Friday 1 March 2024

Miss Waldron’s Red Colobus. The Lowe-Waldron specimens were NOT the first to reach London’s Natural History Museum


From McGraw 2005

Willoughby Lowe’s entry in the Eponym Dictionary of Mammals has a couple of unkind sentences:

He is also notorious for having shot eight specimens of Miss Waldron’s Red Colobus Piliocolobus badius waldroni in Ghana in 1933…The colobus was already rare and is now thought to be extinct. 

There is no evidence that in the 1930s where suitable forest remained the red colobus it was rare. Indeed in the 1970s large troupes were observed. Lowe did warn—correctly—that further loss of forest would have devastating effects on it and other species. In addition, providing series of specimens was exactly what the Museum instructed collectors to do since it was realised that it is important to study variation between individuals from the same location. Lowe’s specimens (12, not 8, are shown in the catalogue) were joined later by 26 collected by Angus Booth (1927-1958) (on whom more in a future article) who was warning even more vociferously about the dangers of deforestation which was proceeding apace and the ease of hunting the red colobus for meat.

Willoughby Lowe clearly believed that he and Fannie Waldron had discovered the monkey named for Miss Waldron. However, looking at the catalogue of the Natural History Museum that was not the case. Two had arrived more than 20 years earlier.

The job of sorting the collection of mammals brought back by Lowe and Waldron in 1934 and 1935 from the Gold Coast (Ghana) was given by Martin Hinton, who was in charge of mammals, to Robert William Hayman (1905-1985). Hayman made a mistake in the introduction to his paper. He had just Lowe on the 1933-34 expedition with Miss Waldron also present in 1934-35. That is incorrect. Waldron and Lowe travelled together on both.

Hayman named the red colobus as a new subspecies, Colobus badius waldroni. In his description he compared these specimens from Goaso with the nominate subspecies:

A red Colobus closely related to the Liberia and Sierra Leone form, Colobus badius badius Kerr, but differing in the distribution of the black in the pelage. In true badius the black extends from the front of the forehead back over head, neck, shoulders, and upper arms, all the back, and thence on to the outer side of the thighs as far as the knee. The black on the outside of the thighs extends behind to meet the white perineal area. The tail is dark reddish on the basal half, darkening to black on the apical half.

In this new Gold Coast race a series of eleven specimens (six adult male skins and skulls and one immature male, three adult female skins and skulls, and one adult female in alcohol) shows the following constant differences:—The forehead is dark red, deepening on the crown to black. The black either does not extend to the thighs at all, as in the majority, or, at most, as in one specimen goes no further than part way down the centre of the outside of the thigh, not passing behind to the back of the thigh. The tail is jet-black above and below throughout.

One of the males, represented by a skin and skull was chosen as the type (holotype) for the new subspecies: ‘Collected at Goaso, Ashanti, December 31, 1933, by  Willoughby P. Lowe’

Hayman went on to note:

The whole series exhibits little variation in colour. In one female skin the thighs have a thin median streak of blackish brown quite narrow, and the back of the thigh is broadly red, as is the whole of the thigh in the rest of the series. In the British Museum Collection is a very young skin from Bibianaha [Bibiani], Gold Coast (Spurrell,, which exhibits all the characters of this race with the sole exception of the tail, which is mainly reddish below, although completely black above from root to tip.

I have much pleasure in connecting with this remarkably handsome discovery the name of Miss Waldron, who contributed much to the success of the expedition.

The specimens collected by Spurrell in 1912 are currently listed in the catalogue as Colobus (Piliocolobus) badius waldroni.

Hayman must have realised that Spurrell’s red colobus was of the same subspecies he had described from the new series of specimens. Bibiani is less than 30 miles from Goaso.

Thus we have a situation in which the name of the original collector was overlooked as an eponym. I can see two or three reasons why one of the later specimens was selected as the type and why Miss Waldron’s name was chosen. The greater range of adult males and females brought back by Lowe and Waldron enabled a greater certainty of separation from the nominate subspecies. Secondly, I suspect museum employees were told when possible to honour Fannie Waldron the best they could. Money was very tight in the 1930s. Grateful thanks and recognition may have brought funding for further trips from Miss Waldron. She had at least part-funded the first expedition and probably wholly funded the second. A fish, a bird and a mammal were duly named. Thirdly, Spurrell (Herbert George Flaxman Spurrell, 1877-1918, on whom more in a later article) was dead.

This excellent comparison is from McGraw (2005)

In terms of taxonomy, Reginald Innes Pocock (1863-1947) changed the generic name in the paper that followed immediately on from Hayman’s in Proceedings of the Zoological Society. As can be seen it was Pocock (working on mammals in the Natural History Museum after retiring as Superintendent of London Zoo) who asked Lowe to bring back a specimen preserved in spirit from the second expedition:

On his return in 1934 from the first of his two recent expeditions to Ashanti, Mr. Willoughby Lowe informed me that he had shot female examples of the Red Colobus (Procolobus badius waldroni) showing a swelling of the external genitalia which he had never previously observed in any representatives of this genus. The swelling in question, obviously similar in its nature to that of Baboons (Papio), Mangabeys (Cercocebus), and of some species assigned to Macaca, was clearly indicated, although in a shrivelled and otherwise distorted condition in two of his dried skins. Since the phenomenon in question had apparently never been described, or even recorded, in any representative of this family of Monkeys, I begged him, before starting on his second expedition to the same district, to bring back, if possible, a specimen of this monkey preserved in alcohol or formalin. To his kind acquiescence in this request I owe the opportunity of describing and figuring not only the catamental swelling in question, but other external characters of a Red Colobus for the most part previously known only from dried skins. A comparison of the characters with those of examples of the Black and White Colobus Monkeys recorded in my paper on the external characters of the Catarrhine Monkeys (Proc. Zool. Soc. Lond, 1926, pp. 1479-1579) suggests that the two main divisions of these monkeys may be generically distinguished; and since the name Colobus belongs primarily to the "Black and White" section, typified by polykomos (=ursinus), the red section may be provisionally assigned to Procolobus, its oldest available name, with verus van Bren. as the the type species.

Rochebrune…also proposed the generic name Piliocolobus for one or two different kinds of "red " species, and Allen selected badius as the type. It remains to be seen if there are generic differences between Procolobus and Piliocolobus.

Fast forward to 2024 and the accepted generic name is Piliocolobus.

Hayman’s retention of Miss Waldron’s Red Colobus in the species now known as Piliocolobus badius, but as a new subspecies, seems entirely reasonable since there is a series of red colobus forms along the forest regions of West Africa which have become delineated by river systems and geological features. However, it was inevitable given the species concepts being employed on some groups of mammals in the latter part of the 20th century and beyond that P. badius would be split. Thus we have Piliocolobus waldroni (or infuriatingly so as to supposedly agree with the rules of nomenclature on using the correct gender of the latin genitive eponym in specific names, Piliocolobus waldronae. That stupidity seems to have been stopped but it is under that name that the species is described in the Primate volume of Handbook of the Mammals of the World published in 2013. Equally infuriating is the fact that the ‘split’ was based on the mitochondrial genome of a single individual. As an aside, I do find it amazing that in no other branch of science are decisions made by one individual or school of thought so readily accepted as gospel by others in essentially the same field. While there may be a little to and fro, by and large and for a time at least, the new designation is just accepted.

