Frogs of China. Alice Boring’s Life and Work. 2. Legacy: Liu Cheng Chao

Alice Boring made good use of her time on long leave in 1928-29. From correspondence with the leading American herpetologists of the day and by working at the American Museum of Natural History in New York she had, by the time she sailed back to China, compiled a checklist of amphibians from all parts of China. The checklist was published with Nathaniel Gist Gee (1876-1937)*. According to her biography she was instilled with what was then the American way of doing taxonomy: collecting a wide range of specimens over a wide area in order to determine the degree of variation within and between species. She established a network of people who sent her specimens as well as sending students and collectors on collecting trips to various parts of China. Because museums depend on access to specimens she sent and exchanged amphibians for study

Geoffrey Herklots at the University of Hong Kong, was, to go right back to the opening paragraph of Part 1, part of that network that existed throughout China and to the U.S.A. He sent collections of frogs to Peking for her to identify. In turn she published the findings in Hong Kong Naturalist.

Boring was a founder member of the Peking Natural History Society and it was in the the Society’s bulletin and a handbook that she published most of her work. However, it was not all taxonomy as some biographers have implied. One paper was on Bidder’s organ while another was on seasonal changes in the reproductive organs of frogs and toads, both continuing the sort of work she was doing in Maine before she left for China.

Liu Cheng Chou
from Contributions to the History
of Herpetology

Boring’s major legacy was a student Liu Cheng Chao (1900-1976). Not officially her student, Alice Boring provided advice and encouragement for Liu’s interest in amphibians at Yenching. After graduation, Liu lost his herpetological papers and books in 1931 when forced to leave Northeastern University at Mukden when the Japanese moved in. Boring took great trouble in  arranging for him to study for a Ph.D. in the U.S.A. funded by the Rockefeller Foundation. He went to Cornell to work with Albert Hazen Wright (1879-1970). Back in China, Liu was first at Suzhou University but the Japanese advance took him to Chengdu in Sichuan and the West China Union University. It was from there that he made 11 long field trips to the hills and mountains on the edge of the Tibetan Plateau. On one he was seriously ill with typhus fever. In September 1946 Liu returned to the U.S.A. and was based at what was then called the Chicago Natural History Museum, now the Field Museum of Natural History.

Liu was with some big names of classical herpetology in Chicago: Karl P. Schmidt, Clifford H. Pope and Robert F. Inger. Pope, of course, had travelled extensively in China and written a book on the reptiles. During his year there Liu wrote his major work Amphibians of Western China. Much more than a catalogue of amphibians it describes the field trips, the peoples he encountered and the geography. He discusses the adaptations of adults and tadpoles in relation to habitat as well as the history and cultural significance of the various species. It is not surprising that after this tour-de-force Liu was described as China’s most prominent herpetologist.

By the time Amphibians of Western China was published in 1950 (Liu’s Preface was dated 1 November 1948) he had returned to Yenching University as head of department. In 1951 he moved to Chengdu as President of Sichuan Medical College. He became a member of the Chinese Academy of Sciences. He and his wife Hu Shu Chin (Shu-qin) (1914-1992) later published Chinese Tailless Amphibians in 1961.

Liu acknowledge the part Boring had played in his career in Amphibians of Western China:

My former teacher at Yenching University, Dr. Alice M. Boring, has my grateful remembrance for her continued encouragement and help during the war years. 

In 1945 Liu named a spectacular amphibian, Vibrissaphora (now Leptobrachium) boringii in Alice Boring’s honour. This species is known as the Emei Moustache Toad because the males in the breeding season have sharp tubercles around the upper lip. These spines are used in defence of their nests. A frog full of vim and vigour named for a ‘dame full of vim and vigor’. And, surely, it should have the common name, Boring’s Toad?

