on this species.
Photo: JonRicfield from here.

Sand (still Zoond then) left Cape Town in 1933; before doing so he took a Ph.D. from that university. He then went to Cambridge to work with James Gray FRS (later Sir James) then Reader in Experimental Zoology who was just beginning his famous work on animal locomotion. It was their joint work on the locomotory rhythm of the dogfish published in 1936 that was on ETBF’s reading list.

Alexander Zoond changed his name by deed poll on 9 March 1935 to Alexander Sand. The notice in the London Gazette gave his address as 9 Storey’s Way, Cambridge, his occupation as ‘biologist’ and stated that he was a naturalised British subject. I have not been able to work out in what capacity Sand was working with Gray. His second, this time Cambridge, Ph.D. was awarded in 1938. He must have been in statu pupillari to have been awarded a Ph.D. since those paid by the university were (and I think still are) disbarred. Did he have some form of external fellowship or did he fund himself?

In early 1935 he joined the staff of the Marine Biological Association’s famous laboratory at Plymouth as Physiologist. There he worked on the sensory physiology of fish including the lateral line, stretch receptors in the muscles and the ampullae of Lorenzini. The research on the auditory labyrinth he did with Otto Löwenstein (1906-1999, FRS 1955) was—and still is—of relevance in all vertebrates and clinical conditions. What happens in the semicircular canals during movement of the head was sorted out for the first time in experiments that would have been impossible in mammals.

Although I am not going to describe here the work on sense organs in fish, I thought it worth quoting the following paragraph that indicates the sensitivity of the lateral line and why Sand worked in the cellar of the Plymouth laboratory ‘which is hewn out of solid Devonian limestone’:

To demonstrate the delicacy of the response he formed a light hammer of a rubber stopper which was allowed to fall through a distance of 8 cm. on to a rubber sponge lying on the edge of the table about 3 feet away from the prepara­tion, the shocks imparted to the table being quite imperceptible to the human touch. 

There are a few traces in the newspapers of Sand’s time at Plymouth. The Western Morning News of 28 December 1936 reported that a fire had been discovered in Fardel Manor on the Cornwood-Ivybridge road which was occupied by Sand at the time. Large beams underneath a hearth in one of the upstairs rooms were smouldering. One of Sand’s children was due to sleep in the room. The fire brigade had to cut away the floorboards before extinguishing the fire ‘with the aid of patent extinguishers’. The medieval house, now Grade I listed and about 9 miles from the MBA laboratory, was once the seat of the Raleigh family.

By the time of the special census in 1939, the Sand family had moved a couple of miles to Gerston, Harford, now part of Ivybridge. Sand is shown there (as a research biologist) with his wife, Edith G Sand (born 19 April 1909), two children (records still closed) and Elizabeth B Reidy, born 1914, described as ‘paid companion help’. Russell wrote that he had two sons.

Motorists in Britain lived for decades in fear and dread of leaving their car without lights at night. Bored policemen pounding their beats were always on the lookout. Sand was fined ten shillings (50p) at Totnes Petty Sessions for ‘leaving a motor car without lights’ at Redworth Terrace, Totnes on 7 January 1940. How being fined for leaving a car without lights tied in with the wartime blackout regulations which demanded no lights, I do not know. Normal for Devon, maybe.

Russell then notes that Sand did all he could to get into the armed forces as quickly as possible after the outbreak of war.  He was commissioned on 1 May 1940 in Special Branch of the Royal Navy Volunteer Reserve. Promotion to Lieutenant followed in October 1940. He, like many biologists in the war, was involved in radar. Russell wrote:

After a time on the mainland in the north of Scotland he was posted as Officer in Charge of the station on the Shetlands. In 1941 Sand came south to lecture on radar for some months before being posted to a cruiser in which he was in the Indian Ocean. He contracted malaria in the Persian Gulf and when recovered went on leave to South Africa where his wife and two sons were living for the war years. He was then posted to a monitor and was present at the Sicilian and Salerno landings, his ship being hit. After a period of unemployment he was seconded for research at the Naval Department in the Medical Research Council’s Laboratory at Hampstead.