Becoming bogged down in arcane taxonomic niceties does not answer the pressing question of whether any of Miss Waldron’s Red Colobus remain alive. The devastating losses caused by destruction of its forest habitat and by hunting for human food have been such that the species has, after searches, been declared extinct but for that conclusion to be challenged after local hunters across the border in the Tanoé Swamp Forest of Ivory Coast (Côte d’Ivoire) have claimed to have seen it, to know its call, and have produced skins as evidence. However, there seems to have been no further positive news in the past 15 years. The IUCN Red List (where it is listed as a species) has it as ‘Critically Endangered (Possibly Extinct)’. Whether it is a true biological species or a form of P. badius is immaterial to the conservation message: when you lose habitats you lose their inhabitants. The whole ecosystem (for want of a better, non-teleological, term) is gone. The history of how this came about was outlined in a paper by Oates, Struhsaker & Whitesides in 1997:

Three subspecies of forest primate are known only from southwestern Ghana and parts of neighboring Côte d'Ivoire to the east. These are the white-naped mangabey (Cercocebus atys lunulatus), the Roloway guenon (Cercopithecus diana roloway), and Miss Waldron's red colobus (Procolobus badius waldroni)…The rainforest area where these endemic primates occur has undergone very rapid development since World War II. Logging activity has been more intense than in almost any other part of tropical Africa, and many people have moved into the region to cultivate the land as it has been opened up. Logging, farming and human population growth in the region have been accompanied by the increased hunting of wild mammals and larger birds for meat, much of which has been traded out of the immediate area for sale in towns.

Beolens B, Watkins M, Grayson M. 2009. The Eponym Dictionary of Mammals. Baltimore: Johns Hopkins University Press

Hayman RW 1935. On a collection of mammals from the Gold Coast. Proceedings of the Zoological Society of London 1935, 915-937.

McGraw WS. 2005. Update on the search for Miss Waldron’s Colobus monkey. Internatikonal Journal of Primatology 26, 605-619.

Oates JF, Struhsaker TT,  Whitesides GH. 1996/97. Extinction faces Ghana's Red Colobus monkey and other locally endemic subspecies. Primate Conservation (17), 138-144.

Pocock RI. 1935. The external characters of a female Red Colobus Monkey (Procolobus badius waldroni). Proceedings of the Zoological Society of London 1935, 939-944.

Tuesday 27 February 2024

A Plaintive Cuckoo devours caterpillars in Hong Kong

AJP spotted this Plaintive Cuckoo (Cacomantis merulinus) in the New Territories of Hong Kong on Christmas Eve last year—an unusual sighting at that time of year. It was devouring caterpillars of the Red-based Jezebel (Delias pasithoe), a very common butterfly during the winter months.


The Lowe-Waldron Expeditions to the Gold Coast in 1933-34 and 1934-35

One of Fannie Waldron's photographs of the expeditions

In following up my tracking down of Fannie Waldron (1876-1959) as the person after whom Miss Waldron’s Red Colobus was named, I have obtained a copy—the only one I could find on sale—of Willoughby Lowe’s final book The End of the Trail. The expeditions in 1933-34 and 1934-35 with Fannie Waldron were Lowe’s last collecting trips and his account is the final chapter of the limited edition of 400 copies published in Exeter by the printers, James Townsend & Sons, in 1947.

Frontispiece from
The End of the Trail

The strange thing about this chapter is that Lowe does not refer to Fannie Waldron; she is always ‘my companion’. Indeed, the only clue that she was there at all is in the caption to the photographs: ‘Photo by F. Waldron’. Having mentioned her by name (wrongly as Fanny) in his earlier account of the birds they collected, the omission does seem odd. Did Fannie wish to arrange anonymous and was annoyed by the earlier public acknowledgement of her presence and funding? Had there been gossip about their travelling together as, for the time, an elderly spinster and a married man?

What does emerge from Lowe’s account are some of the jobs which Fannie undertook. However, there is no indication whether she worked as a collector of specimens per se, or did any of the arduous jobs of skinning, curing, pickling and preparing the specimens for shipping with them to London. Lowe comments on the time needed to prepare a monkey skin (the collected numerous individuals of seven species Goaso) and it would appear that he alone did the preparation:

…I was fully occupied attending to the preservation of these creatures [monkeys], and no one knows the amount of labour required, until they have tried, to preserve a monkey properly. The hands and feet are particularly troublesome as every finger must be turned back to the nails, the the skin is coated with a tough muscular fat as far as the digits, every particle of which must be removed with scissors, or the specimen will be ruined.

Museum staff praised Fannie for the collection of fish (caught by the locals in a fish trap and by poisoning a stretch of river) and I suspect she took on the job of preserving specimens in spirit.

We do know from Lowe’s account some of things Fannie did: she took the photographs; she raised a young Woodford’s Owl (fate unknown) and in London (Lowe lived in Exmouth, Devon) she delivered a lungfish cocooned in dried mud to London Zoo and blood samples from what was to become her eponymous monkey and the other species to the Wellcome Bureau for Scientific Research. Those blood sample proved important in the study of yellow fever as I shall describe in a further article.

On Miss Waldron’s Red Colobus, and other monkeys,  it would seem from reading his  account that some were shot for Lowe by the local hunters, partly because his large supply of cartridges had been held up by muddle at customs and they used their own cartridges until he could repay them in kind. Lowe though noted:

Never shall I forget hunting my first specimen [of Miss Waldron’s Red Colobus], when I waded and struggled nearly up to my waist in filthy decayed vegetable matter, tormented with insects, but luckily getting what I was after.

Lowe was greatly alarmed by the rate of destruction of the forests:

But there is yet another sad and serious view of these forests and their hidden secrets, their destruction. It is obvious to every traveller that vast forests in different parts of the world have been, and still are being, destroyed, and West Africa is no exception. Man with even crude axes, aided by fire, has done more to destroy once fertile lands than most people conceive.

We have already enough desert and arid areas in the world and it is high time those in authority should exercise their persuasive powers to prevent further destruction. Probably the forest region of West Africa is already more than half destroyed. Fires rage yearly from the dry northern regions through the savannah to the edges of the forest, into which it gradually creeps. The interior of the forest is cut, burnt and cultivated, and Forestry Officers told me they are in many places fighting in their last trench. With the destruction of the trees go the fauna and flora; and a hideous country, once beautiful and yielding fruit and products for mankind, is all that remains.

The second trip, a year later, concentrated on the northern parts of the Gold Coast. Lowe explained:

Contrary to our intentions we decided to make a second journey to Ashanti, and also to visit the Northern Territories of the Gold Coast. This decision was made because, after a cursory examination of our zoological collections, it was found that there were many animals and birds not represented, and others which could not be properly determined without more material. This being the case arrangements were immediately made to have the same servants and lorry, plus a trailer, and to travel through the extensive dry northern part which we had left untouched.

Travelling by lorry was not comfortable.

West Africa was not a healthy place. Even thirty years ago with most of the benefits of modern medicine I was told by somebody who worked there as a wildlife consultant that visitors who stayed for more than short periods often became ill, not from any specific recognised disease but just generally debilitude. It would seem that was the case 60 years earlier because Lowe noted how the Gold Coast took its toll on the District Officers and other expatriates involved in administration and commerce:

I feel I cannot refrain from some remarks on the term these men have to do under such trying conditions. West Africa used to be a nine-months’ tour, then it was raised to a year, only to be altered again to 18 months, and now there is talk of making it two years!