A plate from Liu's book showing the species named for Alice Boring
I have added a red arrow to show the sharp tubercles

Another plate from Amphibians of Western China

A modern photograph of Boring's Toad
from Hudson & Fu 2013

*Yenching University had, incidentally, originally lined Gee up for Boring’s job when he was free to take it. That is why she was only offered a two-year appointment.

Anon. 2014. Boring, Alice M. In Contributions to the History of Herpetology (Volume 1, revised and expanded), Edited by Kraig Adler, pp 107-108. Society for the Study of Reptiles and Amphibians.

Anon. 2007. Hu, Shu-qin (1914-1992). In Contributions to the History of Herpetology (Volume 2,), Edited by Kraig Adler, pp 207-208. Society for the Study of Reptiles and Amphibians.

Anon. 2014. Liu, Cheng-chao (1900-1976). In Contributions to the History of Herpetology (Volume 1, revised and expanded), Edited by Kraig Adler, pp 123-124. Society for the Study of Reptiles and Amphibians.

Fu L. 2017. Nathaniel Gist Gee’s contribution to biology in modern China. Protein & Cell 8, 237-239 DOI 10.1007/s13238-016-0318-x 

Hudson, CM, Fu, J. 2013. Male-biased sexual size dimorphism, resource defense polygyny, and multiple paternity in the Emei Moustache Toad (Leptobrachium boringii). PLoS ONE 8(6): e67502. doi:10.1371/journal.pone.0067502 

Liu C-C. 1950. Amphibians of Western China. Fieldiana: Zoology Memoirs Volume 2. Chicago: Chicago Natural History Museum

Ogilvie MB, Choquette CJ, 1999. A Dame Full of Vim and Vigor: a Biography of Alice Middleton Boring: biologist in China. Amsterdam: Harwood.

Zheng. 2015. Alice M. Boring: a pioneer in the study of Chinese amphibians and reptiles. Protein & Cell 6, 625-627 DOI 10.1007/s13238-015-0165-1 

Frogs of China. Alice Boring’s Life and Work. 1. Biography

As we explored the Hong Kong University Compound after our arrival in November 1965 we came across the lily pond. Seeing the occasional plop as a frog jumped into the water, we decided to continue the exploration in the evening. As well as collecting two nightwatchmen as spectators, several frogs were caught. We had no idea what they were so the next day tried to identify them. Fortunately, there was a tiny collection of offprints in the old zoology department’s office. Some were the checklists given by John Romer but others were from the Hong Kong Naturalist series of the 1930s. How those offprints had survived or had been obtained later we had no idea since the then newly-opened science building had been stripped of everything during the Japanese Occupation. However, they were there and as well as identifying the frogs as Günther's frog, Sylvirana (formerly Rana and Hylarana) guentheri, we became acquainted with the name of the author of the key paper, Alice M. Boring.

Alice Middleton Boring*

Only a couple of years ago when I read her short biography in Contributions to the History of Herpetology did I realise that she was the doyenne of amphibians in China. I also found a book on her life had been published in 1999, A Dame Full of Vim and Vigor. That provides a highly detailed account of her entire life together with background on political and military events unfolding in China during the 20th century. After reading it I see why the authors chose that title.

Alice Boring, as well as living in China in interesting times—very interesting times—was, in the early decades of the 20th century, part of the cutting edge of biological research in the U.S.A. Then, in China, she realised that it would be impossible to pursue such studies and turned to describing and cataloguing the amphibians of China. It is for these latter works of scholarship that she is still remembered today, rather than for the original research on cytogenetics with world leaders in their fields of her earlier years.

From a school in Philadelphia and part of a largely Quaker family, Alice Middleton Boring, gained admission to the Quaker-founded women’s college Bryn Mawr in Pennsylvania. Teaching at the college were Nettie Stevens (1861-1912) who amongst some confusion about credit at the time appears to have discovered sex chromosomes. There was also Thomas Hunt Morgan (1866-1945) who went on to win a Nobel Prize for his work on genetics. After a Ph.D. under Stevens she worked in Würzburg (with Stevens) where they annoyed Theodor Boveri as “the purest blood suckers” (i.e. visiting scientists who visit for a very short time, take up the host’s time and resources and emerge with only superficial findings) and then in Naples.

