Aviculture, Art and a Ghost Pig—An Entertaining Diversion

In my last post I listed a number of prominent British bird keepers of the early decades of the 20th Century. I was trying to work out where Falcon Close, the house of Captain Reginald Waud was situated when a Google search threw up a link to a Reginald Waud and the sighting of a ghost pig in Hampshire. Intrigued, I dug a little further. Were these Wauds one and the same person?

I now know that the Reginald Wauds were one and the same—Louis Reginald Waud (1873-1960), the scion of a very wealthy worsted spinner of Bradford in Yorkshire. In the 1930s Waud was member of the Council of the Avicultural Society*, a Fellow of the Zoological Society of London and a member of the British Ornithologists’ Union. Sometime he moved from Falcon Close at Woolton Hill, near Newbury to Bradley Court also near Newbury but at Chievely over the border in Berkshire. He was a dairy farmer and local newspapers documented his success at local agricultural shows.

I was hampered by Waud sometimes using his forenames in reverse order and of his associate being known, possibly as an initial error, by two names. Reversal of forenames was not uncommon; indeed my father in law always, as far as he knew, had his reversed which gave him difficulties in later life with legal papers.

Like many bird keepers of the time, he allowed nesting birds to leave their aviary and forage in the wild for food to feed their young. He had a Hawk-headed Parrot (Deroptyus accipitrinus) which flew in the grounds all day and returned to its cage for the night. It attacked and saw off an owl.

During the First World War, Reginald Waud volunteered for the Artists Rifles at the age of 41. He was appointed to the 10th Service Battalion of the South Staffordshire Regiment, a reserve force; he was promoted to Captain in early 1915. Records show that he was transferred to the 8th Battalion of the Manchester Regiment in August 1915. I have been unable to find out in what capacity he served since there were reserve as well as an active unit in the 8/Manchesters, the latter serving in Gallipoli. The next record I could find is in the London Gazette in 1918. He is described as having served in the South Staffordshire Regiment and having the rank of Honorary Captain. Does this mean he was serving by then serving as in some sort of civilian role within the army? He was though in that same announcement appointed as Captain in the army proper for duty under the Director of Railways from 14 January with seniority from 23 November 1914, the date shown at his enlistment with the Artists Rifles. Having finished reading Haig’s war diaries last year I now realise what a successful railway system had been set up behind the front line to bring materiel from the Channel ports.

Shortly after the armistice Waud married Ann Bower (1881-1972) in Westminster. He is shown on the marriage certificate as a Captain in the South Staffordshires. Waud contributed to the parish war memorial at Welford (Hoe Benham falls within that parish).

Waud was referred as Captain Waud after the War. I have often wondered why so many soldiers did so, since it was—and still is—considered infra-dig to carry a rank below that of Major into civilian life. I have read that they did so in order to indicate that they had served their country at a relatively advanced age and were considered sufficiently able to reach that rank. Waud had no need to establish his middle-class credentials as some very unrespectable Captains did.

Before the War, Waud was an artist. A graduate of the Royal Academy Schools, he attended from February 1896 until January 1901.

I must now jump to Reginald Waud’s friend, Osmond (sometimes Oswald, sometimes Osmund—even in the same document) Pittman (1874-1958). Pittman was also an artist at the Royal Academy Schools from July 1897 until July 1902 (he exhibited there in 1901, 1902 and 1904). He married Waud’s cousin, Caroline Mabel Waud, in 1908.

It was shortly before Pittman’s marriage when Waud and Pittman were living at Hoe Benham, a hamlet and, again, near Newbury, that Pittman saw ‘the ghost pig’. The story has been repeated numerous times in books on ghosts and the paranormal, sometimes with the details wrong. However, here is the original version:

                                                                                         Hoe Benham, Newbury
On November 2nd, 1907, I was painting in the Studio with my friend Reginald Waud; the model was our servant dressed as a widow, and we were waiting for Miss Miles to join us. At 10 o’clock I knew the milkman had come by the dogs barking in the cottage at the top of the garden. So I said, “I will take the milk in,” and went up to the cottage. After putting the jug in the pantry and shutting the cottage door, I looked up the road and saw Miss Miles coming down with her easel and paint-box. Following quite close behind her was a large white pig, with a long snout. I went down to the Studio and said to Waud, “What do you think Miss Miles is bringing down with her this morning, instead of her Chow? A large pig!” We roared with laughter, and he said, “Call out and tell her not to bring her friend in, and to be sure to shut the gate, as we take a great pride in our garden.” The moment Miss Miles appeared I opened the window and shouted out, “What have you done with your companion?” She was very surprised and said, “ My companion, what do you mean?” Then I told her what I had seen following her. She immediately said, “If a pig were trotting behind me, I must have heard it. Besides, there is a very easy way of finding out, for I passed the milkman in the lane and he must have seen it, but I shall go and look for myself.” When she came back she said: “There is no trace of a pig anywhere.” We made enquiries all over the village; no one had seen a stray pig. There is only one white one in the place, and this one, its owner assured us, could not possibly have got loose without his knowing it. At the present time there is a notice out forbidding owners to allow pigs to stray, under penalty of a fine, as there have been cases of swine fever. We enquired of the milkman next morning. He remembered passing Miss Miles, as he usually met her about the same time. He most emphatically said there was no pig to be seen anywhere on the road.

