Joan Procter, Arthur Loveridge and the Pancake Tortoise. 2. Joan Procter in London: Structure of the shell and the question of defensive ‘inflation’

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Not content with gross anatomical description, Joan Procter threw whatever modern technique she could find to discover as much as she could about the Pancake Tortoises, now known as Malacochersus tornieri, sent to London by Arthur Loveridge, the dead ones at the Museum and the live ones at the Zoo. She used X-rays and fluorescent screen x-ray equipment provided by the surgeon Sir John Bland-Sutton (1855-1936) of the Middlesex Hospital. He was a keen supporter and vice-president of the Zoological Society of London.  She collaborated with Richard Higgins Burne (1868-1953, elected FRS 1927) who was Physiological Curator at the Royal College of Surgeons on the structure of the jaw. Burne was also a key member of the Zoological Society in the 1920s and 30s.

Boulenger had commented on the first specimens sent by Loveridge to confirm the pliable nature of the carapace and plastron. Instead of solid bone underlying the epidermal shields as in other tortoises, Joan Procter found areas with no bone at all, especially, as suggested by Loveridge in the centre of the plastron. Deep sutures between the shields also indicated a marked degree of mobility.

Figures from Joan Procter's paper showing the bony
carapace with the large areas (hatched) lacking bone

The bony plastron showing the lack of bone in the
large central area

In all, Miss Procter was able to work on 23 dead specimens, preserved in spirit, as well as handling the two live ones at the Zoo. Another unusual feature, apart from the areas of the carapace and plastron without bone, the tortoise is the appearance of teeth on both jaws. These teeth which are not true teeth are part of the jawbones with an overlying continuous horny sheath.

By studying young specimens and comparing the development of the carapace and plastron in other tortoises she concluded that the adult Pancake Tortoise resembles to a great extent the young of other species in which the bone then continues to grow and fill the gaps. In other words, the fenestration seen in the adult is caused by arrested development of the bony shell rather than by breakdown of a complete bony structure.

Procter noted the great variability in the shells of the Pancake Tortoise—the subject of a recent paper describing the variation between individuals in more detail.

A question that Joan Procter addressed, whether the bone of the carapace is derived from the skeleton or from the skin I will not deal with further here but will return to in the future since the question has been the subject of research for getting on for two centuries and there is recent work suggesting that old views are wrong. However, before turning to a functional problem it is worth considering how Procter argued on the mechanism evolution of the shell of the Pancake Tortoise. It is difficult for those of us who first got to know the history of theories of evolution in the 1950s to appreciate just how common Lamarckian explanations were in the 1920s or of how powerful and combative some of the individuals like Ernest MacBride FRS, who rejected both natural selection and modern genetics until his dying day, were in the zoological circles of London. Joan Procter sat on the fence:

     It can be argued on the one hand that the flattened carapace is brought about by the habit of living beneath stones and squeezing into rock-crevices. This habit, induced by environment, would be bound to have a modifying effect; for, during youth, the development of a domed and solid carapace would be interfered with by the constant application of pressure, and in a sufficient number of generations the ability to form a normal carapace might be lost altogether. The fact that the Burrowing Tortoise, T. polyphemus, has a thin or fenestrated and somewhat flattened carapace supports this view. Could this be proved experimentally, it would furnish a convincing argument in favour of the heritance of acquired characters. 

     On the other hand, it can be equally well maintained that an inherited tendency to the arrest in development is orthogenetic, brought about either gradually or as a mutation, and that the furtive habit of hiding beneath stones was the natural result, since the tortoise no longer possessed adequate protection from enemies. 

     Possibly both principles come into play, the reduced armour and loss of ribs being orthogenetic, and the depression and relative condition of the vertebrae being subsequently induced by the rockdwelling habit.

Apart from the developmental origin of the bony shell, the subject that has stirred interest in the Pancake Tortoise has been the question of ‘inflation’.

