arthur james cain - Biographical Memoirs of Fellows of the Royal ...

ARTHUR JAMES CAIN
25 July 1921 — 20 August 1999
Biogr. Mems Fell. R. Soc. 54, 47–57 (2008)

ARTHUR JAMES CAIN
25 July 1921 — 20 August 1999
Elected FRS 1989
BY BRYAN C. CLARKE FRS
Institute of Genetics, Queens Medical Centre, University of Nottingham,
Clifton Boulevard, Nottingham NG7 2UH, UK
Arthur Cain was a polymath, and one of Britain’s leading evolutionary biologists. A strong
believer in, and advocate for, the importance of natural selection, he greatly influenced evolutionary
thought during the period from the 1950s to the 1980s. He wrote important papers in
many areas of biology, and he was a noted historian of science.
LIFE AND CAREER
Arthur was born in Rugby, Warwickshire, the son of James Peter Cain and Margaret
Susan, née Goodall. His father was of Manx and Welsh descent, and his mother came from
Wiltshire. He wrote firmly that he had ‘no distinguished relatives on either side’*. His
secondary education took place at the local boys’ grammar school in Rugby, the Lawrence
Sheriff School, which he entered in 1931. There he came under the influence of an outstanding
teacher, Wilfred Kings. Kings managed very effectively to generate enthusiasm for
biology among his pupils. Cain wrote, ‘I am sure whatever I have achieved in field work
owes a lot to his patient encouragement. He was an ideal master for those who really wanted
to learn.’ Several of Kings’s students became well-known biologists. They included John
Harper FRS, Gerald Thompson, who was the founder of the Oxford Natural History Film
Unit, and A. F. L. Clowes, who gained an open exhibition at Magdalen College and became
Reader in Botany.
When discussing his intellectual development, Arthur drew attention to the fact that the
town of Rugby had an unusually good public library, as a result of the international scheme
of apprenticeships at the British Thompson Houston factory. He also mentioned an abundance
* Unattributed quotations are taken from Arthur’s Personal Record deposited in the Library of the Royal Society.
doi:10.1098/rsbm.2007.0034 49 This publication is © 2008 The Royal Society

50 Biographical Memoirs
of wildlife within easy reach, much of it suitable for maintenance in aquaria. He enjoyed the
local cement quarries, where he searched the spoil heaps for Jurassic fossils, and occasionally
trespassed into the quarries themselves.
In 1939 Arthur won a Demyship (the most prestigious scholarship) at Magdalen College,
Oxford. He was tutored by J. Z. Young (FRS 1945), and emerged with first-class honours in
zoology after a course that was truncated because of World War II. He then served as a Second
Lieutenant in the Royal Army Ordnance Corps, and later transferred to the Royal Electrical
and Mechanical Engineers. He was promoted to Captain in 1942. A large part of his wartime
service was carried out in the Orkney Islands, where he was responsible for anti-aircraft radar,
protecting the ships of the fleet. December 1945 brought him back to Oxford for some further
tuition by Peter (later Sir Peter) Medawar (FRS 1949, and Nobel laureate), and for work
towards a DPhil, which he finished in 1948. During this time, he met and married, in 1947,
Joyce Perrott, who was reading mathematics at St Hilda’s College. Their daughter, Celia, and
their son, David, were born in 1948 and 1951, respectively. He was appointed Departmental
Demonstrator in Zoology, and in 1949 became a university demonstrator (equivalent to a
lecturer) in animal taxonomy.
He spent six months in 1950 working with Ernst Mayr (ForMemRS 1988) at the American
Museum of Natural History, and lecturing at Harvard, Yale and several other eastern universities.
In 1954 he was made Curator of Zoological Collections in the University Museum at
Oxford, alongside his lectureship. From 1958 to 1961 he was also Lecturer in Zoology at St
Peter’s Hall, Oxford.
He led two expeditions, one to the Solomon Islands in 1953 and one to Guyana (then British
Guiana) in 1959. The trip to the Solomons enabled him to study the correlations between geographical,
ecological, and morphological variation among the endemic birds. He also visited
New Guinea. In British Guiana he examined the comparative ecology of closely related bird
species within its tropical forests. Because of his experience in the tropics he was chosen, in
1961, to represent Oxford University on the Inter-Universities Committee for research at Lake
Kariba. He acted as the scientific secretary to this committee in the UK, and during this period
he made several trips to Kariba and Johannesburg. In 1965 he went to the Serengeti National
Park, once more working on avian ecology. In 1983 he joined an expedition to the Dominican
Republic, where he concentrated on the ecology and distribution of terrestrial molluscs.