Returning to the U.S.A. in 1910, Boring got her first job in the agricultural college of the University of Maine where her boss was Raymond Pearl (1879-1940). It would take a book to describe Pearl’s life and career, his many wrong-headed ideas as well as some better ones. Suffice it to say that Boring collaborated with Pearl on reproduction and the plumage pattern of domestic chickens. She advanced rapidly through the ranks to be Associate Professor by 1913. Then, in 1918 at the age of 35, she upped sticks for China on a two-year appointment.

The job she went to was a new one. Rockefeller money had bought out the old Union Medical College in Peking from the joint control of British and American missionary societies and renamed it the Peking Union Medical College. There was though pretty open warfare between the missionaries and Rockefeller employees even though in the end the board that ran the college had input from both. Boring arrived at the time of expansion into a new pre-clinical school. She was fiercely anti-missionary, taught there for two years and at the end of her contract returned to the U.S.A. as professor of zoology at Wellesley College.

Boring was apparently frustrated by Wellesley. Zoology was treated as a poor relation. She entered into negotiations to return to China. It was intended that a new university organised on American lines, Yenching, formed by merging various colleges in Peking (I will stick to the Wade-Giles transliteration instead of the Pinyin ‘Beijing’) should act as a pre-clinical feeder to the Peking Union Medical College, and should take over that function entirely from the College. The university itself was eager not to give Boring a long-term position since they had somebody else lined up for the permanent job. Eventually, she was reassured about her many concerns by the head of the university, John Leighton Stuart (1876-1962) and after making sure she would not be on a missionary’s salary, and that the position at Wellesley would be kept open for her return, she was all set to go: “It would seem like a bit of real contribution to civilization as opposed to teaching in America”. She did not return to Wellesley, preferring Chinese students eager to learn to American young ladies keener to acquire a Mrs as a pre-nominal than a bachelor’s degree after it.

Alice Boring arrived at Yenching University in 1923. She  landed at a time of considerable political and military turmoil which continued until and after her final departure 27 years later. Like her boss, Stuart, Boring took an anti-Western stance. Anti-missionary initially, her position became more nuanced when she argued that the christian groups were also opposed to imperialism and capitalism. She—and Stuart who was born in China, the son of American missionaries—supported the political aspirations of the students, as anti-Western and anti-Japanese rioting flared up and as battles raged between warlords.

Yenching University ca 1937*

Boring was remembered as an austere and tough but kindly task-mistress in preparing students for admission to the medical school. Reading the accounts of her student laboratory sessions with regular tests, my impression is one of overteaching and more like a school class than what would have been expected at the time in an English university. She certainly demanded high standards but also ensured the students were looked after and became proficient in the couth and culture required to move in Western circles and to spend time in the U.S.A. for postgraduate studies. Students who succeeded clearly thought highly of her, both for her knowledge and as a mentor even though she was known by students of both sexes to favour the men. That she was prim, proper, fierce and opinionated there is no doubt. But beneath the prim exterior was something very kind and a little more wild. On a camping trip in Mongolia soon after her arrival in China she wrote: ‘…we take shower baths in the morning by rolling naked in the long wet grass’. Her brother wrote of her as enthusiasm as a ‘menace to society’.

Teaching clearly occupied a great deal of her time (she was in her green lab coat and spectacles nicknamed the ‘Green Grasshopper’ by her students) in Peking. Research was a different matter. She decided that it would be impossible to continue her interest in what was then seen as modern biology. Instead she turned towards the description and taxonomy of the amphibians and reptiles of China. During a long leave at the University of Pennsylvania in 1928-1929 she was in contact with the leading herpetologists of the day to learn all she could about the field and what she needed to do in order to become proficient. At the same time she avoided most of her old acquaintances in cytology and genetics who would undoubtedly have looked down their noses at a move to the world of dead frogs and museums. Pearl though, with whom she was in contact, was supportive. She had already met and helped Clifford H. Pope (1899-1974) on one of his numerous expeditions to China. Further assistance and support was obtained from G.K. Noble (1894-1940) and Albert Hazen Wright (1879-1970).