The suggestion that Pancake Tortoises inflate to jam themselves more effectively into crevices between rocks came from Loveridge (see previous post of 22 November 2018):

The tortoise takes full advantage of this flexibility, as I soon found on trying to remove one from beneath a boulder. It inflated its lungs sufficiently to obtain additional purchase against the roof and floor of its retreat and, bracing its strongly clawed feet—some of the claws were over half an inch long—used them as struts so as to render its extraction extremely difficult.

Joan Procter described the animal’s characteristics thus:

In general appearance it looks as if it had been crushed in youth and had only survived by a miracle. When taken in the hand alive it has a boneless feeling which is uncanny; both carapace and plastron react to pressure on the abdominal region with a springy motion, and the animal is able to inflate itself to a slight degree.

The key question, does the Pancake Tortoise inflate itself and thereby jam itself into a space between rocks, was tackled by Leonard Ireland and Carl Gans (1923-2009), then of the University of Michigan; their paper was published in 1972. They noted that only Robert Mertens, in a paper published in wartime Germany, had questioned the occurrence of inflation: wedging yes; inflation no, he had concluded.

Carl Gans

The background to their work was that Gans had recently worked with George Hughes (1925-2011) in Bristol to sort out the method of respiration in the Spur-thighed Tortoise, Testudo graeca, itself then a matter of controversy. This tortoise and other chelonians were found to draw air into the lungs using muscles that acted in a manner akin to a diaphragm in mammals but acting within the confines of a fixed frame, the shell. There was no mechanism to force air into the lungs (as in frogs, for example). To Ireland and Gans it seemed unlikely that inflation of the body occurs during a threat. They monitored the pressure in the lungs of the Pancake Tortoise while trying to pull it backwards out of an artificial dark crevice 10 mm higher than the depth of the shell. The tortoises attempted to dig their claws of their forelimbs into the floor and rotated their forelimbs outwards. Those actions raised the front of the body and wedged it in place. The hind limbs were stretched out to the rear such that the claws tended to engage in any irregularities in the floor. The authors remarked that the wedging action was most effective; they gained the impression that the forelimbs would have to be broken before the tortoise could be pulled free of a rocky crevice in the wild. At no time during this pulling and wedging in response did pressure within the lungs rise consistently. There was no inflation. Wedging was purely mechanical and achieved by the positioning of the limbs.

Although inflation of the body cavity appeared to have been excluded as part of the mechanism by which the Pancake Tortoise wedges itself into rocky crevices or under boulders of the kopjes on which it lives, that knowledge never found itself into much of the popular literature or have reached what was then the more scientifically-isolated world of the museums. Books and papers still appeared without any reference to the work of Ireland and Gans. Some make it appear that the degree of inflation is enormous, with the impression created that the tortoise is the next best thing to a puffer fish.

However, was the wedging action of the limbs shown by Ireland and Gans the only way the Pancake Tortoise can fasten itself into crevices? Their test apparatus was arranged such that the ceiing was 10 mm greater than the height of the tortoise’s shell. Calculating the depth (i.e. the distance between the outsides of the plastron and carapace) from the photograph they show and the approximately median length of the carapace of the animals they studied, any ‘bulge’ from inflation would have to reach a 23% increase in the depth between plastron and carapace in order to reach the ceiling of the artificial crevice, a surely impossible ask.

The wedging action also leaves the limbs exposed. While offering protection in a relatively wide crevice, would it not be better to seek a narrower crevice in which the legs and head could be withdrawn. In those circumstances, movement outwards of, say, the soft area of the plastron by only a millimetre or so would really jam the tortoise in place. This is the mechanism proposed by Moll and Klemens in a paper published in 1996 but which I have not yet seen. They suggest that drawing the legs into the shell can ‘inflate’ the soft area of the plastron thus supporting the original observations of Loveridge in the field and of Boulenger and Procter in the museum and zoo. How would that work and can predictions be made that could be tested experimentally?

