Carrington Bonsor Williams, 7 October 1889 - 12 July...

1889 - 12 July 1981Carrington Bonsor Williams, 7 October

Sir Vincent Wigglesworth, F. R. S.

1982, 666-684, published 1 November281982 Biogr. Mems Fell. R. Soc.

Email alerting service

herecorner of the article or click this article - sign up in the box at the top right-hand Receive free email alerts when new articles cite

http://rsbm.royalsocietypublishing.org/subscriptions, go to: Biogr. Mems Fell. R. Soc.To subscribe to

on July 14, 2018http://rsbm.royalsocietypublishing.org/Downloaded from on July 14, 2018http://rsbm.royalsocietypublishing.org/Downloaded from

http://rsbm.royalsocietypublishing.org//cgi/alerts/ctalert

http://rsbm.royalsocietypublishing.org/subscriptions

http://rsbm.royalsocietypublishing.org/


on July 14, 2018http://rsbm.royalsocietypublishing.org/Downloaded from


C A R R IN G T O N BONSOR W IL L IA M S

7 O ctober 1889— 12 July 1981

Elected F .R .S . 1954

By S ir V i n c e n t W i g g l e s w o r t h , F . R . S .

C a r r i n g t o n B o n s o r W i l l i a m s was born in Liverpool on 7 O ctober 1889. His distinctive C hristian names were wasted on him , for he was always known, by jun iors and seniors, by friends and doubtless (if he had ever had any) by enemies, and by his wife, as C. B. His father was Alfred W illiams, banker, his m other Lillian Bonsor W illiam s K irkland). T h e first seven years of his life were spent in his hom e over the bank, at one of the busiest street corners in Liverpool. T h e sole house door led straight on to the pavem ent and there was not a living tree w ithin a mile. Perhaps by perversity he developed a consum ing interest in natural history w hich persisted throughout his long life, only thinly disguised by his role as a professional entom ologist. His parents were not scientifically m inded; in the home there was m ore talk of music. But to com pensate for the absence of country he and his elder sister were given books on natural history, they kept an aquarium , and on Sunday m ornings they could feed the seagulls from the landing stage on the River M ersey.

A bout 1897 the family moved to the Cheshire coast where in those days there were still open fields. D uring a sum m er holiday at Beddgelert, when he was about twelve years old, he and his sister were introduced by the local doctor to the world of caterpillars. T hey returned home laden with elephant hawks and woolly bears and he became a keen butterfly hun ter and m oth collector, in the unusual bu t interesting environm ent of the coastal sand dunes. It was a red-letter day when they were taken by a friend to a m eeting of the Lancashire and Cheshire Entomological Society, which consisted of ‘elderly men, nearly all over tw enty years of age’. A friend put up his name for m em bership and he was duly elected. It was only some years later he learnt that the m em bers thought they were voting for his father and were dismayed when they discovered the dim inutive size of the new m em ber.

C. B. was educated at a preparatory school in Cheshire and then at Birkenhead School from 1903 to 1908, where he obtained a school leaving

667



scholarship to university and an entrance scholarship at Clare College, Cam bridge. He had no idea that any branch of biology could qualify one for a profession and since school science included only chem istry and physics he planned (rather vaguely) to become a chem ist. But one big reason for wishing to go to Cam bridge was its proxim ity to W icken Fen— the home of the swallow tail butterflies— where he was to spend many happy days and nights collecting. For the tripos Part I he took physics, chem istry, botany and zoology. His interest in the life sciences developed rapidly and in his final year he took the D iplom a of A griculture.

It was the heyday of the rediscovery of M endel’s laws of inheritance. He came in contact with W illiam Bateson who was a leading figure in these developm ents, and Bateson got him to assist in some of his experim ents by breeding caterpillars. He realized the extent of his good fortune in 1911 when Bateson had been appointed the first D irector of the John Innes H orticultural Institu tion at M erton and C. B. was offered a research studentship in entomology.

J o h n I n n e s H o r t i c u l t u r a l I n s t i t u t i o n :S y s t e m a t i c s a n d B i o l o g y o f T h y s a n o p t e r a

W illiams rem ained at M erton for nearly five years, living in an atm osphere of genetics and under the influence of the wide outlook and culture of Professor and M rs Bateson. T hey were visited by young geneticists from all over the world. He rem em bered vividly the Russian biologist Vavilov, who later rose to fame and died mysteriously during the time of the Lysenko scandal. D uring this period (1914-15) he held a six m onths’ Carnegie Travelling Scholarship— ‘six glorious m onths in the U nited States and the W est Indies’—officially seeing how Americans coped with insect pests bu t gaining varied experience and making new friends. In New York he met T . H. M organ, at that time a friendly rival of Bateson and the great protagonist of the ‘chrom osom e theory’ of inheritance, which Bateson did not then accept.

W illiam s’s own researches at the Institu tion concerned chiefly the Thysanoptera; he wrote on the biology and systematics of the order, as well as their economic im portance; and in collaboration with J. D . Hood of the U nited States, described new species and made some revisions in classification.

T hen came W orld W ar I; C. B. was in training at the London School of Tropical M edicine, to assist the R .A .M .C . in identifying pathogenic bacteria and protozoa; largely engaged in ‘that unheroic branch of w arfare’, the exam ination of the stools of dysentery patients. T hen one day he was called from his microscope to see a representative of the Colonial Office who asked him to go out im mediately to the W est Indies to investigate a serious insect pest of sugar cane. In those days there was no English sugar beet crop and cane sugar was urgently needed. H ad

668 Biographical Memoirs



Carrington Bonsor Williams 669

there been no war, and had he been a m illionaire, he would have planned to go back to the W est Indies— w hich he had visited for a few weeks in the previous year— and here was a kindly G overnm ent offering to free him from the dysentery stools and to pay him a salary to go. Five weeks later he landed in T rin idad and was to live in the W est Indies for nearly six years.