Well as ghost stories go you will probably admit to not being terribly impressed by this one. Pigs are surprisingly agile and could have through a hedge or over a bank in no time. Did anybody look for trotter prints? And would anybody confess to owning an escaped pig and risk a fine under the swine fever regulations? But as I said the account has been repeated time and time again. But who, you may ask, reported the event and why? It was none other than Clarissa Miles herself, in a paper published in the Journal of the Society for Psychical Research. The above account was signed by Pittman, Waud, Miles and the maid, Louisa Thorne. In fact it was only part of the story of the experiments and happenings at Hoe Benham.

Clarissa Frances Miles, born in Malmesbury, Wiltshire to a very wealthy family, was well known in psychic research circles: she was a dowser, an active member of the Society for Psychical Research (along with a number of Fellows of the Royal Society) and member of the Alchemical Society, Her experiments on ‘thought transference’ over long distances with her friend, Hermione Ramsden were, and still are, discussed in relation to evidence of  telepathy. Indeed some of these experiments were described in the same paper as the ‘ghost’ pig and involved Reginald Waud’s studio. Thoughts argued to have been transferred from Miss Ramsden from Miss Miles and then depicted by Miss Ramsden were of Waud’s new studio at the bottom of his garden. But that was not all.

Miss Miles continued:

On February 4th, 1908, I returned to Laburnum Villa to finish one or two paintings I began last autumn. I also wished to get further evidence of the strange animal forms that are supposed to haunt this lane, as I was told last November, after the episode of the phantom pig, that so many of the villagers had been witnesses of these remarkable apparitions under different forms, such as a cat, a dog, or a rabbit, or other animals. Mr. Pittman and Mr. Waud had never seen anything weird or out of the ordinary till I came down last autumn and began thought-transference with Miss Ramsden; they had only heard quite casually very vague accounts of something strange that had been seen years ago, to which they had paid but little attention. The villagers are very reticient [sic] and will not talk much for fear of being laughed at, but the history of the phantom pig overcame all difficulties on that score as regards my enquiries. 

The villagers all tell the same story to account for these apparitions. They put them all down to “Tommy King.” He was a farmer, who lived over a hundred years ago. He hung himself in an outhouse, and his spirit is supposed to haunt the spot, appearing in these strange shapes and making uncanny noises. The farm buildings and outhouses were all demolished on the occasion of the sale of the property by Mr. Dreweatt to Sir Richard Sutton in 1892; only a cottage remains, which is still called King’s cottage, and a disused well close to the lane, which is called Tommy King’s chalk well. In former years they used to dig out chalk from the side of the hill. I have looked at the Register of the Parish Church and there are entries of the deaths of two Tommy Kings, one in 1741, the other in 1753, so it must be one of these two…

Then Clarissa Miles produced signed statements of sightings. One, in January 1905, of a huge black dog that changed shape into a donkey standing on its hind legs with huge glowing eyes was signed by William Thorne. In 1897, William Thorne and John Barrett saw a ‘curious looking animal, snowy white’ which seemed too large for a cat, ‘more the size of a terrier with a fluffy coat’. Albert Thorne had seen an animal five feet long, the shape of a calf, with large glowing eyes.

There were also stories of ‘summat like a sheep’ that vanished when approached with a stick. After describing other strange sightings by villagers Clarissa Miles described what had happened in February 1908 when she was again staying at Laburnum Villa, Wickham Heath.

On Sunday, February 16th, 1908, I arranged to join Mr. Waud and go to the Church with him at Hoe Benham to 3 o’clock service. All the morning I felt in a weird, trance-like state, the condition I get into when I have visions of coming events, changes likely to affect my future. I described these sensations both to Mr. Pittman and to Mr. Waud. After Church I felt compelled to walk towards Tommy King’s chalk well, as I wished to see the road along which the team of horses came, which were so terrified at the white thing bobbing up and down over their heads [one of the stories from two locals]. We thought we would go and have tea with Mr. and Mrs. Le Mesurier, who live at the end of the lane just opposite Halfway Inn. To reach them from Hoe Benham we always take a short cut across the fields… After tea, Mr. Waud and I walked back. On passing Tommy King’s well an overpowering sense of suffocation seized me, I felt the presence of some awful being. This evil spirit seemed to follow me up Pound Hill. I walked back. On passing Tommy King’s well an overpowering sense of suffocation seized me, I felt the presence of some awful being. This evil spirit seemed to follow me up Pound Hill. I felt it longing to do me some bodily injury, there was such deadly malice and hate in the air. I described all this to Mr. Waud. We constantly looked around but could see nothing. These sensations continued all the way back, but grew worse as we walked up Benham Hill. After the cottage was passed a shiver went down my back and spine, the same kind of shock I experience when dowsing and walking over water. Suddenly quite close to us on the road the stillness of the evening was broken by an unearthly scream which seemed to end in a moan. As the sound seemed so very close to us, we both looked round to see if anything was near which would have produced it, but nothing whatever was visible. We both were speechless for a minute,— the effect was so awful. We hardly spoke till we reached Laburnum Villa, the time being about six o’clock. It was only the next day, in comparing notes, that we found this dreadful wailing cry was heard on the exact spot where the phantom pig was seen by Mr. Pittman following me on November 2nd, 1907. 