Respiration in tortoises is very different to the process in other land vertebrates. The lungs are attached to the carapace and are inflated and deflated by a diaphragm-like structure separating the lungs from the other internal organs. As the limbs are withdrawn into the confines of the shell there is additional inward pressure on the body cavity and, indeed, movements of the limbs are known to take part in the movement of gases into and out of the lungs. For sufficient pressure to be built up inside the body cavity to push upon the gap in the plastron the lungs would also need to be full or probably very nearly full of air. The flow of air outwards from the lungs in response to strong and sustained withdrawal of the legs and head would have to be stopped. The other alternative to stopping the outflow from the lungs would be for the lungs to be emptied completely by the build-up of pressure in the body cavity. Because tortoise lungs are so large and so capacious it seems unlikely that the decrease in volume of the body cavity would be sufficient for the lungs to be emptied and for the the covering of the plastron to be pushed outwards.

Proceeding on the assumption that the lungs are held full or close to full, the most likely scenario is that the glottis is closed and that the pressure of gases inside the lung increases, as assumed by Ireland and Gans. The other possibility, that the ‘diaphragm’ between the lungs and the rest of the body cavity is held tight such that fluid pressure changes are not transmitted to the lungs seems unlikely since that membranous diaphragm would appear to be much more compliant than the horny layer overlying the ‘hole’ in the plastron.

If this mechanism, the movement outwards of the leathery centre of the plastron brought about by  withdrawal of the legs and neck, does operate the corollary is that the tortoise ceases to breathe. And here we come to the known ability of tortoises to hold their breath for long periods. As anybody who has tried to anaesthetise a chelonian using a gaseous anaesthetic will affirm, the tortoise usually remains unanaesthetised by simply holding its breath for tens of minutes while keeping its legs and head drawn into its shell. I have seen such a tortoise after about 30 minutes suddenly thrust its head out and forcibly exhale, and I do mean forcibly. The sudden exhalation was as if pressure was first applied to the lung and then the glottis opened. The best analogy I can think of is the movement down the runway of an aeroplane first held in check by the brakes until the engines reaches full power.

Other factors would come into play. Terrestrial tortoises have large bladders. If full, less inward pressure would be needed on the body cavity from the active pulling inwards of the legs and head; if empty, more. The legs and head would have to virtually seal the gap between carapace and plastron, otherwise the soft skin would be expected to bulge outwards rather than leathery plastron.

This latter point brings me to suggest that an experimental test of the bulging plastron hypothesis is needed; in other words an extension of the approach of Ireland and Gans to see what happens in narrower crevices. I am not entirely convinced by the present arguments in favour. The reason I suggest more work is needed hinges on the properties of the horny material that covers the large hole in the bony plastron. The late bob Davies had a pair of Pancake Tortoises in the early 1990s. When I handled them, I found the horny covering of that hole to be thick and relatively inflexible. However, there was ‘give’. A slight push on the centre would move it inwards by a few millimetres—the ‘springy motion’ described by Procter when she handled the first living examples at the Zoo. The question of whether sufficient pressure can be exerted on the body cavity to make that horny material bulge outwards even by a few millimetres needs to be answered.

Joan Procter’s legacy, judged at the time to have been a tour de force, stills leaves intriguing questions about the Pancake Tortoise.


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Ireland L, Gans C. 1972. The adaptive significance of the flexible shell of the tortoise, Malacochersus tornieri. Animal Behaviour 20, 778-781.

Mautner A-K, Latimer AE, Fritz U, Scheyer TM, 2017. An updated description of the osteology of the Pancake Tortoise Malacochersus tornieri (Testudines: Testudinidae) with special focus on intraspecific variation. Journal of Morphology 278, 321-333.

Procter JB. 1922. A study of the remarkable tortoise, Testudo loveridgii Blgr., and the morphology of the chelonian carapace. Proceedings of the Zoological Society of London 1922, 483-526.