T h e W e s t I n d i e s ; s u g a r c a n e p e s t s , b u t t e r f l y

M IG R A T IO N , A N D M ARRIAGE

T h e pest in question was Tomaspis saccharina, the sugar cane froghopper, w hich had been an increasing cause of sugar cane blight for many years. J. C. Kershaw had recom m ended in 1913 that an attem pt should be m ade to introduce an efficient parasite from outside T rin idad for biological control. T he appointm ent of W illiam s was intended to im plem ent this recom m endation— but on an alm ost hopelessly lim ited scale. C. B. explored the sugar plantations of Surinam , British G uiana, Tobago, G renada, Costa Rica and Panam a. In m any places froghoppers of varied species were attacking sugar cane— but always the parasites present were those already active in T rin idad . He could only point out that the search should be extended outside central America, perhaps preferably in Africa. He did establish that soil conditions had a close relation w ith insect damage; and in his final year he discovered by chance that M osaic virus disease had recently been in troduced into the island and he was able to initiate a cam paign of continual inspection and continual destruction of infected stools w hich was ultim ately successful. ‘So the problem that was my official work was largely unsolved, bu t I saved the island many thousands of pounds by an accidental discovery outside my proper sphere. W hat bad organization!’

Tw o other events of great im portance to him occurred in the W est Indies. In British G uiana he saw his first m igration of butterflies: thousands of yellow Pierids, the Cloudless Su lphur Phoebis eubule flying southeast, day after day for at least a fortnight; and later he was able to see a sim ilar flight in T rin idad . So started w hat became initially his chief hobby, the study of butterfly m igration, w hich eventually became bound up with his official work.

T he second event was his m arriage in 1920 to Ellen M argaret Bain, the daughter of a British W est Indian planter, John Purdie Bain and his wife M ary Rebecca Olton. T h is was a trium phant m arriage. Almost fifty years later C. B. was still celebrating it by dedicating his great book on Patterns in the balance o f nature to his wife ‘in gratitude for her wisdom in the art of living’. T hey had three sons; the eldest became a professional statistician in the governm ent service; the second an engineer (having been told by his headm aster that no boy was encouraged to study biology if he was sufficiently intelligent to learn physics); the th ird entered scientific



medicine, having spent his early career at the Virus Research Station in Uganda.

In 1921 the Williamses returned to England on a somewhat belated honeymoon and while at home he accepted a post in the Entomological Section of the M inistry of A griculture in Egypt.


E g y p t , T a n g a n y i k a , E d i n b u r g h

On the voyage to Egypt C. B. w itnessed a m igration of Painted Lady butterflies northw ards across the M editerranean, which renewed his interest and brought home to him the fact that m igration of butterflies is not confined to the tropics. 1 he pink boll worm Gelechia gossypiella, introduced into Egypt about 15 years earlier, overshadowed all other problem s in cotton production in the Nile Valley. At the outset he was occupied in research on the control of this insect. But after one year he succeeded to the post of D irector of the Entomological Section, a large organization concerned not only with research and advisory work bu t with actually carrying out control measures. These included the fum igation of nearly a million citrus trees w ith hydrogen cyanide gas under great tents night after night to control scale insects. His work was therefore mainly adm inistrative. But alongside such duties C. B. was studying the relation of insects to the climatic conditions in the Egyptian desert, and showing that their survival depended on their utilizing retreats in which the ‘m icroclim ate’ was very different from the climate as m easured by the meteorologist.

T he practical significance of insect m igration was brought home to C. B. while in Egypt, where the Greasy Cutw orm Agrotis ypsilon attacks w inter crops of grain and sometimes young cotton in the spring. M uch effort was spent on trying to poison the m oths w ith sugary baits each year in early spring. But it turned out that all these m oths are im m ature and all m igrate northw ards to Europe before they lay their eggs.

After six years in Egypt, in June 1927 he left for Amani, the old G erm an botanical and agricultural station in the hills of northeast Tanganyika which had just been reopened by the Colonial Office. One of his first jobs was to put four miles of road throughout the forest, with two bridges. ‘I then felt how valuable a good training in entomology can b e .’ He spent only two years at Amani which proved to be an insect paradise: butterflies scarcely ever stopped m igrating past the house.

In his second year they experienced a locust invasion, one swarm of which was estimated to contain about ten thousand million individuals. Among other investigations called for was a study of how locusts bring trains to a standstill by getting crushed and greasing the rails under the driving wheels— a study that entailed lying at full length on the cow catcher of the locomotive in transit.

He made the interesting observation that the swarms were accom




panied by the black hun ting wasp Sphcx aegyptiaeus which preys on the locusts, paralysing them and depositing them in burrow s as food for their offspring. But when the swarm moved on the wasps went with them , leaving behind hundreds of unfinished burrow s and scores of paralysed locusts lying by the waysides.

In 1929 C. B and his family came home on leave with every intention of return ing to East Africa. Indeed all their goods and chattels were left in Amani. But he was offered the Steven lectureship in A gricultural and Forest Entom ology in the U niversity of E dinburgh , and—with the education of three growing boys now becom ing a m ajor problem — this was an opportunity not to be missed. So they settled down to life in a city w here—with milk delivered at the door, letters arriving three times a day and regular lecturing hours— they ‘became almost civilized’. T hey greatly enjoyed their relations with the students, to whom they were ‘at hom e’ on Sunday evenings. It was perhaps the best tim e in their lives and the family never quite forgave him for leaving Edinburgh. But in 1932, after giving a short course of lectures at the U niversity of M innesota, he was appointed head of the Entom ology D epartm ent at the Rotham sted Experim ental Station.

