Monkeys in late Victorian London. An astonishing number arrived at the Zoo

 Although difficult to quantify, anybody who has read Hannah Velten’s Beastly London or Leonard Robert Brightwell’s The Zoo Story, will realise that in the years before the First World War, Britain was awash with exotic animals imported for sale as pets or exhibition by travelling showmen as well as by the comparatively few zoos. For example, Brightwell wrote of London Zoo:

A steadily increasing flow of exhibits poured in from all sources, princes, hunters, dealers and even schoolboys. The animal trade may for one reason or another gradually change its centre of activity. Liverpool has always been one such base of its operations, but in [Abraham Dee] Bartlett’s time the Mecca was in the notorious Ratcliffe Highway, now the eminently respectable St. George’s Road, East, a stone’s-throw from Tower Bridge. The writer has vivid memories of dealers in this quarter, great names that vanished quite suddenly during the Edwardian era. There on the skirts of dockland the great Jamrach and his equally famous rival Hamlyn kept their crowded shops with crazy stable-yards behind, These stables were old warehouses, three stories high and packed with animals of every kind. When a lion roared the whole of one of these aged and rotten buildings rattled in a manner threatening instant collapse. But some spectacular deals were made here, and there are endless stories told of these kings of zoological commerce…

When I totted up some of the numbers of animals that arrived at the Zoo in the 12 years between 1883 and 1895, I was astonished to find over 1,300 individual primates of 166 species (using the identifications and taxonomy of the time). That is a rate of arrival of more than two per day, week in week out, year in year out. Numbers varied from a single Gorilla and one Aye-Aye to over 100 each of Crab-eating Macaques and Rhesus Macaques. The vast majority were presented by private individuals.

With primate arrivals running at over a hundred per year, a demographer is not not needed to point out that the available accommodation would soon be full to the brim and that many of the animals would not have reached their expected lifespan of, in many cases, several decades. Without looking at the records for each year, some will have gone to other collections while some will have been killed as they became unmanageable (the K.B.O. often left unexplained in reports but an abbreviation of Killed By Order). We do though know that the death rate of primates in the indoor accommodation of the time was high. Tuberculosis was, just as in the human population of London, endemic and an improvement in housing and care was the aim of the incoming Secretary in 1903, Peter Chalmers Mitchell.

There was a joke about old-style zoology; a zoo was just a place to keep animals until they could be studied properly—when dead. The Zoological Society’s Prosector throughout the period in question was Frank Evers Beddard FRS (1858-1925). Beddard was Prosector from 1884 until he retired in 1905. He had the responsibility of dissecting suitable specimens and, guided by a set of rules established by a committee, for distributing specimens and parts of animals to interested parties, including university departments, museums and medical schools. He also hosted scientists who wished to work in the prosectorium. Even as late as the 1950s there were disputes as to who was to receive priority in the distribution of specimens. How such matters were decided  on a day-to-day basis when Beddard was in charge is not clear. As well as specimens in whole or in part going to the destinations mentioned some skins went to taxidermists including the famous Edward Gerrard. Details of who got what and when were kept in a book now in the archives at the Zoo.

Important scientists worked in the dead house, as the prosectorium was usually called, and there is a strong case to be made for the view that the only scientific use for an animal in the Zoo was when it was dead. Earlier in the 19th Century, Richard Owen the great anatomist, anti-evolutionist and parasite on royalty, had discovered the parathyroid gland—in an Indian Rhinoceros. Clinicians worked there too. For example, the neurologist Wilfred Harris (1869-1960) applied because he needed to study the brachial plexus of a monkey in order to determine the components of the Vth cranial nerve. His aim was to change surgically the innervation the muscles of a young child who suffered from local paralysis caused by poliomyelitis. After his retirement in 1935 Harris returned to the Zoo to make a systematic study of the brachial plexus throughout the vertebrates, including the Giant Panda. He produced a book on that one topic.

Another visiting worker in the Prosectorium was Peter Chalmers Mitchell. In 1903 he was appointed lecturer in zoology and botany at the Charing Cross Hospital Medical School. His research at the zoo was on the anatomy of birds, particularly of the alimentary canal. Again, he had plenty of specimens at his disposal. I will return to the Beddard-Mitchell relationship in a future article.