Eric Thomas Brazil Francis (1900-1993). Zoologist

This story starts in the 1999 when Kraig Adler was looking for photographs to illustrate the biographies of herpetologists for his series, Contributions to the History of Herpetology. I told him that I had a distant shot of E.T.B. Francis but that the University of Sheffield was sure to have a portrait of him somewhere. Unfortunately, the university could not find one. Then, more recently and while looking for something else, I noticed that the Society for the Study of Reptiles and Amphibians had published in 2004 a reprint edition of ETBF’s book from 1934, The Anatomy of the Salamander with an introduction by James Hanken^†. I also knew of but had not seen the dedication by the late Carl Gans to ETBF in volume 19 of Biology of the Reptilia published in 1998. When I did see that I realised I could add and correct some information to that gathered by Carl Gans from former colleagues. With all the volumes of Biology of the Reptilia now being online, I will not reproduce the dedication here. Jim Hanken kindly sent me a copy of his introduction to the reprint of The Anatomy of the Salamander which explains why ETBF is so highly thought of and why his book is still used today.

The frontispiece to The Anatomy of the Salamander is from a
paper by E.G. Boulenger.
According to Hanken, the chromolithograph is by John Green

ETBF loomed large in the life of zoology students at Sheffield. Our time there (1962-65) was at the crossover between classical zoology and what were then more modern approaches, so we still had to know vertebrates and many invertebrate groups backwards and inside out while getting to grips with comparative physiology, experimental biology, genetics and the like. Comparative endocrinology was the department’s major research activity, driven by Ian Chester Jones who had been appointed to the chair in 1958, and we were exposed to its influence from day one, with a lecture on the pituitary given, we discovered later, by Chester Jones wearing a sweater in need of darning and carpet slippers. A suitable culture shock was delivered to students accustomed to being taught by begowned grammar school teachers.

In our time at Sheffield, for the non-special part of Special Honours, the second and third years were combined for lectures. Thus ETBF gave his lectures on vertebrate zoology every second year, I think over two terms. As Gans picked up from the people he asked, ETBF was highly respected for his broad knowledge of the animal kingdom. That was the way then; university staff were expected to know a lot about a lot not just a lot about a tiny field. His knowledge of marine invertebrates emerged on a field course to the biological station at Rovinj in what was then Yugoslavia at Easter 1964. He was the man who knew what all the odd-looking things were we collected and where to find them in the dense German books that catalogued the fauna of the Adriatic. He was the man who also fixed students up with jobs in labs near their homes during the summer vacation; he knew lots of people throughout Britain and what they were doing.

St Mark’s Square, Venice, early April 1964. Sheffield University Zoology Field Course at Rovinj in Yugoslavia (now Croatia):

A day trip to Venice.

Enlargement from a 4 x 4 cm transparency showing ETBF and Oskars Lusis.

Eric Thomas Brazil Francis was born on 3 August 1900 in Hackney in the East End of London, the son of Thomas Brazil Francis and Emile Anne Tourtel. At the 1901 Census, ETBF was seven months old and the family was living at 22 Bishop’s Road, Hackney, London. In 1912 this street in the east end of London was renamed Killowen Road; the house is a three-story terrace house. Thomas Brazil Francis, born in Peppard, Oxfordshire, was a pork butcher’s assistant. His mother, born in Peckham, London, was the daughter of Thomas J Tourtel, a printer’s reader and widower, born in Guernsey, who, with three sons and a daughter, also lived at 22 Bishop’s Road.

At the 1911 Census, the family lived in a very different environment. Thomas, Emilie, ETBF (age 10 and at school) and Emilie’s sister were living at Chalk House Farm, Great Kidmore End in Oxfordshire. Thomas was manager of a game farm (I presume a pheasant shoot).