In 1964 he became Professor of Zoology at the University of Manchester, and in 1968 he
moved to the Derby Professorship of Zoology at the University of Liverpool, where he stayed
until he retired in 1988.
Cain served as Editor of the Biological Series for Hutchinson’s University Library from
1967 to 1975. He was an Associate Editor of Malacologia, and a member of the Advisory
Board for Bolletino de Zoologia. He was appointed as Advisor in Biology for the Science
Research Council from 1972 to 1975. In 1976 he became President of Section D (Zoology)
of the British Association for the Advancement of Science. He became a Founder Member
of the Society for the Study of Human Biology, President of the Oxford Genetical Society,
Zoological Secretary and President of the Systematics Association, Member of the Council of
the Linnean Society and Fellow of the Institute of Biology. He was chosen to be the founding
President of the European Society for Evolutionary Biology in 1987, and to be an Honorary
Member of the Malacological Society in 1991.
He made lecture tours in Australia, Canada, Israel and the USA. In 1986 he was Herbert
Spencer Lecturer at the University of Oxford. He received the Scientific Medal of the

Arthur James Cain 51
Zoological Society of London in 1962, and its Frink Medal in 1990. He was awarded the
Linnean Medal for Zoology in 1985, and the Founder’s Medal of the Society for the History
of Natural History in 1995. He had a long association with the Academy of Natural Sciences
in Philadelphia, and was made a Fellow in 1976.
SCIENTIFIC CONTRIBUTIONS
Arthur studied for the DPhil under the supervision of Dr John R. Baker, who was then a
lecturer in the Department of Zoology. Arthur told how he came to study under Baker. Peter
Medawar had suggested that he should go round the staff to see what they were all doing.
I started with the room next door, where was John R. Baker FRS, and got no further—he’d just
invented a histochemical test and as I was extremely fond of test tube chemistry I stayed with
him. I could not have done better. He taught me an extremely practical and critical approach to
the subject and inspired me with his wonderful enthusiasm and incredible range of knowledge and
competence. From then on, I was free of the superstition that if a paper looked scientific and had
awe inspiring chemical and other formulae, it was necessarily any good.
His work with Baker involved studies on the histochemistry of lipoids and on the structure of
the Golgi apparatus. ‘John Baker should have been co-author of some of my papers in lipoid
histochemistry, but he wouldn’t hear of it. He was a man of complete integrity.’
Arthur’s thesis resulted in 12 papers, one of which, published in 1947 (1)*, on the use of
Nile Blue in the examination of lipoids, became a ‘Citation Classic’ in 1989. Arthur was asked
to write a commentary about it, and characteristically observed (19):
Surely there are far better techniques for separating lipoids now; the paper should have been
superseded years ago. I’ve never regretted my work on lipoid histochemistry; it showed me that
papers can be decorated with recondite structural formulae and based on apparently scientific
procedures and still be worthless. This paper had no impact on my further career. What it all really
shows is the heterogeneity of the class of citation classics. To call this paper a classic (however
qualified) because people have gone on citing it (I hope while improving or rejecting it, but I fear
not) is downright misleading.
While he was studying for the DPhil, Arthur became a mentor of Philip Sheppard (FRS
1965), who had just returned from being a prisoner of war. Arthur wrote:
He appealed immediately to my love of natural history, and showed me how he and his super visor,
E. B. Ford FRS, were tackling the problems of evolution and natural selection practically in the
wild.
Sheppard was probably the biggest single influence on me. I think our own findings of massive
selection in Cepaea have influenced me most, though the fact that Ford, R. A. Fisher and
Sheppard were finding selection on the apparent trivialities of wing-pattern variation in the scarlet
tiger moth was nearly as important. I’m still working out some of the consequences for evolutionary
theory, and have rejected most of Mayr’s and Stephen J. Gould’s ideas in consequence—they
are not founded on actual studies of selection.
After his DPhil, Arthur turned his attention to evolution and systematics, interests that
lasted him for the rest of his life. The transition may have been helped by his histochemical
* Numbers in this form refer to the bibliography at the end of the text.

