Saturday 28 May 2022

Mavis Gunther: infant feeding and the great personal tragedy in her life, the death of her husband, the zoologist Rolfe

This is a story that for me began in the early 1970s. At a meeting, as I recall of the Society for Endocrinology, I was approached by a little old lady. She had been sent to talk to me by Alfred Cowie, later to become a good friend, whom I learnt later she regularly badgered for information on the endocrine control of lactation. She said to me something along the lines of, ‘All these people are poisoning all the infants in Britain by making up milk too strong. You know something about milk osmolality and tonicity and things, don't you?’. She explained that some mothers and helpers crammed as much baby formula (already too rich in sodium) into the supplied measuring scoop as possible in the belief of the more food the better. When mixed with the correct volume of water, the milk would have a much higher osmotic concentration and possibly lead to dehydration as well as having a long-termdeleterious effect on a newborn’s kidneys. Thus began a short experiment in which I crammed as much baby formula into a measuring scoop as possible to see just how overconcentrated the milk could be made*.

Mavis Gunther at a symposium at the
Ciba Foundation, London, 1965

The little old lady was Mavis Gunther and I use the term in the sense that in all sports and any card game it is wise to beware of the little old lady because at tennis you will soon be 6-0 down, at golf all her putts will drop and at cards or any board game you will be toast. Mavis asked pertinent questions, did not suffer fools, especially her fellow clinicians and other health professionals, gladly and had trenchant views on obstetric practice and the care of infants. She was frustrated by the attitude of of her fellow physicians and, realising that knowledge of lactation had very rarely come from that source, that is why at meetings she sought out those who were studying lactation funded in Britain, like most of mammalian reproductive biology, as part of agricultural research.

As a result of our contact and conversation I asked her to give a paper on human lactation and infant feeding at a symposium I organised for the Zoological Society of London in November 1976. She was then 73 and I think that was the last time we met.

Shortly before she sought me out, Mavis had written a book, Infant Feeding, published by Methuen in 1971. She then produced a revised edition for a paperback published by Penguin in 1973. It was highly influential and her views on mother-infant interactions are still referred to today. I remember reading it at the time but in the 1990s when I was extolling Mavis’s virtues and searching for a copy of the cover with which to illustrate a lecture, I could not find a second-hand copy. I had to borrow a copy from the National Lending Library. The difficulty of getting hold of the book was mentioned by participants when I talked about Mavis at a seminar organised by the Wellcome Foundation in 2007. Was it because so many copies had been sold and held onto or had the print-run been insufficient to meet demand? However, a couple of weeks ago I found a copy of the Penguin edition for sale and paid the £4.56 including postage. It has only taken 49 years.

By now those who know anything about reptiles will be asking the question that I asked 50 years ago. Was she related to the German-born Albert Charles Lewis Gotthilf Günther FRS (1830-1914) of the Natural History Museum in London? ‘Yes’, she replied, ‘He was my late husband’s grandfather’. Only recently have I discovered that her husband, Eustace Rolfe Gunther (known as Rolfe), to whose memory her book was dedicated, was a zoologist and that his death was an utterly tragic event amongst all the tragedies in wartime Britain.

Mavis Gunther (née Carr)

However, before turning to Rolfe, I must return to Mavis’s life. She was born Mavis Hilda Dorothea Carr on 17 June 1903 in Bromley, Kent. She was educated at Bedales School and then Cambridge for pre-clinical medicine, graduating in 1925. She qualified at the Royal Free Hospital, then in the centre of London in 1928. In 1929 she married Rolfe. In 1935 she was co-author of a paper on the genetics of epiloia (now known as tuberous sclerosis) a rare condition. She was in the Research Department of the Royal Eastern Counties Institution for the Mentally Defective in Colchester working with Lionel Penrose (1898-1972, elected FRS in 1953). She had young children, born in 1930, 1933 and 1937. Described as being in general practice in 1939 Register (an emergency census), the Medical Directory for 1940 indicates that Mavis did a number of medical jobs around London, covering activities from antenatal clinics to mental disease. She was a local councillor in the late 1930s but resigned immediately after the death of her husband.