Russell went on to say that the research at Hampstead was on the physiology of diving and that in the course of his duties he received training in diving himself. It was, said, Russell, likely to remain confidential. G. L. Brown (FRS 1946, knighted 1957) was in charge of the research. The Physiological Society always maintained the tradition of using just initials and surnames in its internal communications, never honours or titles, so Sir George Lindor Brown FRS was always G.L. Brown. His biographical memoir contains more details of the research and includes Sand, along with a while string of well-known physiologists and naval officers, as members of the laboratory:

Brown discovered at the Lister Institute, and commandeered, the long-disused chamber that J. S. Haldane, the father of J. B. S., had used a generation earlier in the research that gave divers their first reliable decom­pression tables. The chamber was transported to the garden of the Institute at Hampstead, a hut was built around it, a new compressor was installed, and Brown and his associates began to acquire the somewhat esoteric expertise needed for doing human experiments under pressure. It was Brown’s first opportunity to recruit a research team of his own. H. P. Marks, Macintosh and Collison of the Hampstead staff were already available; they were joined by F. Dickens, C. B. B. Downman and 20-year-old H. B. Barlow; A. Sand and W. D. M. Paton arrived a little later. S. Gay French, H. M. Balfour and other naval medical officers, along with Ellis, were occasional members of the group; others who sometimes took part included Haldane, D. Williams and R. G. Bickford.

Sand was elected to the Royal Society on 16 March 1944.

But then came tragedy. on 11 July 1945 at 58 West End Lane, Hampstead, Alexander Sand shot himself in the heart with a pistol. The famous coroner, Sir Bentley Purchase recorded the verdict of the inquest on the death certificate, ‘dec[ease]’d did kill himself not being of sound mind’. I have been unable to find any press reports that might shed more light on the events leading to his death. There is no mention on the death certificate of service in the navy (occupation shown as .F.R.S. and Doctor of Science’). However he is listed as having died while still serving in the RNVR. His nominal ‘ship’ (all sailors are attached to a ship, even if that ship is a shore establishment) was H.M.S. Victory, the shore establishment at Portsmouth.

Russell concluded his obituary notice:

The whole Plymouth staff were anxiously looking forward to the day when they would once more be reunited at the laboratory and the blow was bitter when it was known that Alec Sand would not be numbered among those returning. Having reached the height of his research powers it was anticipated that he would open up new avenues in the little explored field of the sensory environment of marine animals. All alike felt keenly the loss of a friendly and inspiring colleague, and a gap has been created in the ranks of physiologists that it will be hard to fill. 

Thomas Alan Stephenson (1898-1961, FRS 1951), Lancelot Hogben’s successor in Cape Town until returning to Britain in 1940 as professor of zoology at Aberyswyth, introduced his obituary of Sand in Nature as follows:

In the completely unforeseen death of Alexander Sand at the age of forty-three, comparative physiology suffered a very severe loss. Not only was his later work distinguished in a remarkable degree, but so much more of the same calibre might have been expected from him in the next twenty years. 

UPDATED 11 October 2020

*Hogben claimed he had forgotten the name of his predecessor, ‘a dedicated necrophilist’. It was John Dow Fisher Gilchrist (1866-1926), a marine biologist born in Scotland, who, in his time, seems to have been far more modern in research, but not in teaching perhaps, than Hogben indicates.

1. David Slome (1906-1995). Details here.

2. I can find no further information on Joyce Eyre.

Alexander RMcN. 2001. Otto Egon Lowenstein. 24 October 1906 - 31 January 1999: Elected F.R.S. 1955. Biographical Memoirs of Fellows of the Royal Society 47, 357-368.

Hogben, LT. 1998. Lancelot Hogben. Scientific Humanist. Woodbridge, Suffolk: Merlin Press.

MacIntosh FC, Paton WDM. 1974. George Lindor Brown 1903-1971. Biographical Memoirs of Fellows of the Royal Society 20, 41-73.

Stephenson TA. 1945. Dr Alexander Sand. Nature 156, 383-384.

Russell FS. 1948. Alexander Sand 1901-1945. Obituary Notices of Fellows of the Royal Society 5, 803-815.

Wells GP. 1978. Lancelot Thomas Hogben 9 December 1895 - 22 August 1975. Elected F.R.S. 1936. Biographical Memoirs of Fellows of the Royal Society 24, 183-221.

