Sunday, 16 July 2017

Maxwell Knight, MI5’s Greatest Spymaster. A second biography of the MI5 officer, amateur naturalist and broadcaster

Like London buses, you wait for ages and then two come along. Well, not quite because there is a gap of 33 years between these two but the subject and the subject matter are identical: Maxwell Knight.

The publication in 1984 of the book by Anthony Masters (1940-2003) The Man Who Was M came as a shock to those of use of a certain age who knew Maxwell Knight as a populist amateur naturalist and animal keeper who was a regular on BBC radio and, sometimes, television and who instilled an interest in young listeners and readers that helped start them in careers as professional biological scientists. The shock came from knowing what Knight did earlier in life because he had been a highly successful agent runner for the government security service MI5 and had achieved notable successes in infiltrating himself and, later, his agents into fascist organisations with links to nazi Germany in the 1930s and 40s and into communist organisations with links to and control from the Soviet Union.

Since 1984, additional information has emerged, several books have been published and a number of websites contain information. One book had material from it available to Masters. That one was a sensationalist and, it has been claimed, a rather fanciful account by Joan Miller, an associate of Knight in MI5. Another deals with a particular case, The Kent-Wolkoff Affair. The new material, some from recently declassified MI5 files, has been incorporated in the new book, Maxwell Knight, MI5’s Greatest Spymaster by Henry Hemming.

In many respects, therefore, the new book is like a second edition of the old. It kills a few canards and presents a more balanced view than was possible with material available to Masters in the early 1980s.

Maxwell Knight though remains an enigma: a chameleon by trade, from adopting the persona of a fascist activist in the 1930s to the avucular broadcaster in the 1960s; an inhabitant of a secret world who courted publicity (a castaway on Desert Island Discs for example); an impotent thrice-married womaniser; a gentleman burglar; a naturalist with an amateur interest in the occult.

Maxwell Knight retired early—at the age of 55—from MI5 on the grounds of ill health in 1961. By then though he was already well known by BBC audiences. Hemming writes:

Max—he was no longer ‘M’—was by 1961 one of the BBC’s most prolific broadcasters. In the 1950s alone, he featured in at least 306 original radio broadcasts, he had no fewer than 20 books published in this one decade, he appeared on television more than 40 times, excluding repeats, he gave lectures throughout the country and he wrote numerous magazine articles, all on the subject of natural history. His rich, reassuring voice was synonymous by 1960 with radio programmes such as The Naturalist, Country Questions, Nature Parliament and Naturalists’ Notebook. Max also popped up on Woman’s Hour, did schools programming and featured on television programmes such as Look and the panel show Animal, Vegetable, Mineral. When children became junior members of the London Zoo in 1965, one of the advertised benefits was the chance to attend “film shows and lectures given during the school holidays, when you can meet famous animal experts such as David Attenborough, Maxwell Knight, and Peter Scott”.

His secret and public lives overlapped.

Hemming does make the connexion between Knight’s two lines of work. John Le Carré (who worked under Knight in MI5) illustrated two of his books. But it is links with the Zoological Society of London that are intriguing. Hemming notes that Knight was a member of council at the same time as Ivor Montagu (1904-1984; zoologist, film-maker and table tennis enthusiast) ‘a Soviet agent who had been followed around London thirty-none years earlier by Eric Roberts, acting on Max’s instructions’. There were other former security and secret service officers around the Zoo in the later decades of the last century. One was Gwynne Vevers† but there were others. However, the mix is even more interesting. Gwynne’s father, Geoffrey Marr Vevers (1890-1970), Superintendent of the Zoo until 1948 was a strong supporter of the Soviet Union and its political system, before and during the Second World War (he edited Anglo Soviet Journal) as was his boss, Sir Peter Chalmers Mitchell (1864-1945), Secretary of the Society from 1903 until 1935. There were also links to Ivor Montagu through the distribution of Soviet propaganda films. Many of the individuals involved must have been of interest to MI5, however innocent or humanitarian the motives of those involved since a number of the network of organisations were covertly controlled either directly or indirectly from Moscow, and as we now know some individuals were active Soviet agents. The biographies of Knight stress how he argued strongly within MI5 that the threat of Soviet espionage within the British government was real. He expressed his views in an internal memorandum, ‘The Comintern Is Not Dead’ and history showed with Burgess, Maclean, Philby, Blunt et et al. that he was right. I cannot help but think that at the Zoo, Maxwell Knight was  initially mixing business with pleasure.

Closer to home on the other side of the fence was the sister of the photographer who illustrated another of his books. The great photographer and cinematographer, Wolfgang Suschitzky, (1912-2016) who had set out in life determined to be a zoologist, worked a great deal at London and Whipsnade zoos, his photographs there illustrating many articles and books. His sister, Edith Tudor-Hart (1908-1973), was most certainly of interest to MI5 since she was a major talent spotter and recruiter of the Cambridge spy ring as well as acting as a courier and supplier of photographic equipment to Soviet spies. She also worked as a photographer and one of her photographs, of children exploring a rock pool, appeared in the August 1939 issue of Animal and Zoo Magazine.

Both Masters and Hemming posed the question of what drove Maxwell Knight to pursue (some said in a pushing manner) his second career as a broadcaster and writer. Money seemed to be the answer. He retired early from MI5 on medical grounds. I do not know what the pension arrangements were then but my estimate is that with his thirty years service he would have been on something like three-quarters of half his final salary. Given that he was not in the top echelons of MI5 and had an expensive lifestyle in rented property it is easy to see why a second stream of income was needed.