Within weeks of Rolfe’s death in 1940, she and the children left Liverpool on board Canadian Pacific’s R.M.S. Duchess of Atholl which crossed the Atlantic unescorted after having arrived in convoy carrying Canadian troops. An article behind a ludicrously high paywall which I have only seen in part, indicates that a posthumous letter from her husband urged her to seek safety for the children in Canada. Friends apparently invited her to work in the Department of Obstetrics and Gynaecology at the University of Toronto; while there she was supported by the the Banting Research Foundation for research in human lactation. Some of her work she described in a paper in the Canadian Medical Association Journal.

Mavis’s daughter reported that they were in Canada for four years. Sometime before 1946 she joined Robert Alexander McCance’s (1898-1953; FRS 1948) famous team in Cambridge and its equally famous mission to Wuppertal in Germany after the end of the war. McCance knew food had been and would be short in Germany and that various experts could study the effects of undernutrition and to help the local population. As one of the experts with previous experience, Mavis Gunther studied the volume and composition of breast milk.

At some time she was awarded the M.D. degree by Cambridge.

It appears that Mavis was a member of the Medical Research Council’s staff but I have not been able to find if this employment began with the Wuppertal mission or earlier. We do know though that in 1948 she was seconded the University College Hospital, London, where she was given accommodation and facilities. She expanded a clinic for women with problems breastfeeding and it was recalled: ‘Mavis Gunther used to go round the women each day talking to them about breastfeeding and breast problems. She was known, perhaps irreverently, to many of the students as "The Breast Queen", but her work in her gentle persuasive way was invaluable to the mothers in the days when formula feeding was not very reliable. She helped thousands of babies in the first weeks of their lives and is remembered with affection by women who had their babies during her reign’.

The journalist Katharine Whitehorn (1928-2021) in her Foreword to Mavis’s book wrote of her experiences in the maternity ward of University College Hospital:

Into this overheated atmosphere there came only one voice of utter kindness and sense: the Breast Lady, who came calmly round and sorted us all out, and got a great many of us happily established at breast feeding who could never otherwise have managed it.

At some stage she must have moved from obstetrics to paediatrics and to a lecturer’s position. I think she must have retired from University College around 1972. I only remember writing to her at her home address in Esher, Surrey.

Mavis Gunther collaborated widely in pursuit of improving infant feeding and understanding the problems faced by mothers and infants, while gathering the evidence on which advice should be based. She was no breast-feeding zealot, arguing that mother’s must be free to choose breast or bottle depending on their individual circumstances. She published research on a number of topics including allergies to milk, the possible causes of ‘cot death’, comparisons of breast and bottle feeding, colostrum and milk as a source of antibodies, mastitis, sore nipples, human milk composition, the use of mineral additives in milk formula from different manufacturers and breast pumps. In obstetrics she studied the transfer of blood between baby and placenta immediately after birth. She stressed the importance of considering the mother and child as a unit, and to be supported as such. She argued that the behavioural initiation of breast feeding is not an instinctive process and that the mother needs some degree of instruction either by observation during life and/or by the help of family, friends, doctors and midwives. She based part of that argument on following up early observations on Chimpanzees in captivity by contacting London Zoo and learning of their experiences in the 1930s and early 1940s which suggested that only those brought into captivity late succeeded in breast-feeding their young.

At the 1976 symposium at the Zoo I mentioned above, she described her contacts with Geoffrey Marr Vevers (1890-1970) who, with a human obstetrician, described the birth and rearing of the baby chimp ‘Jubilee’ (named for George V’s 25 years on the throne) in 1935. Vevers, was then Superintendent and like Mavis, medically qualified. Listening was Geoffrey Vevers’s son, Gwynne (1916-1988) who amongst his numerous responsibilities at the Zoo had overall charge, wonderfully delegated, of scientific meetings and publications.

Mavis Gunther died on 30 June 1997. She was buried alongside he husband, at Heacham, a town in Norfolk, not that far from where Rolfe was killed, where his parents had a house, and where he was born.

