Thursday 15 April 2021

The Frog Man. R Maxwell Savage: The Forgotten Doyen of British Ecological Herpetology Part 2

R. Maxwell Savage
from Beebee 2010

In my last post I described the work done by R. Maxwell Savage on the relations between local climate and weather on the time of annual spawning of the Common Frog, Rana temporaria, in Britain. I also noted that I was delighted to find that Trevor Beebee, who found reference to Savage’s often seminal work on amphibians lacking and information on his life absent, had launched an appeal for information. As a result a biography appeared in Herpetological Journal in 2010, 25 years after Savage’s death. More information on Savage’s professional life has emerged and it throws considerable light on the approaches Savage took in studying the Common Frog at all stages of development, from the formation and workings of frog spawn and the life of the tadpole to the triggers for breeding. 

There cannot be many scientists who have had papers in Nature for their amateur as well as their unrelated day job. For that matter, there cannot be many scientists who have received research grants from the Royal Society and a PhD for their spare-time pursuit. R. Maxwell Savage had all of these distinctions as well as the Stamford Raffles Award from the Zoological Society of London in 1967.

Ronald Henry Maxwell Savage* was born in Wood Green, London, on 2 May 1900, the third child of the company secretary of an explosives company. The Savage family had a coat of arms and Savage appears in Fox-Davies’s book, Armorial Families†. From Queen’s College Cambridge (1918-21) he graduated in Natural Sciences. He was a chemist and worked for his entire professional life, 1921-1965, for S. Maw Son & Sons at Barnet in Hertfordshire. Founded in the 1820s the company manufactured surgical instruments, medical kits, as well as common pharmaceuticals, at a large factory built when the form outgrew its London premises. eBay has products made by Maw: infant feeding bottles; bedpans; invalid cups; inhalers; toothpaste; surgical gear and bandages. Field dressings were supplied to British and allied forces and, from published papers, dressings were a particular concern to Savage from the 1930s to the 1950s. Some of the testing and improvement of surgical dressings was done in collaboration with surgeons at the London teaching hospitals. Means of sterilizing dressings, the performance of thrombin-containing dressings designed to speed up blood-clotting and improving the absorbency and holding capacity of dressings were the subjects of some of the papers Savage published in medical journals. He was also involved in the use of chlorophyll as a deodorant—a craze for a while in the 1950s when we had chlorophyll toothpaste, chewing gum, soap, shampoo, lotions etc. With arguments raging on the efficacy of some of these products Savage demonstrated, using proper test procedures and statistical analysis, that ‘chlorophyll on cotton pads does reduce the odour of decomposing blood, the probability of our results arising by chance being less than one in 2,000 million’.

Savage was, in short, the epitome of the brave new world of industrial research in the 1930s. 

His approach to the development and improvement of products, was quantitative, experimental and observational; it included bioassays, testing and quality control that depend on rigorous statistical treatment. It is these attributes that he carried over into his research in the field and in his home laboratory on the Common Frog and other amphibians. He did experiments and used statistical techniques on his field data that put him years ahead of his time—and in so doing probably made much of his work incomprehensible to biologists of the day. I can just imagine the expression on the faces of those who listened to the papers he gave at the Zoological Society in the 1930s. Members of the audience would not, to put it politely, have been familiar with the concepts he presented. It is perhaps not surprising that he complained that his work had been met with ‘polite incredulity’.

I have made a list of his publications from his day job and of those on his research on amphibians;  it is appended below.

In the 1939 Register, the emergency census taken as preparation for war, Savage was living with his wife and incapacitated mother at Derwent Avenue, Mill Hill. He is described as ‘Chemist: analytical and research. Works Manager: surgical dressings’. He was chief chemist for Maw Son & Sons. It was the field dressings made by Savage’s team that were issued to soldiers in the field as ‘First Field Dressing’ packs that could be brought out of a battledress pocket and applied quickly to a bullet or shrapnel wound. Maws supplied many of the field dressings, first aid kits and surgical dressings to military and civil defence organisations and it is clear that as Savage’s research on surgical dressings went on into the 1950s that he was seeking continuous improvement in the company’s products.

British Army First Field Dressing Pack
by S Maw Son & Sons

Savage wrote of how he first became interested—and how that interest was encouraged—in research on amphibians. As a young man he came back from a holiday in France with some Yellow-bellied Toads, Bombina variegata. He was encouraged by Hampton Wildman Parker (1897-1968) then in charge of reptiles and amphibians at the Natural History Museum in London to publish some of the observations he had made and to do more. Savage noted the incongruity of a taxonomist encouraging research which was far from taxonomic but Parker was a Cambridge graduate, like Savage, in natural sciences which included biological subjects and chemistry. It seems possible that they had met in Cambridge because Savage graduated in 1921 and Parker (although three years older) in 1923. The paper on B. variegata was published in 1932 in Proceedings of the Zoological Society (PZS), since retitled Journal of Zoology.

