The Duke of Bedford’s Zoological Exploration of Eastern Asia in the 1900s—2. Anderson, Sowerby and Ward

We know the following were participants in the 1904-11 Exploration. The leader was Malcolm Playfair Anderson and the following were involved for longer or shorter periods of time: Arthur de Carle Sowerby in 1908 and then Frank Kingdon Ward and Dr J.A.C. Smith after Anderson's return to China in 1909. A great deal is known about and has been published about Sowerby and Kingdon Ward but virtually nothing of Smith, although I now have more information on him than was available previously; he will be the subject of a ffurther article in this series.

Malcolm Playfair Anderson

Malcolm Playfair Anderson
1879-1919

Anderson’s father, Melville, Professor of English Literature at Stanford from 1891 to 1910, had the sad job of writing his son’s obituary for Condor, the ornithological journal. Malcolm Anderson was born in Indiana in 1879. For two years in his early teens he was in Germany with his mother and siblings. He returned to the U.S.A. highly unimpressed with both the educational system and militarist schoolboys amongst whom survival depended on ‘force of arms and fists’.

He graduated from Stanford in 1904 with a degree in zoology. However, from the age of 14 he had been on several collecting expeditions in the U.S.A. Somehow he must have come to the attention of Oldfield Thomas at the Natural History Museum in London for he was immediately employed by the Zoological Society of London to conduct the exploration being paid for by the Duke of Bedford, so his father wrote. However, I am by no means convinced the Zoological Society was involved, with both Anderson and his father seeming to conflate the two organisations. The confusion may have been because the Duke was President of the Society at the time.

He collected, he wrote and he photographed through Japan, Korea and China. This is his description of collecting a wildcat on the island of Tsushima, midway between Japan and Korea:

From our hilltop we wandered on down a trail singing and whistling, till we met a man on horseback and another on foot. The horseman dismounted and bowed, then passed on. The man on foot paused and addressed us. He was a funny old fellow with a bald head. Orii answered, told him our business, and asked him about animals. His answers were not very clear, as he had had too much sake. However we gathered that one of his neighbors had recently killed a wildcat, and the old man thought possibly it was still unskinned. WC asked directions and were told to go to the next village and ask for his house (his name, he said, was ‘Man of the Shining Head’ and he was distinguished as the best drinker in his village). We found the owner of the cat (which was skinned). I bought the skin for two yen, and asked about the body. It had been given away. We went to the neighbor who had received it and found that he had eaten part of the cat and buried the head and vertebrae. To dig the thing up was the work of a moment, and we were in possession of scientific evidence of the presence of the wildcat on Tsu-shima.

The cat is regarded as a subspecies of the Leopard Cat, Prionailurus bengalensis, locally critically endangered and the subject of active conservation measures.

In 1908, he travelled to Europe and worked on his collections but returned to China in 1909 to continue the exploration, to ‘the desert of the north beyond the Great Wall, and in the mountains on the border of Thibet’.

I do not know how the finances of the Exploration worked but Anderson is said to have sold the skin of the last confirmed wolf seen on Honshu to the Natural History Museum in London for 8.50 yen. However, is it not possible that having paid two hunters for the carcass, he was simply reclaiming expenses?

After his work on the exploration Anderson made two collecting trips to South America, the second with his wife whom he had married in 1913. He was ill after the second expedition but managed to write some articles on his travels. In 1918 an appeal was launched to encourage men in the U.S.A. to work in the shipyards. Anderson volunteered but he was killed by a fall from scaffolding from a shipyard in Oakland on 21 February 1919. He was 39. His only child, a son, died in infancy.

Oldfield Thomas named three species after Malcolm Anderson: a vole, Myodes andersoni; a shrew-mole Uropsilus andersoni and a niviventer, Niviventer andersoni.

I have found three articles by Anderson, published posthumously in Natural History in 1920, on experiences in China. They are listed below.