Rolfe Gunther

Eustace Rolfe Gunther

In order to describe the tragic events which led to Eustace Rolfe Gunther’s death, I have drawn on an account given by his daughter, Rosalind, for the Dictionary of Falklands Biography. According to an article in a village magazine recording her death in 2021, aged 90, she did so because the records are closed until 2040 and she believed there had been an official cover up. Indeed, the reports of the inquest in the local press are perfunctory. I have also drawn on the obituary written by Alister (later Sir Alister) Hardy, who worked closely with Rolfe, and my own delvings into the happenings in a Norfolk village. This is what Rosalind wrote:

…As a member of the Territorial Army he was called up at the outbreak of war and commissioned 2nd Lt in the 72nd Anti-aircraft Regt RA. In May 1940 he was stationed near North Walsham in Norfolk. When approaching Barton Turf on foot to check that someone was not signalling to the enemy at sea he called at a cottage to read his map and obtain directions to the suspect house. The resident, a Special Constable, directed him on foot, cycled to the house by another route and borrowed a gun from the newly armed Home Guard. Gunther reached the house. The Constable returned with some further Home Guard members and after asking Gunther to hand over his gun, accidentally shot him in the upper leg.

Rolfe Gunther bled to death in Norwich hospital on 31 May 1940.

The local newspaper’s account of the inquest carried no detail of the examination of witness, just the conclusion of the local coroner: ‘died from a gunshot wound caused by a rifle Inadvertently discharged’. 

It does not need much deductive power to envisage the reaction in the first year of the war of the Special Constable to the appearance of an armed officer with a German surname at his door purportedly looking for a potential German spy. It may not have crossed his mind that the last thing a German spy might do would be to use a German surname. A Special Constable, for those readers not in the U.K., is a part-time, unpaid voluntary police officer. I discovered to a degree of amazement that the tiny village of Barton Turf had three special constables in 1939 but it really does seem like the whole affair was covered up since, war or no war, whichever one of the three got things so wrong, should have been charged with manslaughter.

I can see why there were other unanswered questions. For example, why was Rolfe, an officer in a searchlight regiment (not anti-aircraft, as indicated above) investigating somebody suspected of signalling to the enemy of sea? And by his action might not the zealous constable have tipped off the suspect that he was under investigation?

To add to the tragedy, Mavis was called by the police and drove to the hospital; when she arrived Rolfe had just died.

Hardy for an obituary in Nature wrote: ‘In 1937, distressed by Great Britain's unpreparedness for war, Gunther joined the territorials as a sapper and was commissioned a year later. On inquiry about the best use a man of his training might be, he was advised to enter the searchlight service, and his keen powers of observation were of particular use in training spotters’. His territorial unit was the 30th (Surrey) anti-aircraft battalion of the Royal Engineers (later transferred to the Royal Artillery), the unit in which, by coincidence, my father served from 1941). Gunther was commissioned in the 72nd Searchlight Regiment, Royal Artillery, in February 1939.

Hardy in the obituary and in another I suspect he also wrote for The Times, praised his friend and colleague Gunther highly:

Educated at Winchester and Caius College, Cambridge, he was appointed as zoologist in 1924 to be one of the original members of the scientific staff of the Discovery Committee set up by the Colonial Office to investigate the resources of the antarctic seas, particularly in regard to the factors, biological and physical, governing the great whale fisheries of those waters. 

It was my privilege to be closely associated with Gunther on the R.R.S. Discovery on her voyage of 1925-27, on the smaller ship R.R.S. William Scoresby, and afterwards in the joint authorship of an extensive report upon the ecology of the antarctic plankton. He was a man of sterling qualities. Working with him day and night, often under the difficult conditions presented by the Southern Ocean, one was continually impressed by his deep sense of duty, his devotion to his work, and his tireless energy. His enthusiasm was always combined with a scrupulous regard for accuracy: both in the field and the working-up of data. After working at the nets and water-bottles for thirty-six hours on end, except for odd moments snatched for hurried meals, it was only the fear of being inaccurate in the readings and recordings, not fatigue itself, which persuaded him to rest. He had a love of the sea and the open life; he was a real deep-water oceanographer, with the determination to bring back results. 

