Rovinj Marine Biological Laboratory. The Sheffield Zoology Field Trip in 1964. Part Two. The official history 56 years later

When I wrote Part One (30 May 2015) I was aware that E.T.B. Francis had written an account for the University of the first field trip to Rovinj, then in Yugoslavia, from Sheffield’s zoology department. I had written up a report on the reptiles and amphibians we found during the afternoons we were not in the lab, out and about doing the marine biology or visiting places of wider interest. He caught me in the corridor to say that he had sent the report in to the university’s Gazette. However, until a couple of months ago I had not seen what he had written. Then I found on eBay a copy of the Gazette published in October 1964. When it arrived I was pleased to see that it contained the article. With it in hand I have been able to put names to places we had visited and to draw a map and after further Google searches, to caption more accurately the photographs I had taken. At the end of this article I have also added a series of explanatory notes.

First though, I should remark that the trip to Rovinj was revolutionary for the early 1960s. Language students had to spend time abroad but for undergraduate science students a field course in continental Europe was something very special.

Rovinj Marine Laboratory

This is Francis’s complete account from the Gazette:

ROVINJ, EASTER 1964

For many years past it has been a tradition amongst zoological departments that honours students should spend some time during the Easter vacation at a marine station to study ecological and faunistic problems associated with animal life in the well defined habitats of the sea shore and shallow seas. Such students, at the same time, get the valuable experience of examining and identifying living animals collected personally from their natural habitat.
     Considerable emphasis has always been placed on the importance of such studies by the department of zoology at Sheffield, but this year a difficulty arose since the laboratory at Robin Hood’s Bay belonging to the University of Leeds—to which the second year honours students would normally have gone—is in the process of rebuilding and enlargement and was therefore unavailable. Accordingly Professor I. Chester Jones decided on a bold experiment and set arrangements in operation which resulted in a party of 31 students and 7 staff travelling to Rovinj where the Jugoslav government has a research station on the Istrian coast of the Adriatic.
     Two laboratories, one large and one small, were made available by the Director of the Station, Dr. D. Zavodnik, and the Station’s research vessel. Bios, was chartered on several occasions. Thus it was possible to study the faunas characteristic of several types of substratum—hard bottom, secondary hard bottom, sand, mud, shingle—found at moderate depths off shore, as well as the littoral fauna of the shore itself. The use of the Bios also enabled samples of plankton to be collected and studied, thus giving the opportunity to examine forms, both larval and adult, specially adapted to a floating life at the surface of the sea.
     The fauna as a whole proved exceptionally rich, both in quantity and quality, northern species such as are found round the coasts of Britain mingling with others characteristic of Mediterranean waters, the one sometimes supplanting the other in characteristic ecological associations.
     Morale and enthusiasm were high: staff shared with students the excitement of examining alive species which had hitherto been unknown, even to the most experienced members of the party, outside the pages of specialist textbooks. Over 180 species were individually identified and at least seven or eight new records were added to the fauna lists. It is very doubtful whether so rich a zoological experience could have been obtained in the same period of time around the shores of Britain—certainly not from one station alone.
     In addition to the faunistic work a serious and extensive study was made of the speciation of the limpets occurring locally, including a chromatographic analysis of their pigments, in order to compare the forms found at Rovinj with those previously studied in former years from British waters. The Jugoslav government has a station for the culture of oysters and mussels at Limski Fjord, some eight miles north of Rovinj, and the party was able to visit this and to see something of the techniques used.
     As a side-line two devoted herpetologists in the party spent every spare moment of their time in the study and collection of the local amphibia and reptiles. In all they captured some 103 individuals comprising four species of lizards, four species of amphibia and one species of snake.
     Outside the strictly zoological purposes of the excursion, occasions were made to take advantage of such cultural opportunities as the neighbourhood offered. Thus the party took a day off to visit Venice, travelling each way by a specially chartered bus and spending five hours in that unique city. On another occasion a half-day was spent visiting the ancient Roman city of Pula, where there is still a well preserved first century amphitheatre attributed to Augustus, and other relics of Roman times. Even the coach journeys to and from the railhead at Ljubljana on the outward and homeward journeys were utilized. The limestone caves at Postojna were visited on the outward journey. These are famous not only for exceptionally fine stalactite and stalagmite formations, but also as the home of the blind cave-salamander, Proteus anguinus, which the party was able to see alive. On the homeward journey a stop was made at the deserted medieval village of Dvigrad near Kanfanar to visit the church of St. Anthony and to see the very fine and vivid fifteenth century frescoes, and at Porec to visit the sixth century basilica of St. Euphrasia with its fine mosaics, comparable with those of Ravenna in brilliance.
     This remarkably successful expedition would have been impossible but for the enthusiasm, energy and organizing ability of Dr. F. J. Ebling, who not only planned the whole enterprise beforehand, but shouldered the day to day responsibility for its successful conclusion. He deserves great credit and the gratitude of all who benefited from his efforts.
     At the Rovinj end the way was prepared by Anton Perusko and his wife Gillian (nee Gillian Glen, B.Sc. Sheffield 1959). In spite of having to care for two young children, Gillian Perusko met the party at Ljubljana and escorted it to Rovinj and throughout the whole visit acted as general guide, interpreter and liaison officer. Anton Perusko made use of his official connexion with the “Auto-Kamp” Enterprise to smooth away many spiky corners connected with the accommodation, transport and such like matters. Without their invaluable help and full co-operation the venture could never have succeeded or even been contemplated.
     Much friendliness and co-operation was shown by Dr. Zavodnik and his staff at the laboratory as well as by the staff of the “Jadran” Enterprise at whose hotel the party stayed. At the Jadran Hotel it was introduced to many interesting national dishes; in fact, with the exception of milk-fed lamb, which is traditional for “Big Friday” and Easter Day, no major dish was repeated on the menu throughout the whole period. In the interest of international relations the Jadran Enterprise organized a dance on Easter Saturday to which many local people came to meet the English party of whom they had been told by the national radio and press.
     Valuable as the zoological assets of the excursion proved to be, it is clear that these are by no means the only entries on the credit side of what must remain for all who participated a most memorable experience. 

