Vieillot’s Crested Fireback: a colour plate of a pheasant from 1955

In the days when colour printing was extremely expensive, the Avicultural Society had special appeals for funds to support the appearance in Avicultural Magazine of the occasional colour plate. A well-known bird artist was then commissioned. Although the whole run of the Society’s magazines can be found online, the plates rarely see the light of day. Therefore I decided to show one, now and again, on this site.

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This plate from the November-December 1956 issue is Vieillot’s Crested Fireback, a pheasant, also now known as the Malay Crested Fireback, from the Thai-Malay Peninsula and most of Sumatra. It has gone from being considered a subspecies, Lophura ignita rufa, to being split off as a full species, Lophura rufa, in 2014. Jean Delacour (1890-1985), the famous aviculturist and ornithologist, provided a short account of the birds to accompany the colour plate. It is found in lowland forest but now classed as ‘vulnerable’ because of the loss of habitat.

The plate was the work David Morrison Reid Henry (1919-1977) in 1955. He signed his work as D.M. Henry and was an artist favoured by the Avicultural Society for the plates published in this period.

The mystery of ‘Martha’ the last Passenger Pigeon

In a previous article I noted that a Wikipedia entry on Charles Otis Whitman of the University of Chicago stated that he sent the last Passenger Pigeon named ‘Martha’ to Cincinnati Zoo in 1902. Martha died on 1 September 1914 and the species became extinct. The claim that Martha came from Whitman is not supported by some contemporary accounts but can be deduced from another. This is confusing because both accounts arose from the same source.

The 1948 Version

There is no doubt that both Whitman and Cincinnati Zoo bred Passenger Pigeons. Both were unsuccessful in maintaining a breeding stock. An account of Martha’s origins and demise, written by William C. Herman, was published in The Auk. Herman wrote:

The man who, better than anyone else, is qualified to contribute the picture of the final act of this great tragedy is Mr. Sol A. Stephan of the Cincinnati Zoological Gardens in Cincinnati, Ohio (now still living at the age of 97) and from him the present writer obtained much of the information here presented.

Mr. Stephan took charge of the Cincinnati Zoological Gardens in 1878, previous to which time the Zoo had Wild [Passenger] Pigeons on exhibition and had had some success in raising them in captivity. At that time there were four pairs in the aviary which had been purchased in a western locality at a cost of $2.50 per pair; and others were added to this number. At this early date, Wild [Passenger] Pigeons were still to be found in large numbers, and some had visited the beech trees on the grounds that later were taken over by the Cincinnati Zoo. Even at this time Mr. Stephan realized that the Wild Pigeon was becoming scarce. As far back as 1857 a bill to protect it had been proposed by a committee of the Ohio Senate, but no action was taken. As prices became higher, Mr. Stephan became very anxious for the preservation of the species, and he determined to breed the birds in captivity in order to have them for purposes of exhibition. Their habits were carefully observed, with special attention to the diet. He learned that they thrived best on a mixed food of cracked corn, wheat, crackermeal, cooked liver, and eggs. While recent discoveries have been made in the feeding of birds and mammals in captivity, the diet used by Mr. Stephan seemed to be adequate.

Herman then went on to describe the breeding quarters and the nesting. Each nest contained one egg and the birds bred only once a year. Stephan was reported here as having bred 14. The birds became very tame and lived for 10-15 years. Eventually he was left with two birds, one male, one female. Herman continued:

When other zoological gardens learned that the Cincinnati Zoo had the only remaining pair, they offered prices which rose from $100 to $1000 for the pair. At this time Mr. Stephan was not anxious to dispose of such a valuable asset and wisely kept them, for they attracted ornithologists and other scientists from distant parts. The female of this last pair had been hatched in the Cincinnati Zoo in 1885, and was named 'Martha,' in memory of the wife of a friend of Mr. Stephan. This female pigeon became the last survivor of the billions that in flights once darkened the sky.

Martha and her mate built but one nest, and the lone egg that she laid in this proved to be infertile. Since both birds of this last pair were so old that it was apparently impossible for them to rear any young, eggs from Martha were placed under incubating Rock Doves (domestic pigeons), in the hope that the latter might hatch and rear the young to maturity. This failed because the eggs were infertile.

The male of the pair died having been in captivity for 26 years. Martha lived on and died aged 29 (‘an unusually old age in captivity’). She was found dead. Her skin was sent to the United States National Museum in Washington, D.C. Mr. William Palmer, the taxidermist, removed the skin. Dr. Robert W. Shufeldt made an anatomical dissection and published a description of this (Auk, 32: 29-41, pls. 4-6, 1915), giving the cause of death as advanced age’.

The 1908 Version

An account of Whitman’s work with Passenger Pigeons was published in The Auk in 1908 in an article written by Ruthven Deane (1851-1934), a leading amateur ornithologist of his day. Whitman had obtained his birds, as part of his vast study of the evolution of pigeons, from a David W. Whitaker of Milwaukee. A flock of 15 had been bred from a single pair ‘of young birds which he received from a young Indian who trapped them in Shawano County in northeastern Wisconsin’ in 1888. From 1896 to 1897 Whitman bought, in dribs and drabs, the whole flock. During 1897, 9 young were hatched but only four lived. in 1998, he gave 7 birds back to Whitaker. Again eggs were layed by his birds but only 2 hatched. Gradually the flock became depleted as even fewer of the eggs hatched while adults were lost or died. By the end of 1907 all of Whitman’s Passenger Pigeons were dead. He was left with two male Passenger Pigeon x Ring Dove hybrids which he found to be infertile.

Whitaker had four live males in 1908 which died presumably over the next few years.

What is clear from Deane’s paper is that Whitman sent a female bird to Cincinnati Zoo in 1902. But then things get confusing. Deane reported:

For years we have known of the Passenger Pigeons in the Gardens of the Cincinnati Zoölogical Co., and I am much indebted to Mr. S.A. Stephan, General Manager, for the following report of their flock, in a letter written November 9, 1907. 

The original flock, which came from Michigan in 1875, consisted of twenty-six birds, about half males and half females. A short time later, however, five or six of these escaped. They have bred from time to time and we have raised about twenty-three birds. In no instance has more than one egg been deposited at a time. At the present time our flock has been reduced to three, one male from the original flock, now about twenty-three years old, one male, which we raised, is about eighteen years old, and one female that we obtained from Prof. Whitman's flock in 1902, which is about twelve years old. We have never detected any particular disease which has caused the decrease of the flock, but have attributed it in most cases to old age. 

Deane concluded: 

The remnants of the Milwaukee and Cincinnati flocks now number but seven birds (6♂︎, 1 ♀︎) with little or no chance of further reproduction.

This is the version of the story that appeared win the world’s press after the death of the bird in 1914. Further media attention followed the publication of the paper describing the dissection of the bird at the Smithsonian by Robert Wilson Shufeldt (1850-1934) who was medically qualified. In it he wrote:

On February twenty-first, 1914, Mr. S. A. Stephan, General Manager of The Cincinnati Zoological Company, of Cincinnati, Ohio, wrote me that " Our Passenger Pigeon has been promised to the Smithsonian Institution when it dies. This bird is a female and now about 29 years old, and the last one of a flock of eight that we got in 1878." I have since learned that it was hatched in the Garden.

Shufeldt, himself a colourful character with a certain notoriety, noted two things that seem odd. First he wrote:

From Mr. Stephan, who wrote me on the 7th of September, 1914, I learn that "our female passenger pigeon died September 1st [1914] at 1 P. M. of old age, being about 29 years old." It was almost immediately packed in ice and shipped to the National Museum at Washington, D. C., where it was received in fine condition on the fourth of that month.