In 1931 on the R.R.S. William Scoresby, Gunther led a highly successful expedition to investigate the Peru Coastal Current (sometimes called Humboldt's Current) and published in 1936 a comprehensive report on its physical, chemical and biological aspects. Later he again visited the Antarctic on a whale-marking expedition to study migrations, and made valuable observations on the swimming and breathing habits of whales (a paper now in the press). Much of his time before the War was spent in working on the material collected during a trawling survey, partly carried out under his direction, on the extensive banks lying between the Falkland Islands and South America. It is to be hoped that all the work he put into this will eventually be published. 

In addition to his wide interests in zoology and oceanography, Gunther was always delighted to record any unusual natural phenomenon ; his recent letter in NATURE on the ice storm in Wiltshire is an example of this. Colour and scenery were a great joy to him, and he did splendid water-colour drawings, both sea and landscape, as well as accurate colour studies of marine animals. Many will treasure his privately printed "Notes and Sketches made during two years on the Discovery Expedition". 

It was characteristic of Gunther's capacity for work that his long leave after the 1925-27 expedition medicine in the University of Pennsylvania, on should have been spent in the Department of Biochemistry at Cambridge, undertaking researches on the fatty and vitamin content of plankton (published in collaboration with G. Collin, J. C. Drummond [later Sir Jack who also died in tragic circumstances] and T. P. Hilditch). His published work, while extensive, is no real measure of his industry. He was always being attracted by side branches which he felt it his duty to explore, and only when he had carried them a long way did he realize he was being taken too far from the main issue; reluctantly they were put on one side for some later available time--alas, now no more. 

…He has been a worthy upholder of the tradition set by his father, his grandfather, Dr. A.G.L.G. Gunther, F.R.S., and his great-uncle, Prof. W.C. Mcintosh, F.R.S. 

In 1941 Eustace Rolfe Gunther was posthumously awarded the Polar Medal in Bronze for ‘good services between the years 1925-1939 in the Royal Research Ships “Discovery II” and “William Scoresby”'


Amongst the many people I have met Mavis Gunther was one of the most memorable—and one of the most worth remembering.

*About thirty years later I met a paediatrician at a meeting who knew of this work and I asked him if  overconcentration had proved to be a problem. He said no, the infant could cope. However, I now find that there have been cases of dehydration linked to overconcentrated baby formula and I see all sorts of warning about making up the powders properly.

Friday 13 May 2022

Reticulated Giraffe

Kenya is the only country in which it is possible to see three geographically distinct forms of the Giraffe. The Masai subspecies (Giraffa camelopardalis tippelskirchi) is in the south; further north in the west is Rothschild’s Giraffe (G. c. rothschildi); in the north-east is the Reticulated Giraffe (G. c. reticulata). Here is a photograph I took in 1991 in Samburu of two Reticulated Giraffes feeding in the evening. I still watch giraffes with amazement at how they cope with the thorns.

Looking back at out trips to Africa over the years, I realised we have seen five of the nine forms.

Monday 9 May 2022

Find Mr Aharoni and my Ostriches: Lord Walter Rothschild Sends the Future President of Israel on an Errand

Until I re-read Miriam Rothschild’s biography of her uncle, Lord Walter Rothschild, I had no idea that Israel Aharoni was one of Rothschild’s collectors in the field. Aharoni is of course known as the man who, in 1930, organised an expedition to find Golden Hamsters near Aleppo on behalf of Saul Adler for research on kala azar or leishmaniasis, a nasty and often fatal disease caused by a protozoan parasite. Adler was looking for a local species to replace the Chinese Hamster which had to be imported, proved difficult to breed and susceptible to a bacterial infection. Aharoni got his local guide, Georgius Khalil Tah’an, to talk to the local sheik. The latter hired labourers to dig holes in a local wheat field. Eventually, having destroyed the crop in much of the field, they found, eight-feet down, a complete nest comprising a mother and her eleven young. Until very recently, all the Golden Hamsters in laboratories and kept as pets were derived from those individuals.