B u t t e r f l y M i g r a t i o n

Ever since his experience in British G uiana in 1916 the study of butterfly m igration had become a m ajor interest for C. B., what he regarded as a hobby, bu t which entailed a continuous flow of publications from 1917 to 1970. T hese com prised personal observations; collected records from the very scattered literature; reviews of published records; and observations by the num erous friends all over the w orld whom he inspired and indoctrinated with his own enthusiasm . D uring the tim e in E dinburgh he brought all this material together in his book The migration of butterflies (473 pages) 1930. Except for the devotee this is not a very readable book. It is a catalogue of everything that has ever been published (and recovered) on m igrating butterflies. But it established C. B. as the world authority on the subject. A nd it is not only a catalogue for it form ulates all the major problem s: the origin of the m igratory habit; the question of w hether there is a return flight by the species; the relation of flight direction to the wind; the evidence for continuous flight in a constant direction; the cues utilized in orientation, and above all the ‘m ystery of butterfly m igration’.

T he 1930 book was only a m ilestone on the way. Publication along the same lines continued until C. B. was m ore than eighty years of age. He stim ulated interest in the study of m igration in British butterflies, enlisting the enthusiastic support of Capt. T . D annreu ther from 1931 onwards (later followed by R. A. French at Rotham sted) in producing a series of A nnual Reports on im m igrant insects in Britain.

In 1958, some years after his retirem ent, he produced a second book,



Insect migration, in the New N aturalist series. T h is is a highly readable account of the same m atters as before, w ith an additional th irty years of experience and research (C. B. him self had had fu rther personal experience in the Pyrenees, in W est Africa and in South America). T he book covers also in a com parative fashion all other m igratory insects and it is still alive with the old enthusiasm and th irst for factual knowledge.


R o t h a m s t e d E x p e r i m e n t a l S t a t i o n : f l u c t u a t i o n i n

INSECT ABU NDA NCE

W hen W illiams joined the staff at Rotham sted in 1932 Sir John Russell was D irector. It was Russell who, some ten years earlier, had had the brilliant intuition to invite R. A. F isher to join the Station in order to provide statistical guidance on the in terpretation of field trials. T h e result was a revolution in the design of trials which spread throughout the world and involved every branch of experim ental science. It was not surprising that Russell should welcome the proposal that the new head of E n tomology should study the effect of w eather conditions on insect activity and abundance, and particularly to try to assess the part played by the physical environm ent in causing mass outbreaks of pests. W ith the coming of statistical m ethods, and notably the use of m ultiple regressions, it had become possible to extend the num ber of variables studied simultaneously, to consider their interaction and so to gain in wider application what was lost from the accuracy of laboratory control.

T h is approach appealed to C. B. as a field entomologist; so ‘instead of disappearing in a constant tem perature room in the cellars’ he was able to pu t his light traps in the field and to count the num ber of insects which had been caught during the night. T he num ber ranged from nothing to over 70000 in a single night. Each night nature presented a different complex of environm ental conditions, many of which such as tem perature, wind, hum idity, cloud cover and barom etric pressure— could be m easured by standardized meteorological m ethods. After a few m onths it was possible to analyse some of the activity effects, and after four years, with over 1400 replications, it was possible to com mence a study of population changes from year to year. ‘T he whole work was carried out under natural conditions in the field, and no insect was interfered with in any way up to the m om ent that it entered the tra p .’

T he prim ary object of the collections was as a sampling m ethod to obtain data that could be used in statistical analysis. He realized that statistical formulae based on the assum ption that variation about a mean is equal and opposite on an arithm etic scale give unrealistic results when applied to insects whose num bers are varying geometrically. In 1937 he wrote of the advantages of handling the logarithm s of the num bers caught rather than the actual figures— in order to reduce the effects of exceptionally large catches on a single night and to obtain curves which more nearly




approached norm al curves of d istribu tion . T h e benefits of using logarithm s was well shown in a later paper on the analysis of sentence length as a criterion of literary style (1940, Biometrika) in w hich the transform ation produces a norm al instead of a skew distribu tion .

T h e total num ber of insects caught in the light trap in different years varied from 100 thousand to 400 thousand. T h e results as published contain a vast quantity of data on the relative num ber of different orders of insects caught, the sex ratio (which varied greatly in different species), in tim es of capture during the night, the phenology of different species, and the effect of the height of the trap above the ground. C onsistent results are recorded on m any of these subjects bu t no general conclusions of im portance em erge (1939).

In his paper of 1940 {Trans. R . ent. Soc.) and after a second four year spell of trapping his paper of 1951 ( P. R. Soc. Lond.) the results were analysed in great detail to show the effect of w eather conditions on insect activity and to attem pt the forecasting of changes in the insect population. He concluded that the catch depended chiefly upon two factors ‘activity’ and ‘popula tion’. Regression form ulae were obtained by which population changes could be estim ated from a knowledge of previous w eather conditions only, rainfall being the m ost im portant factor in sum m er and tem perature in w inter. Population changes as estim ated from the trap catches and as calculated from the rainfall and tem perature of three previous m onths were closely similar. Between 50% and 60% of the variance of the populations could be accounted for. T hese results refer to the total population of all insects; results for single species were not worked out.

(I t should perhaps be pointed out that in this work ‘activity’ is referred to as a simple param eter; bu t when, largely through encouragem ent by C. B., the actual causes of changes in the ‘activity’, that is, the presence in the air of aphids, was explored in his D epartm ent by C. G. Johnson, this was found to be the outcom e of a highly complex interaction of many factors, biological and meteorological.)