But external appearances were deceptive. He was with William Palmer of the museum who skinned the birds for taxidermy. Then, after a ‘late lunch’, Shufeldt began his dissection but had a surprise:

I found, on the right side of the abdomen, a slit-like opening, one-half a centimeter in length, which led freely into the abdominal cavity, and from which blood was oozing. This opening I enlarged in order to withdraw the viscera for the purpose of making a photograph of them, previous to proceeding with the dissection of the organs within. This has been my practice for a great many sears. 

Much to my surprise, I found a quantity of blood (not clotted) in the abdominal cavity, and the right lobe of the liver and the intestine almost entirely broken up, as though it had been done with some instrument. As to the intestine, it was missing altogether, while the right lobe of the liver was in scattered fragments The firmer organs were apparently intact, but did not occupy their normal positions.

Later in the paper he continued:

There was no evidence whatever of the presence of the intestine in any part of its continuity save a piece about 8 mm. in length, where it emerged from the gizzard and the ragged margin surrounding the abdominal boundary of the vent. All the portion referred to was not in the abdominal cavity. The entire right lobe of the rather large liver was in a disintegrated condition, showing its internal structure, and exposing the organs usually concealed by it to view. The heart was in its normal position, while the gizzard was rotated to the left side. I discovered no blood clots or parasites of any kind in the abdominal cavity 

The inference from this description and the very detailed of how the bird was received and handled at the Smithsonian is clear: somebody had done something to the specimen in Cincinnati. Had somebody poked and pulled inside the abdominal cavity with a long pair of forceps, the only way I can think of by which the whole intestine and chunks of liver could be removed through a half-centimetre slit? But who, and why?

Also of interest with respect to the alleged age of the birds is:

The feet were of a deep, flesh-colored pink, clean and healthy, while the claws presented no evidences indicative of unusual age, though not a few of wear. 


Two different accounts


The reader of these accounts is therefore in some doubt as to which version, if either, is correct. Both accounts are from the same source, Sol Stephan. In 1908 he wrote that the last surviving female was one sent by Whitman in 1902. The wording in Deane’s article can be taken to imply that this was one of the birds obtained from Whitaker in 1896-7. By contrast, in Shufeldt’s and Herman’s accounts written with information supplied by Stephan after the death of the last bird in 1914, Stephan stated that the female had been hatched in the Cincinnati Zoo in 1885, was named Martha in memory of the wife of a friend, and had died aged 29.

There is a great difference in the age of the last Passenger Pigeon at death, 29, if it the one bred at Cincinnati. If we calculate from Stephan’s account from 1907, then, had the bird come from Whitman, it would have been approximately 19 at its death, much closer to the 10-15 years lifespan quoted for other birds kept in captivity.

Different accounts pre-date the bird’s death. For example, in William T. Hornaday's Our Vanishing Wild Life (1913), the bird is described  as ‘twenty years old in 1912’.

A key question is of course, was possible to know the identities and history of the individual birds at Cincinnati Zoo? Were they ringed (banded in the U.S.A.)? Shufeldt made no mention of one being found at autopsy. I have looked at two photographs of Whitman’s birds that can be seen online and can see no trace of a ring.

All publicity is good publicity: A possible explanation

Salvator A ‘Sol’ Stephan was, in his younger days, a showman, working for a travelling menagerie and also as a representative for Carl Hagenbeck, the famous animal dealer of Hamburg. He arrived at the soon to be opened Cincinnati Zoo in 1875 with an elephant. Persuaded to stay on to settle in the animal, he retired 62 years later as Superintendent and General Manager. He died in 1949 aged 100.

Cincinnati Zoo had a difficult time in its early decades. It entered receivership in 1898 and in order to prevent liquidation the shareholders gave up their investments of nearly a quarter of a million dollars. Then it ran as a business for two years. The Cincinnati Traction Company, hoping apparently to use it as a marketing tool for its streetcars, operated the zoo until 1917 (in some accounts 1915).

I wonder if the opportunity was taken, with the death of the last Passenger Pigeon, to make sure that the attendant publicity (for which Sol Stephan was known) showed the Zoo in the best possible light. What better than a record longevity and the local breeding and care of a bird named Martha?

On the name there is even confusion in the literature. Stephan wrote that she was named after the wife of a friend. Other secondary sources say the name is after Martha Washington, with the male of the final remaining pair as George.

I cannot think of any other explanation, other than a marked loss of memory, for the difference in accounts given by Stephan in 1907 and 1914. In other words, my take on the origins of ‘Martha’ is that she was, most likely, the bird sent to Cincinnati by Whitman in 1902. Shufeldt also remarked that the information he was given by Stephan did not tally with what he was told later:

This bird is a female and now about 29 years old, and the last one of a flock of eight that we got in 1878. I have since learned that it was hatched in the Garden.

As outlined above, I have the feeling from reading between the lines that Shufeldt was suspicious over what he was told about the age of the bird and the circumstances of its death and preservation.

Errol Fuller dealt with the story of Martha’s death in his 2014 book on the Passenger Pigeon:

The truth is that we don't actually know when Martha died, at least not with any degree of exactitude. This lack of definitive information is largely due to differing accounts given by the main keeper at the zoo, Salvator "Sol" Stephans, and his son Joseph. Both kept changing their story. One of their accounts states that Martha died in her cage at precisely one o'clock on the afternoon of Tuesday, September 1, 1914. Another suggests her death might actually have occurred some four hours later. One version of the story relays the romantic idea that she died surrounded by a group of grieving keepers. Another maintains that just Sol and Joseph were with her. Yet another is less poetic and indicates she died alone and was found lying on the floor of her cage by an assistant keeper named William Bruntz. The truth hardly matters, of course. Dead is dead.

Furthermore, any serious attempt to go through written zoo records to solve the mystery is impossible; they were destroyed by fire in 1963. What we do know is that the first day of September was a swelteringly hot day with high humidity levels; perhaps it was all just too much for a frail old bird. So what else is there? The answer is, not a lot. It seems certain that Martha was born in captivity, but where? Mr. Stephans' recollections were as varied on this subject as they were on others. Perhaps this was due to captivity, but where? Mr. Stephans' recollections were as varied on this subject as they were on others. Perhaps this was due to poor note keeping; perhaps it was forgetfulness or just a lack of knowledge concerning past events that at the time of their occurrence would have seemed unimportant.

 

What would happen now?

The accounts by ornithologists and aviculturists describe the failure of the faltering and early attempts at what would today be described as a captive breeding programme for a highly endangered species. A useful discussion might be had on whether, with the vast increase in knowledge of wild animal husbandry, we could have done any better in 2022. What stands out in reading the accounts is: the failure of eggs to hatch (still a major problem), which could be related to infertility, or to poor conditions for incubation; the decline in fecundity as the females aged; the degree of inbreeding. All Whitaker’s and therefore Whitman’s birds were descended from a single pair of young birds which had been hand reared. My bet is they were from the same nest. Therefore, inbreeding depression could account for some of Whitman’s findings (a phenomenon he described as ‘weakness of the germ’) and that explanation appeared in print in 1913. The group originally kept in Cincinnati was larger and therefore more genetically diverse. Nevertheless that population also declined over the years. Nutrition may also have been inadequate in captive birds. Wallace Craig recorded the behaviour of Whitman’s birds. He wrote:

A word as to the care of Passenger Pigeons, in case we may be so fortunate as to find some still living. Professor Whitman kept his in the same pen with other species, supplied with the pigeonstaples of mixed seed, grit, oyster shells, salt, and plenty of green food such as lettuce. After he had had his flock many years, he discovered that they would greedily devour earthworms, and when abundantly supplied with this delicacy the birds improved so much in health and vigor that Professor Whitman thought if only he had known of this diet early enough he might have saved his stock from dying away.