Aharoni’s name appears in the Rothschild biography in relation to ostriches, and the involvement of Rothschild’s colleagues in the Zionist movement in getting them—well their skins—to the museum in Tring. After the delivery of the Balfour Declaration in 1917 on the establishment of a Jewish homeland in Palestine, Rothschild always on the lookout to further his zoological obsession decided it was time to involve the future president of Israel, Chaim Weizmann, then a chemist working in Britain, and Nahum Sokolow, also in London, in finding in Palestine two ostriches that he owned but whose whereabouts he had lost touch with during war. With nobody else able to go to Palestine on Rothschild’s behalf it was down to Weizmann to track down Aharoni when he was in the country in 1918; Sokolow apparently knew Aharoni. Weizmann eventually succeeded in finding Aharoni and the two ostriches in October 1918. Miriam described what happened next:

…by the end of November the schoolmaster/naturalist [Aharoni] was again corresponding with Walter. The ostriches were safe and within a year had arrived at Tring. The chick had grown up and in 1919 described it as a new sub-species, Struthio camelus syriacus Rothschild. It was one of the skins retained at the Museum when the bird collection was eventually sold to America.

Later, Miriam added in a footnote, Aharoni, ‘lost his head and began demanding exorbitant sums of money for further specimens. Matters were smoothed over and good relations re-established and he continued collecting for several years for the Tring Museum’.

It was actually the skins of the two birds that arrived at Tring. This is what Walter wrote in the description of the birds as a new subspecies. The paper was given to a meeting of the British Ornithologists’ Club at Pagani’s Restaurant in Great Portland Street, London, on 14 May 1919:

The fact that Ostriches inhabited the Syrian Desert and Arabia has been known for a long time—in fact, there are several passages in the Bible relating to this bird...Some years ago, Mr. J. Aharoni received a number of Ostrich eggs from his Arab hunters from the Syrian Desert, and sent them to Tring. These eggs agree with those of the North-African Ostrich in being smooth and not pitted, but are much smaller and more highly polished than those of that bird. I at once urged Aharoni to procure for me some adult Syrian Ostriches. He managed to procure a pair of nearly full-grown young ones alive, and as soon as they were fully adult he skinned them and sent them to me. 

It was not surprising that Rothschild was concerned about finding his ostriches. Already known to be rare by the early 1900s, sightings and remains of what came to be known as the Arablan Ostrich became more and more infrequent. The subspecies is now considered extinct with the last record—of a dying individual—in 1966.

Aharoni went on to find other specimens of the ostrich: in 1927 a female and two chicks were obtained in Saudi Arabia. They are in the museum (see below) in Tel Aviv.

Aharoni was collecting for Rothschild at least by 1909. The eponymous Aharoni’s Eagle-owl was described as a new subspecies, Bubo bubo aharonii, by Rothschild and his curator at Tring, Ernst Johann Otto Hartert, in 1910. They wrote:

We have received one pair from Mr. Aharoni in Jerusalem. They were obtained on the Wadi Suenit, in the valley of the Jordan in Palestine, on April 5th or 6th, 1909…We have named this most interesting form in honour of Mr. Aharoni, who is an enthusiastic naturalist.

It would appear there has been confusion in the past over the use of the common name, Aharoni’s Eagle-owl with the scientific name of another ‘new’ subspecies that Rothschild and Hartert named in the same paper, Bubo bubo interpositus. At some stage, the two subspecies must have been lumped but why into interpositus and not into aharonii I do not know. I have not been able to find when the two subspecies were lumped together and why the common name of one was linked with the scientific name of the other. It is, however, easy when looking up B. b. interpositus in the original paper to overlook the description of B. b. aharonii that comes later. An example of the result of this confusion was in Whose Bird published in 2003—but sorted out in The Eponym Dictionary of Birds that appeared in 2014.

Incidentally, Aharoni’s initial is sometimes shown as J (by Germans like Hartert) or Y, rather than the english English, I. for Israel. The name is given as J Israel ben A. Aharoni in the Eponym Dictionary of Birds.