52 Biographical Memoirs
work on the neurons of snails, which coincidentally drew his attention to the extraordinary
colour polymorphism in Cepaea nemoralis. Sheppard was interested in the significance of
genetically determined polymorphisms in natural populations, and Arthur introduced him to
the variation in Cepaea. This resulted in a long-term collaboration and friendship between
them. At the time, the variation was regarded as neutral, and unimportant to the survival or
reproduction of the snails. This view was reinforced by the work of Maxime Lamotte (Lamotte
1951), who, in a large and influential paper, fitted the frequencies of Cepaea morphs to the
equations of random genetic drift. Cain and Sheppard found it difficult to believe that the
variation was neutral, and they tested Lamotte’s hypothesis by studying the extremely various
habitats in which C. nemoralis occurs (woods, hedgerows, grasslands, and habitats containing
mixtures of herbaceous plants). In the region of Oxford they found repeatable patterns of
morph frequencies, with higher proportions of yellow and banded shells in grasslands and
rough herbage, but with higher proportions of brown, pink and unbanded shells in woodlands.
The differences were highly significant. Cain and Sheppard interpreted these regularities as
being due to natural selection by thrushes, which are major predators of Cepaea. The thrushes
have a convenient habit of smashing snail shells on stones, around which may accumulate the
remains of tens or hundreds of broken shells. Later experiments, using snails that had been
marked on their shells, allowed Cain and Sheppard to demonstrate that thrushes did indeed
exert differential selection on the morphs. Their 1954 paper on Cepaea (3) has become a classic
in ecological and evolutionary genetics.
Arthur’s studies on Cepaea reached a new level of complexity when, in 1963, he and John
Currey discovered curious patterns of gene frequencies on the Marlborough Downs (15).
These patterns, which they called ‘area effects’, involved relatively uniform morph frequencies
over areas much larger than those occupied by single populations. The frequencies bore
little relationship to the habitats as defined by vegetation. Similar patterns were subsequently
found in other locations in the British Isles. There seemed to be two regional alternatives,
one with clear associations between vegetable habitats and gene frequencies, and the other
with patterns of area effects. Cain and Currey interpreted the latter as being manifestations of
variation in microclimates, arguing that the frequency of brown shells increased where there
was ‘ponding’ of cold air. Other workers, for example Goodhart (1963), argued that the area
effects were due to random genetic drift. Later work by Robert Cameron and his colleagues
(Cameron & Dillon 1984) has indicated that such patterns are the results of changes in land
use. In the regions where area effects are found, C. nemoralis has been restricted in the past to
a few refuges within which differentiation occurred, either by natural selection or by random
genetic drift. Subsequently, the snails would have repopulated intervening habitats, spreading
out to form the area effects. It is still a matter of debate whether the differentiation in the
refuges has been largely due to selection or to drift, but the existence of area effects itself is
certainly not evidence against selection.
In regions of Britain where the colours of shells in the Cepaea populations are adjusted
to the backgrounds of their habitats, Cain and Sheppard found that the colours of the snails’
soft bodies were similarly associated, albeit only in shades of brownish grey. This was further
evidence of natural selection. During and after their work on C. nemoralis, Arthur and his colleagues
performed ecological and genetic studies on several other species of pulmonate land
snails, including Cepaea hortensis, Theba pisana, Trichia striolata and Helix aspersa.
Arthur soon became interested in the shapes of snails as well as their colours. He considered
the ratio between shell height and breadth, examining the ratio in many gastropod taxa.

Arthur James Cain 53
He showed that variations in shape between species within higher taxa are often bimodal. By
surveying the entire molluscan faunas of islands and regions, he was able to relate particular
shapes to particular habitats and ways of life. He could then estimate the extent to which
‘shape niches’ are filled in terrestrial ecosystems. As Laurence Cook remarked, ‘The basic
information was available to all, but Arthur had the insight to make it matter in evolutionary
biology’ (Cook 2000).
In 1954 (4) and 1956 (7) Cain and Sheppard published two important critiques of
Dobzhansky’s and Li’s views on polymorphism, pointing out that the terms ‘adaptive polymorphism’
and ‘adaptive value’ can be very misleading when respectively they mean ‘selected
polymorphism’ and ‘selective value’. Selection does not necessarily lead to adaptation,
particularly when selective values are related to frequencies in the population. The ‘average
adaptive value’ (w – ) is in no sense a measure of adaptation. Unfortunately, modern students of
molecular evolution have followed Dobzhansky’s usage, and equated selection and adaptation
to the considerable detriment of clear thinking. Cain and Sheppard’s two papers on this subject
should be recommended reading today.