Although this is not a book review there are a few statements of a zoological nature in Hemming that range from dubious to an outright howler. In drawing a parallel with his handling of animals and his brilliant handling of his agents, his ability to get a wild-caught toad to feed from the hand is mentioned: ‘Few wild-born toads will feed from a human hand. Fewer still are happy to do this after so little time in captivity’. In my experience this is not true. A hungry toad fresh from the wild will readily accept an earthworm dangled in front of its nose.

Then there is a photograph showing ‘Knight with his favourite pet. Goo the cuckoo’ with quite clearly a young Greater-spotted Woodpecker on his shoulder. The correct photograph for that caption is plate 27 of Masters. Finally, surely every schoolboy knows the plural of mongoose is not ‘mongeese’.

Now we have had two biographies of Maxwell Knight (the latter produced with little reference to the former) how long until we get a third? Will we eventually know what service he performed for the King which led to his receiving in 1931 a gold cigarette case inscribed with the royal cipher and his name?

*Charles Henry Maxwell Knight, 1900-1968. See also the website M: Maxwell Knight and the Frightened Face of Nature

†A BBC producer, Winwood Reade, responsible for many of Maxwell Knight’s broadcasts was the third wife (out of four) of Gwynne Vevers; she was interviewed by Masters,

Hemming H. 2017. Maxwell Knight, MI5’s Greatest Spymaster. London: Preface

Masters A. 1984. The Man Who Was M. The Life of Maxwell Knight. Oxford: Blackwell

Clough B. 2005. State Secrets. The Kent-Wolkoff Affair. Hove: Hideaway

Miller J. 1986. One Girl’s War. Dingle: Brandon

Thursday, 6 July 2017

Komodo Dragons in the 1930s: a zoo quest before ‘Zoo Quest’ with links to Adolf Hitler, nazi spy scares, the FBI, a cuckolded husband and John F Kennedy

I was mystified when I came across an article in the June 1939 edition of Animal and Zoo Magazine. I have read a fair bit both scientific and popular on the natural history and history of the Komodo Dragon. Some of the popular books and articles I found to contain misprints, incorrect references and misinterpretations and none that I had found covered the story in that article from 1939.

Reading the article, entitled ‘Island of Dragons’ was the start of a search for more information but one which seemed at first attempt to lead nowhere. The article was written by Dr Paul Fejas (note the spelling ‘Fejas') and describes his attempts to capture Komodo Dragons and, under licence from the Dutch colonial government, to ship one to Stockholm Zoo and another to Copenhagen. He caught nineteen, chose two of moderate size that he thought would withstand the long journey, and released the rest. He noted that the two reached their destinations.

But who was the writer? Google searches revealed nothing but came back suggesting I meant ‘Fejos’ not ‘Fejas’. Eventually I realised that the magazine article was by Dr Paul Fejos (1897-1963). A typographical error had led me on a wild goose chase. I should not have been surprised because in the same article ‘Paranus’ instead of ‘Varanus’ is used a couple of times.

from Dodds (1973)

It was Dr Paul Fejos who collected the dragons while on Komodo to film them, thereby predating the BBC’s Zoo Quest and its same objectives by nearly twenty years. His own remarkable history is exceeded by that of Inga Arvad (1913-1973) his then wife—the subject of a recent book—a Scandinavian beauty queen, journalist and actress who interviewed Adolf Hitler and other leading nazis, who was suspected of being a German spy by the paranoid but thorough F.B.I., who became, while married to Fejos and being bugged by the F.B.I., the yet-to-be President John F. Kennedy’s lover and who, later, became a leading British politician’s short-term fiancée. 

Paul Fejos rewrote his own history so there are a number of alt-facts, i.e. lies, myths and legends, out there. His version appears in a biography that verges on a hagiography published shortly after his death; even the author of that—a friend of Fejos—did not not know what was truth and what was self-made myth. The truth seems more prosaic but nonetheless remarkable. 

Paul Fejos was born in Hungary in 1897. According to a Wikipedia biography which seems to sort the myths from reality in his early life, he served as a medical orderly in the Austrian Army on the Italian Front in the First Word War while a medical student. He graduated from the Royal Hungarian Medical University in Budapest (now Semmelweis University) in 1921 but developed a fascination for cinema and theatre, directing films, plays and operas. He never practised medicine. He left Hungary in 1923 and reached the U.S.A. via Vienna, Paris and Berlin. After manual work in a piano factory he found a job as technician at the Rockefeller Institute for Medical Research in New York working for Jacques Jacob Bronfenbrenner (1883-1953) on bacteriophages. He left there for California and the hope of getting to Hollywood. The story of how he got started as a director there involves being picked up while hitchhiking by a rich man who wanted to be a film producer.

The cinema buffs’ websites describe Fejos’s successful career as a Hollywood director. But he tired of the place and returned to Europe to make early sound films. By 1934 he was in Denmark working for Nordisk (now making some of the Scandi series shown on BBC4 on Saturday nights).. It was during this time that he married, as his third wife, Inga Arvad, whom he recruited to star in one of his films. But he was tiring of fiction and while trying to get out of his contract with Nordisk persuaded them to send him to Madagascar with a cameraman but without Inga to make a documentary. Although unsuitable for a full-length feature film, the footage was used to make a series of documentaries which, by being factual rather than staged stories, earned the respect of anthropologists.