Two evenings after, on February 18th, I walked to Tommy King’s chalk well with Mr Pittman to try if any more strange phenomena could be seen. It was a brilliant moonlight night. No evil beings were there to disturb the stillness of this evening, all was calm and peaceful. I tried automatic writing, and two sentences were written twice over; they came with such lightning speed, no human hand could have written that pace alone. The words were, “I am in hell, pray for me, I am in hell, pray for me”; nothing more. On walking back we looked around and across the lane a white band of light appeared;—the same sparkling effect I have sometimes seen at sea, from the deck of a yacht on a dark night, shimmering on the ocean. It was of a quite different quality from the moonlight. This luminous effect undulated up and down, backwards and forwards, and seemed to come towards us, till it appeared to be only five yards away; it was about a foot from the ground; it then receded and disappeared. We both saw it; most strange and mysterious it appeared in the brilliant moonlight. Another evening a little later I walked down with Mr. Pittman but nothing of special interest occurred. But the pencil wrote automatically, “I am happier.” 

     On Benham Hill where so many strange apparitions have been seen, one evening when it was half daylight, half dark, I distinctly saw a white shadowy form which appeared like drapery standing against the gate. No one else saw this. 

     On Thursday, February 27th, my last evening at Hoe Benham, I dined at the cottage, and Mr. Pittman and Mr. Waud walked back with me. We were laughing and talking. Suddenly our conversation was interrupted by this unearthly moaning cry, which came wailing across the stillness of the evening. This doleful sound seemed to come over our heads from the roof of the Chapel which is exactly opposite Laburnum Villa. We all heard it. It was a repetition of the groan Mr. Waud and I heard the other evening preceded by that awful scream.
                                                                                          (Signed) Clarissa Miles.
                                                                                                       Reginald Waud.
                                                                                                     Osmund Pittman.
Mr. Waud adds:
On Monday, February 24th, whilst Pittman and I were taking tea in the cottage, two distinct taps were heard on the door just behind us. Both dogs started barking. I got up, thinking some one had called. I went to both doors and not a sign of any one or anything was to be seen. It was about 6.15.
                                                                                                       Reginald Waud.

Clarissa Miles even died in strange circumstances. She lived in a flat at 59 Egerton Gardens, South Kensington, London (and was doing so at the time of her visits to Hoe Benham in 1907-08). The lease was due to end on 14 August 1930 but she did not renew it, having visions that something was going to happen. She died on 13 August, one day before the lease was up; she was 70†.

Osmund/Oswald Pittman continued working as an artist and illustrator. His and Reginald Waud’s pictures appear at auction and examples can be found online.

A Louis Reginald Waud painting. Was
the model Louisa Thorne?

By Osmond Pittman

Another by Osmond Pittman

And what of the happenings at Hoe Benham in 1907 and 1908? Did they disappear when Clarissa Miles no longer visited? Did later residents see strange animals or strange lights?

While I can think of lots of alternative explanations (alcoholic beverages sold at the inn at the end of the lane to Hoe Benham could be involved) including an albino badger, and the ‘fantasy prone personality’ of Miss Miles, I will leave the reader with the story as it emerged at the height of the spiritualism and paranormal boom in the early years of the 20th Century.

But if anybody fancies a walk one night along the dark lane towards Hoe Benham, here is a satellite view of the area Miss Miles described from Google Earth for to to find your way:

But I may not join you.

This modern Google Earth view shows Hoe Benham. I have added the site of King's Barn from an 1890s Ordnance Survey.

The circle shows my estimate of the location of Reginald Waud's House. There is a house there with a building in the garden that could have been his studio and a lane (obscured by the trees) runs alongside to the garden to join the wider lane that passes the cottages and Benham Farm where the Thorne family worked, mainly as agricultural labourers. That wider lane leads to Wickham Heath where Clarissa Miles was staying.

My suggested location also accords with there being children from the cottages playing further along the lane and who, when questioned, said they had not seen a pig.

The distance from the circle to Laburnum Villa is approximately half a mile.

At the bottom of the map can be seen the A4, the major route between London and Bristol at the time.