—§§§§§—

As unsatisfactory as the different accounts of the death of what is still presumed to be the last Passenger Pigeon (Ectopistes migratorius) are, they are immaterial to the extinction of the species that occurred over a relatively short period. I started this series of two articles with the achievements of Oscar Riddle in discovering, firstly, the hormone prolactin and, secondly, the latter’s effect on the pigeon crop sac. Because of Riddle’s close links to Charles Otis Whitman, I became fascinated by the history of the extinction of the Passenger Pigeon and of those who kept and bred the bird in captivity. I still have much more to read.

Martha's feathers were not in very good condition when she died and
her skin was not displayed by the National Museum in Washington for
many years. However, it is now on display, as described here.
Photographed in 2015 by Ph0705.

Deane R. 1896. Some notes on the Passenger Pigeon (Ectopistes migratorius) in confinement. Auk 13, 234-237.

Deane R. 1908. The Passenger Pigeon (Ectopistes migratorius) in confinement. Auk 25, 181-183.

Fuller E. 2014. The Passenger Pigeon. Princeton and Oxford: Princeton University Press.

Grant W. 1913. Recollections of the Passenger Pigeon in captivity. Bird Lore 15,  93-99

Herman WC. 1948. The last Passenger Pigeon. Auk 65, 77-80.

Shufeldt RW. 1915. Notes on the Passenger Pigeon (Ectopistes migratorius) lately living in the Cincinnati Zoölogical Gardens. Auk 32, 29-41.

 

What—and who—connects the last Passenger Pigeons with the discovery of a hormone?

Oscar Riddle was a scientist who came good. After an inauspicious and tedious early research career he is remembered for discovering the hormone prolactin. Prolactin from the pituitary gland is essential for lactation as well as being involved in a whole host of physiological processes in vertebrates, acting both through the blood stream as a hormone and as a local controlling factor within tissues and organs.

Oscar Riddle
from here

Around 1930 Riddle became interested in trying to find the stimulus to milk secretion. He knew of then recent research which showed that aqueous extracts of the anterior pituitary will induce milk secretion in rabbits. Attempts by others to isolate and identify the chemical factor had, however, failed. With the assistance of a chemist he had taken on, Robert Wesley Bates (1904-1994), and of Simon W. Dykshorn, a research student, he succeeded in isolating a compound that had the stimulatory effect on the mammary gland but was not the same as the putative hormones present in anterior pituitary extracts which affected other tissues and organs. Riddle had a new hormone, and the name he gave it, prolactin, still stands.

Working on something mammalian was a major diversion for Riddle. He had for over 20 years   worked almost exclusively on pigeons, particularly in the 1920s on metabolism, reproduction and endocrinology. He was therefore well prepared to ask the simple question, is the secretion of crop ‘milk’ in pigeons also stimulated by his newly-extracted hormone. Yes, was the answer and Riddle, Bates and Dykshorn soon developed the pigeon crop sac as a bioassay for prolactin activity. The lasting importance of this discovery is sometimes overlooked by modern endocrinologists who all seem to be based in the increasingly inward looking sub-discipline of clinical endocrinology. Because they had to wait until the 1970s for a reliable method of measuring the concentrations of prolactin in blood, the bioassay is of no interest. However, it continued to be used, certainly into the 1980s, by those looking for prolactin-like hormones produced in other tissues, the physiologically very important placental lactogens produced by the placenta, for example.

On 26 December 1932, shortly after the publication of their work, Simon Dykshorn, variously described then as a research student or research fellow, was found shot dead in woods, where he had gone hunting, about a mile west of the Cold Spring Harbor Laboratory on Long Island, New York, where Riddle had worked since 1912. Dykshorn was 30.

After this tragedy, with Bates and Ernest Lester Lahr (1897-1982), Riddle went on to further purify their prolactin preparation and to identify it as a protein.

Riddle's Prolactin Team
From Family Trees on ancestry.com

The reason Riddle had spent much of his time studying pigeons and making little headway for some years was that he completed his PhD in Chicago under the supervision of Charles Otis Whitman. Whitman was trying to find the cause of the pattern of alternate light and dark bars on the feathers of many species of birds. This question arose out of Whitman’s long interest in the evolution of birds. There were many dead ends and misinterpretations, not helped by Whitman’s rejection of both Mendelian genetics and natural selection. His devotion to the study of pigeons was such that he worked increasingly at home where for a period of 15 years he usually had around 500 individuals of 40 species in dovecotes around his house. He hybridised 40 species. Foy years Whitman moved his pigeons each summer to Marine Laboratory at Woods Hole in Massachusetts (where had had been its director) by rail, and then moved them back to Chicago in September. Initially the University of Chicago paid for the pigeons to be moved but then refused and thereafter Whitman and his wife paid.

Riddle valued highly the companionship and guidance of Whitman. He wrote: ‘Whitman became nearer to being a father to me than anyone I have known.’ Whitman died suddenly in 1910, aged 57. Riddle undertook to write up Whitman’s work. However, the new regime in Chicago, not impressed by Whitman’s research, did not want his disciple. Whitman though had been a influential figure in American zoology and In 1912, the Carnegie Institute of Washington took on Riddle as a research associate, together with Whitman’s pigeons and papers. By 1913 all were ensconced at Cold Spring Harbor. There is something odd in that the director of the lab, Charles Benedict Davenport (1866-1944), must have acquiesced to Riddle’s appointment—perhaps he had no choice—but did nothing to provide him with adequate facilities. Whitman’s views were the exact opposite of the main thrust of work, Mendelian genetics, at the lab and it must have come as some embarrassment when Whitman’s work was eventually pulled together over the next 6 years and published in three volumes, when the title of the first was Orthogenetic Evolution in Pigeons. Whitman really believed he had demonstrated orthogenetic evolution or evolution along pre-determined pathways. However, Davenport wrote an extensive obituary of Whitman. Perhaps they were friends and Davenport felt under some obligation to provide some means of concluding Whitman’s work while not appreciating the direction Whitman was heading. In the meantime Davenport was earning opprobrium of his own as one of the leading American exponents of eugenics. His views were so extreme and his notions from Mendelian genetics so warped that other eugenicists disowned him. He was, and remained, a supporter of the Nazi party in Germany before and during the Second World War. Perhaps then there is a simpler explanation for Davenport’s lack of support for Riddle: was neither a Mendelian geneticist nor a eugenicist who did not fit into Davenport’s organisation.

Whitman with his pigeons
From here

Most of the immense amount of work to sort and write up Whitman’s notes fell to Riddle. But even this labour of love ran into difficulties. For some reason, Whitman’s widow did not allow Riddle access to all of her husband’s papers. The very long and arduous episode in Riddle’s life must be one of the best illustrations of ‘no good deed goes unpunished’. The publications were not well received, particularly the one on orthogenesis. Reviewers of that volume pointed out that many of his ideas had already been, in kinder words, shot down in flames. The three volumes can be found online and even a quick look provides some appreciation of the amount of work and the level of detail involved. Two artists were engaged to provide the colour plates and other illustrations. The cost to the Carnegie Institute must have been enormous.

Volume 1 of the posthumous works of Whitman

Eventually, Riddle, by developing interests in how birds work and into the burgeoning fields of metabolism and endocrinology, pulled away from Whitman’s ideas on evolution. He was by 1930 very well equipped to discover something really important.

And it was Whitman who provides the link I referred to in the title of this article. Lord Walter Rothschild in his 1907 book, Extinct Birds, quoted from notes sent to him by James H. Fleming of Toronto”

For all practical purposes, the close of the Nineteenth Century saw the final extinction of the Passenger Pigeon in a wild state, and there remained only the small flock, numbering in 1903 not more than a dozen, that had been bred in captivity by Prof. C. O. Whitman, of Chicago; these birds are the descendants of a single pair, and have long ago ceased to breed. It was in an effort to obtain fresh blood for this flock that I started a newspaper enquiry that brought many replies, none of which could be substantiated as  records of the Passenger Pigeon, and many referred to the Mourning Dove. I am aware that there has been lately wide-spread and persistent rumours of the return of the pigeons, but no rumour has borne investigation, and I feel that Prof. Whitman's small flock, now reduced (in 1906) to five birds, are the last representatives of a species around whose disappearance mystery and fable will always gather.