F r e q u e n c y d i s t r i b u t i o n : t h e l o g a r i t h m i c s e r i e s

In an unpublished thesis of the U niversity of M innesota in 1928, G arthside had shown that random catches of insects, taken by sweep nets or by traps, contained a great majority of species represented by very small num bers, only a very few species by very large num bers. T he frequency distribution on an arithm etic plot of abundance showed a typical hollow curve. It happened that in 1942 A. S. C orbet published a paper on his collection of butterflies in M alaya which showed a similar relation between the num ber of species and the num ber of individuals of these species. He consulted R. A. Fisher, who was then working in W illiam s’s D epartm ent, about the nature of his frequency curve which



did not give a good fit w ith an hyperbola. As the result of his light trapping over a span of four years W illiams had some 15 000 identified m oths representing about 240 species com parable with Corbet s data but far more num erous and taken w ithout any bias in relation to frequency.

On theoretical grounds Fisher suggested that the true series would not be represented by the hyperbolic series

«i, 2wi>iwi>iwi>but m ight be a logarithm ic series, the expansion of — loge (1 —x) in which the num ber of groups with 1, 2, 3, 4, etc. units per group (i.e. individuals per species) is given by

tl i , ~2M 1 ^ > 3 ^ 1 ^ > 4 ̂ 1

where n x is the num ber of groups with one unit, and x is a constant (for the sample) which is less than unity.

L ater it was found more convenient to use the alternative form

olx, \a x 2, ja x 3, . . . ,

where a is the original n l divided by x.T here are two known param eters, N (the total num ber of individuals)

which equals ocx/( 1 —x) and (the num ber of species) w hich equals — aloge (1 — x ).Hence the values of a and x can be derived and the whole series calculated.

W hen C. B. analysed the data from his 15 000 M acrolepidoptera taken in the light trap they showed an extremely close fit to F isher’s formula. He suggested that the constant a, which indicates the slope of the line relating the num ber of species in the sample with the logarithm of the num ber of individuals in the sample, should be term ed the ‘index of diversity’, which is low when the num ber of species is low in relation to the num ber of individuals, high when the proportion of species is high.

F urther applications suggested that the frequency d istribution of species with different num bers of individuals, especially with very large num bers, is better represented by a logarithm ic-norm al d istribution (as suggested by Preston (1948) and supported by Fisher and by Bliss) than by the log series. But C. B. still found that in some cases the logarithm ic series gave a better fit.

T he jo in t paper by Corbet, W illiams & Fisher (1943) in which the logarithm ic series was form ulated by Fisher proved a turning point in C .B .’s career. He was enthralled by the power of this form ula in providing a simple pattern for complex data; and he set about trying it out on data of all sorts. He showed that U dny Y ule’s ‘characteristic’ ( of diversity is proportional to the reciprocal of the ‘index of diversity’ (a) (1946).





One of the first exam ples tackled was the d istribu tion of parasites on a series of hosts. Patrick Buxton had recorded the num bers of head lice in the hair shaved from prisoners adm itted to a goal in South India. A total of 7442 lice were obtained from 461 heads of hair that contained lice. T h is gives an arithm etic mean of 16.1 lice per head. T he m axim um infestation was 385 lice. C. B. was very excited to discover the close agreem ent betw een a calculated logarithm ic series and the actual findings. T he calculated num ber of heads w ith one louse was 107; the observed value was 106. As C. B. pu t it: ‘So N ature was only one w rong’ (1944).

O ther examples concerned the num ber of species of plants in quadrats of grassland of different sizes (data of Blackman and others); the species of ground vegetation in Tectonia forest in Java (data of A rrhenius); the grouping of species in genera— for exam ple, the d istribu tion of the species of the O rthoptera of the w orld into genera by K irkby shows a very close agreem ent w ith the calculated logarithm ic series; and the same was true in m any other groups of insects and in families of flowering plants— all these and many m ore examples in the 1944 paper.

The concept o f diversity

It is of course obvious that the num ber of genera am ong which a given group of species is divided will depend upon the judgem ent of taxonom ists. T hese may be ‘sp litters’ or ‘lum pers’, bu t provided the principles on which the taxonom ist is w orking are consistent the resulting frequency distribu tion will still follow the log series—only the constant a w hich is determ ined by diversity will be changed. ‘T he differences in diversity are properties of the m inds of the workers. As a large num ber of scientists w ithout any differences of opinion would be disastrous, we m ust welcome the different approaches— provided that in each case the principles have been applied with consistency.’ C. B. does adm it that in the days when he was a system atist of the Thysanoptera he was influenced by convenience and deliberately sought reasons for dividing large genera. In his book on Patterns in the balance o f nature (1964) he writes: ‘I think that insect taxonom y benefited when I discontinued work in this field!’

It had been a general belief that congeneric species tend to exclude one another by com petition in habitats of low environm ental com plexity. But it was shown by W illiams (1947) that in com paring the fauna and flora of contiguous areas, there is a decrease in generic diversity associated with a decrease in ecological com plexity— as revealed in the m ore frequent occurrence of pairs of species belonging to the same genus. Analysis of M oreau’s records of East African bird com m unities confirm ed this relation. C. B. inferred that while closely related species are probably m ore suited to similar physical environm ents, they are m ore likely to exhibit biological com petition. But observed results indicate that the



advantages of close relationship are on average greater than the draw backs (1951).

Extensive observations have been made of the Specific D iversity of populations and most of this work suggests that diversity is low when, as in polar latitudes, the physical conditions of the environm ent are severe, high when, as in the wet tropics, physical conditions are favourable and the balance is chiefly determ ined by biological com petition. W illiams considered the conception of D iversity as a m easurable property of a population to have been one of the more im portant developm ents of quantitative biology.