Fejos was hooked by his new interest in anthropology. He was commissioned by the Swedish Svensk Filmindistri to make a whole series of ethnographic films in Asia. In 1937 and 1938 he and Inga travelled in much of the Far East and made films in Indonesia, the Philippines, New Guinea, Ceylon and Thailand during 1937 and 1938, including the 13-minute The Komodo Dragon (Draken på Komodo). Although Inga was supposed to be radio operator and ‘script girl’ she was left at the Helena May Institute in Hong Kong when the film crew set off for Komodo. But the first landing on Komodo was a disaster—allegedly. Fejos’s motor boat lowered from a freighter into a strong current hit a reef and split in two. The three men (Fejos, camera and radio operators) swam ashore and managed only to retrieve rope and torches. They could find no water and felt doomed. But the ship that that dropped them appeared on the horizon during the night. It had gone round the island because of the state of the tide rather than sail away through the strait between Komodo and Sumbawa. Using a torch from the top of the tree they managed to signal SOS and the ship’s boat picked them up the next day. Another—this time successful—landing was made a few weeks later, with Inga being left on the neighbouring island of Sumbawa. I will return to the filming and capture of the dragons later.

Whilst in Singapore, the film party was invited on board the yacht Southern Cross by her owners, Axel and Marguerite Wenner-Gren. According to Inga’s biography, the meeting was engineered by Fejos’s sound recordist who contacted the Wenner-Grens to say that fellow Scandinavians were in Singapore making films about the East. That meeting set Fejos off in a completely new direction.

Axel Wenner-Gren was said to be one of the wealthiest men in the world. He owned Electrolux, early manufacturers of washing machines and refrigerators, and was one of the founders of Saab. Fejos and Wenner-Gren became great friends and the latter decided to finance a filming expedition to Peru in 1939. It started in Maldonado (yup, been there, tick) in the Peruvian amazon and went badly. Peruvian soldiers accompanying part of the large expedition were inveigled into helping one tribe involved in a tribal war. At least one death resulted. But while in Cusco Fejos heard reports of buried cities and obtained further funding from Wenner-Gren to explore the area along the trail to Machu Picchua and beyond. Here the expedition uncovered, mapped and photographed large and small Incan cities and roads. He also continued, between phases of the main expedition, his ethnographic filming.

While all this was going on Inga had been left in the U.S.A. Her links as a budding journalist with Hitler and other leading nazis and Wenner-Gren’s suspected support for nazi Germany (including using his vast South American holdings to advance German infiltration in South America) created the perfect storm. The F.B.I. kept a close and paranoid eye on both of them as well as on Fejos’s expedition.

It was while Fejos was in South America that Inga began her affair with John F. Kennedy. He was serving in the Office of Naval Intelligence. The F.B.I. bugged there every action and it is thought the head of the F.B.I., J. Edgar Hoover, used the material to remind the then President Kennedy that he had it and that he wished to remain in post; he did.

Fejos and Inga Arvad were divorced in 1942. In the meantime, Wenner-Gren in dispute with the U.S. Internal Revenue Service, set up the Viking Fund in New York to support anthropology. He endowed the fund with $2.5 million worth of shares in Electrolux and Servel (fridges and air conditioners) and appointed Paul Fejos to run it.

Eventually it was realised that neither Inga Arvad and Wenner-Gren (who had his assets frozen by the U.S. and British governments) were nazi sympathisers. The Viking Fund was re-named the Wenner-Gren Foundation for Anthropological Research in 1951, Fejos remained as its head until his death in 1963. The Foundation continues to support research in anthropology.

I do not know how history has treated Paul Fejos’s reputation in his final career in ethnology and anthropology. The fact that he supported the Jesuit Teilhard de Chardin (1881-1955) and his crazy notions on human evolution I could not fail to hold against him.

But back to Komodo. The article in Animal and Zoo Magazine describes how he caught the dragons and in notes published in his biography, Fejos wrote of his second landing:

     A little later I went back to the blasted island, because I decided that I wouldn't allow it to lick me. But I went back with two native sailboats and many fresh coconuts. We drank coconut water during the whole time and were able to work. Also, I brought some extra coolies with me, and we trapped the dragons in traps which we constructed on the island. They were box, gravity-fall traps; a skeleton of a box was made with wood, and then chicken wire nailed on it all around. We put a dead goat inside, and about three days later when the goat smelled to high heaven, then the dragons came down, one after the other.
     The Komodo dragon was even then a protected animal; nobody was allowed to catch or shoot one. But when I told the Java officials what we had seen, they asked me please to try to catch one for the zoo in Java, if I went back. They gave me permission to capture or kill two animals for myself and to capture one for them, which I did. When I came back, they asked me how many were on the island, and I said, “I haven't the slightest idea, but from what I saw and the frequency of the encounter, maybe three or four hundred, maybe more.

'Having slipped a noose around your dragon's neck your difficulties
are only just starting, as the picture below shows'
Animal and Zoo Magazine
'Though equipped with formidable teeth the dragon's most powerful weapon
is his scaly tail'
'The wire trap is set with its inviting bait of goat or deer, and when caught,
the dragon still needs careful watching, for it is very cunning.

     Of the two we captured and kept, one went to the Zoological Gardens in Stockholm, and one went to the Zoological Gardens in Copenhagen, where in due course they died, not from illness or climate, but from visitors. Some stupid visitor threw beer caps into the place and they ate them. One of them had a perforated intestine; we performed an autopsy on him later. The one which went to Stockholm was 14½ feet long, the Copenhagen one 13 feet.