Rothschild’s correspondent seems to have overlooked the Passenger Pigeons at Cincinnati Zoo, where they had also bred and where the last one, ‘Martha’, died in 1914. The entry for Whitman on Wikipedia states that he sent Martha to Cincinnati in 1902. However, some contemporary evidence indicates that was not the case while another account—from the same source—suggests that the last living Passenger Pigeon was one received from Whitman. I will return to Martha and her origins in another article.

Oscar Riddle was elected to the U.S. National Academy of Sciences in 1939 for his work on prolactin. It was as a result of his earlier career with Whitman that he realised the importance of studying the effect of prolactin on the pigeon’s crop. The results of that second discovery were then developed into a vital bioassay that was used—and could well be still used—throughout the world for studying the biological activity prolactin-like compounds in all vertebrates.

Riddle was a major player in his later life in promoting the study and teaching of evolution in the USA where belief in the supernatural then—and still does—trumps rationality. He really did, after a shaky start, come good.

Oscar Riddle died on 29 November 1968, aged 91.

Female Passenger Pigeon by K. Hayashi
K. Hayashi and K. Toda were the artists commissioned to provide the illustrations
to Whitman's posthumous works. This is from Volume 2

Corner GW. 1974. Oscar Riddle 1877-1968. National Academy of Sciences. Biographical Memoirs: Volume 45. p 448-488. Washington, DC: The National Academies Press. doi.org/10.17226/568

Davenport, CB. 1917. The personality, heredity and work of Charles Otis Whitman. 1843-1910. American Naturalist 51, 5-30.

Shedding light on gulls’ eggs: how light exposure triggers different development in the last egg of a clutch

In one of those quirks of circumstance a photograph of a much smaller than usual Black-headed Gull appeared on the Ayrshire Nature Facebook page. A short time later, news of a new paper offering a possible explanation arrived in my inbox.

In gulls the young hatch in the order in which eggs are laid. Small young gulls are those that hatched last and are at the receiving end of intense competition for resources from their earlier hatched siblings. These last hatched chicks are known to exhibit a resilient “junior phenotype” ‘characterized by accelerated hatching, increased begging behavior and a slower growth rate’. Is there some cue from the outside world that causes them to develop these physiological and behavioural features?

Francisco Ruiz-Raya, Jose Noguera and Alberto Velando of the University of Vigo in Spain considered that light might be the trigger for changing the pre-hatching development to the junior phenotype. As senior birds hatch, the parents spend less time incubating and more time feeding them. The last egg is therefore exposed to light for longer. To test the hypothesis they studied Yellow-legged Gulls (Larus michahellis), which lay three eggs per clutch at intervals of 1-3 days with incubation beginning after the second egg is laid. Junior birds typically hatch 1-3 days after their siblings. The difference of exposure of the eggs to light was marked. Third-laid eggs spent 82% of their time exposed to high light intensities during a six-hour period two days before hatching. By contrast senior eggs spent only 0.05% of their time. If light is the trigger, then the signal is loud (well, bright) and clear.

Yellow-legged Gull (Larus michahellis) in the Gulf of Olbia (Sardinia, Italy)
Gzzz, CC BY-SA 3.0 <https://creativecommons.org/licenses/by-sa/3.0>, via Wikimedia Commons

By manipulating the exposure of eggs to light it became clear that the junior phenotype of the can indeed be triggered by their exposure to light while still in the egg. The authors also found corresponding physiological changes associated with different patterns of gene expression. Somehow light exposure at perhaps a critical time before hatching results in the production of chicks with different physiological and behavioural characteristics from the same clutch of eggs.

Ruiz-Raya F, Noguera JC, Velando A. 2022. Light received by embryos promotes postnatal junior phenotypes in a seabird. Behavioral Ecology doi.org/10.1093/beheco/arac079

Impala in the Masai Mara of Kenya

 

This male Impala (Aepyceros melampus) in the Masai Mara of Kenya was engaged in 'tongue flicking' when we photographed him in September 1991.

Tokay Geckos in Hong Kong: wild and/or feral populations?

Tokay geckos are known to occur on the island of Lantau, in the mainland New Territories and on Hong Kong island. The new paper on the trade in Tokay geckos in Hong Kong I referred to in a previous article found suggestions that at least one of the populations in the New Territories contains the red-spotted morph, indicating a non-Hong Kong origin and likely the result of introduction either by escape of captive geckos or deliberate release. The authors concluded:

Our genetic analysis shows that TCM [traditional Chinese medicine] trade in Hong Kong is not likely supplied by local wild populations, but by imports from Southeast Asia. Additional information from SIA [stable isotope analysis] supports our genetic findings, as we detected very little overlap in isotopic signatures between TCM and Hong Kong wild tokays. Trade also leads to mixing of populations through introductions related to the live pet trade. We found that at least two Hong Kong wild tokay populations were likely established from released pets. 

The conclusion that Tokay Geckos are not now collected from the wild in Hong Kong suggests a change has occurred since the first discovery of Tokay Geckos in the wild on Lantau Island on 23 February 1950 since that discovery was only made when the source of live geckos for sale in a snake shop came to light.

The grand old man of Hong Kong herpetology, John Dudley Romer (1920-1982) wrote in the journal Copeia:

On February 23, 1950, I purchased a live adult Gekko gecko from one of the snake-dealers, and on making enquiries regarding its origin, was informed that it had been received from Tung Chung. A few days later I visited that locality with some friends and enquiries on arrival at once revealed that this gecko is well-known to the local Chinese. One live adult specimen was obtained from a woman who had apparently been keeping it to sell in the city. Since time was short we engaged one of the villagers to show us exactly where the geckos were found, and after an hour or two of stiff climbing, he finally halted on a rocky hillside and pointed out several narrow clefts in some particularly large rocks. It was then only a few minutes before we saw one of these giant geckoes in a cleft in one of the rocks.

John went on to suggest that the gecko may have been more widely distributed in Hong Kong since it had not been found in the typical habitats of this species, in northern Burma for example: “houses, huts and trees and at nights are often seen on fence posts. In forests they live in holes or on the bark of trees.” He concluded:

It seems likely that their absence from dwellings and apparent confinement to the rocks on lonely hill-sides in this colony may be related to their extermination from other types of habitat.

In other words, he was suggesting that the population on Lantau was a relict of a once much wider distribution across Hong Kong. The discovery of a second population on Lion Rock in the New Territories could give credence to such an explanation.

When the second edition of Hong Kong Reptiles and Amphibians was compiled and published in 1998, the populations on Lantau and on Lion Rock were considered native; it seems that conclusion still holds.

Those arriving or leaving Hong Kong by air will, if sitting on the side nearest the mountains of Lantau will be passing the site John Romer described. However, his fishing village of Tung Chung is now  the nearest town to the Chek Lap Kok airport with a population of over 1000,000 and still rising. If tourists ever return Hong Kong, the Ngong Ping Cable Car links Tung Chung to the Po Lin Monastery and Big Buddha. The once difficult to reach village is now just another stop on the MTR.

The page on the Tokay Gecko from Karsen, Lau and Bogadek's
Hong Kong Amphibians and Reptiles

Dufour PC, Miot EF, So TC, Tang SL, Jones EE, Kong TC, Yuan FL, Sung Y-H, Dingle C, Bonebrake TC. 2022 Home and hub: pet trade and traditional medicine impact reptile populations in source locations and destinations. Proceedings of the Royal Society B 289:20221011 doi.org/10.1098/rspb.2022.1011 

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

Romer JD. 1951. The occurrence of the lizard Gekko gecko in the colony of Hong Hong. Copeia 1951, 80.