Monkeys were still coming into London in considerable numbers in the decades after the First World War. Hamlyn’s Menageries Magazine, published by the importer and dealer John Hamlyn, make fascinating reading since the issues show the numbers of animals he, and sometimes his remaining rivals, had in stock or were being shipped. For example, in January-June 1919, he received 426 primates (some just described as ‘monkeys’). By then Jamrach’s had ceased trading but other dealers were on the scene. George Chapman, with premises on Tottenham Court Road, apparently bought out the Hamlyn business after John Hamlyn’s death in 1922. Chapman was a major supplier of primates and other animals (a final Thylacine, for example) to London Zoo including the  >100 Hamadryas Baboons for Monkey Hill in 1925.

The old records also show why the breeding records were so poor and why self-destructive behaviours were so common in zoo monkeys. Many, perhaps most, had been hand-reared in their country of origin, kept as house pets and then passed on when their owners could not cope with them. They did not know what they were when they arrived. They were simply crazy, mixed-up monkeys who may not even have recognised others of their species. Psychopathies, poor nutrition and infectious diseases were the norm in these unfortunate animals kept in the 19th and much of the 20th century.

The old Monkey House at London Zoo demolished early 1920s
There was no outdoor accommodation

Opened in 1903 this house was replaced by present Reptile House in the mid-1920s

Brightwell LR. 1952. The Zoo Story. London: Museum Press.

Felger EA, Zeiger MA. 2010. The death of an Indian Rhinoceros. World Journal of Surgery. DOI 10.1007/s00268-010-0603-4 

Nieman, E. 2001. Wilfred Harris (1859-1960). In, Twentieth Century Neurology: The British Contribution, edited by FC Rose. Singapore: World Scientific

Velten H. 2013. Beastly London.  London: Reaktion.

Zoological Society of London. 1896. The Vertebrated Animals Now or Lately Living in the Gardens of the Zoological Society of London. Ninth Edition. London: Longmans Green.

Edward Bles (1864-1926). Part 2. Gentleman zoologist. Frogs, plankton, embryos and protozoa

Edward Bles was a zoologist with a substantial private income. He did not need to work but did so anyway, eventually in his own private laboratory. As described in Part 1 he was the first to describe the development of Xenopus in detail. Indeed much of his his work was in embryology at a time when this field was a leading and often controversial aspect of zoological research. Confusing for genealogists, he was known as Edward Jeremiah Bles but his birth was registered as Jeremiah Edward. Therefore, some documents do not fall immediately to hand when searching the historical records.

Bles was born in Salford near Manchester in 1864, the son of Abraham Jeremiah Samuel Bles (1838-1909) and Esther Polak. Abraham and his brother, David Samuel, were born in The Hague; their father established S.D. Bles & Co, merchants and shippers, in Machester largely for the Dutch trade. The Bles’s were leading lights in the Jewish community in Manchester as well as looking after the interests of the Dutch. Abraham was Dutch Consul.

In 1876 the Manchester Courier reported that young Bles had passed the Government Science Examination. Aged 14, Edward Bles was sent to a school in Hanover and at 18 started work in the family business. An interest in science which developed at school in Germany led to his joining the Manchester Microscopical Society (still in existence) of which he became Secretary. Such clubs brought amateurs and professionals together and it was there that Bles fell under the influence of Arthur Milnes Marshall FRS (1852-1893) who, in 1879 at the age of 27 had been appointed to the new chair of zoology in Owens College (later incorporated into what is now the University of Manchester). Bles therefore became a student at the college. In 1890 Bles published with Marshall papers on the development of amphibians, in this case the kidneys and fat bodies, and the blood vessels.

From Owens College, he moved to King’s College, London, graduating with a B.Sc in 1890. His obituarist (see below) noted that he spent time at the Naples marine laboratory but returned to Manchester, as junior demonstrator in zoology. In the summer of 1892 he was working at the Plymouth laboratory of the Marine Biological Association on plankton. He is shown in the resulting paper as honorary research fellow at Owens College. Bles must have been well known in the Plymouth laboratory because he was appointed Director in April 1893. But this was at a time of financial stringency. The Director had a heavy administrative load which prevented personal research. These factors resulted in a rapid turnover of Directors. Bles left in 1894. However, there may have been other reasons. In his second report as Director he noted that the issue of the Association’s journal was late: ‘Unforeseen circumstances affecting myself have caused a further postponement’. Were these ‘unforeseen circumstances’ and leaving Plymouth related to illness, which dogged him in later years, or to the fact that he married Bertha Bachmann of Augsberg in Dusseldorf, Germany on 12 November 1893?