Arthur de Carle Sowerby

In his account of his work on the Clark Expedition, Sowerby explained where he went on the Duke of Bedford’s Exploration:

Before going into details of the present work, it might be well to say something about that already done in the same districts. In connection with the Duke of Bedford’s Exploration of Eastern Asia in the early part of 1908, Mr. Malcolm P. Anderson and the writer had already made collections of mammals in Shensi [Shaanxi Province], at Yen-an Fu and Yü-lin Fu. We also collected in the mountains of Shansi (Shanxi Province], north-west of T’ai-yüan Fu.

As well as a biography published in 1956, Sowerby was the subject of a major article in the Journal of the Hong Kong Branch of the Royal Asiatic Society for 1998-99. Keith Stevens introduced it as follows:

Although the lives of many Western expatriates who lived in China and experienced the excitements and horrors of travel and the exoticism of the old civilisation cry out to be recorded, most expatriates lived mundane, cliché-ridden existences, apart that is from the occasional excitement caused by the troubles and emergencies of the times, brigandage, rioting and war. They never or only very rarely ventured far from their Treaty Fort and certainly not into the dark hinterland of China. Should they have ventured anywhere at all it would have been to hunt or shoot in the immediate area of the Port or go to a nearby beach or classical tourist site, such as Nanking or Soochou. And of all only a mere handful of those who did venture far afield have left sufficient records to enable a portrait of their life to be disentangled and recorded. Arthur de C. Sowerby was one such venturer.

Arthur Sowerby was born in China to missionary parents in 1885 and lived in interesting times: through the fall of the Manchu Dynasty; the era of the War Lords; Nationalists versus Communists; invasion, occupation and internment by the Japanese. He retired first to U.K. and then to the U.S.A. where he died in 1954. Towards the end of the First World War he served in the Army where, in view of his complete fluency in Chinese dialects, he served as an officer in the Chinese Labour Corps. By this time he was suffering from arthritis which plagued him for the rest of his life.

Stevens describes Sowerby as an explorer and author but as a man who needed to earn his living, at least, perhaps, until he married his second, American, wife in 1922. The dire state of his finances was noted in his correspondence with Clark. There are gaps in the record in which it is not known what he was about. Educated in China, Bath and the University of Bristol, he did not complete his degree course but ran away to sea after a failed romance. He worked his passage to Canada, stayed there for a while before returning to China and his parents in 1905.  He taught at the Anglo-Chinese College in Tientsin but it is thought he only stayed there for the year before he joined the Duke of Bedford’s exploration. He was paid by Robert Sterling Clark for the duration of that expedition and for his collecting expeditions over the next 20 years, and may well have continued teaching in Tientsin.

In late 1911 he took part in the Shensi Relief Expedition to rescue and lead to safety foreign missionaries caught up in the aftermath of the revolution. An account of this successful mission through country where banditti were rampaging was written by Sowerby for an appendix to a book on the fall of the Manchu Dynasty.

Manchuria and parts of Mongolia were the subject of four expeditions. After his demobilisation in 1919 he stayed in Edinburgh for a year to complete his five-volume work, The Naturalist in Manchuria. By 1921 he was off back to China via the U.S.A. where further expeditions followed, again through bandit-ridden country.

For around 25 years he lived in Shanghai. He produced and published a monthly periodical, the China Journal. He served as honorary director of the Shanghai Museum, one of the activities of the Royal Asiatic Society. He was president of the China Society of Science of Art which was incorporated into the Royal Asiatic Society. He later served as president of the latter. He had his publishing business and was a director of other companies. He took an active part in the Shanghai International Settlement, the residents’ associations, for example. 

A jerboa was named after Sowerby by Thomas as Dipus sowerbyi. It is now recognised as a synonym of Dipus sagitta, the Northern Three-toed Jerboa.

Frank Kingdon Ward (publishing as Kingdon-Ward)

Frank Kingdon Ward
1885-1958

I will not dwell on the explorer, plant collector and author, Francis Kingdon Ward (1885-1958) simply because there is a whole website devoted to his life and further information elsewhere. However, he took part in one expedition of the exploration because of his extraordinary family connexions. Ward’s father was Professor of Botany at Cambridge. The early death of his father left Frank needing to find paid work after two years at Cambridge rather than the three needed for a degree. A family friend, none other than one of the authors of the Wade-Giles system of transliteration of the Chinese language into roman characters—and Professor of Chinese at Cambridge—Herbert Allen Giles, pulled strings in Shanghai and Frank Ward soon found himself teaching at a school in that city.