Israel Aharoni (1882-1946) had arrived in Palestine in 1901 (or 1902 in other accounts). He was born in Vidzy, now in Belarus. His father, a rabbi, died before Israel was born, and his mother when he was two. Brought up by his grandmother, he apparently fled but then studied at Prague university. Accounts online of his life vary slightly but it would seem that he was a schoolteacher in Rehovot before in 1904 he moved to Jerusalem. There he taught languages at a school and an art school. With specimens from his many research and collecting trips in the Middle East, as well as his writing on ‘biblical zoology’, he established a zoological museum in 1925 under the auspices of the World Zionist Organisation. The collection is now housed in the Steinhardt Museum of the Hebrew University in Tel Aviv. He also lectured at the Hebrew University.

During the First World War, Aharoni served the Ottoman army as zoologist, apparently in anti-locust control. For six months in 1915 a swarm of locusts devastated Palestine, Mount Lebanon and Syria. It is perhaps not surprising that Aharoni, then working for the enemy, was not in contact with Rothschild. On the collapse of the Ottoman Empire, Aharoni continued with the same work but with the British army until 1921. That would explain why Weizmann was unable to contact Aharoni until October 1918.

Aharoni's good relations with the Ottoman rulers and later with the non-Jewish populations of Palestine and surrounding countries enabled him to travel more easily and to recruit the help of local bigwigs on his expeditions, as he showed with his search in Syria for Golden Hamsters. Earlier he had explored the area of the Dead Sea and Transjordan. The butterflies he collected were to presented to a Sultan of the Ottoman Empire. He travelled and collected in what are now the countries of Israel, Jordan, Turkey, Iraq, and Saudi Arabia.

In Haasiana, a biennial newsletter of the national natural history collections of the Hebrew University published in 2014, the bird collection which was begun by Israel Aharoni with birds he collected in 1906 is described:

Aharoni was an avid naturalist and a multi-lingual expert in near east and other (e.g., East European) languages. In his autobiographic book, “The Memories of a Hebrew Zoologist” (1943), he stated that “Even before my arrival to the land of Israel, my double life-long goal was clear to me: a) The study of wild animals in their natural habitat, in the birth place of each one of them, and b) The study of the original name of each creature, whom the desert dwellers (who live on their hunting and did not change their culture and way of life since the days of “Abraham” our father) are calling each living animal known to them”…

…Aharoni was responsible for important zoological discoveries, specifically (but not limited to) the region’s avifauna. Aharoni collected many Northern Bald Ibises (Geronticus eremita) discovered in the Syrian Desert. This species is nowadays at the brink of extinction, surviving in the wild only in two locations: in southeastern Turkey under a semi-natural setting (the birds are captured during winter to reduce their mortality), and in southern Morocco in several colonies consisting of about 500 birds. In 2002, a tiny population of less than ten individuals was located in the Syrian desert in the exact same locations where Aharoni had traced the birds a hundred years ago… Yet, since this re-discovery, the population has further declined and is now believed to have become extinct…

When the Hebrew University of Jerusalem was founded, Aharoni was already a renowned teacher and scientist and, consequently, he became one of the first professors of the newly established university. He founded a collection that was meant to become a zoological museum of local wildlife at the university’s campus in Mt. Scopus. Aharoni wrote several books for students of zoology, providing important contribution to the development of the study of local zoology. His autobiographic book provides ample information about his teaching and, especially, his expeditions in the Middle East, as well as a rich description of different aspects of his personal life during several decades since his immigration to Israel. His animal collection was maintained by himself with the help of his daughter, Bat-Sheva Aharoni, a scientist in her own right. 

Israel Aharoni (holding the skull of a roe deer)
with his daughter Bat-Sheva. ca 1940

Anon. The Bird Collection, including the collection of Israel Aharoni. 2014. Haasiana No 7. July 2014. p 3-9.

Beolens, B. & Watkins, M. 2003. Whose Bird? London: Christopher Helm.

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

Rothschild M. 1983. Walter Rothschild. The Man, the Museum and the Menagerie. Balaban (Paperback published in 2008 by the Natural History Museum, London).