R e t i r e m e n t

W hen C. B. retired from Rotham sted in 1955 he prom ptly m igrated to Inverness-shire where, taking a house on the m oorland at K incraig, at the foot of the Cairngorm s, he set about repeating the light trap (and suction trap) observations in an environm ent as different as possible from that of H ertfordshire. It was not an easy environm ent; it was in some ways like a return to the early days in the tropics— apart from the tem perature. T he work was continuous, requiring regular attendance to the traps and their captures, as well as taking unbroken meteorological records on a fair range of instrum ents. T he work was supported by the A gricultural Research Council. He was single handed, though his wife threw herself eagerly into the unrem itting task of recording the biological and m eteorological data, the traps being operated virtually every night for the four years from May 1955 to A ugust 1959. T he results published in the Philosophical Transactions in 1961 confirmed in broad outline, in respect to the prediction of ‘activity’ and ‘population’ changes, those obtained at Rotham sted (see above).

It soon became evident that it would be a good plan to concentrate on the Simuliidae (‘black flies’) of which some 17 500 specimens belonging to ten species were captured. Later the trapping was repeated on a small scale at Selkirk in south east Scotland, where the W illiams later settled. A series of papers described the nocturnal flight of Simuliidae; the phenology of the different species; and in collaboration with other workers the favoured hosts of the different species were determ ined by the use of precipitin tests—and m uch m ore new inform ation about the ‘black flies’ was reported.

They moved to Edinburgh for a year or two after Kincraig, and then to Selkirk and finally to K irkcudbright. C. B. continued to w rite on his favoured topics. As already described, he published Insect migration in 1958. In 1960 in a paper on ‘T he range and pattern of insect abundance’ he made a speculative attem pt to extend the logarithm ic series to estim ate the insect population of the world. He arrived at a figure of three million species (something approaching one million have been nam ed at the




present time) and a basic figure for the total population of insects in the w orld at any tim e of 1018.

In 1964 he published Patterns in the balance o f nature which is a detailed review and discussion of all the fields of quantitative biology with w hich he had been occupied th roughout his research career— in short, on m athem atical synecology. It is a rem arkable book in w hich are foreshadow ed very m any of the argum ents w hich are being advanced at the present tim e in studies of m ultispecies populations, island biogeography and the effects of com petition on evolutionary generic relationships— often w ithout acknow ledgem ent.

S c i e n t i f i c a s s e s s m e n t

In his early days at C am bridge and at M erton C. B. had been a close friend of Patrick Buxton, who was some two years jun io r to him and they had published a small am ount of jo in t work. T here are m any points of sim ilarity between them . Both were keen naturalists and both were attracted to statistical studies of ecology and the relation of climate, in particu lar of ‘m icroclim ate’, to the life of insects in the hot deserts; and both tu rned to the scientific study of all sorts of incidental topics. Both became applied entom ologists, Buxton in the medical field and C. B. in the agricultural.

As an agricultural entom ologist C. B .’s heart was not in coping with the day to day problem s of the farm er. He was inspired by the m ore remote am bition of understanding the factors behind the sudden changes in abundance that tu rn insects into pests. In his presidential address to the Association of A pplied Biologists in 1947, at the height of the excitem ent about D D T , he predicted that the excitem ent would soon die down and that eventually applied entomology w ould be concerned with the p rediction and prevention of outbreaks on the basis of integrated knowledge of meteorology and insect ecology.

His claims to fame lie in the two fields of butterfly m igration and statistical synecology. T he study of butterfly m igration was a passion. By m arshalling all existing published observations, recruiting friends all over the world to make careful new observations and analysing the results in a long series of publications he became the authority on the subject. M any of the problem s which he form ulated in his 1930 book had been clarified by the tim e of the 1958 book: the relations between wind direction and m igration, the existence of return flights, the optom otor response to other moving insects, the cues to orientation offered by the sun and the pattern of polarized light in the sky, and the ability of insects to estim ate the time of day. All this had made many aspects of butterfly m igration intelligible; bu t it had not dispersed the ‘m ystery’ of long range m igration. Indeed I sometimes felt that if the m ystery had been resolved the subject m ight have lost some of its charm!




Some of the most illum inating work has come from inside his own D epartm ent at Rotham sted, where C. G. Johnson has pointed out that there is commonly an explosive period of scattering in the early adult life of most winged insects; m igratory flight seems to be the prim e locomotive act in many forms. O ther insects are caused to leave their environm ent by stim uli which past evolutionary experience has shown to be a w arning of bad times to come. In many other ways this work has illustrated the link between spectacular m igrations and the every day behaviour of what are usually regarded as non-m igrant species.

In the field of statistical ecology C. B .’s patient labours of light trapping alongside meteorological records gave repeatable results from which predictions of abundance could be made. C. B. W illiams has been described as ‘the first real quantitative em pirical ecologist’— a naturalist who was num erate. But it should be rem em bered that C. B .’s statistical ecology dates forty to fifty years ago. T he subject has become more sophisticated since that time. Considerations of insect behaviour in the widest sense have largely transform ed the term s in which these m atters are now discussed.

In 1952, a few years before he retired, W illiams argued that the use of population data for control would in the future, require a national if not an international scale of organization. T he m ost tangible outcom e of this argum ent has been the establishm ent in later years of the Rotham sted Insect Survey, which developed a dual series of traps: light traps as used by C. B. and suction traps of the type developed by C. G. Johnson. T he traps have been located throughout the British Isles and in recent years have been extended to the continent of Europe. T his cooperative organization and the sophistication in the interpretation of the results by L. R. Taylor and others far outdistance what C. B. accom plished—but the initial steps and the original conceptions were his.

D i s t i n c t i o n s

C. B. W illiams obtained the Cam bridge Sc.D . He received the O rder of the Nile 4th Class for service in Egypt. He was President of the Royal Entomological Society of London, President and an H onorary M em ber of the Association of Applied Biologists and President of the British Ecological Society, and an H onorary M em ber of the entomological societies of Egypt, the N etherlands, Brazil and Chile. He was a m em ber of the A nti-Locust Advisory Com m ittee of the Colonial Office and of the Scottish Com m ittee of the N ature Conservancy.