I myself have seen a little of the world and of varying climates and, I repeat what I have stated before, that no one should do more than a year’s work in really bad climates.

Lowe does not describe the work needed to prepare specimens for the museum. For that we have to rely on the Museum’s own Handbook of Instruction for Collectors which appeared in various editions over the years. One, from 1902—shortly after the discovery of the Okapi—can be found here I had, and still may have, a copy I bought in the Museum’s shop in the late 1950s which dated  from the early decades of the century. He does, though, describe the hard and frustrating life of the collector in a tropical forest, expressing thoughts not dissimilar to those of wildlife watchers of the 21st century:

The reader may exclaim-what an interesting and exciting place for work! And yet, though these and many other animals and birds exist, a naturalist's work in dense evergreen forests can be deeply disappointing and trying. Anything may exist, but to find it, even after years of experience, is terribly difficult. Sharp eyes are required, keen ears, and the matured field-craft of a lifetime, as well as the patience of Job, for almost every day seems a failure to get what you want and to find what you know is there.

People who have never tried to make a collection of natural history objects in such a place as the Ashanti forests little realize the difficulty in a piece of unspoilt jungle. The sweat, labour, discomfort, exhaustion, and the daily disappointment of seeing an unknown creature for the fraction of a second that may not be seen again for a generation. Think what it would mean to find it again, and perhaps get familiar with its life history. No! naturalists have plenty of work ahead of them.

I have plotted the places visited in the Gold Coast (Ghana) in the two expeditions. The map shows the coverage of the country they achieved.

Willoughby Lowe and Fannie Waldron spent a few days in Accra with the Governor before their ship left port for home. While there they visited Achimota College, now the University of Ghana. It struck Lowe that ‘the buildings and general lay-out seemed excellent’ but ‘it was all several centuries ahead of the times’. But it was to Achimota College and a tragic death that the story of Miss Waldron’s Red Colobus continues. And, as a preview, incidentally it was not Lowe and Waldron who first sent this monkey to the Natural History Museum in London, as they thought to be the case.

A photograph of Lowe taken
by Fannie Waldron

Sunday 18 February 2024

Showing off in Hong Kong: a male Fork-tailed Sunbird displays his attributes

AJP spotted this Fork-tailed Sunbird (Aethopyga christinae) displaying to females and seeing off other males at Tai Po Kau last week.

It is difficult to believe that this common and widespread bird in Hong Kong was once uncommon. It was only first recorded in 1959, again at Tai Po Kau, which for years remained its stronghold. We were amazed when returning to Hong Kong on 1997 for the first time in 29 years to find a pair nesting outside our window at Robert Black College in the university compound. 

Monday 12 February 2024

Sooty-headed Bulbul in the hills of Hong Kong

AJP took this photograph of a Sooty-headed Bulbul (Pycnonotus aurigaster) in Hong Kong a couple of weeks ago. This is  the third most common bulbul in Hong Kong and noticeably much less common than when we lived there in the 1960s. I think the clue is in its habitat: ‘scrubland and thinly wooded hillsides’. The hills certainly fitted that description in the 1960s following the gathering of anything for firewood during the Japanese occupation but since then the trees have grown considerably, thus reducing the suitability of the hillsides for this species—and also cutting off former familiar views for walkers on Hong Kong island, for example. The common name that used to be used in Hong Kong was Red-vented Bulbul, a name that also applies to another species, P. cafer, largely confined to the Indian subcontinent.

Saturday 3 February 2024

What happens to moths and butterflies during very cold weather in Hong Kong?

AJP was up and about early last week after a very cold night (around 6°C at sea level). The butterflies and moths in the hills of the Tai Po Kau forest were lying on the road either recovering or dying from the overnight cold. This Tropical Swallowtail moth, Lyssa zampa, did not make it.

It can be difficult for visitors to imagine Hong Kong being cold if they are there in the summer with temperatures over 30°C and humdity at 90%.

Wednesday 31 January 2024

Leafbirds in Hong Kong

AJP spotted these leafbirds at Tai Po Kau in Hong Kong during their recent cold weather. The species now goes under the name of Greyish-crowned Leafbird (Chloropsis lazulina) but was considered a subspecies of Hardwick's Chloropsis, Leafbird or Fruitsucker, Chloropsis hardwickii melliana. They live in high forests and the cold weather may have brought them down to the hills. They appear to have been taking nectar from the flowers.

They were once considered to be very rare in Hong Kong. Herklots noted that when he wrote his book in 1953 only one had ever been seen been seen—in Lam Tsuen Valley on 27 January 1934. The species occurs in south-west China down to Vietnam. They now have uncommon resident and winter visitor in the field guide. We have never seen one in Hong Kong either when we lived there in the 1960s or on our visits since.

Tuesday 30 January 2024

Miss Waldron’s Red Colobus. Finding Miss Waldron

This article started out as a follow-up to a previous one which touched on Willoughby Prescott Lowe (1872-1949), a major animal collector for the Natural History Museum in London and others. On looking him up in the Eponym Dictionary of Mammals and in the Eponym Dictionary of Birds I found the following:

…He is also notorious for having shot eight specimens of Miss Waldron’s Red Colobus Piliocolobus badius waldroni in Ghana in 1933. Miss Waldron also worked at the museum and accompanied him on this trip. The colobus was already rare and is now thought to be extinct.

But who was Miss Waldron. What was she doing in the Gold Coast—as  Ghana then was—in the 1930s. In the 2014 edition of the Eponym Dictionary of Birds she was described as:

Miss Fanny Waldron [dates not found] was an employee of the BMNH [British Museum Natural History]. She accompanied W.P. Lowe on his expedition…in 1934-35 when she was well over 60.

Intrigued by the lack of information I then found that others on the online Bird Forum had also failed to find out anything about Miss Waldron and that in one index she has been described as Frances since, presumably, Fanny was sometimes used as an informal derivation from the diminutive ‘Fran’, as in ‘Peggy’ for ‘Margaret’. Since Lowe and senior staff of the Museum used ‘Fanny’ I can only assume the indexer had not realised that an informal name would just not have been used in a formal report or paper in the 1930s.

I am pleased to say that I have found who Miss Waldron was. Readers will soon realise that much of the information given above is wrong while little is right or even half-right.

Finding Miss Waldron

I soon found her by searching through shipping records in and There she is, along with Willoughby Lowe, on board Elder Dempster Line ships, travelling between England and Gold Coast (now Ghana). Fortunately these records give her age and address in London.

With that information I was able to widen the search and obtain a fairly comprehensive picture of Miss Waldron. She was indeed a ‘Fanny’ but not ‘Fanny’ with a ‘y’ but ‘Fannie’. Fannie was the name registered at birth and was the one she used when filling in her own forms for a ship’s manifest, for example. However, census enumerators and those writing about her, in the ornithological literature for example, have used ‘Fanny’. It would seem her own family who had to fill in some census forms for themselves also forgot and used ‘Fanny’ on occasion. But her name was simply Fannie Waldron. Even the record of probate after her death notes her name as ‘Fannie otherwise Fanny’.

I have included some detail of Fannie and her family in the hope that others may have further information or photographs. Current family members are often unaware of what their antecedents got up to, as evidenced by the number of people contacting me to say they had found an article of mine while seeking information on family history.