Thomas Brazil Francis’s father was James Francis; he married Phillis Brazil in 1872 in Oxfordshire. James Francis is difficult to trace in the censuses since he was never present with his wife. However, there is a James Francis who fits the bill; he was a baker born in Kidmore End. Phillis Brazil was the daughter of Thomas Brazil (born about 1820) who, in 1881 was living about six miles from Kidmore End. He was a farmer employing three men and one boy; his wife was also called Phillis, and their daughter, by then Phillis Francis, and grandson, Thomas Brazil Francis, were also present on the night of the census.

Looking at the records, it is clear that the Brazils (originally, apparently, of Irish origin) were, over the years, butchers in London and the area of Oxfordshire north of Reading or farmers, also in that area north of Reading, over the county border in Berkshire.

In the 1921 Census Eric Francis, aged 21, was working as a fruit and poultrey farmer in the employment of his father, shown as Manager of a Game Farm and also as a fruit and poultry farmer. He graduated with an external London degree in 1929, suggesting that he was a late starter to university life. The University of Reading gained university status in 1926, indicating that he must have registered as a student before then but that once registered with the University of London, the degree he received was also a London one. His PhD, awarded in 1933, is a Readsing degree.

We know from his preface to The Anatomy of the Salamander written in August 1933 that he had worked in the laboratory of Professor F.J. Cole FRS (who wrote a remarkable historical introduction to ETBF’s monograph) at Reading. While there he prepared and donated specimens to Cole's museum. I have written about these here. In 1933 he was appointed assistant lecturer in Sheffield.

ETBF married Vera Christine Davison (born 26 September 1901) in 1938 in the Northumberland West Registration District. According to the 1911 Census, she was born in Hadley Wood, Enfield. However, at the census she was with her sister at her grandmother’s house, 8 Washington Terrace, Tynemouth which may account for why they were married in that part of England. Her grandmother, Annie E Ewart was a widow, aged 65, and a schoolmistress.

The Sheffield telephone directories from 1944 to 1983 show the Francis's address as 120 Brooklands Crescent, Sheffield 10.

Their son, Eric David Francis, was born in 1940. He was a classicist and his suicide in 1987 while employed by an American university must have been a great blow to ETBF and his wife, then aged 87 and 86.

At Sheffield, ETBF progressed from Lecturer to Reader in Vertebrate Zoology. Carl Gans stated that ETBF retired in 1973. However, this is wrong (see John Ebling's appreciation below). He retired in 1965 and I suspect 1973 was the year in which he gave up appearing in his old department.

ETBF died in 1993 in Sheffield, aged 93; Vera Christine Francis died aged 98 in 2000.

It would be easy to believe that ETBF was a dyed-in-the-wool comparative anatomist of the E.S. Goodrich kind. In that respect, The Anatomy of the Salamander probably counted against him; it was descriptive and descriptive zoology was out. His interests were actually wide and fully embraced experimental biology. Thus, in Scientific Research in British Universities 1960-61, his research activities are listed as:

• The conductive system of the vertebrate heart 
• The salivary enzymes of amphibia
• Water relations in reptiles
• Host reactions to parasites with special reference to the gut
• Nutritional requirements of intestinal worms with special reference to larval stages 

By the 1962-63 edition, the list had shortened to:

• The conductive system of the vertebrate heart
• Water relations in reptiles
• Host reactions to parasites with special reference to the gut

The work on the heart and on the salivary enzymes appears in the list of publication that accompanies Garl Gan’s dedication. I was well aware of his work on water relations in reptiles. As a new student walking along the zoology corridor to the large lab at the end, the offices/small labs were on the left. ETBF’s door was open (I never remember it closed) and he had a number of glass aquaria/vivaria on an iron stand on the right. In them could be seen a few reptiles; the most noticeable was a Stump-tailed Skink (Tiliqua rugosa). At some stage I learnt that he was measuring water losses across the skin and, once when I went to see him, he and his technician were actually doing so. All I can remember is that they were holding a small cylinder against the flank of a skink and saying they were not having much luck. He told me that he really wanted to look at cutaneous water losses in chamaeleons and needed some animals. I imported two species from Kenya during the 1964 summer vacation and he had about ten of them. I have been unable to find any reference to publication of his physiological work on reptiles. Earlier he had supervised work on rats in drift mines done by Graham Twigg, adding an ecological dimension to his interests.