*He joined the Society in May 1913. Confusingly, for several years he was listed as P. or S. Reginald Waud; the error was eventually corrected.

†I have been unable to find an entry in the registers for her birth. She was christened on 7 May 1860, probably suggesting her birth was earlier in that year.

Chequers, the Prime Minister’s official country house once had storks perched on its chimneys

In this Age of Wilful Ignorance, the only general agreement seems to be that the intellectual ability of our elected politicians is at an all-time low. But when the duplicitous and incompetent meet at the Prime Minister of the day’s official country house to make cunning plans, do they realise it was once the home of a major collection of birds kept by a previous inhabitant and that free-flying storks rested on its chimneys?

Keeping birds spanned all strata of Victorian and Edwardian society. The super-rich had large aviaries containing exotic birds or lakes housing exotic waterfowl, often of species that even today are often difficult to see in the wild, let alone catch and bring back alive during a long sea passage. The poor, on the other hand, had their singing canary or linnet in a cage.

Chequers or Checkers Court was handed to the nation in 1921. In the late 19th until the early 20th Century, the house was owned by the Astley family, descended from the Plantagenet kings of England.

The Reverend Hubert Delaval Astley (1860-1925) had a large collection of birds in later life at his house at Brinsop Court in Herefordshire. However, he makes it clear that he began in Chequers, then occupied by his brother. Indeed he had a photograph of ‘my beautiful old home’ as the frontispiece to his book, My Birds in Freedom & Captivity (Dent, London, 1900), an exemplar of sickly Victorian literature.

Chequers - from Astley's book. Subseqent alterations to the house can be
seen by comparing this to a modern photograph

Astley, who served as President of the Avicultural Society which then served the posher end of bird keeping, wrote:

I could not resist introducing as a frontispiece my beautiful old home in Buckinghamshire, of which my mother was heiress (now belonging to my brother, Mr Frankland-Russell Astley, a house restored in the reign of Queen Elizabeth—who there for a time incarcerated Lady Mary, the sister of Lady Jane Grey—and where once Lady Russell, one of the daughters of Oliver Cromwell, presided as mistress. On its chimneys many a stork has, of late years, rested; and in its grounds many another of my birds has walked and flown.

Hubert Astley with his Trumpeters at Brinsop Court

Casey Albert Wood (from here)

Astley’s birds featured in a paper published in 1926 written by a Canadian ophthalmologist who had visited the well-known aviculturists in England. He was there to see what could be learned from their methods of housing and caring for exotic birds that could be applied by bird keepers in North America. That meant he toured a number stately and upper middle-class homes. Casey Albert Wood (1856-1942) had served with the U.S. Army in the First World War and was well known for his work on the comparative anatomy of the avian eye and as a clinician. He described the accommodation for birds at Brinsop Court as well as those belonging to: a Captain Waud of Falcon Close, Woolton Hill in Hampshire; the Duchess of Wellington at Ewhurst Park, Hampshire; Mr Shore-Baily at Westbury, Wiltshire; the Marquess of Tavistock at Warblington, Hampshire; Alfred Ezra at Foxwarren Park, Surrey; Herbert Whitley of Paignton, Devon (later Paignton Zoo); Miss Ethel Chawner at Forest Bank, Hampshire; the late Mrs Dalton-Burgess (Clifton, Bristol) who is described as having the ‘largest, private indoor aviary in England’.

Wood illustrated his article with photographs of Hubert Astley and of Brinsop Court. These photographs were also used for a paper in a French magazine on Astley’s collection by Jean Delacour.

Wood CA. 1926. Lessons in aviculture from English aviaries. Condor 28, 3-30.

Malnutrition: Classic experiments from the 1970s—and what happened next

John Waterlow at 93 (from Millward, see below)

I was reading the recently-published biographical memoir of John Waterlow CMG FRS (1916-2010) written by Joe Millward, when I realised I needed to look something up—the outcome of a conversation I had with John over lunch at his club, the Savile, a couple of years before he died.

He was one of the authors of a paper in an important series from the 1970s written by members of his department at the London School of Hygiene and Tropical Medicine. Most of his work had centred on the nature, treatment and consequences of infant malnutrition, notably in Jamaica from 1954 until 1970 when he returned to London.

During the 1990s I became interested in maternal effects, and especially those where some change in the mother affected not just her offspring but those of one or more subsequent generations, that is, intergenerational effects. The field, then largely ignored, is now the huge band-wagon of epigenetics (using the term epigenetics in its wide sense and not the narrow definition employed by some molecular geneticists who restrict its use—and thereby cause confusion—to the switching of genes on and off by methylation).

Some non-genomic effects are now known to be transmitted from the father as well as from the mother. Thus, parental effects can be defined as arising when the parent makes a contribution to the phenotype of its progeny over and above that which results from the genes it contributes to the zygote. Thus in subsequent generations, if the effect persists we have grand-parental effects and so on. The essence of such an effect is that it involves no difference in the genome—the basis of genetic inheritance.

