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, 12.6.20.1), 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.