FANNIE WALDRON was born on 5 June 1876 in Hungerford, Berkshire. She was the daughter of Walter Brind Waldron (1840-1913) and Marian Wood (1847-1917). The Waldrons came to own large estates in Patagonia and in Kenya, as well as farming in England. Walter and Marian had eight children: Mary Elizabeth in 1868, Clara in 1870, Edith in 1871, Walter George in 1872, Ruth Marian in 1874 (named as a contact by Fannie when entering the USA), Fanny in 1876, Hilda in 1878 and Dorothy Brind in 1890.

In the 1881 Census, Walter in shown as farming 1600 acres and employing 37 men, 10 boys and 8 women. The farm was Poughley Park near Lambourn. It was part of the Hungerford Registration District and it is where Fannie was born.

By 1891 the family was living in Peasemore House in Berkshire. Fannie was still living with her parents then as well as in 1901 and 1911. By the latter Census, the Waldrons had moved to 15 Portarlington Road, Bournemouth; Walter is described as ‘chairman and director of companies’. That record shows one of the children had died.

By the time of the 1921 Census, both Fannie’s parents had died. She can be found, as ‘head of household’, at The Broadway, Birkenshaw, Totland on the Isle of Wight. On census day she had three visitors from Bournemouth and a servant.

In 1924 shipping records show her address as 12 Southwell Gardens, London SW7. From then on, and probably until 1950, she was living at 26 Moore Street in Chelsea, London. Today this address is assessed as ‘a 5 bedroom freehold terraced house spread over 3,283 square feet, making it one of the largest properties here—it is ranked as the 5th most expensive property in SW3 2QW, with a valuation of £5,918,000’.


The following is a list of Fannie Waldron’s international travels from 1899 to 1938. It may be incomplete since only some shipping records can be found online. I have included them all because although I found no collecting activities associated with trips others than the two with Willoughby Lowe in 1933-34 and 1934-35 other records may turn up in the future that could be fitted to the earlier voyages.

1899 Chile. Although only listed as ‘F Waldron’ it seems highly likely this was Fannie since the family had farming interests in Chile and had used Punta Arenas as their base. She arrived in Liverpool on 26 August 1898 from Punta Arenas on board Orissa (Pacific Steam Navigation Company).

1920-21 Hawaii. On 27 October she arrived in Quebec on board Canadian Pacific’s Empress of France having departed from Liverpool on 21 October. She crossed the US border the same day at St Albans, Vermont. Then she can be found arriving in Honolulu on the Union Steam Ship Company of New Zealand’s ship RMS Niagara on 17 November 1920 from Vancouver. Nearly three months later she left Honolulu for San Francisco on board Matson Line’s Matsonia, arriving on 16 March 1921. For immigration she was described as 5’ 8” with brown hair and grey eyes. She travelled with her maid, Dorothy Wales, aged 22.

1924. Amazon Cruise to Manaus, Brazil. Departed Liverpool on Booth Steamship Company’s Hildebrand

1926-27 India and Ceylon (Sri Lanka). She departed from London on board P&O’s Rawalpindi on 12 November 1926. The next record is of her leaving Colombo on P&O’s Moldavia which arrived in London on 11 March 1927. She actually disembarked at Plymouth, before the ship sailed on to London.

1930 Trinidad. She left Avonmouth on 11 January 1930 on board the Elders & Fyffes ship Ariguani bound for Trinidad. She returned on the same company’s Camito which arrived at Avonmouth (Bristol) on 27 February.

1933-34 West Africa. Departed Liverpool for Takoradi, Gold Coast (Ghana), on board the Elder & Dempster Line ship Appa on 29 November 1933 with Willoughby Lowe. The return trip was on the same line’s ship, Accra, which arrived in Liverpool on 20 March 1934 from Takoradi.

1934-35 West Africa. Departed Liverpool for Takoradi, Gold Coast (Ghana), on board the Elder & Dempster Line ship Adda on 28 November 1934 with Willoughby. Returned on the same line’s Apapa to arrive in Plymouth on 18 March 1935. Lowe embarked at Accra; Miss Waldron at Takoradi.

1938 West Indies. A cruise from Dover (returning to Plymouth) to Kingston, Puerto Colombia, Curacao, Trinidad, Barbados on board the Royal Netherlands Steamship Company’s Colombia.

Reports of the 1933-34 and 1934-5 Expeditions to West Africa

Reports by David Bannerman of the Natural History Museum on the 1933-34 and 1934-35 expeditions appeared in the Bulletin of the British Ornithologists’ Club. David Bannerman (1886-1979) at the time was chairman of the Club and the expert on the birds of West Africa:

Mr. David Bannerman made some remarks on a collection of some 560 birds recently made in Ashanti, Gold Coast, by Mr. Willoughby P. Lowe for the British Museum, and exhibited a specimen of Glaucidium tephronotum tephronotum, the Gold Coast Yellow-legged Owlet, from Mampong. Mr. Bannerman said that no large collection from the forests of the Gold Coast had been made since Governor Ussher had employed Aubinn—a native collector—to obtain birds for him between the years 1867-1872. Although a number of very rare birds were secured in that collection, and finally were presented to the British Museum, none bore any data on the labels, the only locality mentioned being either Denkera (a spot which is not marked on modern maps, but which is situated in lat. 6°15'N., 2°12'W. long., south-west of Kumasi) or "the interior of the Gold Coast." It was imperative, therefore, that we should have confirmation of Ussher’s records, and arrangements were made with Mr. W. P. Lowe, with the aid of the Godman Fund, to spend three months in Ashanti from December 1933 to March of this year. Mr Lowe's collection contains a number of rare species, but a cursory glance has not revealed anything entirely new. By far the greatest prize is a female specimen which had just finished laying of Glaucidium tephronotum tepgronotum Sharpe, which was secured on February 25, 1934, at Mampong, Ashanti…. Volume 54, p 122, 1933-34.

Other rare birds exhibited, of which Mr. Lowe secured specimens, are: The Black Dwarf Hornbill (Lophoceros hartlaubi hartlaubi), a rare species, of which few specimens are known. The Long-tailed Hawk (Urotriorchis macrourus macrourus), three specimens of which were secured at Mampong and Ejura. The Fernando Po or Fraser's Eagle-Owl (Bubo poensis poensis), two adults and an immature one being obtained-the first specimen I have ever seen in immature dress. The Maned Owl (Jubula letti), the first record from the Gold Coast: previously not known from any locality between Liberia and Cameroons. An account of Mr. Lowe's trip, together with an annotated list of the specimens he secured, will, it is hoped, be published in 'The Ibis' in due course. The passerine birds, which have not yet been named, are likely to prove of considerable interest, and the collection as a whole is a valuable asset to our West African material in the British Museum. Volume 54 p 123, 1933-34.

Mr. Willoughby P. Lowe visited the Gold Coast last winter on behalf of the British Museum, and made collections of birds, fish, and mammals in Ashanti. He is leaving again this month for the same destination. Volume 55, p  35, 1934-35.

Mr. David Bannerman exhibited a selection of birds of many species which had recently been obtained by Mr. Willoughby P. Lowe in Ashanti, and made the following remarks :—

I am exhibiting tonight a number of beautiful and interesting birds which Mr. W. P. Lowe and Miss F. Waldron have recently collected from the Gold Coast. It will be remembered that in the winter of 1933-34 the same travellers made a large collection in the forests of Ashanti on behalf of the British Museum. I made some mention of the results of that trip at our April meeting in 1934…

This year they again visited the Gold Coast in the hope of adding a number of species to their former collection. In this they have been successful. They have not only secured a number of interesting species which they failed to get the year previously, but have been instrumental in making two or three discoveries of particular interest which I propose to mention to you now.