There is a photograph of the one sent to Copenhagen but by this time stuffed and on display in the museum. 

from Dodds (1973)

Fejos’s collecting activities are not mentioned in The Living Dragon by Dick and Marie Lutz. The authors suggest that the Stockholm and Copenhagen specimens were obtained by de Jong in 1937 who collected on Flores and possibly Komodo and who supplied a number of zoos with specimens. Clearly we now know the ones in Stockholm and Copenhagen came from Fejos.

Readers in Britain of a certain age will be puzzled to learn which well-known politician was Inga briefly affianced. It was Robert Boothby, later Lord Boothby, the ambisexual ‘bounder but not a cad’ who was the long-term lover of the Prime Minister’s wife, Dorothy Macmillan, and acquaintance—at least—of the notorious Kray twins.

So having had a celebrity magazine tour of the 1930s and 40s, I found that having searched fruitlessly for the misprinted name above the main article, Animal and Zoo Magazine had go it right, as Fejos, in the list of contents on the first page!

Paul Fejos in 1962
by Robert Fuchs (in oils)
(reproduced in Dodds, 1963)

Dodds JW. 1973. The Several Lives of Paul Fejos. Wnnner-Gren Foundation. (John Wendell Dodds 1902-1989, Stanford University)*

Dodds JW. 1963. Eulogy for Paul Fejos. Current Anthropology 4, 405-406*

Farris, S. Inga. Kennedy’s Great Love, Hitler’s Perfect Beauty, and J. Edgar Hoover’s Prime Suspect. Guilford, Connecticut: LP

Fejas [sic] P. 1939. Island of Dragons. Animal and Zoo Magazine 4 (1, June 1939) 4-6

Lutz D, Lutz JM. 1997. The Living Dragon. 2nd edition. Salem, Oregon: DIMI Press

*When I bought the biography for a song from a bookseller, the eulogy was found folded inside together with a letter which began:

The Wenner-Gren Foundation is pleased to send you with its compliments this copy of The Several Lives of Paul Fejos. This limited edition is being distributed to many of the Foundation’s past and current grantees, to those who have helped the Foundation over the years in the design and execution of its programs, and to others who will want to gain insight into the traditions of the Foundation’s philosophy, style and approach.
Unfortunately, the book bears no signature or plate so I do not know who the recipient was.

Tuesday, 6 June 2017

How did he come to write that?

My eye, in speed reading a review of a biography of Sir Hans Sloane in The Times (Saturday 27 May), came to a sudden halt:

Sloane was no Darwinist; the flora and fauna, animals and artefacts he was to acquire articulated a vision of providential order.

Not only was there an obvious tautology describing the collection but Sloane could hardly have been a Darwinist. Sloane, whose collection formed the basis of the British Museum was born in 1660 and died in 1753. The Origin of Species was published in 1859.

I do hope the Victoria and Albert Museum has a copy of Ernst Mayr’s The Growth of Biological Thought for the author of the review is its new director, the former Labour MP, Tristram Hunt.

Komodo Dragon: extending the saliva story

The previous post generated a number of emails on the use of biologically-active substances in saliva by the Komodo Dragon to help kill its prey, ranging from ‘unnecessary and highly speculative’ to ‘generally supportive but needs more work’. I was also told that the post seemed somewhat harsh. It was not meant that way but I think my style in writing it reflects the sort of debate that has arisen and the tone adopted by some authors and commentators. The comments/replies on a paper* are as robust as any I have seen in print for a long time.

Much of the controversy centres on use of the terms ‘venom’, ‘venomous’ and ‘venom glands’. I will not go into the arguments further here but there is a long-standing problem in naming structures or molecules for presumed or first-discovered function, potential function or partial function. The names of hormones and growth factors are well-known nightmares in this regard. There are actually two hypotheses, one an evolutionary question on the stage at which reptiles acquired the capacity to produce venom—the Toxicoferon Hypothesis—and the related but distinct question of how the Komodo Dragon (and possibly other extant and extinct monitor lizards) kills its prey. This latter question is susceptible to morphological, chemical, physiological and pharmacological observation and experiment as demonstrated by Bryan Fry and his international team in their original paper. I will restrict myself to expanding the dragon story.

Typical Komodo Dragon habitat on the island of Rinca, part of the Komodo
National Park, with gullies and open areas of grassland

My own view is that there is a chain of events by which biologically active substances in saliva could be used to decrease the time between initial attack and death of the large mammal prey. At present, though, I see a gap in knowledge between a scenario which describes what could happen and one that describes what does happen. I also see it as a gap that could be filled and which, if filled, would subdue further objections and a great deal of speculation on alternative roles for biologically active substances in the Dragon’s saliva.

There is strong evidence in favour of saliva doing something—and something deleterious to the life of the prey. The presence of a relatively large lumen in the mandibular gland where secretion can be stored is highly suggestive of the use of a ready store of saliva when a dragon attacks prey, and a store that can be delivered to the base of the teeth. The authors of the original papers also showed by direct biological assays that dragon saliva has components that act to cause vasodilatation and decrease blood pressure and to contain substances that are potent anticoagulants.