I had, until I found the book online on research in British universities yesterday, forgotten that he also worked on the physiological effects of gut parasites on the host. I now remember, seeing in the animal house cages of mice (?) and somebody in a white lab coat telling me all about them. But I cannot remember who that was; it wasn’t ETBF, it wasn’t his technician who I think was called Martin. Was it a PhD student or an MSc student? I cannot find any reference to the work being published but it could have been, without ETBF as a co-author.

I can add a publication to the list given by Gans. ETBF was a contributor to A Dictionary of Birds* published in 1985, when he was 85. That volume was remarkable for the inaccuracy of some of the contributions on physiology but ETBF’s articles do not fall into that category!

Gans only knew ETBF in his later years but his correspondents noted that he had during his working life seen a transformation of the way university departments operate. Such change did not seem to suit him. At a student event, my now wife spoke to Mrs Francis for some time. The latter was apologetic and said that such occasions were much better for students in Professor [L.E.S.] Eastham’s days as head (1931-58) while the former received the strong impression that ETBF was unhappy with the direction in which universities, including Sheffield, and academic zoology were moving.

E.T.B. Francis was highly respected and remembered not only as a zoologist but as an Englsh gentleman.

-------------------------------

Note added:

I have found in the University of Sheffield Gazette this note marking Francis's retirement in 1965. It was written by F.J.G. (John) Ebling:

ERIC FRANCIS has devoted virtually the whole of his academic career to the University of Sheffield. Having obtained an external degree of the University of London in 1929, he spent four postgraduate years under G. F. COLE and N. B. EALES at the University of Reading, getting his Ph.D. in 1933. Subsequently, he was appointed Assistant Lecturer and Lecturer at Sheffield, being promoted to Senior Lecturer in 1946 and given the title of Reader in Vertebrate Zoology in 1954.

     The pattern for the meticulous approach to his subject and to his life which Francis maintains, is to be found in his book, “The Anatomy of the Salamander", published in 1934. This work of nearly 400 pages contains 84 figures in 25 plates, nearly all drawn—very beautifully—by the author, an index of 21 pages, and 840 references, bearing an introductory note which reads: "All the works quoted in the bibliography have been personally examined except those marked with an asterisk”. Only 32 are so marked!

     At Sheffield Eric Francis collaborated with the late FRANCIS DAVIES of Anatomy in a study of the conducting system of the heart of the salamander, and this was followed by a fine series of anatomical, histological and experimental studies of other vertebrate hearts by the two friends and other co-workers. In 1961 Francis returned again to the Amphibia with a paper on the function of the salivary secretions in frogs, toads, and—of course—the salamander; currently he is working on the water relations of reptiles.

     The scope of his research does not reflect the whole range of his scientific interests. As well as his authoritative teaching of vertebrate zoology, he has maintained a flourishing research school in parasitology, a discipline we shall now lack. During the past two years, as early in his career, he has taken a most active part in departmental courses in Marine Zoology, appearing as at home among worms, molluscs and sea-anemones as among backboned animals, fully prepared to give a talk on sea-squirts at an hour's notice, and shaming his collaborators by his extensive knowledge of the marine plankton.

     Francis, above all, is a scholar. As the pendulum swings between the view that research must be predominant and the counter pressure to turn universities into teaching factories, he suffers no dichotomy of inclination or duty; in his example scholarship grows by research and propagates through the teacher. Among his younger colleagues he is known for his unfailing friendship and courtesy, his great sense of humour and his unflagging zest for life. He is especially fond of music and is always to be found at chamber and symphony concerts; he has an insatiable interest in art and archaeology; he is a card-carrying cricket supporter; he enjoys good food and is an undefeatable traveller, who can reach any railway dining car in under four minutes from the gong while his fellow travellers have long before succumbed to languor in their compartments.