The work on Cepaea convinced Arthur that it is always a mistake to assume that a character
is selectively neutral without a careful study of its biological effects and their relations
to the ecology and habits of the organism. Moreover, a prima-facie belief in neutrality can
hold up progress because it discourages further observations and experiments. On occasion
Arthur’s enthusiasm to make this argument led him apparently to imply that the null hypothesis
should be selection rather than neutrality, a strategy that is of course impracticable. Arthur
was inspired to accumulate evidence about the power and delicacy of natural selection, and
he summarized his views trenchantly in a widely quoted article (16), which was reprinted in
1989 (18).
After his experiences in British Guiana, the Solomon Islands and the Dominican Republic,
Arthur became very interested in the process of adaptive radiation. As he wrote to Ernst Mayr,
‘My visit to the neo-tropical region has had a reinforcing effect on what I saw in the Solomons
and Australia, and ADAPTIVE RADIATION is now written firmly all over my frontal lobes.
I see it as the most outstanding of all evolutionary phenomena.’* He was drawn to work on
the Australasian and Oceanian birds (6, 9), an interest that led him to analyse how taxonomic
decisions are reached, to study the methods of Linnaeus, and to consider the philosophy and
history of pre- and post-Linnaean developments in systematics (10, 12, 13, 20). He made an
early attempt, with Geoffrey Harrison, to create a method for numerical taxonomy (11, 14).
Arthur gave much attention to the concepts of species (2) and genera (8), and was inspired
in 1954 to produce an excellent short book, Animal species and their evolution (5), reprinted
by Princeton University Press in 1993 with a new ‘Afterword’. His convictions about the
power of natural selection persuaded him that convergent evolution is common enough to
make the construction of phylogenies from anatomical characters extremely insecure. He
became a campaigner against Hennig’s cladistic arguments as well as against ‘neutralists’ who
attributed the larger part of organic evolution to the random sampling of genetic variants. He
also took strong exception to Stephen Jay Gould’s advocacy of chance and ‘contingency’ in
the evolutionary process.
* This letter, with most of Cain’s scientific correspondence, is deposited in the library of the American Philosophical
Society at Philadelphia, alongside the papers of his friend and colleague, Professor Philip Sheppard FRS.

54 Biographical Memoirs
PERSONALITY
Arthur Cain cared passionately about science, and his enthusiasm was very appealing, particularly
to younger students and colleagues. John Turner writes:
I was very deeply impressed by his erudition and powers of communication, and must have spent
excessively long tea breaks sitting at Arthur’s table in the library along with his graduate students
lapping it all up. He became in a way my unofficial supervisor. … I never burdened Arthur with
any questions about my own research and he probably never knew what I was doing. It was just
general education. I loved his interest in the history and philosophy of biology. He was a fountain
of quotations: what impressed me was his memory for prose.
Laurence Cook writes:
I first met Arthur when interviewed as a prospective research student. After some wary sparring on
both sides I mentioned that the animal I proposed to study was at the edge of its range, and therefore
likely to show clear evidence of ecological limiting factors. ‘Ah, peripheral populations’, he
said, and I was treated to a 40-minute monologue on all aspects of the subject. After that I was in,
and learned that this was one of his favoured patterns of communication. Generations of research
students remember Arthur holding forth and developing his ideas during tea and coffee breaks.
A vast range of biological topics was covered and the enthusiasm was infectious. The ebullience
… had its negative side, he was sensitive to disagreement and tended to take it as personal criticism.
This had one compensation, because to get over a perceived setback he would take up a
completely different topic.
He talked, as an equal, with experts in many obscure animal and plant groups. His style of
supervision was a sort of benign neglect. He was there to help if asked, but otherwise left one
alone. His help was given freely, and it included the gift of some of his data. However, he never
asked to put his name on any of the papers. It is a pity that this example is not followed today.
David Parkin, who was a research student of Arthur’s in Manchester, has written about the total
extent of his supervision. In the first year Arthur said, ‘I think you ought to work on Arianta
arbustorum, David.’ In the second year David was given 35 minutes of supervision, and in the
third year Arthur said, ‘I’m glad you got something useful out of that piece of work.’
Cain’s passion about science quite often got the better of him. He tended to view people
in black and white. They were either brilliant or monstrous, and at one time or another most
of his colleagues had been both. At meetings and seminars he was liable to explode, and then
storm out angrily. At other times he would be entirely charming.