Boring returned to Yenching to enjoy the new laboratory she had designed and the formal opening of the university. However all was not sweetness and light. First there was the need to raise cash after the Wall Street Crash of 1929; Boring helped but reluctantly. Second there was Stuart, a widower since 1926, who attracted a coterie of female fans. Boring emerged as top dog after what can only be described as a cat-fight with her former friend and house-mate Grace Boynton who taught English. While Boynton was on leave, Boring began to have Sunday dinner with Stuart. When Boynton returned she joined them. Boring asked her to leave. The resultant mutual loathing lasted for many years even though in the end they came to some form of accommodation if not renewed friendship. Students saw Boring and Stuart walking hand in hand over the bridge on campus and tongues began to wag. One staff member noted that Boring became, in effect, social secretary and hostess to Leighton Stuart. She rode with him and travelled with him to make up a foursome on a camping trip. Members of the English department wrote a poem about her ‘something about the power behind the power behind the power behind the throne’. Whatever happened between them though, Stuart avoided permanent commitment. Indeed, during the Japanese occupation he brought one Alice Gregg to the campus. Gregg described herself as “just Stuart’s Shanghai Sweetie”. Grace Boynton described Boring as “frantic” but that Stuart “never seemed aware that AMB was green with jealousy”.

Leighton Stuart's house, Yenching University*

External political forces continued to have major effects on the university campus and student body. After Chiang Kai-shek became established as top warlord in 1930 to head a Nationalist government, Japanese encroachment and occupation of Manchuria was not met by firm action. Student protests demanding action against Japan erupted. Some faculty members backed the students; others Chiang Kai-shek. Boring met Madame Chiang (a former Wellesley student) and was impressed. Time after time, some incident sparked off protests and strikes. Eager to ensure the students did not waste all their time on patriotic protests, Boring once found herself on the wrong side of them when she attempted to hold a lecture during a demonstration.

1937 saw a major shift. Japanese forces pushed into China and occupied Peking. At the time Boring and other staff members of Yenching were on local leave 70 miles south of Peking. They made their way back via Canton and Hong Hong and the university carried on operating with the Japanese in occupation while to the south were the Nationalists and the Communists. Students from the south crossed the lines to and from Yenching, eager to get the best education in Yenching while becoming open to accusations of being unpatriotic. There was a degree of rapprochement between Boring and Boynton with the latter becoming the conduit for information from Yenching to Boring’s family in the U.S.A. since she chose to be evacuated (until returning to unoccupied China) while Boring was insistent on staying on.

After the attacks on Pearl Harbour and on Hong Kong, the foreigners in Yenching were interned, first on campus and then into houses left empty by American diplomats. Finally on 25 March 1943 they were moved to a camp—foul it goes without saying—in Shandong. However, she was not there that long because Alice Boring was down for repatriation in exchange for Japanese internees. She sailed with other Yenching staff on 24 August on board Teia Maru (crowded, dirty and short of food and water) for Goa, then belonging to neutral Portugal. A neutral ship, the Swedish M.S. Gripsholm then carried the 1,440 Americans and Canadians to Jersey City.

Alice Boring clearly felt it hard to be back in the U.S.A at the age of 60. Nobody, least of all her family, was really interested in her life in China and like so many long-term expatriates she remained wrapped up in Chinese affairs. She was a supporter of Chiang Kai-shek, even while recognising the shortcomings of his regime. A couple of temporary teaching jobs occupied some of her time. It is not surprising that she headed back to Yenching as soon as it was possible to do so.