During the long school holidays he visited Java and Borneo but after two years, another school friend—this time Oldfield Thomas—saw him recruited for the Duke of Bedford’s exploration for another collecting expedition. He wrote that trip up, the approximate route of which is shown on the website, in his On the Road to Tibet. He was accompanied by Dr J.A.C. Smith.

Thomas named a vole, Eothenomys wardi and a shrew, Blarinella wardi for Ward.

Ward went on to many further and alarming travels and much plant-hunting in Assam, Burma, China and Tibet. He served in the Indian Army in the First World War, as a spy in Tibet, and taught survival techniques in the Second World War.

Anderson MB. 1919. Malcolm Playfair Anderson. Condor 21, 115-119

Clark RS, Sowerby A deC. 1912. Through Shên-kan. London: T Fisher Unwin

Sowerby RR. 1956. Sowerby of China : Arthur de Carle Sowerby F.R.G.S., F.Z.S. Kendal: Titus Wilson.
The author was Richard Raine Sowerby (1887-1968) a solicitor, farmer and local historian. He described himself as a distant kinsman of Arthur Sowerby.

Stevens K. 1998-9. Naturalist, author, artist, explorer and editor and an almost forgotten president: Arthur de Carle Sowerby 1885-1954: President of the North China Branch of the Royal Asiatic Society 1935-1940. Journal of the Hong Kong Branch of the Royal Asiatic Society 38, 121-136

Ward FK. 1910. On the Road to Tibet. Shanghai: Shanghai Mercury

Articles by Malcolm Playfair Anderson in Natural History Volume 20:
For the sake of his ancestors, pp 92-99;
The discovery of the Chinese Takin. pp 428-433;
A winter journey in northern China. 517-534.

UPDATED 5 September 2019

The Duke of Bedford’s Zoological Exploration of Eastern Asia in the 1900s—But some sources get the wrong duke

When I was looking up stuff on the rodents of China before and after our trip to Sichuan in November 2017, I came across references to Oldfield Thomas (of his eponymous pika) and to the Zoological Society of London’s exploration of eastern Asia paid for by the Duke of Bedford. Reading further I realised there were errors—amplified of course by repetition on the internet—in some of the secondary accounts.

11th Duke of Bedford

The Duke of Bedford from 1893 until his son inherited the title on his death in 1940 was the 11th Duke, Herbrand Arthur Russell (1858-1940). He was President of the Zoological Society of London from 1899 until 1935 and is well known for gathering the last live specimens of Père David’s Deer after the Boxer Rebellion and building up a herd at Woburn, his country seat in Bedfordshire.  Less welcome was his rôle in introducing the Grey Squirrel from North America.

Unfortunately, some accounts by authors, who should have known better, of the exploration paid for by the 11th Duke, attribute the naming of the exploration to his son (whom he loathed and was loathed equally in return), the 12th Duke, Hastings William Sackville Russell (1888-1953). Hasting Russell who inherited the title in 1940 (before then he used the junior title of Marquess of Tavistock) was a cranky pacifist, fascist and famous aviculturist.

The personalities and lifestyles of these two Dukes of Bedford is described by the 13th Duke, John Ian Robert Russell (1917-2002) in his book, A Silver-plated Spoon. He had a difficult, to put it mildly, relationship with his father and grandfather. Of his grandfather he wrote:

In the midst of all this magnificence my grandfather lived a completely lonely and austere life. He had no contemporaries or friends of his own and most of the people who came to stay were cousins or other relatives. He had very little time for human beings and rarely spoke.

A somewhat different picture emerges from his Royal Society biographical memoir written by Peter Chalmers Mitchell. However, this one does come with a caveat. The 11th Duke, as President, and Mitchell, as Secretary, were in cahoots for over 30 years in running the Zoological Society and Mitchell was a brilliant publicist.