In the bad times, he would retire gloomily into himself. Then it was difficult to get a word
out of him. The transition between enthusiasm and gloom was often quick, apparently precipitated
by some imagined or real slight but also, as his daughter Celia reports, because of
very bad migraines.
John Turner writes, ‘He used to sit … by that pond in the middle of the courtyard at the
Zoology Department, daring anyone to come and speak to him, and then when somebody did,
would complain that nobody wanted to speak to him because he was a failure.’
He applied for the chairs at Oxford and Durham, was offered Durham, but turned it down
because Joyce very much preferred to stay in Oxford. When John Pringle got Oxford, Arthur
contacted Durham saying that he would like to accept after all, only to find it had already gone
to someone else. One can see why he felt miserable about it.
With all these ups and downs, working in his company was never dull. He created a sense
of excitement about the subjects that interested him. He was angered by people who, as he saw

Arthur James Cain 55
Figure 1. Watercolour by Arthur Cain inspired by Out of the silent planet by C. S. Lewis; the painting represents Meldilorn.
(Online version in colour.)
it, had forsaken the true paths of scientific virtue and thereby had become managerially important.
He made his contempt for them very evident, which did not endear him to university
administrations, and may well have hindered the progress of the departments for which he was
responsible. He was certainly not a good administrator himself, but he created an intellectual
atmosphere in which ideas flowed continuously, and were continuously challenged.
Arthur had a very wide range of interests and talents, which included painting (figure 1)
and music (on occasion he would carry about a suitcase containing recorders of various sizes,
which he used to play with my daughter when she was small). He and Joyce shared an enthusiasm
for their garden in Hoylake.
George Davis wrote (Davis 2000):
Those who knew him marveled at his ability for total recall of world-wide species of plants and
animals, his knowledge of geology, to say nothing of his ability to reconstruct the architectural
history of cathedrals of Europe (stone by stone and porch by porch), or to entertain one by sitting
in a rocking chair and ‘reading’ Tolkein’s Hobbit (without the book!), giving flawless renditions
of Shakespeare’s sonnets from memory, or quoting endless poems and doggerel by Edward Lear,
and from the Cautionary Verses of Hilaire Belloc.
Despite Arthur’s tendencies to combat, he was often very perceptive. Here are two examples,
written 20 years ago (17), that illustrate his style:
The contrast between the potential rapidity of cultural … evolution and the slowness of genetic
evolution is certainly striking. But what is of far more immediate importance is that between
cultural evolution and the rate of human reproduction. Without any nuclear holocausts, it is more
than likely that at least the terrestrial parts of the planet will be ecologically and economically

56 Biographical Memoirs
ruined in half a century, simply because cultural evolution is too slow. … as Lawton has said, no
politically viable alternative has been presented to the multiplying myriads of the world. … It is no
good standing in line, singing ‘Woodman, spare that tree!’ to a shifting cultivator with a wife and
children to feed who is about to demolish the last patch of rain-forest in his neighbourhood.
Intellectuals no doubt are individually biased; nevertheless by their … conflicts with one another
and, in science, by appeal to actual experiment, they can progress towards a more objective
appreciation of the world. … What they have to realize is that they are more probably among
the evolutionary eccentrics, having (some of them at least) remained children in their curiosity
and exploration of the world, than representative of mankind in general. Periods in history when
discussion has been genuinely free are very few, very short, and often followed by repression or
downright persecution. … It is more than possible that in Britain we are coming to the end of
one such golden period; Dorothy Nelkin’s book on the conflict of fundamentalism and science
in the USA documents the immense strides made by irrationalism in that country of glorious
freedom. Certainly the culturally retarded politicians that can see in a Vice-Chancellor only a
works manager, and in a university only a source of cheap technological information; who in the
name of efficiency are removing all safeguards to intellectual freedom and concentrating power
in the hands of the few to ensure ‘efficient’ running; and those whose only criterion of value in
appraising research is the amount of money successfully acquired and spent on a piece of work;
are very unlikely to respond to any long-term plans. There will be enough quislings inside the
universities to abet them.
Arthur Cain was survived by his wife Joyce, who died in 2005, their son David and their
daughter, Celia Tew.
ACKNOWLEDGEMENTS
I am very grateful indeed to the following people for their help in the writing of this memoir: Mr David Cain,
Professor L. M. Cook, Mrs C. Gill, Professor J. J. D. Greenwood, Professor G. A. Parker FRS, Professor D. T.