Unfortunately, while Lamarckian in character, there are those who overstate the importance of epigenetics in terms of evolutionary mechanisms, arguing that the generally accepted neo-Darwinian orthodoxy should be changed. But intergenerational effects are usually restricted to a few generations and there is still on evidence that changes in phenotype during a lifetime can affect the DNA of the germ line. I regard parental effects simply as part of the genetically controlled and therefore subject-to-natural-selection suite of options available to a parent to maximise the chances of survival and reproductive success of its offspring.

There is now considerable evidence for important parental effects of considerable importance in such fields as human health and welfare, conservation biology, farm animal health and welfare and in plant sciences. But back to London and John Waterlow’s department in London.

To explore the effects seen in malnourished human populations, the group in London kept rats on a low-protein diet for 10-12 generations. There was a rapid change in size and shape of the animals which stabilised after two or three generations. Males were 40% lighter than well-fed controls; females 30%. There were changes in the weights of various organs except the eyes. Changes also occurred in the endocrine system, in the reaction of the rats to noise and in their ability to learn.

These findings were interesting and important in mimicking the effects in children born in malnourished communities but the effects of rehabilitation to a normal diet were even more so. Mothers previously malnourished were given a normal diet from about two-thirds the way through pregnancy or pups born to malnourished mothers were fostered from birth by well-fed mothers. Physical recovery was fairly rapid but still took 2-3 generations to be complete. However—and this was the point stressed to me by John Waterlow—complex behavioural deficiencies were still evident. In fact he said, as far as he could remember the behavioural effects were not followed up further because the animals had been passed on to somebody in the U.S.A. to explore the long-term effects further—but he could not remember who. Remember he was around 90 at the time of our chat.

It is what happened to that colony of rats that Joe Millward’s memoir prompted me to find out.

Janina Galler in 2016 (Wikipedia)
Photograph: MaynardClark

The work was continued by Janina Galler at Harvard. She described in a chapter of a book published in 2001 the background:

Approximately 30 years ago, I first began working with an important colony of rats that originated with R.J.C. Stewart at the London School of Tropical Hygiene. When Reg Stewart was forced to retire at age 65, I inherited the colony. The rats were then in their thirteenth generation of malnutrition, and I had the privilege of bringing the colony from England to the United States. This animal model was particularly relevant to human populations where malnutrition is usually not a one-time event, but rather a chronic, insidious process present throughout the life span and across generations.

I will not describe the test procedures for behavioural differences suffice it to say that with simple tasks there appeared no difference between well-fed and previously malnourished rats. Complex tasks were a different matter, with a clear deficiency in the previously malnourished animals. The differences in behaviour and ability to cope with the world were not confined to the pups. Mothers paid more attention to previously malnourished foster pups; they seemed to compensate for the inability of the young pups to find their way back to the nest (a standard test).

Janina Galler then compared the findings in rats with those in the long-term studies she was doing on the human population of Barbados in relation to the alleviation, treatment and long-term effects of malnutrition. Her study began in 1973 using data on every child born between 1967 and 1972 who had suffered malnutrition. At that time the incidence of malnutrition increased when the price of sugar decreased. While the physical effects were relatively quickly reversed, differences in behaviour (attention deficit and poor performance in the 11+ exam) were not and indeed persisted, just as in the rat, into, at least, the next generation.

Thanks to that work alongside and in collaboration with Sir Frank Ramsey (1926-2009) on the public health side, malnutrition has been eliminated from Barbados. But clearly the effects persist with the question being: for how long?

Those rats in the studies started in London by Reginald J.C. Stewart have certainly played an invaluable part in demonstrating the immediate and intergenerational—and sometimes cryptic effects—of malnutrition.

Galler, J. 2001. The early experience. In, The Origins of Creativity, edited by K.H. Pfenninger and V.R. Shubik, pp101-116. Oxford University Press.

Millward DJ. 2018. John Conrad Waterlow. 13 June 1916-19 October 2010. Biographical Memoirs of Fellows of the Royal Society 69, 429-448.

Stewart RJC, Sheppard H, Preece R, Waterlow JC. 1980. The effect of rehabilitation at Merent stages of development of rats marginally malnourished for ten to twelve generations. British Journal of Nutrition 43, 403-412.

Stones in the stomach of crocodilians: new light on a weighty problem

Crocodiles have been in the news recently and the topic has been one in which I have a proprietary interest.

Ever since the discovery of stones in the abdomen of fossil crocodilians in 1831 by Geoffroy Saint-Hilaire there has been discussion as to the function they serve. Stones—gastroliths—found in the stomach of living crocodilians have usually been considered to have a hydrostatic function, in other words as ballast to increase the specific gravity of the animal in water, or to assist in the trituration of food in the stomach, like grit in the gizzard of birds.