By disposition the Duke was a rather old-fashioned country gentleman, not fond of company, but taking his pleasure in fishing, shooting, natural history and estate management. Neither hunting nor racing had any attraction for him… In politics the Duke, originally a Liberal, passed over to the Conservative side through Liberal-Unionism, but maintained the traditions of his family by allowing the fullest scope of opinion to others. It was the good fortune of the present writer to be closely associated with him for nearly forty years, directly in connexion with the Zoological Society and zoological affairs, but incidentally in many other respects. He has never known any one more modest about his wide attainments and unusual ability, with a greater sense of duty to his country and to his King, or more conscientiously resolved to accept the responsi­bilities of his wealth and position. He was courteous, almost deferential to every one, and the kindest of friends. 

How the exploration of eastern Asia paid for by the Duke came about I do not know but it would appear that Oldfield Thomas was a driving force (he served on the Council of the Zoological Society). It was clearly not simply a single expedition since various people took part over different periods. I have also not found out why or how the leader and various members were chosen. It ran from 1904 until 1911 and concentrated on mammals. A further complication is that this enterprise overlapped with the Clark expedition of 1908-1909 in northern China. That was led and funded by Robert Sterling Clark, with specimens going to the U.S.A. The complication is that Arthur de Carle Sowerby (1885-1954) worked on both.

Oldfield Thomas

The leader of the Bedford exploration was the American, Malcolm Playfair Anderson (1879-1919). The intention seems to have been collecting for Oldfield Thomas at the Natural History Museum in London, then known, of course, by that awful title, British Museum (Natural History). Sowerby was involved in 1908. Francis (Frank) Kingdon-Ward (1885-1958), a botanical collector described as one of Anderson’s assistants, was involved in a trip through western China in 1909-1910 with Dr J.A.C. Smith, also described by Thomas as Anderson’s assistant.

Thomas described the mammalian finds and new species in a number of papers. In a paper to the Zoological Society in 1911 he explained some of the history of the whole affair:

After the departure of his assistants, Dr. J.A.C. Smith and Mr. Kingdon Ward, down the Yang-tze, with the remarkable collection of mammals which formed the subject of paper No. XIII of the present series, Mr. Malcolm Anderson worked on by himself from September 1910 to the end of March 1911, when he brought back to Shanghai the series on which the present paper is based. 

During this period he explored a number of localities in Western Sze-chuan and N. Yunnan, and has been rewarded by finding a considerable and quite unexpected number of species additional to those previously sent…  

To our great loss and regret Mr. Anderson now proposes to give up the arduous life of the field collector, and this is therefore the last paper to be written on his collections for the Duke of Bedford’s Exploration of Eastern Asia. On this account it is suitable to five a few words of appreciation for the manner in which he has risen to the opportunity afforded by our President’s [i.e. the 11th Duke of Bedford] continued generosity. 

From the commencement of the work in 1904 to the sending home of this last valuable collection in 1911 Mr. Anderson has worked for the Exploration with superb enthusiasm and success, and in the extent to which his collections have revolutionized our knowledge of an extended part of the earth’s surface he has made a record which, so far as I know, has never been equalled… 

In turn Mr. Anderson has visited and collected in Japan and its islands, both the small southern ones and the large northern Hokkaido and Saghalien, Korea, N. China and Mongolia, Shen-si, Shan-si, Kansu, Sze-chwan and N. Yunnan, his collecting-localities being dotted through all the previously most unexplored parts of this little-known area. 

Over 2700 mammals have been collected, besides many birds and, as is natural, a large number of novelties have been found among them.

I will write about what I have found on Anderson, Smith, Kingson-Ward and Sowerby in my next post.

I am, as I shall suggest in the other articles in this series, doubtful of any involvement of the Zoological Society of London. I suspect the American, Anderson, conflated the Natural History Museum in London with the Zoological Society of London since the Duke of Bedford who paid for the whole thing was President of the latter at the time. Anderson may have seen the Duke at the Society's offices in connexion with the eponymous Exploration. It is clear that Anderson was working de facto for Oldfield Thomas at the Museum.