P e r s o n a l i t y

As a person C. B. was greatly liked. He was a popular and successful head of D epartm ent at Rotham sted. In those days the D epartm ent was small, and independent, and he had sufficient time for his own personal




research, the excitem ent of w hich he shared with all his colleagues. All were sure of a welcome in the hom e that he and M rs W illiam s made in H arpenden. C. B. really cared about people and was ready to help in their problem s, both personal and scientific. He encouraged young workers to follow their own ‘hunches’ bu t was always ready to discuss and to catalyse their ideas. H is enthusiasm was infectious and he was a m ost stim ulating and kindly person.

T h is m em oir has em phasized that C. B. was a field naturalist. He him self preferred to do his scientific work in the field; and by encouragem ent and by exam ple he got m ost of his colleagues at R otham sted out into the field. Perhaps they were not encouraged to work in as close contact w ith the curren t problem s of the farm ers as they m ight have been— but the tradition of research work in the field has paid off and some of the greatest recent successes in entom ology at R otham sted had their roots in the W illiams era.

It was strange that his interest in natural history failed to ensure an interest in the individual insect and its behaviour; the insect often became only a un it for a calculator. But his forte was the m athem atical handling of samples and perhaps it w ould have been asking too m uch for the two modes of thought to be com bined. He never seemed really to appreciate the im portant bearing of the work he had initiated on aphid behaviour (by C. G. Johnson) on the basic problem s of insect m igration.

C. B. had a puckish enthusiasm for riddles, puns, limericks and m athem atical puzzles and paradoxes; and was always anxious to share his latest discovery with others. A fter his retirem ent he gave some charm ing short broadcasts on the character and uses of chance and similar m athem atical problem s. His article on ‘T he duties of an adm inistrative entom ologist’ (1929) is a m asterpiece of hum our. He was often bubbling over w ith fun and could be extrem ely am using. He once received a prize from the New Statesman for a novel form of dentistry in w hich the drill was a fixture and it was the patient strapped to the chair who was whizzed round. A nother famous invention was a m ore equitable m ode of providing fair handicaps in racing; the com petitors would all start level bu t at half way during the race they would be stopped and required to run back to the starting point—w here they would all arrive at the same instant.

Patrick Buxton used to tell a story of C. B. being found sitting on a log in the rain forest of T rin idad m urm uring: ‘She loves me, she loves me not!’ as he plucked the limbs off a giant centipede. T h is is not the kind of story that an author can vouch for— but it is not incongruous.

I am indebted to form er colleagues (and successors) of C. B. W illiams at Rotham sted: D r C. G. Johnson, D r C. G. Butler, F .R .S ., D r L. R. T aylor and M r R. A. French for helpful inform ation.

T he photograph was taken by Elliott and Fry about 1948 and was kindly supplied by the Photographic D epartm ent at Rotham sted.




B i b l o g r a p h y ( s e l e c t e d )

Systematics and biology o f Thysanoptera1913 On two new species of Thysanoptera from the W est Indies. Jf. Biol. 8, 209—215.

Records and descriptions of British Thysanoptera. J . econ. Biol. 8, 216-230.1914 A new species of Chirothrips (Thysanoptera) from South America. Entomologist 47, 51-53.

Kakothrips n.g., a division of the genus Frankliniella (Thysanoptera). Entomologist 47,247-248.

1915 The Pea Thrips ( Kakothripsrobustus). Ann. appl. Biol. 1, 222-246.(With J. D. H ood) New Thysanoptera from Florida and Louisiana, jfl N .Y . ent. Soc. 23,

121-138.A new thrips damaging coffee in British East Africa. Bull. ent. Res. 6, 269—272.

1916 Thrips oryzae sp. nov. injurious to rice in India. Bull ent. Res. 6, 353-355.Biological and systematic notes on British Thysanoptera. Entomologist 49, 221-227; 243-245;

275-284.1917 A new thrips damaging orchids in the West Indies. Bull. ent. Res. 8, 59-61.1918 Notes on some Trinidad thrips of economic importance. Bull. Dep. Agric. Trin. Tobago 17,

143-146.1921 A blood sucking thrips. Entomologist 54, 163-164. ̂ . . ,1925 (With J. D. H ood) In H ood’s ‘New neotropical Thysanoptera collected by C. B. W illiam s’

Psyche 32, 48—69.1927 (With J. D. H ood) A synopsis of the Thysanopterous family Urothripidae. Ann. ent. Soc. Am.

20, 1—8.1961 Notes on Thysanoptera. Entomologist 94, 139-140.1972 A graphical method of demonstrating allometric growth, with special reference to the

antennae of Thysanoptera. J. Entomol. (R . Ent. Soc. London) B41, 151-153.

Sugar cane pests and diseases in the West Indies1918 The sugar-cane froghopper in Grenada. Bull. ent. Res. 9, 83-87.

A froghopper on sugar-cane in British Guiana. Bull. ent. Res. 9, 163-173.1919 The food of the mongoose in Trinidad. Bull. Dep. Agric. Trin. Tobago 17, 167-176.

Relation of root-fungus to froghopper blight. Bull. Dep. Agric. Trin. Tobago 18, 52-56. Sugar-cane varieties and froghopper blight. Bull. Dep. Agric. Trin. Tobago 18, 70—83. Rainfall, sugar production and froghopper blight in Trinidad. Bull. Dep. Agric. Trin. Tobago

18, 153-167.1920 (With W m . N owell) Sugar-cane blight in Trinidad: a summary of conclusions. Bull. Dep.

Agric. Trin. Tobago 19, 8-10.The mosaic disease of sugar-cane in Trinidad. Bull. Dep. Agric. Trin. Tobago 19, 30-37.

1921 Report on froghopper blight of sugar-cane in Trinidad. Bull. Dep. Agric. Trin. TobagoMemoir no. 1, 170 pages.

1923 A froghopper damaging cacao in Panama. Bull. ent. Res. 13, 271-274.

Pests of cotton in Egypt1923 The Pink Boll worm in Egypt in 1922. Third Ann. Rep. Cott. Res. B d Egypt pp. 1-7.1925 (With I. Bishara) The survival of the Pink Boll worm in buried seed during the winter in

Egypt. Tech. Bull. Minist. Agric. Egypt. 58, 7 pages.1933 The Boll worms of cotton. Emp. Cott. Grow. Rev. 10, 273-281.1934 The cotton stainer problem. Emp. Cott. Grow. Rev. 11, 99-110.

Insect migration1917 Some notes on butterfly migration in British Guiana. Trans. R. ent. Soc. Lond. 1917, 154-164.1919 A migration of yellow butterflies (Catopsilia statira) in Trinidad. Trans. R. ent. Soc. Lond.

1919, 76-88.1920 Records on insect migration in tropical America. Trans. R. ent. Soc. Lond. 1920, 146-165. 1923 Records and problems of insect migration. Trans. R. ent. Soc. Lond. 1923, 207-233.1925 Notes on insect migration in Egypt and the near East. Trans. R. ent. Soc. Lond. 1924,

439-456.The migrations of the Painted Lady Butterfly. N ature , Lond. 115, 535-537.




1926 Some unsolved problems of insect migration. Proc. 3rd Int. Congr. Ent., Zurich 2, 100-108. Voluntary or involuntary migration of insects. Entomologist 59, 281-288.Further records of insect migration. Trans. R. ent. Soc. Lond. 74, 193-202.

1927 Records of migratory insects, chiefly from Africa. Bull. Soc. ent. Egypt 10, 224-256.A study of butterfly migration in South India and Ceylon. Trans. R. ent. Soc. Lond. 75, 1-33.

1928 Collected records relating to insect migration. Trans R. ent. Soc. Lond. 76, 79-91.1929 Some records of dragon-fly migration. Entomologist 62, 145-148.

Evidence for the migration of butterflies. Bull. Soc. ent. Egypt 13, 193-210.1930 Records of butterfly migration in East Africa. J l E. Africa Uganda nat. Hist. Soc. No. 35,

9-24.Collected records relating to insect migration: second series. Trans. R. ent. Soc. Lond. 78,

139-170.1931 A migratory flight of the butterfly Belenois mesentina in East Africa in 1930. Proc. R. phys. Soc.

Edinb. 22, 35-39.1933 Further collected records relating to insect migration. Trans. R. ent. Soc. Lond. 81, 103-115.

Observations on the Desert Locust in East Africa. Ann. appl. Biol. 20, 463-497.1935 Further evidence for the migration of butterflies. Bull. Soc. ent. Egypt. 1935, 250-261.

British immigrant butterflies and moths. British Museum Pamphlet E.571936 Collected records relating to insect migration. Third Series. Proc. R. ent. Soc. Lond. A l l ,

6- 10.

1937 Butterfly travellers. Natn. geogr. M ag. 71, 568-585.Butterfly migration in the tropics. British Museum Pamphlet E.58.The migrations of day-flying moths of the genus Urania in tropical America. Proc. R. ent.

Soc. Lond. A12, 141-147.1938 Recent progress in the study of some North American migrant butterflies. Ann. ent. Soc. Am.

31, 211-239.The migration of butterflies in India. J . Bombay nat. Hist. Soc. 40, 439-457.

1939 The migration of the Cabbage-W hite butterfly (Pieris brassicae). 7th Int. Congr. Ent., Berlin 1,482-492.

Records of butterfly migration in Africa. Proc. R. ent. Soc. Lond. A 14, 69-74.Some butterfly migrations in Europe, Asia and Australia. Proc. R. ent. Soc. Lond. A 14,

131-137.Some records of butterfly migration in America. Proc. R. ent. Soc. Lond. A 14, 139-144.

1942 (With C ockb ill , D ownes & G ibbs) Studies in the migration of Lepidoptera. Proc. R. ent.Soc. Lond. 92, 101-283, with oz. 600 ref.

1943 Notes on some Monarch butterflies caught in Great Britain. Entomologist 76, 1-3.1945 Occurrence of Vanessa cardui at sea off the West African coast. Proc. R. ent. Soc. Lond. A20,

4-5 .Notes on the fat content of two migrant moths. Proc. R. ent. Soc. Lond. A20, 6-13. Evidence of migration of Lepidoptera in South America. Revta Ent., Rio de J . 16, 113-131. (With G. Beall) Geographical variation in wing-length of Danais plexippus. Proc. R. ent. Soc.

Lond. A 20, 65-76.1946 Migration of butterflies in South America, in Livre de Hominagem a R. F. d ’Almeida, pp.

335-340, Sao Paulo, Brasil.1949 Migration of Lepidoptera and the problem of orientation. Proc. R. ent. Soc. Lond. C.13,

70-84.1950 (With D. L o n g ). Phase colouration in larvae of Lepidoptera. N ature, Lond. 166, 1035.1951 Seasonal changes of flight direction of migrant butterflies in the British Isles. J . Anim. Ecol.

20, 180-190.Migrations of Libytheine butterflies in Africa. Niger Fid 16, 152-159.

1953 Migration and drift of insects in its international aspect. Trans. 9th Int. Congr. Ent.,Amsterdam, 1951 1, 63-68.

1954 Notes on some migrations of butterflies in West Africa. Entomologist 87, 203-206.1956 (With C o m m o n , F rench , M uspratt & W ill iams , M. C.) Observations of the migrations of

insects in the Pyrenees. Trans R. ent. Soc. Lond. 108, 385-407.1957 Insect migration. A . Rev. Ent. 2, 163-180.1970 The migrations of the Painted Lady butterfly ( Vanessa cardui) with special reference to North

Am erica.^. Lepid. Soc. 24, 157-175.1976 The migration of the hesperid butterfly, Andronymous neander Plotz, in Africa. Ecol. Ent. 1,

213-220.