Hong Kong Dragonfly: the female version in gold

 

This one from AJP seems to be the work of a Mr Auric Goldfinger before his demise at the hand of Bond, James Bond. However, it is probably a female Common Blue Skimmer, Orthetrum glaucum. I showed the pruinescent blue male in my last post.

A pruinescent Hong Kong Dragonfly

 A weekend photograph came from Hong Kong of a common species there, the Common Blue Skimmer, Orthetrum glaucum.

The blue colour is not a blue pigment but created by Tyndall scattering of light in wax particles secreted on the surface (pruinescence) of the dragonfly.

A new paper on the vast trade in Tokay Geckos in general and Hong Kong in particular

The numbers of some species of wild animal traded defy the most vivid imagination. I read, in a new paper:

About three million tokay geckos per year were exported from Indonesia from 2015 to 2021, all dead and dried, with an export record of 5,974,550 in 2022.

For how long populations in the wild can be sustained, not just in Indonesia but in other south-east Asian countries from which Tokay geckos are exported, is open to question. The dried geckos are sent to China for human consumption in traditional Chinese medicine.

Some idea of the size of the market for Tokay Geckos (Gekko gecko) in Hong Kong was brought home to me about 20 years ago. Outside a medicine/food importers were bags of goods waiting to be moved inside. Amongst the interesting aromas which surround such establishments were sacks full of dried geckos. Each must have contained several thousand adults.

The recent paper tracks, using one mitochondrial gene, the origin of the geckos imported into Hong Kong. However, Hong Kong is interesting in that there is a wild population. In addition, live geckos were and still are imported both for traditional medicine and for sale in pet shops, although in comparison with the former, the numbers involved must be very small. Such is the concern with the volume of imports that the Tokay Gecko has recently been afforded protection under the Hong Kong Protection of Endangered Species of Animals and Plants Ordinance. Since 2019 it has been listed as CITES II: a species for which international trade should be controlled.

Shopkeepers asked in 2020 about the origins of the geckos they were selling reported China, Thailand and Vietnam, responses borne out by genetic analysis. They said that Indonesian geckos had been but were no longer imported. 88% of shopkeepers believed Tokay geckos to be farmed.

I realised as I read the paper that there must be historical information on where the live Tokay Geckos intended for human consumption in Hong Kong had been collected in the past. When we arrived in Hong Kong in November 1965, K.W. Chiu was working on the skin of Tokay Geckos. He was working for a PhD with Paul Maderson who had started the work in Hong Kong in 1962 but who had moved to the USA earlier in 1965. The geckos, intended for human consumption, were bought from the local snake shop. Suddenly the supply ceased, possibly because of disruption caused by Mao’s ‘Cultural Revolution’ and Chiu needed more in order to complete his work. A letter was sent to the British Embassy in Peking (now pinyinised to Beijing) asking the scientific attaché to help since it was known the animals came from southern China. I cannot remember what happened next but more geckos were secured.

I thought I would see if I could find any mention of where in China the 1960s geckos came from by searching some of the Maderson & Chiu papers. I quickly found the answer: Kwangsi now known as Guanxi Province. Whether Guanxi was then, and is now, the area of origin of all the geckos or whether it was a gathering point of geckos traded from neighbouring Vietnam and nearby north-east Thailand and Laos I do not know but in the genetic analysis these area loomed large and it would seem to me the trade in Hong Kong still relies on traders and trade routes that have endured at least since the end of the Second World War.

However, there is a complication. The new paper treats two morphs (black-spotted and red-spotted) as geographical distinct subspecies (G. gecko reevesii and G. g. gecko). By contrast, the IUCN Red List follows a 2011 paper in assigning the two forms species status and thereby resurrecting Reeves’s Tokay Gecko, G reevesii named by John Edward Gray of the British Museum in 1831. Therefore it is difficult to knowing what occurs where, Guanxi for example, when reading different accounts. However, the IUCN report does state that most records of G. gecko from Nanning, Guanxi are thought to be of animals that had escaped from farms. If—as seems likely—there are gecko farms in Guanxi then there may be all sorts of genetic admixture, just as in the case of Chinese giant salamanders.

The new paper discusses the effect of trade on populations in the countries of origin of Tokay Geckos and on the effect on accidental or intentional releases of live animals in the regions to which they are imported. The authors also consider the origins of the Tokay Gecko in Hong Kong—a topic to which I shall return.

Dried Tokay Geckos in a Chinese medicine shop in Hong Kong
This photoraph taken in 2010  by istolethetv
<https://creativecommons.org/licenses/by/2.0>, via Wikimedia Commons)
is interesting in that it shows the red-spotted form, less common
than the black-spotted form in the Dufour et al. survey.

Dufour PC, Miot EF, So TC, Tang SL, Jones EE, Kong TC, Yuan FL, Sung Y-H, Dingle C, Bonebrake TC. 2022 Home and hub: pet trade and traditional medicine impact reptile populations in source locations and destinations. Proceedings of the Royal Society B 289:20221011 doi.org/10.1098/rspb.2022.1011 

Chiu KW, Maderson PFA. 1980. Observations on the interactions between thermal conditions and skin shedding frequency in the Tokay (Gekko gecko). Journal of Herpetology 14, 245-254.

Two feral birds (Black Swan and Canada Goose) and two feral mammals in one morning

Two species of feral birds loomed large at the mouth of the Doon on 8 September just before the announcement came that the Queen had died. Both species are native to Commonwealth countires from opposite ends of the world. First, offshore was a large flock of Canada Geese (Branta canadensis). They seem to be present in much greater numbers than, say, 10 years ago. Second, feeding in the freshwater outflow were two Black Swans (Cygnus atratus). Native to Australia they are the result of escapes, like the Canada Geese, from waterfowl collections. There are now small - as yet - breeding populations in UK.

Black Swans in the mouth of the Doon, Ayrshire
8 September 2022

Part of a flock of Canada Geese off the mouth of the Doon, Ayr
8 September 2022

Further evidence of introductions during the the anthropocene as I arrived back home: Rabbits (Oryctolagus cuniculus) and a Grey Squirrel (Sciurus carolinensis) in the garden.

A reminder of how Raymond Cowles and Charles Bogert changed how the world viewed body temperature regulation in reptiles

An offprint for sale caught my eye. Shortly afterwards I had my first comfortable look at an important piece of research published in wartime USA that I had quoted in my final-year student seminar in February 1965. I say comfortable because all I had then was an incomplete pre-xerox photocopy that was difficult to read. After I read the paper and looked at the photographs, I looked up the authors both in Contributions to the History of Herpetology and on Wikipedia. Showing how appallingly bad many zoologically based articles on the latter are, the most important work of the two authors does not even get a mention.

Raymond Bridgman Cowles (1896-1975) and Charles Michill Bogert (1908-1992) found from their studies of desert lizards that the poikilothermic animals, usually described as ‘cold-blooded’, are  not simply at the same temperature as their surroundings. Instead, by day they use behavioural mechanisms to achieve and maintain a preferred body temperature. When cool they move into, and orientate their bodies towards, the sun. When too hot, they move into shade and/or into burrows. Later, the preferred body temperature of a particular species was shown by Paul Licht to correspond to the optimum temperature for the activity of key enzymes in the tissues.

Cowles and Bogert’s research was enormously influential not only in changing the prevailing views of how poikilothermic animals work but also in opening up a new field of physiological ecology. Indeed Cowles changed the nomenclature.  Poikilothermic (i.e. variable body temperature) was taken to imply that body temperature was not regulated while homoiothermic was the term used for the internally regulated constant body temperature of birds and mammals. He argued that the term ‘ectothermic’ for animals such as reptiles that rely on external sources of heat to maintain their body temperature was more appropriate, with ‘endothermic’ for mammals and birds which generate their own body heat.

Raymond B Cowles
from Contributions to Herpetology
see below

Cowles started the whole thing off. He was born to an American missionary family in South Africa. There, as a boy, he developed an interest in birds and reptiles and back in the USA his PhD thesis was on the life history of the Nile Monitor. Appointed to the University of California at Los Angeles (UCLA) in 1929 he became interested in the extinction of the dinosaurs. He developed the unconventional idea, after seeing the work of two friends on the importance of body temperature on the activity of reptiles at night and on the heat tolerance of lizards and snakes, that dinosaurs were wiped out not by a fall in temperature at the end of the Mesozoic but by a rise. He realised that he should investigate thermoregulation in reptiles both in the field and in the laboratory and to that end established a field station near Indian Wells in 1939.

As well as his work in physiological ecology, Cowles was horrified by the effects of over-population and campaigned vigorously. He saw the effects when he returned in the 1950s to the valley in South Arica where he had lived as a boy. The land was over-grazed, barren and ripe for erosion and flooding. I find astonishing the fact that his trenchant views on the growth of the human population caused him to become unpopular—even a pariah—in his own institution and among the general American public.

Cowles remained at UCLA from 1939 until his retirement in 1962; thereafter he worked at the University of California in Santa Barbara.

Charles M Bogert
from Contributions to Herpetology
see below

Charles Bogert was a student of Cowles at UCLA in the mid-1930s, and like Cowles had wide interests in natural history. By the time he was collaborating with Cowles on thermoregulation in reptiles he was at the American Museum of Natural History in New York. Unable to afford progression to a PhD at UCLA, he had been appointed as assistant to Gladwyn Kingsley Noble in 1936. On Noble’s death at the age of 46 in 1940, Bogert was appointed to head the herpetology department. There he continued the classical ‘museum style’ systematic studies but continued with laboratory and field work in a number of areas, most notably on frog vocalisations and their biological significance. In 1966 UCLA awarded him an honorary LLD. He took early retirement in 1968, continuing to write but a series of strokes from 1988 and severe arthritis debilitated and depressed him. He killed himself in 1992 at home in Santa Fe, New Mexico.

It has been a pleasure reading Cowles and Bogert’s seminal paper which they described as a ‘preliminary study’. It described the first four years of their work  in the Coachella Valley which they described as ‘a desert of extreme heat and aridity, characterized by sporadically abundant annual, and scanty perennial vegetation’. The style of the publication, which appeared in the Bulletin of the American Museum of Natural History, is more discursive than that of a scientific paper with photographs of the terrain, experimental housing and some of the animals they studied.

Much as been added over the 80 years since Cowles and Bogert were busy in the Coachella Valley but some fundamental questions still remain. For example, although some ectotherms may achieve their preferred body temperature for part of the day, they are still poikilothermic—their body temperature does vary over the course of a day. Since temperature affects all chemical processes, then there are implications for such physiological mechanisms as the internal biological clock, for processes like memory and the action of hormones. In addition, questions of ectothermy and endothermy in dinosaurs—the problem that first attracted Cowles to the thermal physiology of desert reptiles—are still a hot topic.

Cowles RB, Bogert CM. 1944. A preliminary study of the thermal requirements of desert reptiles. Bulletin of the American Museum of Natural History 83, 261-296.

Anon. 2014. Cowles, Raymond B (1896-1975). In Contributions to the History of Herpetology (Volume 1, revised and expanded), Edited by Kraig Adler, pp 116-117. Society for the Study of Reptiles and Amphibians.

Anon. 2007. Bogert, Charles M (1908-1992). In Contributions to the History of Herpetology (Volume 2), Edited by Kraig Adler, pp 178-180. Society for the Study of Reptiles and Amphibians.

Turner JS. 1984. Raymond B Cowles and the biology of temperature in reptiles. Journal of Herpetology 18, 421-436.

Archaeology and herpetology: two species of terrapin at Butrint, Albania

We have been to the archaeological site of Butrint—a UNESCO World Heritage Centre at the southern tip of Albania—twice, in 2010 and 2017. Fascinating as this ancient Greek, Roman, Byzantine and then Venetian city is, there are other delights. The flooded basements of the ruined buildings that were abandoned in the late Middle Ages are home to two of the three species of terrapin in Europe, the European Pond Terrapin (Emys orbicularis) and the Balkan Terrapin (Mauremys rivulata). Judging from the numbers of animals present they seem to be thriving. In 2010 most were in the water but in April 2017 when it was pleasantly sunny in the morning, virtually all were basking.

This is the video of the terrapins I made in 2017:

I have followed the nomenclature in the field guide* on the Balkan Terrapin (or turtle to those readers in North America) both in the common name, Balkan, and scientific name, Mauremys rivulata. Those familiar with the reptiles of Europe will realise that it was known more widely as a subspecies the Caspian Terrapin, Mauremys caspica rivulata but about twenty years ago that species was split. Indeed, the IUCN Red List still has its common name as Western Caspian Turtle.

*Speybroek J, Beukema W, Bok B, Van Der Voort J.2016. Field Guide to the Amphibians and Reptiles of Britain and Europe. London: Bloomsbury.

Wildebeest in the Masai Mara

White-bearded Wildebeest (Connochaetes taurinus) moving across the Masai Mara of Kenya in September 1991.

The early demise of the great physiologist, Ernest Starling: More light on the circumstances of his death on board ship off Jamaica in 1927?

I have been doing some catching up reading. This time it was a book from 2005 on the embodiment of physiological discoveries in the early decades of the 20th century: A Life of Ernest Starling by John Henderson. Starling’s major discoveries are very well explained by Henderson and put into the context of knowledge at the time and of how important they have continued to be. Starling’s made three major advances—the discovery with Bayliss of secretin, the first hormone; his eponymous ‘Law of the Heart’; how blood capillaries work; sorting out the importance of filtration and secretion in the kidney.

A great deal has been written about Starling. But, as pointed out by Henderson, information on the cause and manner of his death at the age of 61 is scant. In brief, The Times of Wednesday 4 May 1927 carried the news:

Our Kingston, Jamaica, Correspondent telegraphs that Professor Ernest Starling, the eminent physiologist, died on board a steamer as it was entering Kingston Harbour on Monday. He was travelling for the benefit of his health. He was buried at Kingston yesterday.

That paragraph was a shortened version of the article published in Jamaica’a Daily Gleaner also on 4 May. Given that in summer time, Jamaica is 6 hours behind London time, the article in The Times may have been printed first. The Daily Gleaner reported that Starling had died ‘on board the Elders and Fyffe’s steamer Ariguani on Monday morning [2 May] shortly before the vessel reached Port Royal’. Starling’s body was taken to the mortuary at St Joseph’s Sanatorium while the United Fruit Company (agents for and owners of Elders and Fyffes and after whom the term ‘banana republic’ was coined) cabled the Starling family for instructions. Things moved at speed. The burial was on 3 May at St Andrew’s Parish Church at Half Way Tree, Kingston, in a torrential downpour. The governor’s ADC and a dozen doctors attended.

One of the mourners was Dr I.W. McLean. He was the ‘informant’ of death to the registrar in Kingston. He actually signed the death certificate two days after the burial. He was a medical officer for the United Fruit Company, giving the address of the company in Kingston. His qualifications were given as ‘MD, Maryland, USA’.

Isaac William McLean's passport photograph 1917

I have now found that the doctor was Isaac William McLean PharmD, MD. He was born in Charlotte, North Carolina, on 18 November 1880. He died there on 19 September 1953. He graduated, as shown on the death certificate, from the University of Maryland. Records show McLean worked for the US government as a physician in the Panama Canal Zone in 1908-1912. In an undated company magazine he is then found at the United Fruit Company’s hospital and health centre in Bocas del Toro, Panama, where he was described as universally popular. By the 1920s (by which time disease had devastated the banana plants in Bocas del Toro) he can be traced to ships leaving Jamaica, for leave in the USA.