Chalmers Mitchell P. 1941. Herbrand Arthur Russell. Duke of Bedford. Biographical Memoirs of Fellows of the Royal Society 3, 499-502

John, Duke of Bedford. 1959. A Silver-plated Spoon. London: Cassell (1960, The Reprint Society)

Thomas O. 1912. The Duke of Bedford’s Zoological Exploration of Eastern Asia.—XV. On mammals from the provinces of Sze-chwan and Yunnan, Western China. Proceedings of the Zoological Society of London (1912), 127-141

UPDATED 30 July 2019

Where the Lion Trod. C.H. Keeling. Zoo history - a new re-publication of the 1984 book, edited by John Adams for the Bartlett Society

My old friend, Clinton Keeling (1932-2007), after a series of misfortunes, some of which it has to be said, were of his own making, reinvented himself as a historian of British zoos*. Beginning in 1984 he produced a series of booklets over the next 19 years with particular attention to a number to zoos that had long been closed. There were two problems why these booklets remain relatively unknown. The first was that they were self-published as typewritten cyclostyled sheets and so never reached conventional publishing and retailing channels. The second was that their content was not obvious from the title (e.g. Where the Leopard Lazed, Where the Macaw Preened, Where the Penguin Plunged). The booklets were known to very few people and the numbers sold must have been very small. Those surviving in circulation must also have dwindled. I suspect the few that I had in the 1980s got bundled with similarly-bound reports and ended in that open-ended receptacle in my office that was sooner or (rarely) later the final resting place of A4-sized documents received from government departments.

The Bartlett Society, which Clin founded in 1984 for zoo history enthusiasts, has made the brave decision to republish the first of Clin’s books, Where the Lion Trod, but this time as hardback. Brave because the information it contains is dated, not surprisingly since sources of information were far harder to access in the 1980s, and brave because it is difficult to predict whether such a venture would be other than a financial drain.

Zoo histories come as two sorts. The first sort which look outwards are produced by social historians and tell you all about the milieu in which the zoo operated, its supposed social significance and why everything is all down to colonialism. The second tell you a lot about the animals kept, the people who set them up and something about the layout and architecture. Clin’s books fall firmly into the latter category; the first sort would have had him foaming at the mouth. His prejudices, both about how animals were kept and should be kept, and about the world in general, shine through so that those who never spent time in his company will get some idea of his diatribes against those who ran zoos in Britain, his loathing of popular culture and his hatred of ‘The Great British Public”.

Clin’s research and musings covered twenty extinct collections/zoos dating from the 13th century (the menagerie in the Tower of London) until 1959 (the date of closure of Maidstone Zoo). Various chapters consider, for example, the long-gone zoos in Liverpool, Manchester before Belle Vue, Edinburgh before the present Edinburgh Zoo, Hull (where a pub still recalls the zoo that closed in 1862), Southport, Cardiff, Halifax, Oxford, Sheringham and Southend.

The original format has been followed so that this hardback reprint bears a striking resemblance to the original typed version. I feel that was an error since I had the feeling of reading a dissertation for a degree rather than a published book. The editor has also added a few notes of explanation or correction as well as a list of known species kept at each establishment; the latter I finds adds little.

I must end with this paragraph on Maidstone Zoo (1934-1959) and its owner, the brewer Sir Garrard Tyrwhitt-Drake (1881-1964):

Sir Garrard was nothing if not a character. When I met him I almost did a double-take in amazement, so astonishing was his likeness to Charlie Chaplin. He invited me into the manor, gave me a glass of beer and after talking to me for a time said ”Hmmm, you know a bit about animals”, which I took to be praised indeed. Then suddenly he said “Well I must ask you to go now Mr Keeling—I want my lunch!” And that was that.

You can buy the book from the Bartlett Society here.

*Those who knew Clin Keeling will realise I risk his posthumous wrath. He never called a zoo a zoo, always a ‘zoological gardens’. I also visit the bot in Hong Kong and also, on occasion, the bot in Edinburgh and the bot at Kew.

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.