I am pleased to say that Mr. Lowe is present with us this evening and will be glad to answer any questions which may be put to him about his experiences. The birds are exhibited in the cases on the table. I am now engaged in selecting those birds which are required by the British Museum and the remainder will go to the Royal Natural History Museum in Sweden. Volume 55, p 126-126, 1934-35.

Mr. Bannerman next described a new race of the from the Gold Coast, which he proposed to name

Anthoscopus flavifrons waldroni, subsp. nov.

Description (adult male).—Differs from A. flavifrons flavifrons in the brighter, more yellowish-olive plumage of the crown, mantle, back, rump, and wing-coverts, and in the paler underparts, which are more yellow, particularly on the breast, than in the typical species. Eye dark brown, bill black, the base of lower mandible and edges of both mandibles bluish-white.

Measurements of type.—Bill 10, wing 55, tail 28, tarsus 13.5 mm. Distribution.—The forests of Ashanti, Gold Coast. Type.—♂︎ adult, Goaso, Ashanti, Dec. 15, 1934. Collectors: W. P. Lowe and Miss F. Waldron. Remarks.—The discovery, after many years, of a race of Anthoscopus flavifrons flavifrons in Upper Guinea is of great interest. The typical species inhabits the Cameroon forests, and no other race of this bird had previously been discovered. I have much pleasure in naming this bird after Miss Fanny Waldron, who accompanied Mr. Lowe to the Gold Coast and who has been instrumental in obtaining many valuable specimens—particularly of fish—for the Zoological Departments of the British Museum. Volume 55, p 131, 1934-35.

Mr. Bannerman finally exhibited an adult, young, and eggs of the Ahanta Francolin (Francolinus ahantensis ahantensis), and made the following observations :—Although the Ahanta Francolin has such a wide range in West Africa, Mr. W. P. Lowe is the first collector to secure specimens of the young and at the same time a clutch of eggs…. Volume 55, p 132, 1934-35.

Mr. David Bannerman exhibited a specimen of a rare Cuckoo-Shrike from West Africa, and made some further observations upon Mr. Willoughby Lowe's collection from Ashanti. He said :—

I have now completed the examination of the Passerine species in the collection which Mr. W. P. Lowe and Miss Waldron made in Ashanti last winter, and the following are worthy of record…. Volume 55, p 154, 1934-35.

Lowe himself wrote up an account the birds seen and collected in the two expeditions. The report (Report on the Lowe—Waldron Expeditions to the Ashanti Forests and Northern Territories of the Gold Coast) extended over three papers in The Ibis in 1937. The first part details the itinerary and the beginning of the bird list; parts two and three continue the list which includes notes on each species. He ended his acknowledgements with:

Finally, I must express my gratitude to my companion, Miss Fanny Waldron, without whose interest and assistance these natural history collections could never have been made.

He clearly did not know that her name was actually ‘Fannie’.

The 1934-35 expedition with Fannie, when he was 64 and Fannie 58 on their return, were Willoughby Lowe’s last. He was said to have been be in a low condition, from which he never fully covered, after his son, aged 21, drowned off Exmouth in 1931.

Photographs from Willoughby Lowe's Report on the two expeditions

Specimens in the Natural History Museum, London

A search of the catalogue shows the specimens collected by Lowe and Waldron in 1933-34 and 1934-35 that have been retained by the museum. In terms of specimens I counted 1 plant, 54 fish, 2 amphibians, 7 reptiles, 5 birds and 90 mammals. Bannerman (see above) and Lowe reported that the majority of birds from the 1934-35 expedition were sent to Sweden.


1. Miss Waldron’s Red Colobus

The most important finding of the two collecting trips was the monkey, now known as Miss Waldron’s Red Colobus (Piliocolobus waldroni, or sometimes as waldronae to reflect, pedantically, the female for whom it is named. It may be extinct but there appears to be some evidence that it may just be hanging on in the Ivory Coast. The first specimen was, I read, shot or obtained by Lowe in December 1933. It was named as a subspecies of what is now Piliocolobus badius waldroni by Robert William Hayman of the Natural History Museum in 1935 (not 1936 as stated in some publications, although the paper may have actually appeared from the printers in 1936, a matter taken into consideration by priority-obsessed taxonomists). I will deal with this species, if indeed it is a species, in a separate article.

2. Anthoscopus flavifrons waldroni

See Bannerman’s description of the subspecies of the Forest Penduline-tit in the Bulletin of the British Ornithologists’ Club above. Again it sometimes appears as waldronae.

3. Barbus waldroni

Described by John Roxborough Norman (1898-1944) of the Natural History Museum in 1935. Currently regarded as a subspecies of Labeobarbus bynni, the Niger Barb. 

Press Coverage

Willoughby Lowe appears to have had an eye to publicity. He announced his plans for the 1933-4 trip to the Western Morning News and Daily Gazette of 4 October 1933:

…In an interview granted to a “Western Morning News” representative, Mr Willoughby Lowe discussed his plans for his latest expedition, on which he expects to be absent from England for three or four months, With a map. spread in front of him, Mr. Lowe indicated that he would land at Takoradi, on the Gold Coast, and, collecting his safarl, would proceed by train to Kumasi, in the interior. Mr. Lowe's party will not be large, and he will be the only white man…

Northwards from Kumasi huge forests, through which, owing to the rapid growth of vegetation, travel is painfully slow. "These jungles have never been properly explored, and it Is here that I hope to get some of my best specimens." said Mr. Lowe. I think these jungles ought to hold mammals, birds, and plants that no one has ever seen. In the swamps there should be rare fishes and reptiles. Previously I have only visited the coast of West Africa, but I have always come back with some new specimens….

The Western Morning News of 31 March 1934 in a long article gave news of his return:

In neither article does Miss Waldron get a mention. By contrast in the press coverage of the return in 1935, both Lowe and Waldron feature. The Yorkshire Post of 30 March, provides an example:

At the Natural History Museum South Kensington. I watched to-day a number of fish from West Africa being classified. These have been collected by Miss F. Waldron and Mr. Lowe. and some may prove to be new to science. Miss Waldron and Mr. Lowe have brought back certain mud or lung fish which, during the heat of Summer live in a state of suspended animation completely encased in dry mud. Even should the Summer drought up particularly severe they can survive well over six months.

The Belfast Telegraph, 18 March 1935:

Several new species were among a collection of 800 birds, reptiles, and monkeys consigned to the British Museum and Zoo which were landed at Plymouth on Sunday. They were brought ashore from the Elder-Dempster liner Apapa by Mr. Willoughby Lowe and Miss F. Waldron, the naturalists, who have travelled 1,000 miles in the interior of West Africa, searching for specimens for the British Museum. One of the most interesting finds was a sleeping fish, which can remain inanimate and apparently dead for months when the rivers dry up. These fish bury themselves in the mud or clay and remain in a comatose condition until the rainy season comes round and floods the dried-up river beds.

This story made the national press as this article from the News Chronicle (18 March):

It would appear that the party had brought some animals for London Zoo. Examination of the ‘day books’ in ZSL’s archives could provide more information. Lowe mentioned in his 1937 report that:

We caught two [Red-necked Buzzards, Buteo auguralis] alive at Ejura, and intended them for the Zoological Gardens, London, but we were so short of space we could not carry them.