     In the Department of Zoology his wisdom as well as his scholarship will be missed. This may well be the last manuscript over which a colleague will request his critical eye, and Francis will soon have written his last testimonials for Sheffield students and given them his last advice about choosing jobs. But he remains in his profession. During next session he will be a member of the teaching staff at Royal Holloway College, and he will continue to examine for the University of London. In wishing Eric and Christine Francis a happy future it would be premature to talk of retirement. More appropriate would be the comment of the captain of the Bios, the research vessel belonging to the Marine Station at Rovinj, Jugoslavia, which Francis recently visited in company with his colleagues and students. We had, on a beautiful spring day, put in to an uninhabited Adriatic island, and because there was no harbour it was necessary to jump from the bow of the ship on to the rocks. Francis performed this operation with a mechanical elegance which, perhaps especially in view of his personal statistics, drew universal admiration and the exclamation "E come un giovane, questo professore!"

F. J. G. EBLING.

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UPDATED: 8 August 2023 and 27 August 2025

†Hanken, J.  2002.  Eric Thomas Brazil Francis and the evolutionary morphology of salamanders.  Introduction to the reprint of E.T.B. Francis, The Anatomy of the Salamander, pp. v–xiv.  Society for the Study of Amphibians and Reptiles, Ithaca, New York.

The reprint edition 2002

*A Dictionary of Birds. 1985. Campbell B & Lack E (editors). Poyser.

Biology of the Reptilia. Carl Gans's Magnum Opus Online

The 22 volumes of Biology of the Reptilia can now be read online, and access is free. The Gans Collections and Charitable Fund, established in the memory of Carl Gans (1923-2009) who ran the whole show from the first volume in 1969 to the final one published in 2010 just after his death, has somehow managed to overcome problems of copyright with five publishers, to scan all the volumes and to present the articles in an accessible form.

Like a number of series, Avian Biology and Lactation, for example, Biology of the Reptilia began as a publication of Academic Press, now sadly part of the Elsevier empire. Much as the effort to make the Biology of the Reptilia available will be appreciated, over the years I have found that the Academic Press series of multi-author books had a certain interest at the time of publication, they soon fell into the category of 'very rarely looked it' —and that includes a long article of mine. I suppose that is because in an active field, a review paper in an edited volume was just a snapshot at a particular time, useful for drawing the old literature together, but soon passing into the category of ephemera as the field moved on. Somewhat paradoxically, I suppose, such volumes will be most useful in fields that are no longer popular or have run out of steam simply because the article will be a more complete statement of knowledge. For those rare beings researching the history of a field of research access to the Gans volumes will be very welcome. All too often publishers (including learned societies) are charging for access to old literature that should be freely available; they have already made their pile from selling books and journals. Charging for access to research decades old and paid for by the taxpayer is outrageous. So the appearance of Biology of the Reptilia online is a case for applause to the Carl Gans charity.

I never met Carl Gans but I did have a correspondence with him over several months. Amphisbaenians were his great interest and he had arranged with the late Harold Fox, the then editor, to have a whole issue of the British Journal of Herpetology (now Herpetological Journal) devoted to that group of lizards, with Gans inviting the authors. When I took over as editor from Harold Fox*, I had a whole issue in the pipeline that required very little editorial work since Carl got the papers in and did what was necessary before sending the whole package to me by airmail. I was extremely grateful because the journal had been printed by Harold Fox's family publishing company and I was establishing a new format with a new printing company to try to bring the journal into the 1970s.

*Harold Fox, 24 October 1922-29 May 2003, worked at University College London from the late 1940s until 1994. The family bequest funds the Harold and Olga Fox Scholarships in Biological Sciences, supporting four students at a cost of £23,000 per annum.