Returning to Yenching in September 1946, exactly one year after VJ-Day, she set about gathering equipment for her course. But the civil war was close. Stuart was made U.S. Ambassador to the Chinese government—an episode that did not end well. The end of 1948 brought Nationalist forces retreating through the campus followed by very polite Communist soldiers who got on well with both staff and students. The initial soft approach of the new regime to Western interests hardened. Political events and family tragedies beckoned Boring home. She left Yenching in August 1950 and died on 18 September 1955, aged 72. Grace Boynton, Alice Boring’s rival for Leighton Stuart’s attention, was given the job best described as ‘historical executor’, sorting her letters and papers on her life at Yenching for archiving.

In Part 2 I will cover Alice Boring’s contributions to herpetology and her lasting legacy in China.

*Photographs from International Mission Photography Archive, University of Southern California

Ogilvie MB, Choquette CJ, 1999. A Dame Full of Vim and Vigor: a Biography of Alice Middleton Boring: biologist in China. Amsterdam: Harwood.

Anon. 2014. Boring, Alice M. In Contributions to the History of Herpetology (Volume 1, revised and expanded), Edite by Kraig Adler, pp 107-108. Society for the Study of Reptiles and Amphibians.

Philip Street—prolific author from the 1950s to the 1970s. Who was he?

Philip Street wrote a number of popular books, aimed at all ages, on animals, marine biology and conservation from the 1950s to the late 1970s. His books were very well received and were widely available in public libraries. However I could found virtually nothing about Philip Street even though one of his books, Shell Life on the Seashore, was reprinted to great acclaim in 2019. From his book, Whipsnade, he is shown as having the degree of MSc and was a Fellow of the Zoological Society. Leo Harrison Matthews, then Scientific Director of the Society wrote in an Introduction ‘he is a qualified zoologist’ and ‘a writer of experience’. The 2019 reprint states that he was a member of the Marine Biological Society, but I think this must an error. There is a Marine Biological Association, but not a society.

After searching genealogical sites and local newspapers, I have now discovered who Philip Street was and something about his life. Philip Arthur Richard Street was born 4 February 1915 in Harrow, Middlesex. He was educated at Harrow Weald Primary School, Harrow County School (now Harrow High School) and then University College London from 1933 until 1938. He graduated with a BSc in zoology in 1936 and with an MSc in genetics in 1938. In the 1939 Register he was shown living with his parents at 68 Woodlands Drive in Harrow as a schoolmaster with two years research experience in biology.

At some stage he taught at his old school but was appointed to teach biology at Kingsbury County Grammar School in 1945. In 1953 he was careers master there. In the 1950s he was living with his wife (married in 1945) at 6 Bellfield Avenue, Harrow.

By the late 1950s Philip Street was in Buckinghamshire, giving courses of 12 lectures for the Workers’ Educational Association on wild animals in captivity. In 1978 he was reported at the Chesham and District Natural History Society speaking on animal navigation. Philip Street died on 14 May 1984 at Aylesbury, Buckinghamshire, aged 69.

Philip Street’s books had—and with the recent interest in his republished one on seashells still have—a great influence in firing an interest in the natural world. Can anybody provide any further information?

Books by Philip Street:

Between the Tides 1952

Whipsnade 1953

The Seashore for Boys and Girls 1954

The London Zoo 1956

Animal Life 1957

Animal Partners 1958. Reprinted 2010

Shell Life on the Seashore 1961. Updated edition 2019

Mammals in the British Isles 1961

Vanishing Animals - Preserving Nature’s Rarities 1963

The Seashore 1965

Animals in Captivity 1965

Survivants de la préhistoire (French) 1966

The Crab and its Relatives 1966

Burst of Spring 1969

Wildlife Preservation 1970

Animal Weapons 1971

Animal Reproduction 1974

Animal Partners and Parasites 1975

Colour in Animals 1977

Animal Migration and Navigation 1976

Poison in Animals 1978

Why is the human sex ratio 1:1 at birth?