Lungfish certainly made it to the Zoo. as desribed in a syndicated article that reached the Motherwell Times (29 March 1935).

The 1933-4 and 1934-5 Expeditions: Unanswered Questions

Throughout her life, Fannie Waldron was not employed but ‘living on private means’. She was not, as is has been stated, or inferred, a member of staff of the Natural History Museum; nor was Willoughby Lowe. Collectors were on short-term contracts of some sort but in this case I do not know if Lowe was paid a fixed sum plus expenses, or if payment was based on the number of specimens. Did London Zoo pay Lowe for the animals it received? Was Miss Waldron included in the arrangement with Lowe? I have been unable to find any information on how Willoughby Lowe and Fannie met or planed their first trip. Bannerman mentioned that the Godman Fund had been used to part finance the 1933-34 trip. The charity, now closed was the Godman Exploration Fund set up to support travel on behalf of the museum. I strongly suspect that given the acknowledgement Lowe wrote in his account of the expeditionsI, the fact that Fannie Waldron was a very wealthy woman together with the three eponyms bestowed by museum staff, she may have largely funded the two expeditions. 

London Circles

Perhaps as a result of the collections made in West Africa  Fannie Waldron must have become known in the zoological circles of London. She was a guest on two occasions at meetings of the  British Aviculturists’ Club (founded in 1946) . The first was on 12 November 1947; the second  on 22 June 1948. The latter was a pretty grand affair :

That is all the information I have found on Fannie until her death. A press report stated that she had moved from London to Goring-by-Sea, near Worthing, on the Sussex coast in 1950. That move would have come when she was 74.


The Worthing Herald of Friday 6 November 1959 announced:


A member of St. John's Guild for the Blind and a well-known supporter of the Church and South American Missionary Societies, Miss Fannie Waldron, of Wayside, Ashurst-drive, Goring, died at a Worthing nursing home last Thursday. She was 85. Miss Waldron came to Worthing from London in 1950 and soon became well known for her regular visits to Gifford House and other homes for disabled and elderly folk. Among the societies she supported were the RSPCA, the Society for Sick Animals and the Putney Home for Incurables. She was also a member of Goring Parish Church. The funeral was at Peasemore Parish Church on Monday.

Peasemore House, Berkshire, had been her family’s house and other members of the family are buried at Peasemore.

Fannie died on 29 October 1959 in a nursing home at 40 Mill Road, Worthing. The probate record is shown below:

The Worthing Gazette (24 February 1960) later provided some details of her will:

…Probate has been granted to her nephews, Stephen W. Brown, of Peasemore, near Newbury, Berks.. and Maurice C. Waldron, of 16 Coleman-street, E.C.

She left an annuity of £300, a legacy of £300, the proceeds from the sale of her residence and certain effects, to her friend Gertrude Aldridge; £100 to St. John's Guild for the Blind; £50 each to South American Missionary Society and Putney Home for Incurables; £25 each to the Society for the Protection of Animals in North Africa, Peasemore Churchyard Fund, the Society for Sick Animals, and the R.S.P.C.A.

Other bequests were. £50 to her friend Irene Wix; £25 each to her friends Mary O'Dea and Teresa Crowley; £25 *as a token of my appreciation for the many kindnesses he has shown to me during my lifetime" to Mr. G. Dixon. of 3 Tennyson-road, Worthing; a few other personal legacies and the

remainder specifically to relatives and The Friends of the Poor and Gentlefolk's Help, Wireless for the Bedridden Society, The Church Missionary Society, and Royal London Society for the Blind.

A Sad Afternote: Pure Fiction

A story based on Willoughby Lowe and Fannie Waldron and their expedition to West Africa appeared in the book Animal Crackers by Hannah Tinti in 2004. At the end of the short story the author wrote: ‘Although loosely based on historical fact, all characters and events in this story are fictional’. However, the work of fiction traduces both Lowe and Waldron and has reinforced my utter loathing of historical fiction.

A Revised Entry for Eponym Lexicographers


Miss Waldron’s Red Colobus Colobus badius waldroni Hayman, 1935 (currently Piliocolobus waldroni)

Forest Penduline Tit Anthoscopus flavifrons waldroni Bannerman, 1935

Barbus waldroni Norman, 1935 (currently a ssp of the Niger Barb, Labeobarbus bynni)

Miss Fannie (sometimes Fanny) Waldron (1876-1959) took part in two joint collecting expeditions with Willoughby Prescott Lowe (1872-1949) to the Gold Coast (Ghana) in 1933-34 and 1934-35 for the British Museum (Natural History). Some specimens were sent from BMNH to the Royal Natural History Museum in Sweden. Live specimens were received by London Zoo. All three of her eponymous organisms are sometimes shown as waldronae. Lowe wrote: 'Finally, I must express my gratitude to my companion, Miss Fanny Waldron, without whose interest and assistance these natural history collections could never have been made'.

Bannerman DR. 1950. Willoughby Prescott Lowe. Vice-President B.O.U. 1943–45 and Union Silver Medallist. Ibis 92, 142–145 

Beolens B, Watkins M, Grayson M. 2009. The Eponym Dictionary of Mammals. Baltimore: Johns Hopkins University Press

Beolens, B, Watkins M, Grayson M. 2014. The Eponym Dictionary of Birds. London: Bloomsbury.

Hayman RW. 1935: On a collection of mammals from the Gold Coast. Proceedings of the Zoological Society of London.1935, 915-937.

Lowe WP. 1937a. Report on the Lowe-Waldron Expeditions to the Ashanti Forests and Northern Territories of the Gold Coast.. Ibis 79, 345-368.

Lowe WP. 1937b. Report on the Lowe-Waldron Expeditions to the Ashanti Forests and Northern Territories of the Gold Coast.—Part II. Ibis 79, 635-662.

Lowe WP. 1937c. Report on the Lowe-Waldron Expeditions to the Ashanti Forests and Northern Territories of the Gold Coast.—Part III. Ibis 79, 830-846.

McGraw WS. 2005. Update on the Search for Miss Waldron’s Red Colobus Monkey. International Journal of Primatology 26, 605-619.  DOI: 10.1007/s10764-005-4368-9 

Norman JR. 1935. A collection of fishes from the Ashanti Forest, Gold Coast. Annals and Magazine of Natural History (Series 10) 15, 215-221.

Thursday 18 January 2024

George Finlayson (1790-1823) on travel, curiosity and science - plus his eponymous bulbul

In a recent article I wrote on George Finlayson’s eponymous squirrel. Finlayson had written notes before his death on board the ship carrying him from Calcutta to London. Sir Stamford Raffles, who edited Finlayson’s account of his travels in south-east Asia as part of the East India Company’s diplomatic/trade mission in 1821-22, thought the notes were intended for inclusion in a book to be written by Finlayson after his return to Britain. In an introduction Raffles included the following, which, apart from the supervention of Charles Darwin, is an apt a statement today as it was over 200 years ago:

In a greater or less degree, there is, perhaps, inherent in the minds of most men, a desire to visit foreign countries,—desire which neither storms nor tempests, deserts, wilds, nor precipices, with all their appalling fears, have been able to counteract or to check. Cast naked and helpless on this earth, man has aspired to scan its limits, to ascertain its bounds, and even to scrutinize its nature: he has risen superior to the contending elements, which might seem to have opposed an insuperable barrier to his restless ambition, to his ever-active, never-satisfied curiosity; and even the great globe itself no longer seems to offer a theatre too great or too extensive for the exertion of his activity. 