Therefore, a good case has been made for local effects of biologically-active substances on the wound inflicted by the dragon—parallel deep slashes inflicted by pulling after the initial bite. Diffusion into the tissues surrounding the wound would increase the rate of blood loss and keep the severed ends of large and small arteries open and the blood flowing from all severed vessels. That’s where the relatively low bite strength of the Dragon which was demonstrated by Fry and his collaborators would be of selective advantage. If the Dragon bites, pulls what have been described as its steak-knife teeth across blood vessels (‘grip-and-rip’), and holds that grip, the last thing it needs is a strong bite. Compression by a strong bite would compress the wound and slow the rate of blood loss. Moreover, it would decrease the rate of movement of substances from the saliva away from the wound into surrounding tissues and into the circulation through venous or lymphatic drainage.

So, as far as local actions of saliva are concerned, they must be present and I can think of experiments that would test how far diffusion would take them into surrounding tissues. 

Central effects, for example, in decreasing blood pressure, would be harder but not impossible to demonstrate, as would establishing the concentrations of substances from saliva in the peripheral circulation of the prey in order to determine whether they reach the levels required to exert the effect that can be demonstrated with particular concentrations in biological assay systems. The latter is not a trivial question with substances that would be appearing in the circulation relatively slowly from a wound and which may be relatively quickly broken down into inactive compounds.

The presence of several compounds known to be part of the arsenal of toxins deployed by classic venomous reptiles adds weight to the suggestion of some systemic role for Dragon saliva in the bitten animal. Others have suggested that may may fulfil other roles in the mouth or in digestion of the prey before ingestion and the action of stomach enzymes; I do not find the latter argument compelling although it could be tested experimentally.

So, while demonstration of central toxic events has not yet been attempted, there must be a local effect of saliva, at least, on the gripped-and-ripped prey. The question that remains on this local action is whether the effect is biologically significant in shortening the life or ability to resist of the animal that has been ambushed, or is so in a sufficiently large number of attacks as to confer a selective advantage.

Both local and systemic actions rely on getting saliva into the wound. Different durations of bites could have different effects. The mandibular gland implicated in storage seems to work in a similar way to that of back-fanged snakes in that the pressure exerted by chewing forces the secretion out. If that is so then a ‘grip-and-rip’ feeding attack could have an entirely different outcome from a quick offensive or defensive nip.

Some commentators have suggested testing the components of Dragon saliva on their natural prey. However, what is their natural prey? Water Buffalo were introduced relatively recently, Timor Deer in antiquity while domestic goats are, well, domestic. The Komodo Dragon and its extinct relatives appear to have been around the Indonesian islands, where they spread from Australia, even before the now extinct dwarf elephant appeared in their habitat so that fundamental mismatch between the bulk and apparent adaptations for killing large animals of Komodo Dragons and the size of potential prey species seems to be unexplained.

Other questions arise like are there differences between the sexes in the size of prey and feeding strategies, adult females being lighter than males? Do juvenile Dragons attack prey that is larger than that attempted by, say, Water Monitors (Varanus salvator) of similar size and, if so, would that make the use of saliva as a chemical weapon to help in subduing prey more likely? In other words, the advantage may come more to juveniles tackling large prey than to adults. Indeed, if other species such as the Lace Monitor or Goanna (Varanus varius) from Australia have a similar array of potential biological agents, which seems to be the case, and in quantitatively equivalent amounts, might the whole arrangement be an adaptation to extend in size the range of prey that can be killed and eaten by some or all monitor lizards?

So, after arguing with myself several times and learning lots about venomous and non-venomous reptiles, that’s may take on the Komodo Dragon’s feeding method in the light of present evidence. In essence, I argue that the available evidence points to a local chemical role for saliva in helping to subdue/kill large prey (while recognising that sheer physical force may often or sometimes be sufficient). As for toxins in saliva having a systemic effect on the prey, I argue that the gap between the prima facie potential effect and actual effect has not yet been filled by experimental data. And as to whether we should call Dragons ‘venomous’ or the gland the ‘venom’ gland…

...I will leave to others.

Whatever the final outcome on these fascinating and enigmatic lizards, the authors of the 2009 paper have done a signal service in examining how it and other extinct and extant monitor lizards kill their prey and have not shied from erecting hypotheses on possible mechanisms and their wider importance in the evolution of reptiles. 

*Fry BG, Casewell NR, Wüster W, Vidal N, Young B, Jackson TNW. 2012. The structural and functional diversification of the Toxicofera reptile venom system. Toxicon 60, 434-448. Weinstein SA, Keyler DE, White J. 2012. Replies to Fry et al. (Toxicon 2012, 60/4 434-448). Part A. Analyses of squamate reptile oral glands and their products: A call for caution in formal assignment of terminology designating biological function, Toxicon 60, 954-963. Kardong KV. 2013. Replies to Fry et al. (Toxicon 2012 60/4 434-448). Part B. Properties and biological roles of squamate oral products: The “venomous lifestyle” and preadaptation, Toxicon 63, 113-115. Jackson TNW, Casewell NR, Fry BG. 2013. Response to “Replies to Fry et al. (Toxicon 2012, 60/4, 434–448). Part A. Analyses of squamate reptile oral glands and their products: A call for caution in formal assignment of terminology designating biological function”, Toxicon 64, 106-112. Weinstein SA, White J, Keyler DE, Kardong KV. 2013. Response to Jackson et al. (2012), Toxicon 64, 116-127.