October 1896 saw him admitted to King’s College, Cambridge at the age of 32. He graduated B.A. (as a research degree—the Ph.D, ‘the German degree’ was not awarded in Cambridge until 1921) in 1898 (M.A. 1907). In 1902 John Graham Kerr left Cambridge to the chair of natural history in Glasgow taking Bles (and his frogs) with him as senior assistant, which would be the equivalent of senior lecturer in English universities.

After 5 years in Glasgow, by which time he had been awarded the D.Sc. degree (by the University of London in 1906) and elected to the Royal Society of Edinburgh in 1904, Bles moved out of academia. First he moved to the Hill House, Iffley, Oxford and then to Cambridge where he and Bertha lived at ‘Elterholm’, 12 Madingley Road—a very large house. It would appear it was there that he further equipped his laboratory and ‘started to breed various species of rare amphibia, a difficult enterprise in which he had the assistance of his devoted wife’.

By that time he was also working on Arcella, a freshwater protozoan, in particular the role and control of the gas vacuoles which regulate its buoyancy. Bles had a marked determination not to rush into print. Although he had virtually completed the work by 1914, his long paper was not published until after his ‘very sudden’ death on 3 May 1926. Bertha, who died in 1960, had again helped Bles with is research and she helped with publication. Over the years it has been widely cited. He had again commissioned Kirkpatrick Maxwell to draw the plates.

One of the plates from Bles's paper
on Arcella which was published
three years aftere hisa death

Here is Bles’s words is a description of a part of this work:

These uniformly positive results strongly support the view, which has long been held, that the function of the gas-vacuoles is to reduce the specific gravity of the Arcella, and float it up to the surface which is rich in dissolved oxygen. But it is now possible to go a step farther. In considering the natural causes, or changes in the environment, which might possibly stimulate gas formation for hydrostatic purposes in Arcella, the first consideration was, what is the most obvious and most important physiological difference between the water at the bottom and the water under the surface-film? This is clearly a difference in oxygen pressure. The analyses of pond-water for the determination of dissolved gases carried out by Knauthe (1898, 1899) and Zuntz (1900) show that pond-water may, by the influence of physical and biological conditions, be entirely deprived of oxygen. Owing to the slow rate of diffusion of oxygen in water, the bottom water of a pond or ditch, exhausted of oxygen, will be replenished only a long time after the surface layers. Hence it will clearly benefit those aerobic or semi-anaerobic organisms which live on the bottom, to have a means of escape which will rapidly carry them from a level of oxygen depletion to a level of oxygen plenty. The principal stimulus to form gas-vacuoles in Arcella and similar organisms which live at the bottom of ponds and ditches, is lack of oxygen. There may be, and probably are, other sets of external conditions which stimulate the production of gas by these organisms, and there are also conditions arising within the cell which stimulate the gas-forming structures. These will be described and discussed later…

It is evident that in Cambridge Bles was well known and respected. Sir Frederick Gowland Hopkins (1861-1947) wrote Bles’s obituary for Nature while he and David Keilin (1887-1963) completed the Arcella paper, the biochemical aspects of which fell into their own interests in cell metabolism.

Hopkins began the obituary in Nature:

By the recent death of Edward J. Bles, zoological science has lost a devoted worker whose qualities of mind and character were of the highest. It is the faith of many of his friends that, but for factors of temperament, and health, he would have become a leader of thought in the subject of his choice. His publications, though of high merit, were relatively few; but his intimates know that they were far from representing all that he accomplished, and are aware of the temperamental restraints but for which he could and would have published much more. He was one of those investigators-deserving sympathy from colleagues with easier standards—who would fain allow publication to wait for perfection, and yet realise even better than others that perfection never arrives. In spite of such inhibitions, or perhaps because of them, his published output is of high value and stamped with the quality of absolute reliability.      For elementary teaching, or, at any rate, for the shackles of departmental teaching and organisation, Bles had some distaste. On the other hand, he was the ideal colleague and one of the most educative influences for the young research worker…

He ended:

Bles was not merely a scholarly biologist in a very wide sense, he was also a man of fine general culture; music, literature, and the arts all made a vivid appeal to him. He had, moreover, a true sense of values and a very beautiful appreciation of the relative importance of things. His knowledge was of the widest, but so philosophic was the cast of his mind that synthetic thought was essential to him. He endeavoured always to see things as a whole. 