Insatiable ambition, boundless curiosity, are to be reckoned among the more prominent of the attributes with which man is endowed. To what mighty ends have they not led? If they have brought upon him, and upon the race, unnumbered evils, they have also had their attendant good. His share of peace, perhaps of happiness, had been greater had he indulged these propensities less; but it is not in his power to resist the unalterable impulse, conferred upon him, doubtless, for the best of purposes. The curiosity that is gratified with inquiring into the laws implanted in organized beings, or into the general phenomena which characterize the material world at large, admits of, and is usually attended by gratification as permanent as it is unmixed; every step is attended with unalloyed pleasure, every new acquisition leads and stimulates to further discovery. 

This disposition of the mind is particularly observable in those who have made nature and natural objects their study. Hence the eagerness with which men engage in them: no one capable of reflection but has at one time or other experienced this laudable curiosity, and wished for the power to gratify it. To this source we must refer the encouragement held forth in the present day to voyagers and travellers, and in general to every one engaged in matters of discovery. It is not extraordinary, therefore, that persons should readily be found eager to enter upon the investigation of new and distant countries, and of the various objects of knowledge which they contain. It is the lot of few to indulge their inclinations this way; and of these few, how scanty is the proportion of individuals qualified for the important task, either by original endowment, by previous pursuits and habits, or by the necessary education, or by a proper train and temper of mind! Fortunately, however, the objects of pursuit are as numerous as the taste of man is various, and something is left even to the most humble intentions. A proper consideration of this matter would lead to the most important acquirements both on the part of the most humbly endowed, and for the benefit of science and knowledge in general. The principle need not be enforced by argument: let us follow it, if possible, with alacrity, and make the most of the opportunities which fall in our way. Let us devote to the task those abilities, however moderate, with which the Almighty has endowed us, and we shall rarely fail altogether of deriving benefit from our exertions. We may rest secure that the labours so bestowed will seldom fail to be duly appreciated; that our observations will be received with candour, and our alignments, if urged with modesty, will rarely fail to be listened to by the circle of our friends and acquaintances, to the approbation of whom no one can be altogether indifferent. It is in this temper of mind that we may hope to avoid a two-fold evil; that of exaggerating the importance of the feeble exertions of an individual on the one hand, and of thinking too meanly of his capacity on the other,—since both are alike hurtful, and alike oppose the acquisition of useful knowledge. 

Since I can find no portrait of Finlayson I include a photograph of his eponymous bird, Pycnonotus finlaysoni, the Stripe-throated Bulbul. He collected it on the mission. Thomas Horsfield, curator of the East India Company’s museum in London and who described Finlayson’s Squirrel, sent the specimen to High Edwin Strickland (1811-1853) who got round to describing it in 1844, over 20 years after Finlayson’s death.

Streak-throated Bulbul in Thailand
Photograph by JJ Harrison*

Finlayson G. 1826. The Mission to Siam, and Hué the Capital of Cochin China in the Years 1821-2. London: John Murray.

Strickland HE. 1844 Descriptions of several new or imperfectly-defined genera and species of Birds. Annals and Magazine of Natural History 13, 409-415.

*JJ Harrison Creative Commons Attribution-Share Alike 3.0 Unported

Saturday 13 January 2024

Bronze-Winged Mannikins: a colour plate from 1969

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. This is the 16th in the series.

– – – – – – – – – –

The artist for this plate was Chloe Elizabeth Talbot Kelly (born 1927) who went on to illustrate a number of field guides. Her paintings of birds appear in art sales. She began painting in 1945 at the Natural History Museum in London.

The article accompanying this plate was written by Colin James Oliver Harrison (1926-2003) who worked for many years at the Natural History Museum’s bird collection at Tring.

The Bronze-winged or Bronze Mannikin (Spermestes cucullata) occurs in grassland across a large swathe of Africa. Large flocks feed mainly on grass seeds but also take small invertebrates during breeding in the wet season. 

Avicultural Magazine Vol 74, 1968

Saturday 6 January 2024

The Army Surgeon’s White Squirrel of Siam

This article could begin as part of a quiz: What mammal of unusual coloration connects a mission to Thailand, when it was Siam, and a Scottish doctor to my parents’ garden in the early 1960s?

The answer is Finlayson’s Squirrel. Callosciurus finlaysonii is unusual for two reasons. The first is that it comes in a wide variety of colour forms in different parts of its range in south-east Asia. The second is that in one of those colour forms the colour of the pelage is entirely white.

Finlayson's Squirrel, photographed in Saraburi, Thailand
Photograph from the project Noah Website*

It was this white form that was seen and collected by George Finlayson in southern Thailand. He was the surgeon and naturalist to a less than successful trade/diplomatic mission from the East India Company to Siam and Cochin China (Vietnam) led by John Crawfurd (1783-1868) and comprising the following members of the Indian Army plus Mrs Crawfurd: Captain Frederick Dangerfield (1789-1828) was assistant head; he had established a name for himself as a surveyor and geologist; Lieutenant Walter Rutherford (1801-1856) was in command of thirty sepoys. As far as I can ascertain all members of the mission—apart from the sepoys—were Scottish or of Scottish descent.

George Finlayson was born in Thurso in 1790. He became clerk to Dr Somerville, head of the army medical staff in Scotland during the Napoleonic wars. He did so in succession to his elder brother, Donald, who Somerville had moved on to the army medical service, such were his talents. This was a time when clerks served effectively as clinical apprentices. George followed Donald into the army. Donald served in the engagements preceding, and at, the Battle of Waterloo but then came to a sad end. He was assistant-surgeon to the 33rd Regiment of Foot but disappeared as the army marched on to Paris. It was thought Donald had fallen to marauders following in the wake of the retreating French army. George took leave to search for his brother but could find no trace. He was so distraught that the army, at Dr Somerville’s suggestion, moved him to the medical staff about to leave for Ceylon (Sri Lanka). Once there he threw himself into the pursuit of natural history. In 1819, he moved to India as assistant-surgeon to the 8th Light Dragoons then stationed at Meerut.

In order to stay in India for Crawfurd’s mission, when his regiment sailed back to Britain after its service in India, Finlayson moved to a different regiment.

After the mission in south-east Asia, George Finlayson quickly succumbed to what he self-diagnosed as phthisis—tuberculosis, as it later became known. He died on the passage from Calcutta to London on the East India Company’s ship, General Hewett, in 1823.

Finlayson had written an account of the Crawfurd Mission. It was published in 1826 with a ‘Memoir of the Author’ written by Sir Stamford Raffles (1781-1826), founder of modern Singapore and of the Zoological Society of London. Raffles clearly thought very highly of Finlayson.

Finlayson had embarked on General Hewett with the specimens he had collected, including a white squirrel, destined for the museum of the East India Company which was in Leadenhall Street in London. The Keeper was another friend of Raffles, Thomas Horsfield (1773-1859). There Horsfield described the squirrel and named it after Finlayson as Sciurus finlaysonii. He began his description with a quotation from Buffon:

Sc. lacteus dorso flavescente, oculis vibrissis palmis plantisque nigris, cauda pilis nigris raris interspersa.

Ecureuil blanc de Siam, Buff. Hist. Nat. VII. p. 256.