Saturday, 20 May 2017

Komodo Dragons. A peaceful morning in the Komodo National Park and an acrimonious debate on reptilian venoms

Last September’s Expedition Cruise from Darwin through the Lesser Sunda islands of East Timor and Indonesia included stops at Rinca and Komodo. Ever since my fourth cousin once removed visited Komodo in 1956 I have wanted to see Komodo Dragons alive and in their natural habitat. They did not disappoint. We saw very large males (hanging out around the kitchen of the ranger station on Rinca hoping for a hand out), females and juveniles (but not the hatchlings which apparently take to the trees to avoid their predatory parents). One female was digging out the nest mound of a megapode, the Orange-footed Scrubfowl (Megapodius reinwardt) in which to lay her own eggs, September being the egg-laying time in their breeding cycle.

When we got back I had a chance to look at and think about the various hypotheses that have been advanced as to how Komodo Dragons—and, possibly, some other monitor lizards—kill their prey. The highly publicised but never properly tested proposal that Komodo Dragons are venomous in the sense that venomous snakes are venomous, i.e. toxins delivered by injection having a very or fairly rapid systemic effect on the prey, seems to be in the process of being discarded in the course of some pretty acrimonious arguments, along with the associated hypothesis of a single, early origin of venom in reptilian evolution—the Toxicofera hypothesis. The other idea, that pathogenic bacteria harboured in the mouth of dragons causes sepsis in prey animals that are bitten but escape the initial attack, doesn’t seem that convincing or special given the likelihood of infection from a bite from any animal, as any postman will testify, especially if that postman were to immerse his bitten ankle in fetid water, as non-native water buffalo do when bitten by a Dragon. If I were to bite a Timor Deer on the leg, there is every likelihood an infected wound that could impede mobility would ensue. 

Given the opposition to the Toxicofera hypothesis and the contentious nature of the evidence for Komodo Dragons being venomous, it seems a pity that the BBC repeated last November the screening of an episode from its 2011-12 season of the Natural World Series called Komodo—Secrets of the Dragon that was devoted virtually entirely to that proposition and its main protagonist. Had this been a programme about a topic other than science-led natural history the BBC would have been falling over itself to show someone with the opposite view, however perverse. As it is, the viewing public in Britain has been left with the impression that the venomous nature of the Dragon is accepted. At the very least the BBC should have been aware of what was going on (by reading Wikipedia for example) and before rescreening it should have added an annex to the original programme. That annex could have  explained the opposing views and evidence—some of which made news media reports*—that have been accumulating since 2009. Adam Hargreaves (Oxford), Abigail Tucker (King’s, London) and John Mulley (Bangor), who produced a devastating critique** of the toxicoferan hypothesis in 2015, should have been consulted and involved. But let’s leave the BBC and its usually excellent (but sometimes spectacularly poor) natural history programmes and return to the mouth of the dragon.

None of the criticisms of the venom hypothesis imply that the composition and quantity of saliva are not of selective advantage to the despatch of prey, subsequent swallowing and digestion or protection of the oral cavity against infection. Nor, indeed, do they refute the idea or evidence that saliva may have local beneficial—to the Dragon—effects on inflicted wounds, like the application of a secreted anticoagulant, for example.

The drooling mouths of some of the male Dragons we saw were impressive. The only comparable example I could think of was my late mother-in-law’s Boxer dogs given the slightest hint of finding something edible. With no obvious sign of food or feeding activity, is Dragon saliva being used for some other purpose like scent marking?

The accumulated data that I have found suggests to me that Komodo Dragons, as originally thought, kill their prey by sheer brute force from wounds inflicted by a very large mouth with very big teeth. In one study†, 17 attacks on large prey were observed, 12 were fatal. Of the 5 that escaped with injuries to their limbs and rump, 1 was quickly attacked and killed by a second Dragon, 2 died within hours, one fled being pursued by other Dragons and one limped away without being pursued. The eventual fate of the two that possibly survived is not known.

One may though ask how, if the Komodo Dragon originally preyed upon the now extinct dwarf elephant, Stegodon, of Flores, as suggested in 1987 by Jared Diamond, those beasts (smaller than a domestic water buffalo) were killed? Would biologically-active substances in saliva have been of selective advantage?

Nowhere have I found (but I am not have looked in the right place) any discussion of the selective benefits of a slow death of the prey to the predator. A snake with a fast-acting neurotoxin can quickly track and swallow its prey without that investment in metabolically expensive venom being lost to another snake or any old predator happening to find the corpse. But let’s say a Dragon does inject toxins with a small bite and the animal runs away to die. There is no guarantee that the investment in venom would pay off. With lots of other Dragons around, the prey could be lost entirely or the original killer would get only a small share in a communal but competitive feeding session. Surely, if monitor lizards are venomous at all then why have they not evolved a more effective venom to ensure a quick kill?

The venomous dragon and other monitor lizards hypothesis has been around now for ten years. I find it sad that so little has been done at the whole animal and tissue levels to test it. I found some of the original observations and experiments‡ unconvincing. But given the captive populations of Dragons and a local abundance on Komodo and Rinca, a rigorous examination of the whole question cannot be beyond the bounds of practical realisation or funding.