The other question was whether the gastroliths had any function at all, but simply there by dint of their being ingested accidentally with their food.

When I came into the picture, as a small bit player, Hugh Cott (1900-1987) then in Cambridge had published in 1961 his work on the Nile Crocodile in East Africa. He made a strong case that the stone ballast enables the crocodile to stay under water and to pull the struggling large mammals they often feed on under the surface. The position of the stones in the gut raises the head end of floating crocodiles and stabilises them against rolling, he posited.

In the early 1960s, it was unresolved how crocodilians acquired the stones. Did they seek them out and swallow them?

I have written before on the American, usually then called the Mississippi, Alligator I had in the early 1960s. My father and I converted the end of a greenhouse to accommodate it. We built a concrete pond and left a basking area. We could not leave the ‘dry’ area as natural soil since the water would have been dirty in no time. We had a fairly small amount of mixed builders’ gravel which we spread across the dry area. The pond was filled, emptied and refilled each day for about a week to wash out soluble salts, the heater was installed and the 51 cm Alligator moved in from its bare temporary tank.

A short time later, while doing something else, I turned round to see the Alligator picking up pieces of gravel (plus some adhering earth) and swallowing them. It picked up each piece with the left side of the jaws held towards the substrate, then, when it had the stone in its mouth, straightened its head and swallowed. Each piece was about 1.5 cm in diameter.

I did not want to disturb the newly-installed animal that day since I wanted him to feed undisturbed. However, several days later I picked it up and could feel the stones in the stomach.

Apparently, I was the first to observe that the acquisition of stomach stones is a deliberate action and later authors have either credited me with that discovery, forgotten all about it or have not been diligent in searching the literature.

My alligator - from its size it was about a year old

Whether the gastroliths serve a hydrostatic function as ballast has been the subject of a good few papers and reviews since the 1960s but with comparatively little experimentation. Similarly, the efficiency of digestion in crocodilians with or without gastroliths has not been tested.

Cott’s views appeared to take a beating from mathematical modelling, the increase in specific gravity caused by a typical weight of gastroliths, as found by Cott in large crocodiles, not emerging as sufficient to have any marked effect on buoyancy. The conclusion was reached that control of buoyancy is achieved by the degree of filling of the lungs with air.

Expanding the lungs to dive could, though, increase the buoyancy to such an extent that the animal could not dive. Therefore, there were suggestions and some evidence, starting with Cott, that gastroliths could enable a greater volume of air in the lungs and therefore enable crocodiles to stay submerged for longer. The recently published paper tested this hypothesis directly—in my old friend, the American Alligator.

Juvenile alligators were used. First, all had any stones washed out of their stomachs. Then the duration of each of 21 voluntary dives in excess of 1 minute they made in an aquarium was recorded over two days. Then, granite stones no bigger than 1 cm were placed in the aquarium at a weight equal to 2.5% of body-weight. Four of the seven animals ingested all the stones. The three that swallowed only some were gently persuaded to ingest the rest by placing them at the back of the mouth.

The dives of the this time ballast-loaded alligators were recorded again. The results were very clear. The average duration of dives increased by 88% (5.9 to 11 minutes) and the maximum duration increased by 117% (11.7 to 25.3 minutes).

From Uriona et al 2019

The authors suggest that because juvenile crocodilians are subject to a high death rate through predation, a behaviour ‘that enhances the ability of juvenile crocodylians to remain submerged may have high selective value. Furthermore, juveniles have cartilaginous tissues that have lower specific gravity than the bone that will replace these tissues in later life, and the rate of depletion of oxygen stores during diving is relatively higher in juveniles than in larger animals making the increase in diving lung volume enabled by lithophagy potentially more important in juvenile animals than in adults.

A question that arises and one I have now wondered about for nearly 60 years after my alligator picked up and swallowed stones is: how do crocodilians know they need to find and ingest stones? Do they sense in some way how high they float in the water or whether they need to use muscles to maintain their characteristic pose on the surface with just the tip of the nose and eyes above the water?

The function of gastroliths in crocodiles has wider ramifications. This is because they are found in those extinct reptiles, the plesiosaurs but not in ichthyosaurs. Twenty-five years ago, Michael Taylor, a plesiosaur expert then at the Leicestershire Museums, wrote an excellent survey and comparative analysis of gastroliths and their possible function in marine tetrapods. I show the entire summary of his paper below because it puts the acquisition and occurrence of gastroliths into context:

Gastroliths or ‘stomach stones’ occur frequently in some, but not all, groups of fossil and living marine tetrapods. Comparative analysis of gastrolith distribution suggests a role in buoyancy control rather than food processing. Once accidental ingestion by bottom-feeding animals is excluded, gastroliths occur in most tetrapods which ‘fly’ underwater with hydrofoil limbs, including plesiosaurs, penguins, and otariid pinnipeds, but not the marine chelonians. They do not usually occur in cetaceans, ichthyosaurs, mosasaurs, and odobenid and phocid pinnipeds, which swim with a caudal fin or the equivalent. Occurrence in amphibious forms is variable; crocodilians often have gastroliths, but nothosaurs and placodonts do not. The correlation of gastroliths and underwater flight is corroborated by a comparative analysis which takes phylogenetic factors into account. There is no correlation with diet. Consideration of function and occurrence in terrestrial forms suggests that the use of gastroliths in digestion would not be useful, and might even be harmful, to a carnivorous marine tetrapod. Gastroliths are more efficient than skeletal bone (as in pachyostosis) in terms of sinking force per unit of added mass or volume. As well as driftwood and ice, marine tetrapods should be considered as a potential source of erratic stones in freshwater and marine sediments. Gastroliths may have evolved by the accidental ingestion of stones, the retention into adulthood of stones used by juveniles to process insect or plant food, or as a compensatory replacement for dense bones habitually filling the stomach. Their presence or absence should be more carefully recorded and further studies should be carried out on their function. 

Over the years I have seen and watched in the wild 9 of the currently-recognised 24 species of crocodilian: American Alligator (A. mississippiensis), Spectacled Caiman (Caiman crocodilus), Yacare Caiman (C. yacare), Black Caiman (Melanosuchus niger), American Crocodile (Crocodylus acutus), Australian Freshwater Crocodile (C. johnstoni), Nile Crocodile (C. niloticus), Mugger (C. palustris), Saltwater (Estuarine) Crocodile (C. porosus). As I have done so, I have pondered the question of whether they have gastroliths (the Spectacled Caiman is rumoured not to have) and whether or not they were making use of their presence as I watched.

It does finally appear that Humboldt—he of the eponymous current and penguin—was wrong. When told during his expedition to South America at the turn of the 19th Century that natives believed crocodilians augmented their weight with stones, he dismissed the notion as an “absurd hypothesis”.

The new paper:

Uriona TJ, Lyon M, Farmer CG. 2019. Lithophagy prolongs voluntary dives in American alligators (Alligator mississippiensis). Integrative Organismal Biology 1  doi.org/10.1093/iob/oby008

My old note:

Peaker M 1969. Active acquisition of stomach stones in a specimen of Alligator mississippiensis Daudin. British Journal of Herpetology 4, 103‑104

Two very useful reviews:

Taylor MA. 1993. Stomach stones for feeding or buoyancy? The occurrence and function of gastroliths in marine tetrapods. Philosophical Transactions of the Royal Society B 341, 163-175.

Wings O. 2007. A review of gastrolith function with implications for fossil vertebrates and a revised classification. Acta Palaeontologica Polonica 52, 1-16.

Western Lowland Gorillas—New research on their social structure and possible implications for the transmission of ebola virus

It is good to see old friends in print. This time it is the Western Lowland Gorillas we were privileged to visit and watch at Ngaga in the Odzala-Kokoua National Park of the Republic of Congo (Congo, Brazzaville) nearly five years ago (see my post of 18 August 2014).

We visited two groups just a few miles apart and it is the interactions that occur between the three group in the immediate vicinity of Ngaga that is the subject of a paper just published in Proceedings of the Royal Society.

I cannot overemphasise the enormous amount of work that went into this study done between 2013 and 2017. Behaviour of the groups was monitored on approximately 305 days each year. The trackers (who move at speed through the forest) located the groups each morning. Every individual gorilla can be recognised by the trackers and researchers. Notes were made of their behaviour for an average of two hours per day, itself a difficult undertaking because of the dense vegetation. And the authors do not even mentikon the sweat bees.

The behavioural work was done by Magdalena Bermejo (who heads the whole project in the Republic of Congo) and Germán Illera, with help from the same trackers who took us from the lodge to the groups of gorillas.

In addition to the behavioural observations, gorillas over a wide area were genotyped from samples of their faeces collected from the nests they build and occupy at night.

Previous observations on Western Lowland Gorillas (Gorilla gorilla gorilla) at bais (open grassy clearings often surrounding a body of water) suggested a very different behaviour from the Mountain Gorilla (G. beringei beringei). Groups of the former met without aggression—in marked contrast to encounters between group of the latter. In Ngaga, an area without bais, these observations were not only confirmed those but indicated a dynamic community structure. In the words of the authors:

Both approaches revealed a social system much more dynamic than anticipated, with non-aggressive intergroup encounters that involved social play by immature individuals, exchanges of members between groups likely modulated by kinship, and absence of infanticide evidenced by infants not fathered by the silverback of the group where they were found. This resulted in a community composed of groups that interacted frequently and not-aggressively, contrasting with the more fragmented and aggressive mountain gorilla (G. beringei beringei) societies. Such extended sociality can promote the sharing of behavioural and cultural traits, but…

The but refers to the disease that has devastated local gorilla and human populations, including the area studied—ebola. The authors make the point that interaction between groups could promote the rapid transmission of the virus:

Social behaviour may thus have greatly contributed to the massive impact of past Ebola outbreaks that have resulted in an increase of the threat level for the species, raising major conservation concerns about population declines in the future. Understanding group dynamics in social species is of utmost importance when coming to model the transmission of pathogens such as Ebolavirus. However, since the high mortality imposed by outbreaks is likely to select against this social behaviour, its persistence in W[estern] L[owland G[orilla] implies that either such massive die-offs may have been rare in the past, or that the associated benefits outweigh the disadvantages. In any case, the peculiar social behaviour of western lowland gorillas is an outcome of its evolutionary history and will definitively impact its fate.