Which translates as: a squirrel, milky with a yellow back; vibrissi, palms, plantar surfaces of feet black, a hairy tail interspersed with rare blacks.

Horsfield continued:

This species is dedicated to the memory of Dr. George Finlayson, (of His Majesty's 59th [2nd Nottinghamshire] Regiment [of Foot],) the naturalist, who accompanied John Crawfurd, Esq. in his mission to Siam and Cochin-China. His health was in a precarious state, from the effects of an Indian climate; and we have to lament that he did not live to return to his native country.

This species has hitherto been mentioned by Buffon alone [in 1789], from the following concise notice in P. Tachard's Travels. “Nous y (at Lonpeen, a village situated in the extensive forests of Siam) vimes aussi des Ecureuils, qui ont le poil parfaitement blanc et la peau tres-noire."—Second Voyage du P. Tachard, Paris, 1689, p. 249.

The following description is extracted from Dr. Finlayson's manuscripts:—“The head and body yellowish white; the head round; the cheeks full; the nose large ; the ears large, plain, not tufted; the iris dark brown; the whiskers long and black; the tail bushy, interspersed with black hairs, and tufted; palms of the feet black. This is an elegant, lively, and active species of Squirrel, almost perfectly white. The body is about seven inches in length, and the tail is equal in length to the body. The eyes are black and lively; and the animal, though white, has not the leucæthiopic habit common in the animals of Siam. It frequents large trees, feeds on their bark and fruit, and is generally seen upon a tall species of Aleurites. One of the specimens was shot by Lieut. Rutherford, on the Islands called Sichang, in the Gulf of Siam."

The India Museum closed in 1879 and the specimens transferred to what is now the Natural History Museum in South Kensington. And there the type specimen—the one shot by Lieutenant Rutherford on the island now known as Koh Si Chang—can be found, catalogued under its original scientific name of Sciurus finlaysonii. In his journal, Finlayson noted that on the group of islands that includes Koh Si Chang:

The [squirrel] is rare, about eight inches in length; an active, lively, and handsome animal.

On the morning of the 13th [August 1822) we landed on the principal island, in pursuit of white squirrels. 

Moving to the 21st century, Finlayson’s Squirrel is now known to vary widely in the coloration of its fur. It varies geographically (the basis of the erection of numerous subspecies, 16 of which are currently recognised). It varies within populations and it varies according to season. Coloration across its range (Burma/Myanmar, Laos, Cambodia, Vietnam and centred in Thailand) varies widely including, as well as white, black, red and agouti. Intergrades between these forms have also been seen where populations meet. An example of variation within a population is in C. f. bocourti, common in Bangkok parks. Individuals may be dark on the dorsal surfaces of the body and forelimbs or entirely white. Variation with season has been described in a population that is maroon-red at one time of year and white at another. Charles Francis in his book published in 2008 made the point that there is a need for more observations in the field and for molecular genetic studies. A key question, of course, is whether all the Finlayson’s Squirrels currently labelled as such are indeed of a single species and whether other forms currently included in different species should be moved to C. finlaysonii. Possible reasons for the great variation in colour and pattern do not seem to have been explored and one can only speculate on the selective forces at play.

George Finlayson realised that white squirrels were present in southern Thailand as well as on the island of Si Chang. It just happens that the formal description of the species was from one shot on Koh Si Chang.

The variations in colour have been shown in several books and papers:

From Francis 2008 - see below

From Wilson, Lacher & Mittermeier 2016 - see below

The coloration of the species varies so much that the name Variable Squirrel has been used and indeed adopted by IUCN for its Red List. Why they should do that when everybody worth knowing has always called it Finlayson’s Squirrel I do not know.

Distribution of Finlayson's Squirrel
Adapted from the IUCN Red List

Horsfield’s use of -ii for the genitive of Finlayson in its specific name of finlaysonii is not the preferred form. Most, and certainly modern usage, would be finlaysoni, the single -i. As the latter it has often appeared in scientific papers and books including Walker’s Mammals of the World in at least some of the editions published since 1964.  However the rules are such that it is also incorrect to change it to the single -i. C. finlaysonii is the original spelling and therefore retained.

Unlike the vast majority of animals that I have kept over the past 65 years I cannot recall how I obtained a female Finlayson’s Squirrel in 1960 or 1961. Had I known then what I know now I would not have done so. Variegated Squirrel Bornavirus 1 (VSBV-1) was first reported in 2015 following reports of the three human deaths in Germany between 2011 and 2013 from of a previously unknown encephalitis. The novel virus was isolated from tissues of a Variegated Squirrel, Sciurus variegatoides, native to Central America, owned by one of those who had died. The virus has since been found in other captive squirrels, including C. finlaysonii and Prevost’s Squirrel (Callosciurus prevostii), in collections in Germany, the Netherlands and Croatia. Those who died were breeders of squirrels, were known to each other and belonged to a squirrel breeding association where they met regularly. Another fatal case occurred in a zoo keeper in Germany, with the virus being found in a Prevost’s Squirrel in the collection. The whole history of the appearance and evolution of the virus has been investigated by studying the molecular epidemiology, together with the activities of the individual keepers and the movement of animals between breeders and from the breeders to private keepers and zoos. Current evidence indicates that a Prevost’s Squirrel was responsible for the primary introduction of the virus.

Prevost’s Squirrels, a beautifully marked species, were commonly available from animal dealers in the 20th century. I have seen them for sale recently in UK and several zoos breed them. I do hope they are all tested, since the fate for those keepers who have caught VSBV-1 has been dire. Fortunately, there have been no cases of human-human infection reported. I do wonder if there have been unreported cases of encephalitis of unknown aetiology from keepers of other species of Callosciurus. For example, In Hong Kong, Pallas’s Squirrel (C. erythraeus) was a popular pet, kept in small cages in small flats and, therefore, in close proximity to the human inhabitants.

Since the discovery of VSBV-1 there has been considerable work in Europe to follow-up these findings with surveys of wild squirrels, zoo workers and the wider human population.

Returning to my Finlayson’s Squirrel of the early 1960s, it was an unsatisfactory addition for the simple reason is that it only appeared from its sleeping box at first light for a short run around its capacious outdoor cage and to eat. Except on rare occasions it stayed in its box for the rest of the day, summer and winter, for years. Only much later did I realise how it could survive on one meal a day. In the wild they are known to eat fruit, seeds, bark, buds, flowers and sap, all of low energy density. I gave the sort of mixture suitable for rabbits or rats: biscuit, wheat, maize, oats, sunflower seed and the like plus fruit like apples and vitamin supplements. All the former have 4-5 times the energy density of tropical fruits. It did not need to emerge for more food and it never or hardly ever did so even in the depths of winter. By contrast, I have seen photographs of tame Finlayson’s Squirrels reared in captivity.

All I remember from around that time is that the Finlayson’s Squirrels were advertised as white. Only much later did I find that not all Finlayson’s Squirrels are white. My guess is that importers wanted white squirrels and that whoever was exporting them from Thailand sorted them accordingly. 

As far as I can recall Finlayson’s Squirrels were only imported into Britain for a relatively short period from around 1960. However, they have now become naturalised as a result of releases/escapes in Singapore, in two areas of Italy from the 1980s, and the Philippines. Genetic evidence suggests they are also present amongst the introduced squirrels of Japan. In short, there is great concern in those regions that they have become a problem as an ‘invasive species’.


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