Having found a dramatic shift in sex ratio at birth in one species of mammal, I have for decades kept an eye on discussions as to whether the human sex ratio at birth differs in some circumstances from 1 male: 1 female. Mammals in theory have two ways of controlling the sex ratio at birth: (i) at conception, or (ii) later by selectively reducing the number of young in utero. We found the latter mechanism at work in the guinea-pig. Since human litter size usually equals the  number of eggs and rarely exceeds 1 and since gestation is relatively long, it always appeared that if there were to be any maternal or paternal genetic control of the sex of the offspring it would have to be at the time of conception, rather than by selective death and reabsorption of embryo or fetus.

In the 1980s I once did the experiment of asking biologists from different disciplines why they thought the human sex ratio is 1:1. The reproductive biologists replied that it was just the result of random segregation of the sex chromosomes. The sex of a human offspring depends on whether it inherits and X or a Y chromosome from its father. Random segregation will, on average, result in a 1:1 ratio. By contrast, the evolutionary biologists said that the sex ratio is explained by Sir Ronald Fisher’s Principle: with the sex of a subject to genetic variation, the the sex ratio will always stabilise at 1:1. I will not repeat the simple explanation that can be found here.

Shifts of the of sex ratio at birth in various animals has been explained in terms of the Trivers-Willard hypothesis: parents which possess a heritable trait that benefits the lifetime reproductive success of one sex will bias the sex ratio towards that sex. There have been various claims in studies of human populations along these lines; for example, male-biased sex ratios in taller, wealthier, high status parents with the offspring likely to be more successful in competition for a mate. However, such claims have been controversial because of the statistical methods used and the results have often not borne out when larger samples were taken from the population.

Both the Fisher Principle and Trivers-Willard rely on there being genetic variation in the sex ratio, in other words that a bias towards one sex or the other is heritable. Thus a key test is to look for heritability in a very large human population. On standard scale of 0 to 1, ‘0’ denotes that a trait is not heritable while ‘1’ all differences in a trait can be explained entirely by genetic variation. Many traits fall somewhere between those two extremes.

But what does determine human sex ratio? It would be predicted if sex ratio is a heritable trait then Fisher’s Principle would apply. By contrast, if it is not heritable then simple Mendelian segregation of the sex chromosomes would suffice as an explanation.

Over the years there have been all sorts of suggestions and claims that the tendency in a family to produce offspring completely or partially biased to one sex is hereditary, and that particular genes could be involved. However, these conclusions have been criticised because the sample sizes were small and the statistical inferences drawn were invalid.

A recent, important paper has tackled the problem by using data from the entire population born in Sweden in and after 1932. That was 3,543,243 individuals and their 4,753,269 children. The results of the analysis were clear. There was no evidence of heritability at all. The calculated heritability was 0. In other words, there was no need to invoke Fisher’s Principle since with no heritability there can be no Fisher.

The authors summed up their results:

In sum, all of our results are consistent with the simple explanation that variation in offspring sex ratio in humans is due to unbiased Mendelian segregation of sex chromosomes during spermatogenesis and unbiased fertilization. The slight excess of male births is likely to be due to a general difference in survival of male and female embryos in the womb, the reasons for which are not yet understood.

Looks like my reproductive biology colleagues were right. Pity I can’t tell most of them; they are long dead.

The question now, of course, is whether the same conclusion, that the human sex ratio at conception is simply the outcome of random segregation of the sex chromosomes, applies to other or to all mammals? And are the statistically-robust demonstrated shifts in sex ratios at birth in some species and in certain environmental conditions all the result of differential loss of embryos and fetuses in utero? I shall permit myself to guess that the answers are ‘Yes’ and ‘Yes’ even though, as with previous human studies, there have been claims that the answer to the first question is ‘No’.