E T B Francis 

The Places

   

My Photographs

I took my ‘Baby’ Rolleiflex 4 x 4 camera. This took 12 exposures on 127 size film. I used Agfacolor CT18 reversal film which is now known to be prone to degradation with time and to be ‘grainy’. Differences in processing may be partly responsible for the poor long-term reputation of this film. Unfortunately, despite being stored in identical conditions, the Rovinj films have survived amongst the least well of the photographs I have stored. The film base has warped, false colours have crept in from the edges and some have faded.

The Journey from the Hook of Holland to Ljubljana via Munich

The Rhine from the train to Munich

Vineyards in Germany from the train

Vineyards from the train

Somewhere in Austria from the overnight train from Munich to Ljubljana

We had several hours in Munich on the way out. The two of us wandered out of the station and found a beer hall which Google Earth shows to be Augustiner Stammhaus. We had steak and chips with, of course, beer. I was in the same establishment seven years later for a dinner during the International Physiological Congress. More beer was consumed.

Postojna Cave 

(2 slides I bought)

Rovinj

M/V Bios - the research vessel

Rovinj: Hotel Jadran (Adriatic) centre

Rovinj from the Bios

Rovinj

Rovinj

Rovinj. The shop on the left sold filigree jewellery for which what is now
the Croatian coast is famous. My wife brought the bracelet (below)

Pula

Venice

The Sheffield party in St Mark's Square

We travelled to and from the bus by vaporetto

Limski 'fiord' north of Rovinj

Film Set for The Long Ships (filmed in 1963) near the end of Limski 'Fiord'

Dvigrad

Church of St Mary of Lukać

Unwelcome Fauna

My wife’s (then girlfriend) abiding memory the Rovinj trip is arriving back with an unwelcome guest. She left the party at Harwich since her parents were living in Suffolk. Drying herself after a bath she though she had developed a new mole. Closer inspection revealed a large blood-swollen  tick. She blamed—and still does—the Munich-Hook of Holland sleeper.

Notes

1. The 31 students were the 2nd years honours group plus a few from 3rd year.
2. The staff were F.J.G. Ebling, E.T.B. Francis, O. Lusis, J. D. Jones, D. Bellamy, L. Hill plus Mr Hancock, the chief technician.
3. D. Zavodnik was one of the authors of the history of the Rovinj laboratory.
4. The hotel we knew in Rovinj as the Hotel Jadran was originally called the Adriatic. It has been refurbished and is again known as Hotel Adriatic.
5. Not mentioned was the visit to the film set built for The Long Ships near the end of Limski ‘fiord’. I read that it is now fallen down. However, The Viking Restaurant was built on the site by the film’s caterer and can be found on Google Earth. It is accessible, as was the film set, by road.
6. The deserted medieval village of Dvigrad is now a major tourist site. My photographs are not in the church Francis mentioned but in the Church of St Mary of Lakuć.
7. Dates, as mentioned in part 1, the date of entry into Yugoslavia was 17 March with departure on 1 April. Remembering the journey, we must have left Sheffield on 15 March and returned, carrying the microscopes we had taken with us, on 3 April.
8. After the appearance of Part 1, I was contacted by several ex-Sheffield students who had been to Rovinj in succeeding years. The Easter field course continued into at least the early 1970s. Len Hill told me in 2015 that they ended because in our time such courses were funded as part of the honours course by grants from local authorities to the individual student. Then funding was given direct to the universities to fund such activities. However, the funds disappeared into the maw of university administrations and funding for these field courses ceased.

Francis ETB. 1964. Rovinj, Easter 1964. 1964. University of Sheffield Gazette, Number 44, October 1944, 74-75.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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.

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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.