I have tried to draw up a list of Bles’s publications; it is shown below.

Bles’s legacy extends beyond his publications. He left the entire residue of his estate (about £44,000) plus his equipment and books to the University of Cambridge. It is difficult to equate the worth of that amount of money to today’s economy but in terms of income value (using GDP/capita as the index) it represents £13 million. Over the years, the Bles Fund has funded the Charles Darwin Chair of Animal Embryology—Bles’s express wish as was its use for ‘the promotion and furtherance of biology as a pure science’.

I have been unable to find a photograph of Edward Bles.

In the final part of this series I will return to Bles’s interest in amphibians and how he came to have a walk-on part in the Kammerer controversy.

Hopkins FG. 1926. Dr Edward J. Bles. Nature 118, 90-91.

Publication by Edward J. Bles (Jeremiah Edward Bles):

Bles EJ. 1884. The remarkable sunsets. Nature 29, 427-428.

Marshall AM, Bles EJ. 1890. The Development of the Kidneys and Fat Bodies in the Frog. Studies from the Biological Laboratories of Owens College 2,133-158 plus 1 plate.

Marshall AM, Bles EJ. 1890. The Development of the Blood-Vessels in the Frog. Studies from the Biological Laboratories of Owens College 2,185-268 plus 3 plates.

Bles EJ. 1892. Notes on the plankton observed at Plymouth during June, July, August and September 1892. Journal of the Marine Biological Association of the United Kingdom 2, 340-343.

Bles EJ. 1893. Director’s Report,—No. I. Journal of the Marine Biological Association of the United Kingdom 3, ix-x.

Bles EJ. 1894. Director’s Report,—No. II. Journal of the Marine Biological Association of the United Kingdom 3, xvii-xx.

Bles EJ. 1898. The correlated distribution of abdominal pores and nephrostomes in fishes. Journal of Anatomy and Physiology 32, 484-512.

Bles EJ. 1898. On the openings in the wall of the body-cavity of vertebrates. Proceedings of the Royal Society 62, 232-247.

Bles EJ. 1901. On the breeding habits of Xenopus laevis Daud. Proceedings of the Cambridge Philosophical Society 11, 220-222.

Bles EJ. 1905. The life-history of Xenopus laevis Daud. Transactions of the Royal Society of Edinburgh 41, 789-821.

Bles EJ. 1905. Notes on the development of Phyllomedusa hypochondrialis. Report of the 74th Meeting of the British Association for the Advancement of Science in 1904, pp 605-606.

Bles EJ. 1905. Bles E J On the hatching of anuran tadpoles and the function…[incomplete]. 6th International Congress of Zoology, Bern 1904. (Compte-rendu des séances du sixième Congrès international de zoologie, tenu à Berne du 14 au 16 août 1904[no further details]

Bles EJ. 1906. The life-history of Xenopus laevis Daud. DSc Thesis, University of London.

Bles EJ. 1907. Notes on anuran development: Paludicola, Hemisus and Phyllomedusa. In The Work of John Samuel Budgett, Balfour Student of the University of Cambridge. Edited by J. Graham Kerr, pp 443-458 plus 6 plates. Cambridge University Press.

Bles, EJ. 1929. Arcella. a study in cell physiology. Quarterly Journal of Microscopical Science 72, 527-648.

Comparative anatomy and physiology of excitatory conduction in the heart: Francis Davies and Eric Francis in Sheffield

‘Have you ever tied a Stannius ligature?’, is a conversation stopper. If the answer is ‘yes’ then you know the person you are talking to studied physiology at some time in the past and that a practical class was concerned with the workings of the frog’s heart—an organ ideal for hamfisted students since it beats spontaneously in isolation and they can learn a great deal about how hearts work in a couple of hours*. However, amphibian and reptilian hearts differ in a number of respects from those of birds and mammals.

One of the key players in the comparative anatomy and physiology of the heart is not remembered by the herpetologists, for example, although his great friend and collaborator is. Francis Davies (1897-1965) was at first sight a classical human anatomist—indeed he became co-editor of Gray’s Anatomy. He was born at Merthyr Tydfil, studied medicine in Cardiff and then University College London. In 1924 he became Senior Demonstrator in anatomy at UCl; he then moved to King’s College London as Reader. In 1935 he arrived in Sheffield as Professor; there he stayed until retirement in 1962. In Sheffield Davies worked on the heart with his friend, Eric Thomas Brazil Francis (1900-1993) who became Reader in Zoology until he retired in 1965.