Zietsch BP, Walum H, Lichtenstein P, Verweij KJH, Kuja-Halkola R. 2020 No genetic contribution to variation in human offspring sex ratio: a total population study of 4.7 million births. Proceedings of the Royal Society B 287: 20192849. http://dx.doi.org/10.1098/rspb.2019.2849

Peaker M, Taylor E. 1996. Sex ratio and litter size in the guinea-pig. Journal of Reproduction and Fertility 108, 63-67.

Fluorescent chameleons, salamanders, newts and frogs...and a budgerigar

An increasing number of animals are being found to fluoresce.  Fluorescence happens when a chemical fluorophore absorbs of shorter-wavelength light and them emits some of the absorbed energy as light at a longer-wavelength. In some biological examples, blue light is absorbed and green light is emitted

Why animals fluoresce is the subject of active research. Does the change to a more noticeable colour mean that fluorescence is sometimes involved in signalling to other animals of the same sex, or of  different sex, or of the same species, or to a predator, or to prey? Is it assisting in camouflage, the fluorescent emission matching, say, the emission from plants in the background?

Perception of fluorescence depends on the properties of the photoreceptors of the animal exposed to the fluorescence, so that in the case of proposed signalling between animals of the same species, it would be a requirement to demonstrate that the visual pigments of the eyes of that species can actually pick up the colour.

There is, of course, the possibility that there is no function of the fluorescence at all, that it is just a by-product of a particular molecule used as a pigment or for some other purpose in the skin or other tissue. This is the same argument as to why some creatures from the depths of the sea have bright colours; the production of brightly coloured molecules used by the animal are simply not selected against since there is no predator present that can see them.

A very well argued paper appeared in 2017 in Philosophical Transactions of the Royal Society by Justin Marshall (University of Queensland) and Sonke Johnsen (Duke University, North Carolina). The authors explained the background to explaining any function of fluorescence in terms of communication between individuals. They set out criteria that must be satisfied in order for such a role for fluorescence to be accepted. The Budgerigar, Melopsittacus undulatus, they found from descriptive and experimental studies, fulfilled all the criteria. In short, both males and females prefer to associate with potential mates that fluoresce.

Fluorescence characteristics of the budgerigar
(a,b) White light and UV fluorescent excitation
photographs of front and back of head showing
fluorescent cheek and crown feathers
from Marshall & Johnsen 2017

More recent studies have shown fluorescence from the bony tubercles of the skull of many species of chameleon—just one more feature added to the list of properties of those extraordinary animals. The phenomenon is particular evident in those chameleons living in humid forests ‘known to have a higher relative component of UV light’. The blue light emitted would be in sharp contrast to the brown and green colours reflected by the surroundings.

from Prötzel et al 2018

  

Even more recently came photographs showing fluorescence in a number of amphibians, this time a green emission from blue light (with some species showing emission in response to UV (ultraviolet)).

from Lamb & Davis 2020

The accumulating evidence for fluorescence in a wide range of animals (and plants) raises so many questions as to the molecular mechanisms involved and possible functions, that the only conclusion at present is the inevitable: more research is required.

In the meantime enjoy the pictures.

Lamb JY, Davis MP. 2020. Salamanders and other amphibians are aglow with biofluorescence. Scientific Reports 10, 2821. https://doi.org/10.1038/s41598-020-59528-9 

Marshall J, Johnsen S. 2017 Fluorescence as a means of colour signal enhancement. Phil. Trans. R. Soc. B 372: 20160335. http://dx.doi.org/10.1098/rstb.2016.0335 

Prötzel D, Heß M, Scherz MD, Schwager M, van’t Padje A, Glaw F. 2018. Widespread bone-based fluorescence in chameleons. Scientific Reports 8, 698. DOI:10.1038/s41598-017-19070-7 

Peter Charles Howard Pritchard 1943-2020. An encyclopaedic knowledge of chelonians and their conservation. How did he start out?