Throughout the 1930s and beyond he published a string of papers on the Common Frog, mainly in PZS. I will not dwell on what he did in this article since I shall write follow-ups on specific aspects, including his book which pulled together his earlier work, as well as on his main hypothesis which occupied much of his time over the next 40 years.

This might though be a good place to put his huge amount of fieldwork in context. In his words:

The area lay on the borders of Middlesex and Hertfordshire, and covered about eighty square miles. Within this area 92 ponds were kept under observation over a period of ten years. None were observed for the whole of this time, for most of the observations were made between 1934 and 1938, although less systematic observations were maintained for another twenty years afterwards, and some of the ponds were known for twenty years before.


The ponds were within driving distance of where he lived, first in Mill Hill and then Hadley Wood.


The approximate area containing the 92 ponds studied by Maxwell
How many survive?

His work on tadpoles was written up for his University of London PhD thesis, The Ecology of Anuran Tadpoles, at Birkbeck College. The date given in the library catalogue is 1950-1951.

Both Savage and Louis Lantz were industrial chemists who worked, as amateurs, on herpetological matters in England. It is evident from Savage’s papers that they were in regular contact. Indeed, Savage collected Painted Frogs for Lantz on the French island of Port-Cros. He was in regular contact with Burgess Barnett—with whom he shared an interest in the mechanism of blood clotting—over the solidification of frog spawn after laying. We also know that Deryk Frazer helped him with summarising phenological data. When Savage was looking at the feeding mechanisms of tadpoles, Laurence Cooper Stuart (1907-1983) of the University of Michigan sent him specimens of a microhylid. Other names are mentioned in his book and it is clear that he was connected with all the key players in his areas of interest, in his ‘amateur’ as well as in his professional life.

Savage thanked his wife, Violetta, née Hetherington, whom he married in 1931 for helping him with fieldwork in the 1930s; they were married in 1931. He also thanked a Dr W.F. Purdy (also acknowledged in one of his papers from Maw Son & Sons) for help with transport and advice on presentation of the figures, and a Mr A. Edwards; I have been unable to find any information on either.

Trevor Beebee mentions that Savage from the 1920s onwards travelled widely on holidays in continental Europe. The fire-bellied toads from France I mentioned above which started his research activity were one obvious result of his travels as were the painted frogs sent to Louis Lantz.

Savage was a Fellow of the Royal Institute of Chemistry and of the Zoological Society of London. He became a member of the British Herpetological Society soon after its foundation in 1947, and was one of my predecessors as editor of its journal, the British Journal of Herpetology (now Herpetological Journal).

On retirement the Savages moved from Hadley Wood to Welwyn where Ronald Maxwell Savage died in 1985.

In the next article in this series I will discuss his book and its reception before moving on to discussion of his big idea, some of his earlier research and of his studies on Xenopus.


†Argent, on a fesse dancetté between four lioncels three in chief and one in base sable, two doves each holding in the beak a branch of olive proper. Mantling sable and argent. Crest—On a wreath of colours, in front of a lion’s jamb couped or, grasping a branch of holly fructed proper, a saltire sable. Motto—“A te pro te”. That coat of arms (a variant of those by branches of the Savage family) appears to have been granted to Henry Maxwell Savage (1861-1938) in 1918.

*He seems to have dropped H[enry] from his initials; he always published under the name R. Maxwell Savage.

Beebee TJC. 2010. Ronald Maxwell Savage, 1900-1985: a tribute. Herpetological Journal 20, 115-116.


Ronald Henry Maxwell Savage

Publications and Radio Broadcasts

Herpetology


Note that the volume numbers of Proceedings of the Zoological Society are ones currently listed on the ZSL publication website. Various other methods of numbering the volumes were used in the past and may be encountered in Savage’s own papers and book.

Savage RM. 1932. The spawning, voice, and sexual behaviour of Bombina variegata. Proceedings of the Zoological Society of London 102, 889-898.

Savage RM. 1934. The breeding behaviour of the common frog, Rana temporaria temporaria Linn., and of the common toad, Bufo bufo bufo Linn. Proceedings of the Zoological Society of London 104, 55-70.

Savage RM. 1935. The influence of external factors on the spawning date and migration of the common frog, Rana temporaria temporaria Linn. Proceedings of the Zoological Society of London 105, 49-98.