Together, Davies and Francis studied the hearts of amphibians and reptiles in order to determine how the signal from the pacemaker that sets the heart rate passes first to the atria and then to the ventricle (single in amphibians and most reptiles) or ventricles (in crocodiles as in birds and mammals). Earlier in the 20th century Sir Thomas Lewis (1881-1945) had worked out what happened in mammals: specialised heart muscle cells form a dividing bundle of fibres (the Bundle of His) that convey the message to contract to all parts of the ventricles from the atrio-ventricular node. However, there are no special fibres from the pacemaking sino-atrial node to the atrio-ventricular node; impulses pass across the atria from muscle cell to muscle cell like a Mexican wave. It was Lewis’s brilliant work which made not only the physiology textbooks; the research explained a number of conditions that account for heart disease.

Conduction in the mammalian heart
The structures shown in BLACK are not present in
amphibians and reptiles.
from my 1961 edition of the classical physiology textbook
'BDS'

Francis had already published his book, The Anatomy of the Salamander, and their first joint work, published in 1941, was on the heart of that species (Salamandra salamandra). Francis and Davies concluded that in amphibians and reptiles there is no special conducting system in the heart responsible for spreading the process of excitation to the ventricles; the waves of excitation pass directly but relatively slowly from heart muscle cell to heart muscle cell. In other words, the Mexican wave of contraction continues across the whole heart in contrast to birds and mammals where a specialised bundle of fibres takes over.

Davies and Francis proposed that the reason for this major difference in the heart between ectothermic amphibians and reptiles, on the one hand, and endothermic birds and mammals on the other, is the the pace of life; heart rates are lower in the former than in the latter. Relying on a Mexican wave is just too slow for high heart rates to be achieved. They also suggested that the development of a special conducting system was a relatively recent evolutionary change. Their views still hold good.

But what about crocodilians with their two ventricles? Do they have a system like that in birds and mammals, or one characteristic of extant reptiles? Davies and Francis had that covered. They showed that crocodilians have no specialised conducting pathways.

Francis Davies and Eric Francis were not exemplars of the dyed-in-the-wool anatomists who never lifted their eyes from the dissecting table; nor did they confine their studies to comparative anatomy. Both stressed form and function. Davies while undoubtedly seen as a human anatomist of the old school, stressed in teaching anatomy to medical students ‘living’ functional anatomy. Francis was a zoological polymath. In their work on the vertebrate heart, they threw every technique then available at the problem of how excitation by the pacemaker is conducted to all parts: gross observation, dissection, serial sections for histology, histochemistry, slow-motion cinephotography, in-vitro physiology and electrocardiography.

Francis Davies had been unwell for some years when when he retired in 1962. He died in 1965. Eric Francis wrote his obituary for the Journal of Anatomy.

ETB Francis's drawing of the Salamander heart from the 1941 paper

*By tying two ligatures Hermann Friedrich Stannius (1808-1883) showed that the pacemaker of the frog’s heartbeat is in the sinus venosus and that impulses pass from there to the atria and then the ventricle. By isolating regions of the heart these two ligatures also showed that the chambers beat to their own rhythm spontaneously in the absence of input from the pacemaker. The first Stannius ligature is tied between the sinus venosus and the right atrium; the second between the atria and the ventricles. As a hoax the late Jim Linzell and I, in response to a letter asking for exhibits for a museum, put a length of cotton thread in an envelope and sent it along with the explanation that this was Stannius’s third ligature which he never got round to using because his wife had sent a message telling him to get home before his dinner got cold. It was dated 1 April. We never had a reply.

Davies F, Francis ETB. 1941. The heart of the salamander (Salamandra salamandra L.), with special reference to the conducting (connecting) system and its bearing on the phylogeny of  the conducting systems of mammalian and avian hearts. Philosophical Transactions of the Royal Society B 232, 99-130.

Davies F, Francis ETB. 1946. The conducting system of the vertebrate heart. Biological Reviews 21, 173-188

Davies F, Francis ETB, King TS. 1951. Electrocardiogram of the crocodilian heart. Nature 167, 146.