The study and conservation of chelonians—turtles, terrapins and tortoises—has lost a major champion with the death on 25 February of Peter Pritchard at the age of 76. He and his Chelonian Research Institute were better known in the U.S.A. and the lands where chelonians breed than in his native United Kingdom. An appreciation of his work, influence and importance can be read on the website of Turtle Conservancy.

From Turtle Conservancy's website

   

I never met Peter Pritchard and often wondered how he had become interested in reptiles. Then I read an appreciation (on what is now a dead link) he had written of Angus d’Albini Bellairs (1918-1990):

Angus was the first herpetologist I ever met. He was the immediate successor to my father as Reader in Anatomy at St. Mary’s Hospital, my father (Dr. J. J. Pritchard*) having been appointed Professor of Anatomy at Queen’s University in Belfast, Northern Ireland the preceding year, and this connection gave me free access to a man I regarded as an intellectual hero.  My (signed) copy of his 1957 book, simply named Reptiles, is dated April 5 1958, when I was just 14 years old, and just a few years later I started to write a book of my own, which I ambitiously entitled Living Turtles of the World.  Despite the clearly schoolboyish flavor of this early draft, not to mention the lack of personal field experience and shortage of library access, Angus introduced me to the concept of peer review (although we were not exactly peers), and he read the whole thing, making gentle suggestions in pencil wherever he saw fit. 

Whenever I was in London, I would find my way to the dusty chambers of St. Mary’s (made famous by Sir Alexander Fleming), and knock on Angus’ door for a conversation on the subject of mutual interest, namely herpetology.  At such times, he would always open a bottle of sherry and bring some small-size laboratory beakers from which we would drink it, as he urged me to pursue an experimental approach to herpetology, by means such as studying underwater respiration in softshell turtles, or scute regeneration in chelonians.  (Somehow, I never became “experimental”, instead concentrating on natural history, taxonomy, skeletal anatomy, and conservation aspects).

Angus also introduced me to other herpetologists, including Miss Grandison, the Keeper of Zoology at the British Museum, a lady who had seemed rather remote and doctrinaire when I wrote to her, but was more like a favorite auntie once encountered in person.  I was also privileged, through her, to meet J. C. Battersby, on his very last day before retiring in 1961; he had been a “boy attendant” of G. A. Boulenger, no less, appointed in 1916.

Angus introduced me to the bizarre militaristic hierarchies at the London Zoo (Regents’ Park), where the “gentlemen officers” (the Curators etc) lorded it over the non-commissioned ranks (head keepers and below), only the former being admitted to such places as the Fellows’ Restaurant.  He himself was, of course, “top of the heap,” a scholar and a gentleman, although unpaid in status as “honorary herpetologist,” and I think his extensive wartime military experience was what prompted him to refer to the Reptile House staff as his “sergeant major,” his “corporal,” etc… 

And that’s how it all began, although his interest in animals was first piqued by being taken as a small boy to London Zoo. Even though his first degree was in chemistry at Oxford, he moved to Florida to work for his Ph.D. with Archie Carr (1909-1987), the doyen of research and conservation of turtles.

The book Pritchard referred to was his Living Turtles of the World, published in 1967—when he was 24—by the infamous Herbert R. Axelrod’s TFH Publications. A completely rewritten survey of the world’s chelonians stretching to nearly 900 pages, also published by TFH, appeared as Encyclopedia of Turtles in 1979. Axelrod himself took a number of the photographs. I have a copy—still a useful reference—on my shelves.

The following video on, and including Peter Pritchard, appeared in 2016.

*Peter Pritchard’s father. John Joseph (Jack) Pritchard. was born in 1916 in Adelaide, South Australia. By the age of 19 he had a first degree and arrived in Oxford on a Rhodes Scholarship to work at Oxford in the Department of Physiology. After becoming medically qualified in London he moved into anatomy, first at UCL, then St. Mary’s before moving to Belfast as Professor. His research was on bone growth and repair, a subject the reader will notice of considerable interest and importance in the life of chelonians. He died in 1979, aged 63.