Savage RM. 1935. The ecology of young tadpoles, with special reference to some adaptations to the habit of mass‐spawning in Rana temporaria temporaria Linn. Proceedings of the Zoological Society of London 105, 605-610.

Savage RM. 1937. The ecology of young tadpoles, with special reference to the nutrition of the early larvae of Rana temporaria temporaria Linn., Bufo bufo bufo Linn., and Bombina variegata variegata Linn. Proceedings of the Zoological Society of London 107, 249-260.

Savage RM. 1939. The ecology of young tadpoles, with special reference to carbohydrate changes in development, and to the function of the envelope. Proceedings of the Zoological Society of London 108, 465-480.

Savage RM. 1939. The distribution of the spawn-ponds of the common frog, Rana temporaria temporaria Linn., over a portion of the London clay and associated drift. Proceedings of the Zoological Society of London 109, 1-19.

Savage RM. 1942. The burrowing and emergence of the Spade‐Foot Toad, Pelobates fuscus fuscus Wagler. Proceedings of the Zoological Society of London 112, 21-35.

Savage RM. 1950. Observations on some natural epizootics of the trematode Polystoma integerrimum among tadpoles of Rana temporaria temporaria. Proceedings of the Zoological Society of London 120, 15-37.

Savage RM. 1952. Malcolm Smith (1951). The British Amphibians and Reptiles [book review]. Journal of Animal Ecology 21, 162-163.

*Savage RM. 1952. Ecological, physiological and anatomical observations on some species of anuran tadpoles. Proceedings of the Zoological Society of London 122, 467-514.

Savage RM. 1955. The ingestive, digestive, and respiratory systems of the microhylid tadpole Hypopachus aguae. Copeia 1955, 120-127.

Savage RM 1956. Thermal function of the envelope of the egg of the common frog Rana temporaria, with observations on the structure of the egg clusters. British Journal of Herpetology 1, 57-66.

Savage RM. 1961. The Ecology and Life History of the Common Frog (Rana temporaria temporaria). London: Pitman.

Savage RM. 1963. A speculation on the pallid tadpoles of Xenopus laevis. British Journal of Herpetology 3, 74-76.

Savage RM. 1965. External stimulus of the natural spawning of Xenopus laevis. Nature 205, 618-619.

Savage RM. 1971. The natural stimulus for spawning in Xenopus laevis (Amphibia). Proceedings of the Zoological Society of London 165, 245-260.


*Footnote to this paper: This paper has been condensed from part of a thesis approved by the University of London for the degree of Ph.D

Professional


Savage RM. 1936. Penetration of heat into surgical dressings. Chemist and Druggist 125(2943), 14.

Savage RM, Chambers WP. 1938. Optimum temperature of formation of a blood clot. Nature 141 287-288.

Savage RM. 1940. Sterility tests on surgical dressings. Quarterly Journal of Pharmacy and Pharmacology 13, 237-251.

Savage RM. 1942. The sterilization of paraffin surgical dressings. British Medical Journal 1942(1), 472-474.

Savage RM. 1944. The sterilising action of steam admixed with air and other gases. Chemist and Druggist 142(3362), 73.

Savage RM. 1945. The sterilisation of surgical dressings. Pharmaceutical Journal 1945, 254.

Chambers WP, Savage RM. 1945. A comparison of methods of analysis of euflavine gauze with observations on the effect of sterilisation. Quarterly Journal of Pharmacy and Pharmacology 12, 237-234.

Savage RM, Bryce DM, Elliott JR. 1952. The water retention coefficient of surgical dressings. Journal of Pharmacy and Pharmacology 1952 4, 944–958.

Bryce DM, Savage RM. 1953. A note on surface-active agents and surgical dressings. Journal of Pharmacy and Pharmacology 5, 911-915.

Bryce D, Savage RM. 1953. Chlorophyll. British Medical Journal 1953(1), 833.

Savage RM. 1954. The sterilization of surgical dressings. Journal of Applied Bacteriology 17, 278-285.

Savage RM. 1954. A statistical study of variation in surgical dressings. Journal of Pharmacy and Pharmacology 6, 843-858.

Savill A, Daynes G, Savage RM. 1956. Bread. Lancet 267, 1071.

Savage RM. 1957. Sterilization of dressings. British Medical Journal 1957(2), 235.

Savage RM. 1962. Sweetened dummies. British Medical Journal 1962(2), 801.

Radio Broadcasts


3 February 1956. Naturalists’s Notebook. Edited by Maxwell Knight. Produced by Brandon Acton-Bond

8 March 1959. The Naturalist. BBC Home Service. Introduced and edited by Maxwell Knight. Produced by Jeffery Boswall



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