I do wonder though whether the dynamic social structure and outbreeding may actually ensure a higher degree of immunocompetence of individuals. In other words, and turning the argument round, whether the social structure is actually protective to individuals exposed to potentially lethal infections.

Here is a link to the video included in the paper.

And here  again are my videos of the gorillas at Ngaga:

Forcina G, Vallet D, Le Gouar PJ, Bernardo-Madrid R, Illera G, Molina-Vacas G, Dréano S, Revilla E, Rodríguez-Teijeiro JD, Ménard N, Bermejo M, Vilà C. 2019. From groups to communities in western lowland gorillas. Proceedings of the Royal Society B 286: 20182019. http://dx.doi.org/10.1098/rspb.2018.2019 

Mycobacterium marinum—not just ‘fish-tank’ disease but a real occupational problem for the 20th Century fishing industry in Hong Kong

I have caught a few diseases from animals over the years. The most irritating and long-lasting was  Mycobacterium marinum, aka fish tuberculosis or ‘fish-tank disease’. In the 1980s I caught a knuckle on the rockwork of a marine aquarium and several weeks later a nasty granular, ulcerating lesion appeared. The local dermatologist thought he recognised what it was and took a biopsy for culture. The dermatologist had read the then recent literature which showed a number of aquarists throughout the western world had caught M. marinum in a similar manner. Cases were also known in which a swimmer had grazed a hand or foot on a sharp edge of a tropical seawater swimming pool. The organism will only grow at the extremities of the human body because it is cool; it cannot survive at internal body temperatures. Unfortunately the bacteriologists had not read the literature and tried to culture the organism at 37° C. Eventually they got the conditions right and confirmed the presence of the organism.

In the meantime I had started a self-cure regime, realising the alternative was a long course of anti-microbials used to treated human tuberculosis. Using glutaraldehyde to kill the tissue and a scalpel to cut each layer of newly dead flesh I gradually got the upper hand. By the time I went to see the dermatologist who had eventually received the report from the bacteriologists, I had the organism in full retreat. He was delighted to find an alternative treatment and said he would write it up along with photographs. A couple of months later, I had won; normal skin completely replaced the ugly lesion. Physiologists, of course, are well-known for doing experiments on themselves—or a handy PhD student.

What I had not realised until recently was that fishermen and their familiies could also be victims of M. marinum. In general the literature deals with case of marine aquarists and swimmers who live in Europe and North America, although other cases, as in a New York fish market, get a mention. While looking for something else entirely, I came across an account compiled by our late great friend Dafydd Evans (1938-2006), a historian as well as former Dean of Law and founder of the Law School at the University of Hong Kong. In his centenary history of the medical school* of that University, he included accounts of work in the various departments. Under Microbiology I found:

The service provided by the Department has benefitted from a constant academic input…Recent examples include the occupational diseases such as the chronic hand diseases of fisherfolk caused by Mycobacterium marinum…

I am now trying to see if I can spot any lesions on the hands of the Hong Kong fisherfolk, who lived virtually their entire lives on their boats, in my own cine films of the 1960s or those of others like Michael Rogge in the 1950s. I do not know how prevalent the problem was, or still is, but clearly of sufficient importance for the local microbiologists to do something about it.

The Hong Kong boat people at the least crowded end of Aberdeen harbour in 1967-68.
How many of those handling fish caught M. marinum?

A view further to the right of the previous photograph showing the end of the harbour and the then new Ap Lei Chau Power Station which opened in 1968. It was closed in 1989.

*I had been looking for a copy of the book in U.K. for another line of research. I then spotted one on eBay from a seller in Norwich. As it wended its way to me I wondered how a bookseller in Norwich had acquired it. The only person in Norfolk who I could think might have had a copy was Brian Lofts (1929-2015), Professor of Zoology at HKU at the time Dafydd wrote the book. Sure enough, the inner cover is signed ‘Brian Lofts”. Also included was an invoice from the University Press stating that the discounted price (HK$120) would be deducted from his salary. Financial control at HKU was always tight!

Evans, Dafydd Emrys. 1987. Constancy of Purpose. An Account of the Foundation and History of the Hong Kong College of Medicine and the Faculty of Medicine of the University of Hong Kong 1887-1987. Hong Kong University Press.