Rothschild W, Hartert E. 1910. Notes on eagle-owls. Novitates Zoologicae 17, 110-112.

Rothschild W. 1919. Description of a new subspecies of Ostrich from Syria. Bulletin of the British Ornithologists' Club. 39, 81–83.

Thursday 5 May 2022

Clashing views on how to recognise species in the 1950s. The case of the Treefrogs

By the middle of the 20th century the museum-based classical zoologists were being looked down on. Their traditional pursuit of describing, naming and cataloguing species that were believed to be newly discovered, or of revising classifications and taxonomy of whole groups of animals, was—and indeed in Britain still is—most definitely out of fashion. The classical museum zoologists, with some notable exceptions, had become wedded to the notion that only morphological characters could and should be used in systematics and taxonomy. What was once a necessary approach as the only one available had become a dogma.

Deryk Frazer, writing in 1983, provided a wonderful example in his recollections:

I remember a scientific meeting of the Zoological Society, where their secretary Lord Chaplin[*] gave a paper on the European and Mediterranean treefrogs (i.e. Hyla arborea†, meridionalis and savigny) where he covered their distribution and probable areas of origin, appearance—including differences in profile of nose and mouth which could only be seen in living individuals, and recordings of the mating calls. His conclusions were quite inescapable, that they were two distinct species, one of which was separated into two subspecies. One learned zoologist then rose to his feet and said that he was not prepared to accept this, because you could not distinguish dead individuals from one another. Nowadays we have the added weapons of sound spectrogram, chromosome picture and the results of hybridisation, for a start.

The attitude of the ‘learned zoologist’ with dead individuals I see being repeated by those of molecular persuasion, however incomplete, inappropriate or misleading their limited data on mitochondrial or nuclear DNA. Both the traditionalists and their modern counterparts remind me of the story of the drunk searching for his house keys under a street light. ‘Was that where you lost them?’ the passerby asked. ‘No, it’s the only place I can see’ was the reply.

This treefrog (Hyla arborea) we saw in Hungary in 2010
decided a shoulder was the ideal place to land

*Anthony Freskyn Charles Hamby Chaplin, 3rd Viscount Chaplin from 1949 (1906-1981) was Secretary of the Zoological Society of London from 1952 until 1955. An amateur zoologist and composer he had a miserable time as Secretary. He was in office when George Cansdale was sacked as superintendent of London Zoo. The presentation to the Society must be that given by Chaplin and Jack Lester on treefrogs and other amphibians which appeared in the account of meetings (Proceedings of the Zoological Society of London 124, 197-197, 1954).

†Recently ‘split’ into four species.

Frazer D. 1983. The British Herpetological Society—a reminiscence. British Herpetological Society Bulletin No 8 December 1983, 10-12.

Tuesday 3 May 2022

A Frog at Butrint in southern Albania: What was it?

Enjoying the archaeological site at Butrint—Greek settlement, Roman city, bishopric, Byzantine and then Venetian, but abandoned in the late Middle Ages—at the southern tip of Albania, greater treats of a herpetological kind were in store. The flooded basements of the ruined buildings are home to two of the three species of terrapin in Europe, the European Pond Terrapin (Emys orbicularis) and the Balkan Terrapin (Mauremys rivulata). In another ruin we spotted a frog—the frog shown in the photograph. At the time (April 2010) I looked it up in the 2nd edition of the Collins Field Guide (2002 and 2004, London: Collins) and wrote it down as a Greek Marsh Frog, Rana balcanica, found in southern Albania, southern Macedonia, south-western Bulgaria and Greece. Nick Arnold, the book’s author, noted though that this species is also known as Rana kurtmuelleri and ‘sometimes not regarded as distinct from Marsh Frog’, i.e. Rana ribibunda.

Frog photographed in the archaeological site of Butrint, April 2010

Since the early 2000s the water frogs have been moved into their own genus, Pelophylax. Therefore the Marsh Frog became Pelophylax ribibundus. The specific name balcanica for the Greek Marsh Frog aka Balkan Water Frog has also gone in favour of P. kurtmuelleri because P. balcanica was quickly determined to be the same species as that described by Gayda in 1940. Gayda’s name had priority. According to the The Eponym Dictionary of Amphibians (Beolens, Watkins, Grayson, 2013, Exeter: Pelagic Publishing) nothing seems to be known of who Kurt Müller was except that he collected the holotype in Albania in 1938.

