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.