*e.g. Zimmer C. 2009. Chemicals in Dragon’s Glands Stir Venom Debate. New York Times, 18 May 2009. Yong E. 2015. A Venomous Fight Among Reptile Scientists. The Atlantic, 2 November 2015

*Hargreaves AD, Tucker, AS, Mulley JF. 2015. A critique of the toxicoferan hypothesis. In, Gopalakrishnakone P (ed), Malhotra M (ed). A critique of the toxicoferan hypothesis. In Evolution of Venomous Animals and Their Toxins: Toxicology. Springer Netherlands, p 1-15. DOI: 10.1007/978-94-007-6727-0_4-1

†Bull JJ, Jessop TS, Whiteley M. 2010. Deathly drool: evolutionary and ecological basis of septic bacteria in Komodo Dragon mouths. PLoS ONE 5(6): e11097. doi:10.1371/journal.pone.0011097

‡Fry BG Wroec S, Teeuwisse W, van Osch MJP, Moreno K, Ingle J, McHenry C, Ferrara T, Clausen P, Scheib H, Winter KL, Greisman L., Roelants K, van der Weer L, Clemente CJ, Giannakis E, Hodgson WC, Luz S, Martelli P, Krishnasamy K, Kochva E, Kwok H, Scanlon D, Karas J, Citron DM, Goldstein EJC, Mcnaughtan JE, Norman JA. 2009. A central role for venom in predation by Varanus komodoensis (Komodo Dragon) and the extinct giant Varanus (Megalania) priscus. Proceedings of the National Academy of Sciences of the USA 106, 8969-8974 doi 10.1073 pnas.0810883106

Monday, 8 May 2017

Burkhardt and Vevers: Family Links in the Early 20th Century

Late last year James Ritchie contacted me after reading my articles on Colonel Valentine Burkhardt (12 April 2016) and Dr Gwynne Vevers (4 October 2016) in order to point out that members of the two families were close friends in the years before the First World War.

I was fascinated to read your article on Val Burkhardt—he was my grandmother's cousin. My grandmother's mother, Marie-Beatrice Siordet (nee Caldwell) was the younger sister of Annie Claudia, Val Burkhardt’s mother. Thank you for your research which has greatly expanded my knowledge of this interesting man. I also note that you have written about Gwynne Vevers and I wondered if you knew of the link between the Vevers family and the Burkhardts?  It is through my great uncle Gerald Caldwell Siordet. Briefly, Siordet was at Clifton College with his cousin Val and then went up to Balliol where he met and became great friends with the Herefordshire artist Brian Hatton. After Oxford, Hatton and Siordet shared a studio in South Kensington. Gwynne's father Geoffrey Vevers [1890-1970] was a medical student and a cousin of Hatton (their mothers were sisters) and the three of them socialised in London just prior to the first war. Hatton and Siordet were subsequently killed in the war. Val B visited the site of Hatton’s death in Egypt and wrote to Siordet about it.

More details can be found in the book by the late Celia Davies, Brian Hatton. A Biography of the Artist (1887-1916) published by Terence Dalton in 1978. However, the book is worth reading for more than those details. It is a fascinating account of Hatton’s early years and his development as an artist. It is as enjoyable and informative of life in Britain the thirty years up to the First World War as Gwen Raverat’s Period Piece.

Brian Hatton was killed on 23 April 1916 in the Battle of Katia, 25 miles east of the Suez Canal. Fifty Royal Engineers, plus a detachment of Worcestershire Yeomanry 180 strong sent to guard them, were sinking a well when they were attacked by 2-3000 Turkish infantry who came across them. He left a wife and baby daughter.

Gerald Siordet was in France when he heard the news. He then wrote to his cousin Val to ask for any more information since Burkhardt was then serving in Egypt. The then Captain Burkhardt replied on 27 September 1916. In the reply (the full text is in the book) Burkhardt stated that he was having a better memorial than the few sticks  and the bottom of a biscuit tin bearing an illegible inscription he found including a separate one for ‘2 Lieut Brian Hatton Worcester Yeo, A fine artist and a gallant soldier’. In a footnote to the letter, Celia Davies noted that after the war the Yeomen were reburied at Kantara on the Suez Canal.

Gerald Caldwell Siordet, artist, poet, critic and one-time tutor to Aldous Huxley was awarded the Military Cross for conspicuous gallantry on the Somme in 1916. He was killed on 9 February 1917 leading an attack on a Turkish position in Mesopotamia (now Iraq); his body has never been found. Val Burkhardt’s letter to him had ended: ‘I hope you are not for Mes[o]pot[amia]’.

Brian Hatton in his London studio (from here)
Gerald 'Jack' Caldwell Sioret (from here)
Geoffrey Marr Vevers (1890-1970) when Superintendent,
London Zoo
(Animal & Zoo Magazine December 1938)

Friday, 31 March 2017

The butterfly’s tongue and the steel rule

‘Why is that there?’, said my four-year old grandson in an accusatory tone when he spotted a Stanley coiled flexible steel rule on my desk. ‘Ah, I need to take a photograph of it to show how a butterfly’s tongue works’, I replied. A look of utter puzzlement and then of disdain on the boy’s face demonstrated his certain knowledge of having a silly grandfather.

L.E.S. Eastham
My wish to illustrate this post with a steel rule was occasioned by a jolt of memory when I was writing a previous one on BEPS—The Invertebrata by Borradaile, Eastham, Potts and Saunders. I had a vivid recollection of the words being said, but not of the face behind the words, during a lecture at Sheffield on insects that in the old days (i.e. before 1958 when Eastham retired as Professor of Zoology) we would have had to have known how the butterfly proboscis works since he, Eastham, it was who worked it out. However, the lecturer added that the only thing about the mechanism that one had to remember was that it works like a steel rule. The more I thought about who the lecturer was, I have the vague notion that it was the Sheffield born-and-bred—and educated—Fred Segrove (1911-2003).