Davies F, Francis ETB, King TS. 1952. The conducting (connecting) system of the crocodilian heart. Journal of Anatomy 86, 152-161.

Francis, ETB. 1965. In memoriam: Francis Davies. Journal of Anatomy 99, 913-915.

Jensen B, Boukens BJD, Postma AV, Gunst QD, van den Hoff MJB, Moorman AFM, Wang T, Christoffels VM. 2012. Identifying the evolutionary building blocks of the cardiac conduction system. PLoS ONE 7(9): e44231. doi:10.1371/journal.pone.0044231 

Frederic Wood Jones. Acerbic anatomist and those who came after him

Frederic Wood Jones
from
Biographical Memoir

In my article on Chapman Pincher and the evolution of the giraffe (1 December 2019) I wrote:

Frederic Wood Jones FRS (1879-1954), a classical anatomist, who had worked on all sorts of biological problems and phenomena from the formation of coral reefs to the lesions caused by judicial hanging, with very strange views on evolution and the value of genetics, wrote to the editor in reply to Pincher. Wood Jones pointed out that he and Robert Broom FRS (1866-1951)—the famous primate palaeontologist and anatomist also possessed of very strange views on evolution by ‘spiritual agencies’—had come up with the same idea as Pincher.

The job of writing Wood Jones’s biographical memoir for the Royal Society fell to Sir Wilfrid Le Gros Clark (1895-1971). Wood Jones had along with Arthur Keith FRS (1866-1955) and Grafton Elliot Smith FRS (1871-1937) inspired Le Gros Clark’s interest in primates and physical anthropology. However, none of the three escaped criticism by their pupil. Lord Solly Zuckerman (1904-1993) noted in his biographical memoir on Le Gros Clark that, in the biographical memoir on Wood Jones, Le Gros Clark had written:

Wood Jones he described as a man who ‘carried his arguments too far and sometimes rather seriously overstated the evidence on which he relied . . .’, as a man who was guilty of ‘needlessly caustic dis­paragement of some of the great biologists of past days’, and as one who was astute in controversy and intolerant of those who disagreed with him. 

But Le Gros Clark had gone much further than in Zuckerman’s examples. Thus:

He was obviously impressed with the precision and. detail of functional adaptations in the animal world, as pre­sumably all zoologists are. But, unlike zoologists in general, he allowed him­self to be so overwhelmed with this outstanding characteristic of living organisms that he was content to ascribe all adaptations to an obscure and undefined kind of directive force, and simply to leave it at that. There was, indeed, a distinct element of mysticism in his attitude to some of the fundamental problems of organic life and, as a result, his more popular writings aroused considerable response from the class of general reader to whom the methods and philosophy of science make little appeal. 

He ended:

A man of restless curiosity and of strong opinions, he sometimes gave an appearance of intolerance towards those who disagreed with him. He was also very critical of the development of new technical methods in anatomical research, even going so far as to imply that any method but that of scalpel and forceps (and perhaps the low power of the microscope) was beyond the proper domain of the science of anatomy. It is little wonder, therefore, that he was out of sympathy with the younger generation of anatomists; so much so, indeed, that he occasionaly [sic] allowed himself to be almost too brusque in his attitude towards them and their work. But although he was possessed of a certain natural acerbity of temperament which impelled him to express his views in somewhat unaccommodating terms, Wood Jones certainly contributed his full share (and more) to the store of comparative anatomical knowledge, and the stimulus which he gave to his students and contemporaries by his lectures and writings can hardly be exaggerated. 

I think it is worth pointing out that Wood Jones, Keith, Elliot Smith and Zuckerman (who, in turn, was critical of Le Gros Clark) came into the biological sciences through medicine. They were all professors of anatomy and clearly ill-equipped in that intellectual tradition, like, sadly, many biomedical scientists of today, to have much worth saying on the mechanisms of evolution or of the importance of the role of evolution in shaping biological systems. The zoologists and geneticists did that. Wood Jones was a man of his tradition and his time.

Le Gros Clark W. 1955. Frederic Wood Jones 1879-1954. Biographical Memoirs of Fellows of the Royal Society 1, 119-134.

Zuckerman S. 1973. Wilfrid Edward Le Gros Clark 1895-1971. Biographical Memoirs of Fellows of the Royal Society 19, 217-233.