Zoond, A. 1933. The mechanism of projection of the chameleon’s tongue. Journal of Experimental Biology 10, 174-185. 

Zoond A, Bokenham NAH. 1935. Studies in reptilian colour response. II. The role of retinal and dermal photoreceptors in the pigmentary activity of the chameleon. Journal of Experimental Biology 12, 39-43.

Zoond A, Eyre J. 1934. Studies in reptilian colour response. I. The bionomics and physiology of the pigmentary activity of the chameleon. Philosophical Transactions of the Royal Society B. 223, 27-55.

Giant Salamanders; Would You Believe They Common Imports into Britain before World War II?

Giant Salamanders have been in the news in the past week. Chinese Giant Salamanders were confiscated by customs and sent to London Zoo. I have not been able to find out whether the salamanders were brought into the country destined for the restaurant trade or to be kept in a public or private animal collection.

I have written about these salamanders before and their being so commonly available that in Hong Kong in the 1960s that they were used for class dissection. All the ones that arrived were about 35 cm long, like the one at London Zoo, and only a sixth of their maximum length. That seems to be about the size favoured by restaurants in China.

Chinese Giant Salamander. Hong Kong 1967
On the roof of the Northcote Science Building

Giant Salamanders are on the menu in China as ‘the fish that walks’. They are bred on a commercial scale but there seems no doubt that wild-caught, i.e. poached, individuals make up a large proportion of those sold. Numbers have fallen massively in the past decades—80% since the 1950s is the current estimate and the species is now classed as Critically Endangered because not only is it a food item worth real money at the luxury end of the market but habitats have been and are being wiped out or ravaged as human habitation expands.

What is not generally known is that giant salamanders were imported into Britain for the amateur herpetologist trade in the 1930s. I found this article in Water Life magazine of 27 September 1938:

September is a good time to review the happenings of the closing season, which has been interesting for several reasons, and to make one’s plans for the winter. First as regards supply. A wealth of species has been imported, among them…Megalobatrachus [the generic name for giant salamanders used for decades before it was decided Andrias had priority]… 

The demand for the Gigantic Salamander must surely now be filled. No less than three of our dealers have had these animals during the last year or two, and a fourth told me recently he is hoping to arrange a consignment at any time now. Considering how commonly these animals are met with in Japan and parts of China, their price remains disappointingly high over here. Since most of the continental zoos and aquaria number their specimens by the half-dozen or more, it is remarkable that the breeding of the species at Amsterdam seventy-odd years ago has never been emulated.

The author did not distinguish between Chinese (Andrias davidianus) and Japanese Giant Salamanders (A. japonicus). Given their once common occurrence in the live-food trade in China, including Hong Kong, my guess is that those imported in the 1930s were from China.

It is also known that giant salamanders (often called Gigantic Salamanders, as above) were imported by European dealers for zoos and for amateur herpetologists in the later decades of the 19th Century. Thus the Reverend Gregory Bateman in his famous book, The Vivarium, published 1897, not only wrote what was then known of these animals but this:

These huge Salamanders may, from time to time, be bought of the larger dealers in wild animals, at prices, according to size, which range from £10 downwards. 

£10 then would be £1000 now (calculated on increase in retail prices). There were some very wealthy amateur herpetologists around in the years before the First World War.

The giant salamanders imported into Europe must have been pretty tough, having survived a long sea voyage through the tropics. Indeed, what struck me about the ones we handled in Hong Kong was how unfazed they were by high temperatures; not at all what might be expected from inhabitants of rocky streams flowing from mountains. Equally, my wife, who did the wrangling (well, somebody had to carry the camera) was surprised by how willing they were to snap at a passing finger or hand.

Finally, I do not know the identity of the author—a prolific and knowledgeable writer on reptiles and amphibians in the 1930s—who wrote under the non-de-plume ‘Amphibius’. It is known that he died between 1939 and about 1950. I have written about him and my search here. I would be most grateful to hear from anybody with more information.

The Outdoor Reptiliary in Britain. 1. London and Other Zoos

This article (part one of a two-part series) is on my other site, Reptiles, Amphibians and Birds: A Historical Perspective of their Care in Captivity. It can be found HERE.

This is an illustration, of the then new reptiliary at London Zoo from The Illustrated London News of 15 September 1928: