Diane McTurk (1932-2016): The ‘Otter Lady’ of Guyana

Diane McTurk with two of her
otters. From Wilderness Explorers,
the ground agents for many
wildlife tours in Guyana

The death in Guyana of Diane McTurk in December 2016 went unreported in Britain. Lack of coverage was surprising in view of her family’s appearance in two works by national treasures: Gerald Durrell’s Three Singles to Adventure published in 1954 and David Attenborough’s BBC Zoo Quest to Guiana in 1955 with the book published in 1956. By contrast, in Guyana and the Caribbean her death was reported widely.

In her later years Diane (pronounced Dee-Ann) became well known for establishing in 1983 a tourism business at the family’s Karanambu ranch in the Rupununi grasslands of Guyana and for rearing rescued orphaned Giant Otters (Pteronura brasiliensis). Not only did she learn how to rear them but successfully reintroduced many of them to the wild. When we were at Karanambu in 2006 Diane told us that the otters had been—and to a some extent still were being—hunted, the orphaned young being left to die. She reared over forty with, fortunately, the supply eventually drying up. The hand-reared otters became incredibly tame, following Diane or her helpers down to the river for a swim, a play with the children watching their mothers washing clothes and to fish. The otters got through enormous quantities of fish caught each day at the river. While the otters were at the river a sharp eye was kept for the pack of wild otters. They are fiercely territorial and would have killed the hand-reared interlopers. A call from Diane soon had them back to heel and they were lead back to the lodge where they had a cosy box to sleep in.

Diane McTurk had an interesting life flitting between a life in Guyana and England. Schooled as an actress but realising she did not have the talent required, she worked in public relations for the Savoy Hotel. And that explained why staying at Karanambu was such an easy experience. The brick building and thatched roof may have been basic but there was everything there that a guest might need—it was a Savoy experience combined with country house weekend dining—hundred of miles from the nearest city. A variety of public relations and business development jobs in Guyana and London followed, as for a time she flitted between the two countries. She was though firmly Guyanan having been treated, she said, by her London contemporaries as just a ‘colonial’.

Politically active—and judging by the photographs and literature in our quarters—a strong supporter of Cheddi Jagan (1918-1997) who fell foul of one of the periodic madnesses of British and US foreign policy in the 1960s, Diane also, it is reported, raised support for the defence of those accused after the Rupununi Uprising of 1969 and who were eventually acquitted.

But what links talking to Diane in the searing heat of the gravel-covered yard at Karanambu (kept clear of plants to reduce the number of biting insects near the buildings) and a former mining village twenty miles inland from where I am writing this? The first McTurk in what was then British Guiana was Michael. Sir Michael McTurk was born in New Cumnock, Ayrshire in 1785. He graduated in medicine from the University of Glasgow in 1810 to become principal medical officer in British Guiana for 34 years. He was responsible for ensuring that Georgetown, the capital and centre of the sugar industry, had a water supply both for the human population and to irrigate the sugar fields. McTurk was concerned with the plight of the slaves in the country; he was knighted 1839 for his efforts on their behalf. After the passing of the Abolition of Slavery Act 1833, he brought in a local measure ensuring the full emancipation of the slaves. He died in 1844.

The next generation of McTurks was in Liverpool with Sir Michael’s son having married into a family of shipowners. On hearing the stories of his grandfather, the grandson, another Michael, took off in one of the family ships to Georgetown. That Michael, Diane’s grandfather, became Protector of the [Amer]Indians for the whole Essequibo region and her father Edward ‘Tiny’ McTurk—known for his association with Durrell and with Attenborough—was born at Bartica. After service in the Royal Flying Corps in the First World War, ‘Tiny’ McTurk decided he needed a proper job so joined the balata industry.

I had previously only associated balata with the covering of expensive golf balls for professionals (back-spin is easier to achieve but at the expense of distance) that were used until very recently. But I now find that In the 1920s balata was booming, being used to cover the underwater cables linking countries and continents, and it came from a tree common in Guyana and the West Indies, Manilkara bidentata. The latex-like material was tapped, like rubber, by cutting gashes in the bark and then left to coagulate in trays. Balata is tough, inelastic and water resistant but can be softened by heating.

So balata was big business and McTurk was given the job of finding somewhere suitable as a collection station. Karanambu did not flood in the wet season and McTurk established the balata station and cattle ranching there in 1927. Diane returned to Karanambu in 1977 and saw that diversification into tourism at a time of dwindling income from traditional activities was essential.

One of our fellow guests remarked as we left Karanambu in its ancient Land Rover in a remark that could have applied to Diane as to the vehicle: ‘They don’t build ‘em like that any more’.

Here is a very short extract from a video I took of Diane with one of her otters in March 2006:

You can see more on the unupdated website of the Karanambu Trust, set up by the McTurk family in 1997, with more information here, here, here and here.

NOTE ADDED 23 July 2021

Emmanuelle Wiecha, a documentary filmmaker based in Montreal tells me that the final Giant Otter Diane McTurk rescued and named 'Buddy' was transferred to the Jacksonville Zoo in Florida and recently sired two litters.

Alpha Helix: a floating laboratory for comparative physiology in the 1960s and 70s. What happened to her?

Anybody reading papers on how animals work from the 1960s and 70s cannot fail to notice that much of the work on certain topics, Bill Dunson’s research on sea snakes for example, was done on board the Research Vessel Alpha Helix in what seemed a series of idyllic expedition cruises in biologically interesting parts of the world. Later, there appeared to be no reference to Alpha Helix. What happened to her?

R/V Alpha Helix

Before looking at what happened after these voyages, it is worth considering her history. The driving force behind her funding, construction and operation was Per (‘Pete’) Scholander (1905-1980), the famous physiologist who worked on survival in the cold and at sea, as well as in other fields like plant physiology. Scholander was then at the Scripps Institution of Oceanography. In 1962 he succeeded in getting funding from the National Science Foundation of the USA (NSF). I am not sure whether the vessel was classified as a ship or a boat but although owned by the Scripps, it was funded throughout by the NSF and operated as a national asset.

The vessel was named Alpha Helix after the structure of proteins discovered by Linus Pauling. I cannot see why because the work done on board bore no relation to the chemical work of Pauling nor was it intended to. Reading between the lines I suspect some political motive.

In Scholander’s biographical memoir, Knut Schmidt-Nielsen wrote:

…in 1966 the ship started on its first cruise, the Billabong Expedition to the Great Barrier Reef. A National Advisory Board, initially chaired by Baird Hastings, evaluated applications for use of the ship. Over the fourteen years the ship remained in service as a floating physiological laboratory, expeditions went to Australia, the South Seas, the Amazon, the Antarctic, the Galapagos, the Bering Sea, and other sites. Several hundred scientists from all over the world have participated actively in these expeditions, and the records of Scripps Institution show that the work on the Alpha Helix has resulted in a total of 547 publications in recognized scientific journals—an impressive record for the relatively modest funds invested in the Alpha Helix.

Details of the various cruises and of the structure of the organisation set up to sort out who was invited and that to run the vessel can be found online (see below). Initially the participants were mainly US based (NSF grantees had priority) but the foreign scientist were on board for many of the cruises. In 1967 Dennis Bellamy (then in Sheffield and later in Cardiff) was one:

…I was asked to be the lead biochemist on Knut Schmidt Nielsen’s Alpha Helix expedition to the Amazon Basin.

So why was the whole concept scrapped in 1980? It seems that interest in using Alpha Helix had waned by the late 1970s—crazy as that sounds now. NSF support was withdrawn and the vessel was sold to the University of Alaska where it was used mainly for oceanographic work. After being in dry dock in 2004-2007, she was sold to a private company and then in 2014 to the Autonomous University of Mexico and based at the port of Ensenada. Alpha Helix is now an oceanographic research vessel. Indeed her whereabout can be traced on Marine Traffic under the name Buque Oceanografico Alpha Heli. This is where she was on 5 May 2018:

From Marine Traffic

Information on the history of Alpha Helix can be found here, here, here and here.

Schmidt-Nielsen K. 1987. Per Scholander 1905-1980. Biographical Memoir, National Academy of Sciences 56, 387-412.

An exotic day out for Sheffield zoology students - 1930s style

Also in the 1930s the first university students were admitted in groups, with a lecturer, at a reduced rate. At this time, for instance, the Sheffield University Zoology Department annually hired a passenger coach, which was slipped from the London express to deliver its complement of students for a day at the newly opened Whipsnade Zoo*.

The method of delivery of the students is interesting since it illustrates a long-gone railway operation. A slip coach (and sometimes coaches) was at the rear of the train. Some distance before the station a guard in the slip coach would pull a lever to ‘slip’ the coupling with the carriage in front, apply a little braking, to make sure the rest of the train would draw ahead and then slowly brake the coach to come to a stop at the station platform, while the express train steamed to its final destination. The whole operation was a routine on some routes but there were hairy moments and a notable crash (the driver thought the slip coach had not detached and stopped to check while his rear vision was obscured by smoke; the slip had detached and so ran into the back of the stopped train). There is a video of a slip operation here. At the end of the day though the slip coach must have been attached to a local train and pulled, presumably, to a station on the line at which it could be attached to the end of the train bound for Sheffield. 

Once at Whipsnade, our predecessors as students by thirty years might have sent a postcard like these shown below by Frederick William Bond (1887-1942) who was not a professional photographer but a member of staff of the accounts department of ZSL.

*Bullough WS, Hamilton F. 1976. The role of education. In, The Zoological Society of London 1826-1876 and Beyond. Ed. Zuckerman. Symposia of the Zoological Society of London 40, 223-231.

Achatina Snail: An invasive species in Hong Kong 2. My fruitless hunt for steroid hormones

In my last post, I mentioned that the accidentally introduced giant snail, Achatina, was very common in the gardens of the University of Hong Kong. The mid-1960s were the heyday of comparative endocrinology. Having established the presence of the same or closely-related steroid hormones from the adrenal glands and reproductive organs in all the vertebrates, the question was whether these steroids were present and working as hormones in the invertebrates. With such a plentiful supply of an invertebrate, in this case a mollusc, outside the lab door, I had the animals.

A well-established method to see which steroids vertebrate endocrine glands synthesise was to incubate the tissue with a radioactively-labelled compound in the early part of the biosynthetic pathway and see what was produced. I used the paper chromatography separation systems made famous by the late Ian Elcock Bush (1928-1986) in his book, The Chromatography of Steroids, published in 1961. We had a good set up for descending paper chromatography in Hong Kong and although newer thin-layer techniques were also in use in the lab, the Bush systems still had a lot to offer, especially for a quick look-see. The chromatography tanks were in a small back room of the old medical school building and Hong Kong being Hong Kong there was no need to heat the tanks. The smell of warm solvents greeted you at the door. It was a glue-sniffer’s paradise if an out and out fire hazard. The dangers inherent of inhaling industrial quantities of benzene fumes were also in the future.

Shandon Panglas chromatography tanks
are now museum pieces or, like this one,
for sale on eBay. This one lacks the trough
at the top for the mobile phase.

To cut a long story short, I could find no known adrenal steroids in the incubation fluids from several possible organs of the snail. Although we were not that interested in reproductive steroids I had run known a number of oestrogens and androgens standards as well. Again, nothing radioactive from the incubates ran in the chromatography systems with them. There were however, very small quantities of a few compounds that just might have been steroids of some sort. However, they were beyond my capacity to determine what they were. Indeed there seemed to be no chemical techniques available at the time that would have enabled me to do so.

The pilot trial on the snail of what after all was a cast-in-the-dark fishing expedition was the end of the line for my only foray into invertebrate life. Other, more productive lines beckoned.

I have though always wondered what those radioactive spots on a paper chromatogram were. Had I thrown the baby out with the bath water? Yesterday, though, I was gratified and breathed a 52-years later sigh of relief when I read of a similar early study on a different mollusc. Alexander Scott wrote in his first review of 2013 (see below):

In some experiments within this study, the gonads and hepatopancreas of this animal [Atlantic Sea Scallop, Placopecten magellanicus] converted up to 70% of added radiolabeled Pregnenolone and 17-hydroxyprogesterone into several metabolites. However, David Idler, who was probably the greatest steroid chemist of his time…was unable to identify a single one of these (apart from a trace of androstenedione)!

Those people working on fish endocrinology, incidentally, were scared of what David Richard Idler (1923-1996), then of the Fisheries Research Board of Canada, would have to say about their work. He was certain to spot any flaws in claims of chemical identification of steroids. He blew a fair number of claims—and their authors—out of the water.

Some time after my quick look-see I saw that several papers had been published suggesting that sex steroids but not adrenal cortical had been found in molluscs and that there was some evidence for some possible involvement in hormonal signalling. However, I had seen nothing to indicate a major rôle for steroid hormones. In order to become less ignorant on this matter I discovered two reviews by Alexander Scott which gathered all the information of steroids in molluscs and subjected the evidence to a pretty devastating analysis.

Historically, there was a problem with proving that a substance found in, or synthesised by, an organism was a particular steroid. Chemical techniques were simply not then available. There was considerable dissent on what was acceptable evidence of chemical identity. Those with a background in steroid chemistry, like David Idler, had the whip hand. Some would write a paper saying that they had identified the compound produced as hormone x because they had run it in two chromatographic systems along with authentic x; others insisted on using more systems, sometimes changing chemically the structure of the putative and known steroid. For studies on steroid synthesis from radioactive precursors, there was a gold standard: recrystallisation of the separated radiolabelled product with added pure compound to constant specific radioactivity but a lot of early work stopped short of this final step and even with its use there were traps for the unwary.

Looking back at the literature on vertebrate steroid hormones an awful lot of poor work was published. However, because evidence of identity was difficult to get, there was also bad luck. I remember being told of one case where radiolabelled product and pure sample ran together in 10 chromatographic systems; in the 11th they separated showing they were not the same.

Alexander Scott, a noted fish endocrinologist, in his two reviews has looked at all the evidence of the presence, formation and possible functions of steroids in molluscs. It is an important topic because of the environmental effects of anthropogenic discharge of hormonal steroids, their metabolites and their chemical analogues. This is how he introduced his first review (references deleted):

The question as to whether vertebrate steroids act as hormones in mollusks is a very important one. Many compounds that behave like vertebrate estrogens are present in the environment (so-called ‘endocrine disrupters’), and if they act in the same way in mollusks as they do in vertebrates then there is genuine cause for concern. The first reports of the existence of vertebrate-type steroids in mollusks appeared in the 1950s. Since then, all but a handful of the 200+ scientific papers and reviews that have been published in this area have a positive message (i.e. they conclude either that mollusks contain vertebrate steroids, are able to biosynthesize them de novo, appear to contain steroid receptor-like binding activity or respond in one way or another when exposed to vertebrate steroids). The sheer ‘weight of evidence’ would seem, on the surface, to make it an ‘open and shut’ case that vertebrate steroids are an important component of molluscan endocrinology. However, if one looks beyond the headline claims (i.e. essentially what is written in the titles and abstracts of many of the papers), a different story emerges – one in which most, if not all, of the positive evidence can be seen to be rather weak (in that the data are open to alternative interpretations). This review paper deals with the strength of the evidence for the presence of steroids in mollusks, for their biosynthesis and for the presence of steroid receptors. Another review paper deals with the strength of evidence for the biological actions of vertebrate steroids on mollusks.

Indeed, in Scott’s detailed analysis the evidence is weak. Steroids found by molluscs may have been taken up and sequestered from the environment, for example. Studies of the sort I did in Hong Kong in 1965-66 if apparently positive can have alternative explanations.

Scott’s second review was concerned with the effects vertebrate sex steroids on molluscs. This is what he concluded:

In assessing the evidence as to whether vertebrate sex steroids (e.g. testosterone, estradiol, progesterone) have hormonal actions in mollusks, ca. 85% of research papers report at least one biological effect; and 18 out of 21 review papers (published between 1970 and 2012) express a positive view. However, just under half of the research studies can be rejected on the grounds that they did not actually test steroids, but compounds or mixtures that were only presumed to behave as steroids (or modulators of steroids) on the basis of their effects in vertebrates (e.g. Bisphenol-A, nonylphenol and sewage treatment effluents). Of the remaining 55 papers, some can be criticized for having no statistical analysis; some for using only a single dose of steroid; others for having irregular dose–response curves; 40 out of the 55 for not replicating the treatments; and 50 out of 55 for having no within-study repetition. Furthermore, most studies had very low effect sizes in comparison to fish-based bioassays for steroids (i.e. they had a very weak ‘signal-to- noise’ ratio). When these facts are combined with the fact that none of the studies were conducted with rigorous randomization or ‘blinding’ procedures (implying the possibility of ‘operator bias’) one must conclude that there is no indisputable bioassay evidence that vertebrate sex steroids have endocrinological or reproductive roles in mollusks. The only observation that has been independently validated is the ability of estradiol to trigger rapid (1–5 min) lysosomal membrane breakdown in hemocytes of Mytilus spp. This is a typical ‘inflammatory’ response, however, and is not proof that estradiol is a hormone – especially when taken in conjunction with the evidence (discussed in a previous review) that mollusks have neither the enzymes necessary to synthesize vertebrate steroids nor nuclear receptors with which to respond to them.

As he points out, there are still a few cases where mollusc tissues appear to have converted known vertebrate hormones into others. Such work needs to be followed up to see if the entire pathway is present or if, fortuitously, an enzyme system in molluscs is doing the job of a specific enzyme in vertebrates.

What is really lacking in all the work is the integrated experimental approach in a single species. The work I first became of that seemed to implicate steroids in the reproductive physiology of a mollusc was that by Herbert Gottfried and Ralph Dorfman in the Giant Land Slug, Ariolimax californicus and published in a series of papers in the late 1960s. They seemed to have established a steroid-synthesising pathway similar to that in vertebrates that led to the production of androgens by the ovotestis. The work also involved experimental manipulation of possible signalling pathways. Unfortunately, this work does not appear to have been followed up but there appears, prima facie, to be a good experimental animal and an interesting and possibly important phenomenon to investigate. Previously, whilst at Sheffield, Gottfried with Oskas Lusis had a paper in Nature describing sex steroid production and metabolism in the eggs of a slug.

Herbert Gottfried* (1934-1977) and Ralph Dorfman (1911-1985) were both then at the Syntex research centre in Palo Alto, California and later at Applied Steroid Laboratories also in Palo Alto. Gottfried had been at Sheffield when I was a student but I do not recall meeting him. He had also been in Hong Kong in 1965 before we arrived introducing the use of gas-liquid chromatography for separating steroids. Ralph Dorfman was famous for his work on steroid hormones—he was elected to the National Academy of Sciences in 1978—including the development of the first oral contraceptives. He was President of Syntex Research. I spoke to him several times in January 1967 at the 3rd Asia and Oceania Congress of Endocrinology in Manila (the first scientific meeting of any sort I had been to) and he invited the Brits and Hongkongers there for drinks in his hotel suite. The bottle of Dimple Haig (named for the shape of the bottle and more popular in the USA and the far east than in Britain) I remember particularly well.

So while it is possible to agree with Scott on the inadequacy of a great deal of past work on steroids in molluscs, there is still the odd twinkle of light to suggest that the role of what might in the circumstances be termed vertebrate-type sex steroids in molluscan physiology can be entirely discounted. Another case of more work—avoiding the many pitfalls highlighted by Scott—being needed.

*Herbert Gottfried was born on 16 February 1934 in Vienna, Austria. In 1947 he became a British National along with his father Isak or Isaac, then a cafe manager in Brighton.  He married Patricia Elizabeth Wall on 31 March 1967 in Monterey, California. He became a US Citizen on 21 July 1970. He died in October 1977, aged 43.

UPDATED 2 OCTOBER 2023

Scott AP. 2012. Do mollusks use vertebrate sex steroids as reproductive hormones? Part I: Critical appraisal of the evidence for the presence, biosynthesis and uptake of steroids. Steroids 77, 1450-1468.

Scott AP. 2013. Do mollusks use vertebrate sex steroids as reproductive hormones? II. Critical review of the evidence that steroids have biological effects. Steroids 78, 268-281.

Achatina Snail: An invasive species in Hong Kong 1. History

The first animals I saw on arriving in Hong Kong on a dark November night in 1965 were giant snails, Achatina or Lissachatina fulica. They were in the dank area around the entrance to the block of flats in what was then called the University Compound. Even after twenty hours (including refuelling stops at Athens, Tehran and Delhi) on a Qantas Boeing 707 and the first sight of Hong Hong harbour from the vehicular ferry, the snails registered. They were big.

by Alexander C Jenner (Wikimedia Commons)

I soon discovered that these East African snails had spread widely in the tropics and were a considerable pest to farmers and gardeners. They were first noticed in Hong Kong in 1941 shortly before the Japanese Invasion in December. Since adults were found it was estimated that introduction occurred around 1937. Laying thousands of eggs over a lifespan of 10 years it is not surprising that its population and range increased rapidly.

The photograph in Jarrett's article
Hong Kong Naturalist vol. 2, 1931

They were first noticed by Vincent Hubert Charles Jarrett (1895-1973), a journalist on the South China Morning Post who was also a naturalist. At the time of the Japanese invasion he was Assistant Editor. Gwulo has a photograph of Jarrett in the 1941 here. He wrote numerous articles, including nature notes, for the newspaper and also for the Hong Kong Naturalist. He described the spread  of Achatina through Malaya and Singapore (he was born in Malacca) where it seems to have been introduced as a food for ducks. He also noted that it had been found in China by Herklots while on a visit to Amoy (now Xiamen) in 1931. Before the snail’s arrival in Hong Kong he raised a vital question:

Latest information from Malaya is that the snails are now found in increasing numbers everywhere, and even wander over the golf courses, where the chagrin of a player whose ball is stopped by a giant mollusc in the grass may be imagined. It is not known whether the question has yet arisen about counting Achatina fulica a natural or artificial hazard. It may be hoped, as a conclusion to this note, that the question will never have to be debated on one of this Colony's courses!

After release from internment at Stanley, Jarrett travelled to Britain in October 1945. I do not know whether or not he returned to Hong Kong. In 1959 he was appointed OBE for his services as ‘Editor of Press Summary’ at the British Embassy in Oslo. He died in Norway in 1973.

Geoffrey Herklots, who remained in Hong Kong as a member of the government after internment, brought Hong Kong up to date with the Achatina pest in an article for the South China Morning Post on 22 May 1946. He reported the snail was now well established on Hong Kong Island. He proceeded to give advice on how it could be killed by gardeners. With characteristic energy he held snail hunts:

On May 19, 126 snails weighing 6-lbs. were collected at Sookunpo; on May 18 and 19 330 snails weighing 20-lbs. at the Protestant Cemetery, and on May 18, 94 weighing 4-lbs. at Government House.

He ended:

These snails are edible. There is a pile of hundreds of empty snail shells at Government House. The obvious inference is that the former Japanese Governor or his staff, regarded them as a delicacy…

It is now known that Achatina snails carry a whole host of human parasites. There will be survivors of the Japanese occupation and their families hoping that a few of the more nasty ones attached themselves to the intestinal walls of the then occupiers of Government House.

What sort of thinkers are today’s zoologists?

Bob Martin in 2013 at the Field Museum
Photograph by Yu Lao (Wikimedia Commons)

Some essays stand the test of time. One such is that written in 1976 by Bob Martin, then running the Wellcome laboratory at London Zoo and now Emeritus Curator at the Field Museum in Chicago. It was entitled A Zoologist’s View of Research on Reproduction and was based on the talk he gave at the symposium of the Zoological Society of London to celebrate its 150th anniversary. He took the opportunity to provide a wide-ranging forward look and asked how things would turn out in the future. Amongst those topics was this one:

At this point, it is relevant to consider some evidence concerning the psychological background to scientific research. In his studies of the mental aptitudes of schoolboys, Liam Hudson[*] (1966, 1968) drew a crucial distinction between "convergent" and "divergent" thinkers. The former performed well on standard IQ tests requiring specific answers to well-defined questions. The latter performed best on "open-ended" intelligence tests requiring imaginative answers to questions which did not prescribe specific answers. This distinction, which is usually one of degree rather than sharp demarcation in individual cases, is in some ways parallel to a distinction between "specialists" and "generalists". As a rule, Hudson's convergent thinkers were more likely to study science at university, while divergent thinkers were more attracted to arts subjects. It is interesting, however, that zoology seemed to attract convergent and divergent thinkers in roughly equal proportions (or people who combined aptitudes in both directions). Different interpretations may be drawn from this observation. It could be said that zoology as a subject attracted a proportion of divergent thinkers because it traditionally lacked the apparently sharp scientific rigour of the physical sciences. At the same time, it is probably true that zoology as a subject has owed many of its major advances to the flair and imagination of those who could both identify and dissect general principles despite the complexities of whole animals in their natural habitats. The question that remains to be answered is: does the future development of zoology depend upon the same aptitudes as the past? For those who view zoology as a domain to be progressively replaced by current approaches to molecular biology, the answer is probably "No". For those who see in zoology a hierarchy of general principles, only some of which may be "explained" in molecular terms, the answer is probably "Yes". Whatever the answer may be, it would seem to be true that the greatest promise for future developments in zoological research lies in a combination of "divergent" and "convergent" thinking. Technical expertise and knowledge of fundamental life-processes should ideally be matched by a comprehensive understanding of whole animals and their interactions in natural environmental systems.

Since 1976 there has been a progressive horizontal slicing of the biological sciences, rather than the vertically integrated, explanatory reductionist approach of the zoologist of the middle of the 20th Century. I warned of this problem in a commentary I wrote in 1989:

It is a strange aspect of scientists’ behaviour that they condemn work at the levels of complexity above that at which they work as out-of-date and as mere natural history. Unfortunately, the divisions and intellectual isolation that lead to these attitudes appear to be increasing as universities reorganize their administrative and teaching structures for the biological sciences. The fashion is for organismal biology to be split from the rest of the biological sciences, while cell biology and molecular biology are being isolated at the other end of the scale. Thus the new disciplines are becoming horizontal divisions corresponding to the level of biological complexity…

It is true that organismal biologists have made more and more use of molecular techniques but the intervening levels like physiology have been pulled either upwards or downwards to an extent such that in Britain at least, comparative physiology has virtually disappeared and explanatory reductionist approaches abandoned.

But what of the people, to return to Bob Martin’s question? Are those working at the molecular and cellular levels of biological organisation more likely to ‘convergent’ and those working  on whole animals in their environment ‘divergent’ thinkers? And are we getting the right people into the right jobs? The latter is an important question because what has not changed is the fact that the subject studied at school, for example biology, will bear very little resemblance to that subject at university level even allowing for the fact that different university departments have different emphases and interests.

*1933-2005. Obituary - see here.

Martin RD. 1976. A Zoologist’s View of Research on Reproduction. In, The Zoological Society of London 1826-1876 and Beyond. Ed. Zuckerman. Symposia of the Zoological Society of London 40, 283-319.

Peaker M. 1989. Commentary: Molecular endocrinology: a welcome extension to, but not a replacement for, endocrinology. Journal of Endocrinology 120, 361-362.

The Complicated History of Monkeys in Hong Kong

There are monkeys in Hong Kong. Forget Hong Kong Island for the time being—in the first part of this article I am concentrating on ‘New’ Kowloon and the New Territories. Unfortunately, the view has been promulgated by some primatologists that these monkeys have been present continuously from before the Second World War, either as original inhabitants or as the result of introduction in the early years of the 20th Century. Sadly, this story seems to have been accepted by the Hong Kong Government’s Agriculture, Fisheries and Conservation Department as the explanation, in part, for the current population of monkeys.

Mainland Hong Kong

The monkeys which at present inhabit the country parks are Rhesus Macaques (Macaca mulatta) or crosses between Rhesus and Long-tailed (also known as Crab-eating) Macaques (M. fascicularis) together with Japanese Macaques (M. fuscata). Because they have been fed by the human population (feeding is now illegal) they make a thorough nuisance of themselves by lining the paths at weekends and stealing food with menaces from passers-by. The males can also be aggressive.

Hong Kong is well within the natural range of the Rhesus Macaque and there are records from the 19th Century. However, it seems that the numbers decreased possibly because of hunting for food as the human population increased. In the early years of the 20th Century Rhesus Macaques were imported and released in the area of the newly-constructed Kowloon reservoir for a bizarre reason. There was concern that fruit from overhanging strychnine trees would fall into the water and poison the public. Because monkeys were thought to be able to eat the fruit without ill effect, some Rhesus Monkeys were introduced in order to reduce the risk of poisoning those drinking the water.

However—and this is the big however—Rhesus Monkeys—and note Rhesus—whether remaining wild or the introduced population appear to have died out completely by the late 1950s to early 1960s. They may have been present in the late 1940s in small numbers. Herklots wrote in his book The Hong Kong Countryside (completed and published after he left Hong Kong in May 1948):

The monkeys that live in the woodland near the Kowloon reservoir are not descendants of the wild stock but of monkeys released during the first world war. During the Japanese occupation of the Colony in the second world war, after the trees had been cut down, the surviving animals scattered. Since the war they have been reported from several districts in the New Territories including their old haunts…

My guess is that the Rhesus Macaques were hunted to extinction or near extinction during the Japanese occupation. Monkeys did feature in Cantonese cuisine and medicine and, moreover, the population was short of food as all accounts of life during that period testify.

The naturalists around in the mid-1960s all seemed to agree that the Rhesus Macaque was extinct in mainland Hong Kong. However, there were monkeys around Kowloon reservoir but of an entirely different species: the Long-tailed Macaque*.

There is also the possibility that Herklots’s book was misleading. When as described above, he wrote that monkeys had been reported in the New Territories after the Second World War it is possible that what had been seen in the late 1940s and early 1950s were not Rhesus Macaques but Long-tailed Macaques. 

In the mid-1960s Patricia Marshall of the Zoology Department wrote in her book, Wild Mammals of Hong Kong (published in 1967):

The group of monkeys living in the Kowloon reservoirs catchment area are long-tailed macaques which were probably released or escaped from captivity during or shortly after the 1939-1945 War. The rhesus monkey (Macaca mulatta) which was once abundant on Hong Kong island and in the New Territories now appears to have died out.

These are photographs I took in December 1967 of the Long-tailed Macaques at the well-known spot near Kowloon reservoir at the 4½ milestone on the Taipo Road:

Long-tailed Macaques near Jubilee Reservoir
on the Taipo Road. December 1967

Later, Rhesus Macaques re-appeared. In 1976, Valentine Lance, also then in Zoology, surveying  the land vertebrates of Hong Kong wrote:

On a bright note in this sad tale is the addition of an animal once lost from Hong Kong and now successfully re-established. This is the Rhesus Monkey (Macaca mulatta). In 1866 Swinhoe noted that they were common on most of the small islands near Hong Kong…and number of them lived in Tai Tam before the second World War. By 1962 they were no longer present in the Colony. Another closely related species, the Crab-eating or Longtailed Macaque (Macaca fascicularis), a more southern common Malayan monkey, successfully established a breeding colony at Kowloon Reservoir (Romer, 1966). This group originated from escaped or released pets. Today there is a flourishing colony of more than twenty individuals in the Kowloon Reservoir catchment area. The rhesus monkeys have only successfully established themselves in the same area in the past few years. Again they are escapees. The criteria for establishing residence is successful breeding. A family with a young baby was seen recently and there are believed to be more breeding pairs in the same area. The two species seem to live fairly amicably side by side in the same restricted habitat. In addition to the colony at Kowloon Reservoir isolated individuals have been seen at Pokfulam Reservoir [Hong Kong Island], Tai Tam Reservoir [Hong Kong Island], and recently at Ma On Shan.

All this seems pretty clear: the present Rhesus Monkey population appears to be derived from animals that escaped or were released in the late 1960s or early 1970s. Unfortunately, as is so often the case, other researchers have not read the relevant literature and ascribe the monkeys to descendants of the pre-Second World War population. Such errors and misinterpretations which omit mention of evidence from the 1960s are, of course, repeated and repeated not only websites, in publications and in newspaper articles but in books on primate conservation. For example, in one relatively recent book it is stated that the introduced Long-tailed Macaques interbred with the existing population of Rhesus. Photographic and observational evidence from the mid-1960s shows that not to be the case; the Long-tailed Macaques were not interbreeding at that time since they alone occupied the Kowloon Hills. Nevertheless, it is true, as David Dudgeon and Richard Corlett remarked in their excellent book on the ecology of Hong Kong, ‘we cannot be certain…that none of the current population are descended from survivors of the original Hong Kong macaques’. The negative is impossible to prove but the probability is highly weighted against a remnant population.

Since the 1970s the population of Rhesus Macaques has increased massively. By 1992 there were around 800 macaques in Hong Kong, 700 of them in the Kowloon Hills and other country parks. But they were not all Rhesus or Long-tailed. Other species kept for some reason either escaped or were released. There are records of Japanese Macaques, Tibetan Macaque (M. thibetana) and a single Pig-tailed Macaques (either M. leonina or M. nemestrina). Hybridisation between three of these species is known, with the production of fertile offspring. (There is a hybrid zone, in northern Thailand, for example, where the Rhesus and Long-tailed meet naturally.) A survey by John Fellowes for the World Wildlife Fund in 1992 found evidence of four species and possible hybrids, as far as they could be identified: Rhesus (pure) 50% of the population; Long-tailed x Rhesus or x Japanese 40%; Japanese (pure) or Japanese x Rhesus 9%; Long-tailed (pure) 1%; Tibetan (0-1%).

The number of Long-tailed Macaques was said to have declined in 1967. Reports in the newspapers blamed trappers for catching the monkeys and selling them as a delicacy for human consumption.  However, there seemed a healthy population in December 1967 when we watched them. In the 1980s in the Kowloon reservoir area, the proportion of Long-tailed in the monkey population fell from 26% to 7% between 1981 and 1987; hybrids between these two from 13 to 3 while the Japanese Macaques fell from 5% to 0.

It appears then that the Long-tailed Macaque population has virtually disappeared together with easily-recognisable hybrids. How much of the Long-tailed genome is present in the present Rhesus population seems to be unknown. Since large traps are available from the sterilisation programme aimed at keeping the population in check, molecular studies are just waiting to be done. But as Dudgeon and Corlett state: ‘It would be interesting to know if the predominance of rhesus macaques reflects a larger founder population (i.e. more releases or survivors) or superior adaptation to an environment which is within the native range of the species’.

Were the Rhesus and other species of monkey recorded escapees or were they released? Explanations I have seen include release by a travelling circus after being prevented from taking the animals to the next destination. Did some escape from food markets? Did well-intentioned individuals buy some from pet shops or markets and release them? Did pet owners release adult monkeys when they turned vicious? Did some escape from private collections?

I do get the impression that the Rhesus Macaques must have been released in substantial numbers, says tens at least, to have multiplied so rapidly and so visibly, rather than in the dribs and drabs of the odd pet escaping or being released and then finding the troupe of Long-tailed Macaques to join. I awoke the other night with a possible explanation. When we arrived in Hong Kong in November 1965, John Phillips (1933-1987, FRS 1981 and later of the University of Hull and Vice-Chancellor of Loughborough University) my PhD supervisor was Professor of Zoology. He told me to get an animal experimentation licence from the government. The government had recently introduced licensing, he said, because some commercial animal testing operations had set up in Hong Kong in order to bypass legislation in their countries of origin and the Hong Kong government wanted to make sure nothing untoward was going on. Yesterday I checked my memory and, indeed, the Animals (Control of Experiments) Ordinance was passed into law on 24 May 1963; it is still in effect. I knew nothing of these laboratories and I don’t think John did either but is it just possible that they had the favourite and cheap primate of the time, the Rhesus Monkey and that after ceasing operations considerable numbers were released into the New Territories?

I know that possible explanation is a long shot but I was intrigued by a paragraph in John Fellowes’s report from 1992. In Tai Po Kau forest reserve, the population seemed to be entirely Rhesus and he wondered if that could reflect a relict population. However, Tai Po Kau was well visited by birders and other naturalists in the 1960s who would surely have spotted a troupe  of monkeys. Is it just possible that the main release of Rhesus Macaques was at or near Tai Po Kau, and that some then made their way to the Kowloon Reservoir area where they found and interbred with the Long-tailed Macaques?

One possibility that I have never seen discussed is that the 'new' wave of Rhesus Macaques from the late 1960s/early 1970s actually arrived from a wild population across the border. By 1985, the nearest wild populations were a long way away in Guangdong or on islands but who knows about the 1960s? Looking at China then from the lookout at Lok Ma Chau, there was just farmland and woods. Could the woods have held a population of monkeys even ones that could have made their way there from Hong Kong in the early decades of the 20th Century?

In summary, the present Rhesus Macaque population of mainland Hong Kong has an interesting history and genetic origin. With molecular techniques already used on two of the species involved†. it should be possible to determine to what extent, if any, the genome of the other macaques (other than the Tibetan which did not appear to have interbred) is still present in the gene pool as well as the size of the founder populations. It is also possible that the geographical origins of the Rhesus Macaques could be determined; are they originally from China or elsewhere in the range, India, for example, which was a major exporter at that time?

Hong Kong Island

Herklots wrote:

On the Island of Hong Kong a monkey family was watched early one morning near Tai Tam reservoir in 1947, and I have had occasional accounts of monkeys having been seen on the Peak and in Deep Water Bay valley. It is possible, but not certain, that these are descendants of the original wild stock [i.e. Rhesus Macaques]…

The monkeys at Tai Tam seem to have disappeared by the early 1960s. Patricia Marshall considered the monkeys on Hong Kong Island, like the Rhesus on the mainland, to have died out. However, monkeys were seen on the Island throughout the 1960s after she wrote her book. There were newspaper reports and a recent request for information from the Facebook Group, Hong Kong in the 1960s, confirmed the presence of small numbers monkeys around various blocks of flats. Lancelot Vance also reported the sighting of individual animals at Pokfulam reservoir and at Tai Tam reservoir. In 1969 a monkey living on Bowen Road was shot by John Romer after it bit a child. There was a considerable fuss in the letters columns of the South China Morning Post. Sightings continued throught the 1970s into the 80s.

The question, as usual, is, were the sightings in the 1960s of descendants of the family group Herklots had seen or of monkeys that had escaped or had been released into the wild more recently? The question is, of course, unanswerable but there were certainly plenty of opportunity for the latter. The British army’s Lyemun barracks had a small zoo; Rhesus Macaques bred there in 1962. Other individuals had wild animal collections in the 1960s and 70s including a Dr Vio who lived on the Peak. The PG Farm, on the site of Ocean Park, a popular attraction for children in the 1960s, had Rhesus Monkeys; at least one escaped—and was recaptured. There were newspaper appeals for people not to keep monkeys as pets. It is fair bet that a few escaped from owners or were released after a first—and usually inevitable—bite.

I have not seen any recent reports of monkeys on Hong Kong Island.

Opportunities for Research

‘Natural’ experiments such as the various introductions of macaques in Hong Kong offer opportunities for research. I have outlined some of the lines which could profitably be pursued and which would throw light on the selective pressures operating to favour, perhaps, the originally native Rhesus Macaque over the closely-related Long-tailed. But as I have also pointed out, we might also get an indication of the origin or origins of the currently highly successful population of Hong Kong monkeys.

And finally…

A photograph of a Rhesus Macaque almost exactly 50 years since I photographed the Long-tailed Macaque within a few hundred metres of this one.

Rhesus Macaque near Jubilee Reservoir catchment. November 2017

*The Hong Kong Government only appears to have realised this in 1966 with the publication of Romer’s note. The Annual Report for 1965: Rhesus monkeys are still found in the vicinity of Kowloon Reservoir in the New Territories, but those which inhabited woods in the Tai Tam area of Hong Kong Island are not known to have been seen during 1965. The 1966 Report had the nearly correct story: Long-tailed or Crab-eating Macaques, a sub-species of those found in Singapore, occur in small numbers in the Kowloon reservoir area and can often be seen near the Kowloon-Taipo Road and at Pipers Hill. These are probably descendants of escaped or released individuals. The indigenous Rhesus Macaques which were present on Hong Kong Island and in the New Territories have now apparently disappeared. No source of information is given in the 1965 Report but the 1966 Report acknowledges the help of Patricia Marshall for the section on wild life.

†Bunlungsup S, Kanthaswamy S, Oldt RF, Smith DG, Houghton P, Hamada Y, Malaivijitnond S. 2017. Genetic analysis of samples from wild populations opens new perspectives on hybridization between long-tailed (Macaca fascicularis) and rhesus macaques (Macaca mulatta). American Journal of Primatology 79. doi: 10.1002/ajp.22726

Dudgeon D, Corlett R. 1994. Hills and Streams. An Ecology of Hong Kong. Hong Kong University Press.
Fellowes JR. 1992. Hong Kong Macaques. Final Report to the WWF Hong Kong Projects Committee. Hong Kong: WWF.
Herklots GAC. 1951. The Hong Kong Countryside. Hong Kong: South China Morning Post.
Hong Kong. Report for the Year 1965. 1966. Hong Kong Government Press.
Hong Kong. Report for the Year 1966. 1967. Hong Kong Government Press.
Lance VA. 1976. The land vertebrates of Hong Kong. In, The Fauna of Hong Kong, edited by B. Lofts. Hong Kong: Hong Kong Branch of the Royal Asiatic Society.
Marshall PM. 1967. Wild Mammals of Hong Kong. Hong Kong: Oxford University Press.
Romer JD. 1966. Long-tailed Macaques in Hong Kong. Memoirs of the Hong Kong Natural History Society 7, 16. I have not been able to track down an easily accessible copy of this paper; therefore I have not read it.
Southwick C, Manry D. 1987. Habitat and population changes for the Kowloon macaques. Primate Conservation 8, 48-49.

Updated 24 April 2018