Having remembered nothing else about the butterfly proboscis, I recently found the original paper by Eastham and Eassa, published in 1955 when Eastham was 62. It is a highly impressive and long paper and involved making serial sections and some physiological studies. It demolished previous theories on how the proboscis worked. Eastham himself provided a short explanation in the later editions of BEPS:

The adults live on the nectar of flowers, and to absorb this a highly specialized proboscis has been formed from the greatly elongated galeae of the maxillae, each being grooved along its inner face and locked to its neighbour…Each half of the proboscis is a tube in itself into which passes blood from the head, and also a trachea and a nerve. Across the cavity of this tube there pass a number of oblique muscles. At rest the proboscis is tightly coiled like a clock spring under the head. When feeding the proboscis is extended and its tip placed in the food source. It is now recognized that the elastic properties of the cuticular wall of the proboscis account for the coiled condition when resting. Extension of the proboscis is brought about by the internal oblique muscle of each galea. These, working in conjunction with a stipital valve controlling the closure of the passage between cephalic and galea haemocoeles, cause the proboscis to develop a dorsal keel along its whole length. The attainment and retention of this new shape depends on the turgidity of the galea tube and the elasticity and flexibility of parts of the cuticular wall. For mechanical reasons it cannot in the keeled position be retained in the coiled state and extension of the proboscis results.In feeding, a complex pharyngeal muscular apparatus causes the fluid food to be sucked into the mouth. The length of the proboscis in many cases corresponds to the depth of the corolla of the flower which the species frequents, and in the Sphingidae (hawk moths) may be greater than that of the body…

In Eastham’s chapter in BEPS there is though no mention of the analogy of a retractable, flexible steel rule, which was used in the original paper and in which a Mr F.W. Adams was thanked for thinking of it:

…a coiled steel rule, when coiled in its case, is flat in section, but when it is drawn out it assumes a curved transverse section, convex on one side and concave on the other, and in this state attains a condition of extension. In this case the rule has to be forced into its case, and bending can only occur when its transverse curvature is flattened out. The condition of rest is one of extension and force has to be applied to coil it up, the rule having been manufactured with these properties.

The difference between a steel rule and a butterfly’s proboscis is, of course, that it is muscular action which produces the curved transverse section for extension and elasticity which returns it to a flat cross section after the muscles relax and thence its recoil.

Small Tortoiseshell with proboscis coiled

Comma feeding with proboscis extended. Eastham & Eassa also explained
how the 'knee-bend' in the proboscis was formed

There is an statement in the paper which sixty-odd years after it was written I find odd:

…As a result of complete analysis of these structures, of observations on the animal during feeding—of numerous operations involving nerve sections and perforations of the haemocoele—a new theory is offered on the proboscis mechanism for which the senior author (L.E.S.E[astham]) alone is responsible.

I can think of several explanations for this statement. Did Eassa disagree is one. Did Eastham think that if the new theory were to be shot down in flames, he was making sure that Eassa could not be blamed? But then the paper was sent to the Royal Society for publication by V.B. Wigglesworth, the name in insect physiology in the 20th Century; he must have been sufficiently impressed. There remains the possibly unworthy thought that Eastham was making sure he got the credit but that goes against Eastham’s reputation in Sheffield amongst those who had worked in his department like E.T.B. Francis, F. Segrove and J.D. Jones as a gentlemanly, paternal figure. And what had Eastham to gain at the age of 62 and three years from retirement? The only possibility is that he was, and perhaps for the last possible time, up for election to the Royal Society. He, like all the other authors of BEPS, was not elected but I do not know—and cannot know until the archives for that period are opened—if he was ever proposed.

But what of Eassa? Who was he, what was he doing in Sheffield and what happened to him?

Youseff Ezeddin Eassa (1914-99) was a famous Egyptian author as well as being Professor of Zoology at the University of Alexandria. There is a website devoted to him and his work. He was a graduate of the University of Cairo and is shown as arriving in Sheffield in 1948 as a Ph.D. student but also as an established playwright and author. However, his Ph.D. thesis (A contribution to the postembryonic development of the head of Pieris brassicae (Linn)) is dated 1949 so he may have travelled to Sheffield earlier than 1948. He must have stayed in Sheffield for some time after 1949 because the Eastham & Eassa paper was sent to the Royal Society in April 1954 and no new address for Eassa is shown.

While in Sheffield he wrote stories and plays that were broadcast on BBC radio. He recalled that his favourite companion at coffee was Hans Krebs, later Sir Hans and then Professor Biochemistry at Sheffield. Eassa was a Fulbright Fellow in Berkeley and Illinois in 1960-61.

Youseff Ezeddin Eassa (from here)

I hope I get the opportunity of explaining to my grandchildren not only how the butterfly feeds, using, of course, the steel rule as an aid, but also of making sure they know that international scientific collaboration was not invented by the EU (as some young academics in Britain clearly believe) and that academics could meet and exchange ideas in university staff (faculty for readers more familiar with the U.S. university system) clubs rather than remain ensiled, as appears to be the case at present, in departmental comfort zones.

Oh, and here is the rule:

Eastham LES, Eassa YEE. 1955. The feeding mechanism of the butterfly Pieris brassicae L. Philosophical Transactions of the Royal Society B 239, 1-43.