McLean stated that the causes of death were ‘Asthenia’ (i.e. tiredness) ‘1 year’ and ‘Heart Failure (15 minutes)’. Starling had complained of tiredness for some time and, as Henderson suggests, McLean may have gleaned that information from those on board the ship. Henderson is fairly scathing about ‘heart failure’ it merely indicating that the heart stopped. But since there was no autopsy there was just a lack of evidence. There has been speculation that he died as a result of secondary tumours from a mass removed along with a longitudinal part of the colon seven years earlier but there is no evidence that his death was related. His friends and colleagues had noted that he easily became fatigued after long experiments. The photographs of him in the 1926 show a figure who looks much older than 60.

The ’15 minutes’ of heart failure suggests that Starling was not simply found dead in his cabin (as has sometimes been supposed) and that passengers as well as crew may have been aware as to what happened, a point I will return to.

The recorded date of Starling’s death is a catalogue of errors and misunderstandings. His gravestone in Kingston has 30 April while the website ‘Find a Grave’ has a short biography and a date of 22 May.

Henderson noted that it was curious no passengers from the ship went to the funeral. However, newspaper reports show the ship had already sailed; having loaded its cargo the Ariguani left Kingston the day it arrived, 2 May. Also, as I note below, only four passengers who had travelled with Starling across the Atlantic left the ship at Kingston, and they may have done so without knowing of the death or the arrangements for the funeral.

The Ariguani, which entered service in 1926, was one of the famous ‘banana boats’ that carried passengers (1st class only) and cargo to and from ports in the Caribbean. Starling decided that a voyage in the warmth would restore him. He booked a 34-day round-trip from Avonmouth docks near Bristol; on 11 April he was seen off by his son. Henderson also thought it odd that the available evidence suggested he was travelling alone since throughout his life he was ‘extraordinarily fond of human company’ and continued: ‘Unfortunately no passenger list has survived for the Elders and Fyffes records in London were destroyed by bombs in the 1939-45 war’. However, the passenger lists prepared for port authorities in Avonmouth of those outbound on 11 April and those inbound on 15 May have survived and are available on the usual genealogical research websites. We can therefore address Henderson’s question as well as to partially answer another: Was medical assistance available on the morning of 2 May when the Ariguani was entering Kingston harbour?

I am showing the names of passengers in the hope that the some of them knew the circumstances of Starling’s death and either wrote of it or told relations the story. For those who think it unlikely that fresh information may be gathered by this means, I have been pleasantly surprised by the number of people who have contacted me as a result of a long gone relation being mentioned in my articles.

The Ariguani’s ports of call were: Bridgetown, Barbados; Port of Spain, Trinidad; Puerto Limon, Costa Rica; Cristobal, Panama Canal Zone; Kingston, Jamaica—and then back to Avonmouth.

Only 20 passengers embarked at Avonmouth on 11 April 1927 and only 5 of those would have been on board when Starling died. Only one of those five was booked, like Starling, to return on the Ariguani to Avonmouth. She was Violet Maud Abbott (aka Maud Vilet Abbott), aged 37, of Beacon Hill, London. There is no indication she knew Starling and is to be found on ships’ passenger lists travelling the world until her death in 1973. In later life she is described as ‘musical director’.

In order of the ports reached before the ship reached Jamaica, this is the passenger list for the Atlantic crossing and Starling’s companions for breakfast, lunch and dinner. For each port are listed those joining the ship who would have been on board when Starling died (all the latter are marked *). Possibly absent are any passengers who travelled only between the ports of call before and including Kingston, Jamaica:

Disembarking at Bridgetown, Barbados:

Harder, Albert Reginald. 46. Birmingham. Traveller

Disembarking at Port of Spain, Trinidad:

Ronniti, Cesare. 36. Italian. Surgeon. Resident of British Guiana

Bennett, Henry Arthur. 23. Monmouthshire. Accountant

Hay, Cecilia Elizabeth Campbell. 27. London. Occupation: none

Macdonell, Sir Phillip James. 54. London. Chief Justice of Trinidad

Macdonell, Lady Alexandrina Sutherland Campbell. 54. wife of above

Stewart, Robert Strother. 48. London. Barrister

Storey, Alan. 30. London. Merchant

Tocker, George. 27. Aberdeen. Engineer

Disembarking at Puerto Limon, Costa Rica:

Pyrenes, Jorge. 21. Costa Rican. Student.

de Padilla, Darine, 23. Costa Rican. Painter.

Disembarking at Cristobal, Panama Canal Zone:

Levy, Edmund Lewis. 56. Merchant. Resident of Salvador

Levy, Sara Lopez Loncel. 45. Wife of above

Collins, Charles George. Penzance. 35. Cable Operator

Travelling to Kingston, Jamaica:

*Abbott, Violet Maud, 37. London. Occupation: none (see above)

*Pontefiore, Harry John. 58. London. Stockbroker

*Pontefiore, Harriet, 47. Wife of above

*Orme, Christopher Guy. 68. Hampshire. Occupation: none

*Orme, Robert William Martin. 18. Hampshire. Occupation: none

Starling, Ernest Henry. 61. London. Professor M.D.

Embarking at Porto Limon, Costa Rica and travelling to Avonmouth:

*Aguilar, Enrique. 18. Student. Costa Rican

*Aurbein, Franz. 30. Merchant. German

*Bogaerts, Celina. 70. Belgian

*Calve Brenes, Virgilio. 37. Tailor. Costa Rican

*Cercelle, Marguerite. 51. Costa Rican. Hotel Business

*Crespe, Santiago. 37. Spanish. Draper

*Crespe, Epifanie. 24. Spanish. Draper

*Dubois, Georgette. 28. French. Occupation: none

*Pinto de Flores. Berta. 27. Costa Rican. Occupation: none

*Prege, Christian K. 33. German. Export Business

*Murtinhe, Adito. 38. Brazilian. Occupation: none

*Murtinhe, Zulay. 16. Brazilian. Occupation: none

*Pinto, Edgar. 18. Costa Rican. Student

*Tovar, Louisa. 45. Belgian. Milliner

*Vedeux, Kurt. 33. German. Purser

Embarking at Cristobal, Panama Canal Zone and travelling to Avonmouth:

*Astley, Betty, 22. Norfolk Occupation: none

*Carr, Stephen D. 21. Carlisle. Commercial Traveller

*Chandler, Irene E. 30 - See Below

*Dawson, Chas W. 25. Cork. Electrician

*Dawson, Frances B. 28. Cork. Housewife

*Garfinkel, Max. 27. [occupation illegible]

*Harris, Elizabeth. 28. Nurse - with the DeLeons of Hampstead

*DeLeon, Eda C. 23. Occupation: none

*DeLeon, May G. 2

*DeLeon, Michael C. 9 months

*Wilson, Nial. 20. Radio Operator

*Nagi, Max. Distressed British Seaman for Repatriation (3rd Class Travel)

*Craine, J. Distressed British Seaman for Repatriation (3rd Class Travel)

*Fitzgerald, E.T. Distressed British Seaman for Repatriation (3rd Class Travel)

Was there a doctor on board? S.S. Ariguani would have had a doctor on board as a member of the crew. Regulations then—and since—specified that a doctor should be carried for a ship carrying more than 100 passengers and crew. Did he attend Starling in that '15 minutes’ of heart failure, and why did he (in all likelihood a he) not sign the death certificate? Or was it more convenient to hand the job to the agents ashore in order not to delay the ship’s departure? Members of the crew are not identified in the statutory passenger lists and there appear to be no record of ships’ crews available (they may be in the National Archives), other than that the captain was John Harrie Howard Scudamore* DSC, RD, RNR. There was also a doctor amongst the passengers who joined the ship at Cristobal. She was Irene Elizabeth Chandler, aged 30 who qualified with the conjoint diploma in 1925. From her address (Frognal Dene, Hampstead. London) I identified her as the daughter of Pretor Whitty Chandler (1858-1941), a solicitor and Master of the Supreme Court. She married Nicholas Dunscombe Dunscombe (1898-1971) in 1937; he was a doctor and barrister working as a Medical Officer of Health in Gloucestershire in the 1940s. The Medical Register for 1940 shows that Irene was awarded the Diploma in Tropical Medicine in 1929 which could well explain her presence on board the Ariguani in 1927; she is shown as having worked later in eye hospitals in London. Irene Dunscombe (1806-1981) died in South Africa. Did she know anything of the circumstances of the death of Starling?