Parkin, Mrs Celia Tew and Professor J. R. G. Turner. Particular thanks go to Professor J. J. Murray for his kindness
in researching the Cain archive at The American Philosophical Society.
The frontispiece photograph was provided by Arthur Cain’s family.
REFERENCES TO OTHER AUTHORS
Cameron, R. A. D. & Dillon, P. J. 1984 Habitat stability, population histories and patterns of variation in Cepaea.
Malacologia 25, 271–290.
Cook, L. M. 2000 A J Cain FRS 1921–1999 Bull. Malac. Soc. Lond. 34, 2–3.
Davis, G. M. 2000 Arthur J. Cain 1921–1999. Proc. Acad. Natl Sci. Philad. 150, 1–2.
Goodhart, C. B. 1963 ‘Area effects’ and non-adaptive variation between populations of Cepaea (Mollusca). Heredity
18, 459–465.
Lamotte, M. 1951 Recherches sur la structure génétique des populations naturelles de Cepaea nemoralis (L.) Bull.
Biol. Suppl. 35, 1–239.
OTHER ACCOUNTS OF ARTHUR CAIN
Claridge, M. 2000 Obituary. Professor Arthur J. Cain FLS FRS. A personal reminiscence. Linnean 16, 39–41.
Clarke, B. 1999 Arthur James Cain (1921–99); naturalist and doughty Darwinian. Nature 401, 872.
Clarke, B. & Parkin, D. T. 1989 Professor Arthur James Cain. Biol. J. Linn. Soc. 36, 1–2.

Arthur James Cain 57
BIBLIOGRAPHY
The following publications are those referred to directly in the text. A full bibliography is
available as electronic supplementary material at http://dx.doi.org/10.1098/rsbm.2007.0034
or via http://journals.royalsociety.org.
(1) 1947 The use of Nile Blue in the examination of lipoids. Q. J. Microsc. Sci. 88, 383–392.
(2) 1953 Geography, ecology and coexistence in relation to the biological definition of the species. Evolution 7,
76–83.
(3) 1954 (With P. M. Sheppard) Natural selection in Cepaea. Genetics 39, 89–116.
(4) (With P. M. Sheppard) The theory of adaptive polymorphism. Am. Nat. 88, 321–326.
(5) Animal species and their evolution. London: Hutchinson’s University Library.
(6) 1955 A revision of Trichoglossus haematodus and of the Australian platycercine parrots. Ibis 97, 432–479.
(7) 1956 (With P. M. Sheppard) Adaptive and selective value. Am. Nat. 90, 202–203.
(8) The genus in evolutionary taxonomy. Syst. Zool. 5, 97–109.
(9) 1957 Range-changes and differential selection in fruit pigeons of the Ptilinopus purpuratus species-group.
Proc. 8th Pacific Sci. Congr. 31, 1393–1412.
(10) 1958 Logic and memory in Linnaeus’s system of taxonomy. Proc. Linn. Soc. Lond. 169, 125–128.
(11) (With G. A. Harrison) An analysis of the taxonomist’s judgement of affinity. Proc. Zool. Soc. Lond.
131, 85–98.
(12) 1959 Deductive and inductive methods in post-Linnaean taxonomy. Proc. Linn. Soc. Lond. 170, 185–217.
(13) The post-Linnaean development of taxonomy. Proc. Linn. Soc. Lond. 170, 234–244.
(14) 1960 (With G. A. Harrison) Phyletic weighting. Proc. Zool. Soc. Lond. 135, 1–31.
(15) 1963 (With J. D. Currey) Area effects in Cepaea. Phil. Trans. R. Soc. B 246, 1–81.
(16) 1964 The perfection of animals. Viewpoints Biol. 3, 36–63.
(17) 1988 Evolution. Plenary address given by the President of the European Society for Evolutionary Biology,
on the occasion of the Society’s founding congress, 26 August 1987, in Basel. J. Evol. Biol. 1, 185–
194.
(18) 1989 The perfection of animals (reprinted from Viewpoints In Biology vol. 3: 36–63, 1964). Biol. J. Linn.
Soc. 36, 3–29.
(19) Validation of a histochemical test—a Citation Classic commentary on the use of Nile Blue in the
examination of lipoids by Cain AJ. Curr. Contents/Clin. Med. (29), 16.
(20) 1995 Linnaeus’s natural and artificial arrangements of plants. Bot. J. Linn. Soc. 117, 73–133.