But is P. kurtmuelleri a valid species, distinct from Marsh Frog, P. ribibundus? One group has contended that it is not, and seeing one member of that group was a main author of the Field Guide to the Amphibians & Reptiles of Britain and Europe (Speybroeck, Beukema, Bok van der Voort, London: Bloomsbury) which was published in 2016, it not surprising that the species is not listed other than to note it had been lumped into Marsh Frog.

However, in fact, lots of others have retained P. kurtmuelleri as a valid species. A key example is the influential Amphibian Species of the World. That site contains a very brief summary of, and a link to, each paper that has appeared on the species. For that reason, I am not providing a list of sources at the end of this article.

I find it very difficult to decide on the basis of current evidence who is right since the status of the water frogs of south-east Europe and south-western Asia is both confused and confusing. However, I am disquieted by those who dismiss studies other than those obtained using the limited analysis of mitochondrial DNA (in this particular case they are those lumping the Balkan Water Frog into Marsh Frog) while dismissing biological evidence which could indicate the Balkan Water Frog should be split from Marsh Frog.

It was Schneider, Sinsch and Sofianidou in 1993 who proposed that their balcanicus, later recognised as kurtmuelleri, should be recognised as a species distinct from the more northerly ribibundus. The grounds for the split was biological evidence: the mating calls of the two forms were different. This is part of the summary of their paper:

The mating call of the lake frogs from Thrace resembles in all parameters that of the Rana ribibunda in the terra typica restricta (Guryev, CIS). Accordingly the lake frogs of eastern Greece belong to R. ridibunda. The mating call of these lake frogs consists of 20 pulses/pulse group and of 7 pulse groups/call on the average. Most of Greece is inhabited by the second taxon, Rana balcanica sp. n. [P. kurtmuelleri] Its mating call is characterized by 27 pulses/pulse group and 4 pulse groups/call on average.

In addition to the analysis of the mating calls, small but statistically significant differences were evident in morphological features when the measurements they made were subjected to discriminant analysis.

One bit of possible evidence to suggest the Balkan Water Frog is a valid species was raised by Schneider and his colleagues. In the early 1980s some work had been done on crossing Marsh Frogs proper with water frogs from what was then southern Yugoslavia, which, they suggest, were in all probability, Balkan Water Frogs. The resulting tadpoles had a high mortality and those that few that did survive to maturity, a low fertility, both indicative of hybridisation between two, rather than within one, species.

Overall, as a ‘lumper’ I can see why the ‘splitters’ have a case in recognising the Balkan Water Frog, Pelophylax kurtmuelleri, as a species, even though even more research is still required.

But back to Butrint in 2010. Was the frog I photographed a Balkan Water Frog? There is another species of Pelophylax living in the southern tip of Albania, the Epirus Water Frog, P. epeiroticus. Both species are highly variable in appearance and hybrids between the two have been found. The Epirus Water Frog was only described (by essentially the same group that described balcanica (kurtmuelleri)) in 1984. A group of Slovakian and Czech zoologists have illustrated the differences in morphology and particularly in coloration within and between the species of water frogs that occur in the south-western Balkans. Looking at the photographs and reading the distinguishing features, it appears to me that the frog in my photograph (not ideal because of the reflections obscuring the tympanum) shows characteristics of the Balkan Water Frog, P. kurtmuelleri. The habitat preferences of the Epirus Water Frog (which is apparently common on the edges of the nearby Lake Butrint) given in the 2016 Field Guide also indicate that the flooded basement of an ancient building are not likely to be inhabited by P. epeiroticus

I thought a second visit to Butrint, this time in 2017 and also in mid-April, would afford me another look at the frogs within the archaeological site. The terrapins were there but not a frog to be seen. The area I had seen the frog in 2010 was nearly dry but for one corner which had a lot of large tadpoles.