Finally, things not only moved quickly in Jamaica. The cable would have been received by Starling’s family in London late in the day of 2 May. On 5 May an announcement of a memorial service on Friday 6 May appeared in The Times. Members of the family, friends and colleagues, including the big names in British physiology were at St James’s Church, Piccadilly for what was, in effect, a funeral service. But the physiologists did not stop there. The 14 May edition of the British Medical Journal had an eight-page obituary section devoted entirely to Starling.

*John Harrie Howard Scudamore DSC, RD, RNR also died on the Ariguani, on 29 December 1935. He was buried at sea. He was born in Plymouth on 18 May 1977. He escorted convoys in converted merchant ships against U Boats in First World War. Shortly before his death he effected the rescue of 490 passengers and crew from the S.S. Rotterdam which ran aground off Jamaica in September 1935. All were taken on board the Ariguani and landed at Kingston the next morning.

Henderson J, 2005. A Life of Ernest Starling. New York: Oxford University Press.

Walking amongst monkeys and Theodore Roosevelt’s policy: ‘speak softly and carry a big stick’

With foreign affairs dominating the news for the first half of the year, now interrupted by incessant coverage of the self-inflicted local difficulties facing the UK government, the doctrine of ‘Speak softly and carry a big stick’ had many an airing. That has always been my motto when walking past groups of habituated or feral monkeys which have the reputation of sometimes attacking passers by or stealing food from their bags or hands. I have wondered in the past if the dictum arose in a part of the world that had to deal with monkeys as well as with evil individuals of another primate species.

When I looked it up, I found that its noted adopter and promoter, Theodore Roosevelt, President of the USA from 1901 until 1909 as well a keen and very well-read amateur zoologist and natural historian, had indeed identified the proverb as having originated in West Africa: 

I have always been fond of the West African proverb: "Speak softly and carry a big stick; you will go far."

The last time I had the opportunity of Roosevelt’s dictum was walking in the New Territories of Hong Kong where large bands of macaques (the origins of which I have discussed previously here) gather to await picnickers, walkers and joggers. At the start and end of a popular walk along a water catchment where monkeys tend to gather close to and on the path, there are piles of sticks gathered from the trees and shrubs, picked up and then discarded by walkers and joggers.

The following short video is of a large troupe of monkeys—around 50 individuals—on a popular path along a water catchment near the Jubilee Reservoir in the New Territories of Hong Kong in December 2017. When we first saw them they were descending a steep hill, covered in concrete at lower levels, to reach the path. They had to paddle through the trickle of water in the catchment and quite clearly did not like getting their feet wet. Later, they lined the path for about 100 yards. I had the ideal implement for the day—a walking pole was my big stick.

Enthusiasm is not enough. Who said that about research?

I never met Sir John Gaddum. He died in 1965, shortly after ill-health forced his retirement, three years before my arrival at the Institute of Animal Physiology at Babraham; Richard Keynes had succeeded him as Director. Many if the pharmacologists Gaddum had appointed were still around and stories of his seven years at the Institute, as well as earlier in Edinburgh, abounded. Most of them concerned his indecisiveness, especially in approving minor expenditure during a time of relative plenty. I read more about Gaddum a couple of years ago while writing a biographical memoir on Len Goodwin. Gaddum was a customer (‘the untidy Dr Gaddum’) at the shoe shop Len’s father managed and there were interactions of the two pharmacologists later in life. William Feldberg (1900-1993) wrote Gaddum’s biographical memoir for the Royal Society; he included 26 notable quotations from Gaddum’s works. Many are concerned with pharmacology and its ramifications but three are of particular note and I wish I had seen them sooner when I was in a position to deploy them with effect.

On research: Enthusiasm is not enough.

It is usually a waste of time to acquire a new research tool and then look around for problems to which it might be applied. 


It will probably always be more important to try a thing out than to argue about it.

Impossible as it is to offer advice to anybody setting out in science in the 21st century by those who operated and survived in the 20th, these three Gaddum quotations are tenets that are timeless.

from Report for 1964-1965, Institute of
Animal Physiology, Babraham, Cambridge
London: Agricultural Research Council

Feldberg W.S. 1967. John Henry Gaddum 1900-1965. Biographical Memoirs of Fellows of the Royal Society 13, 57-77.

Who was the photographer of British Amphibians and Reptiles, Walter S. Pitt?…continued

Walter S. Pitt’s photographs were used to illustrate Malcolm Smith’s seminal book, The British Amphibians and Reptiles published by Collins in the New Naturalist series in 1951. Pitt’s photographs also appeared in magazines on aquaria and there is mention that he was the recipient of one of first four Amazon sword plants to be imported into Britain. He was a member of the Zoological Photographic Club which led me to suspect he might have been a gifted amateur photographer. But I could found nothing further. I asked if anybody had information in a post on here on 22 February 2018. A couple of days ago the President of Bath Photographic Society contacted me to say our W.S. Pitt was unlikely to be the W.S. Pitt—the only possibility I could find—who was a member of that Society in 1888. Today, by chance while looking for somebody else, I found that Walter S. Pitt was a member of the Avicultural Society and that his address was listed. A search in ancestry.com soon showed who he was.

Walter Sydney Pitt was born on 19 November 1900 at 15 Portland Place London. His father was a physician, a consultant at Guy’s Hospital. Walter Pitt became a solicitor in the city (Pontifex, Pitt and Langham of Holborn Circus). He lived in Surrey at ‘Wildwood’, Silverdale Avenue, Walton-on-Thames. He died on 21 February 1983. It appears that he and his wife, Muriel Evelyn Gillard, had no children.

Confirmation that this was the Walter Pitt I was looking for has also appeared since I did the searches in 2018. On eBay at present is a clipping from a 1946 issue Rural Affairs Magazine. The caption to the photograph shows that Pitt won second prize (and £2) in a photographic competition for a photograph of a Greater Black-backed Gull alighting on a rock. From the caption the details of the camera he was using are given. It is described as an Exacta 2¼ x 2¼ fitted with a 8¼ inch Cooke Aviar lens. The film was Ilford HP3  and the exposure 1/1000th second at f/4.5*.

Having identified Walter Sydney Pitt as a gifted amateur photographer, aquarist and aviculturist, the question now is, do his original photographs which illustrated so famous a book, survive anywhere?

Finally, some of the colour plates by Pitt from Malcolm Smith’s book. The newts were photographed in an aquarium, another photographic skill acquired by Walter Pitt.

*The camera is interesting. The negative size shows that is must have been an Exakta 6x6, a camera that only became available in Germany a week or so before the outbreak of war. My guess is it came to U.K. with a serviceman some time in 1945 and that Pitt was trying it out. The lens was a famous Cooke Aviar 210 mm lens made for large format cameras by Taylor, Taylor & Hobson in Leicester. The Exakta’s roll film would have captured just the centre of the image projected by the lens, thereby increasing its apparent focal length. Some special fitting must have been made to adapt the lens to the lens mount of the Exakta. At the time of Pitt’s photograph, the rating of Ilford HP3 film was the equivalent of approximately 250 ASA. In order to use the shutter speed he did (to capture the bird in flight) it must have been a bright if not fully sunlit day when he encountered the gull.