Traps and other techniques1913 The Berlese funnel. Entomologist 46, 273-274.1924 (With T. W. K irkpatrick) A multiple temperature incubator. Tech. Bull. M imst. Agnc.

Egypt 38, 4 pages.An improved light trap for insects. Bull. ent. Res. 15, 57 60.

1935 (With G. A. E mery) A photographic moonlight recorder. J. scient. Instrum. 12, 111 115.1936 (With P. S. M ilne) A mechanical insect trap. Bull. ent. Res. 26, 543-551.

A modified Greenwich night-cloud recorder, used for ecological work. J. Amm. Ecol. 5, 348-350.

1943 A safe method of measuring the wings of set butterflies. Proc. R. ent. Soc. Lond. A 18, 3 5. 1948 The Rothamsted light trap. Proc. R. ent. Soc. Lond. A 23, 80-85.1951 Comparing the efficiency of insect traps. Bull. ent. Res. 42, 513-517.1955 A second experiment in testing the efficiency of insect traps. Bull. ent. Res. 46, 193-204.


General ecology, bioclimatics and insect abundance1923 A short bioclimatic study in the Egyptian desert. Tech. Bull. M inist. Agrtc. Egypt. 29, 18

pages.The cotton plant in relation to temperature and rainfall. Tech. Bull. M inist. Agnc. Egypt. 32, 5

pages.1924 Bioclimatic observations in the Egyptian desert in March 1923 Tech. Bull. M inist. Agnc.

Egypt. 37, 18 pages.(With E. M cK enzie T aylor) A comparison of sand and soil temperatures in Egypt. Tech.

Bull. Minist. Agric. Egypt. 40, 24 pages.1925 Cotton growing in relation to climate in Egypt and the Sudan. Tech. Bull. M inist. Agric.

Egypt. 47, 31 pages.A third bioclimatic study in the Egyptian desert. Tech. Bull. Minist. Agric. Egypt. 50, 32

pages.The seasonal prevalence of fleas in Egypt. Bull. ent. Res. 15, 353-355.

1926 Seasonal variation in pink boll worm attack on cotton in Egypt in the years 1916-1924. Tech.Bull. Minist. Agric. Egypt 67, 12 pages.

1934 Field studies in the relation of insect pests to climatic conditions. Rep. Conf. Cotton GrowingCorp. London 1934, 111-119.

1935 (With F. J. K illing ton ) Hemerobiidae and Chrysopidae in a light trap at Rothamsted.Trans. Soc. Br. Ent. 2, 145-150.

The times of activity of certain nocturnal insects, chiefly Lepidoptera, as indicated by a light trap. Trans. R. ent. Soc. Lond. 83, 523-555.

1936 The influence of moonlight on the activity of certain nocturnal insects as indicated by a lighttrap. Phil. Trans. R. Soc. Lond. B 226, 337-389.

1937 The use of logarithms in the interpretation of certain entomological problems. Ann. appl.Biol. 24, 404-414.

1939 An analysis of four years captures of insects in a light trap. I. General survey sex proportions,phenology and time of flight. Trans. R. ent. Soc. Lond. 89, 79-131.

1940 An analysis of four years captures of insects in a light trap. 11. The effect of weather conditionson insect activity, and the estimation and forecasting of changes in insect population. Trans. R. ent. Soc. Lond. 90, 227-306.The number of insects caught in a light trap at Rothamsted during four years 1933-1937. Proc. R. ent. Soc. Lond. A 15, 78-80.

1947 The field of research in preventive entomology. Presidential Address. Ann. appl. Biol. 34, 175-185.

1951 Changes in insect populations in the field in relation to preceding weather conditions. Proc. R. Soc. Lond. B 138, 130-156.

1953 Graphical and statistical methods in the study of insect phenology. Trans. 9th Int. Congr.Ent., Amsterdam , 2, 174-189.

1954 Phenology: the study of the seasons. A dvm t Sci., Lond. 11, 267-270.The statistical outlook in relation to ecology. J. Ecol. 42, 1 — 13.Some bioclimatic observations in the Egyptian desert. In: Biology of deserts, pp. 18-27.

London: Institute of Biology.




1956 (With B. P. S ing h & S. El Z iady) An investigation into the possible effects of moonlight onthe activity of insects in the field. Proc. R. ent. Soc. Lond. A 31, 135-144.

1957 (With S. El Z iady) On the relative distribution of insects at 5 and 30 feet. Bull. Soc. ent. Egypt41, 663-675.

1960 (With M. F. H. O sm a n ) A new approach to the problem of optimum temperature for insectactivity.^ . Anim. Ecol. 29, 187-189.

1961 Studies in the effect of weather conditions on the activity and abundance of insectpopulations. Phil. Trans. R. Soc. Lond. B. 244, 331-378.

Population studies: competitiondiversity log-series1943 (With R. A. F isher & A. S. C orbet) T he relation between the number of species and the

number of individuals in a random sample of an animal population. J. Anim. Ecol. 12, 44-53.

Area and number of species. Nature, Lond. 152, 264—266.1944 T he number of publications written by biologists. Ann. Eugen. 12, 143-146.

Some applications of the logarithmic series and the Index of Diversity to ecological problems. J . Ecol. 32, 1-44.

1945 Recent light trap catches of Lepidoptera in U .S .A . analysed in relation to the logarithmicseries and the index of diversity. Ann. ent. Soc. Am. 38, 357-364.

1946 Yule’s ‘characteristic’ and the ‘index of diversity’ N ature, Lond. 157, 482.1947 A diagrammatic method of analysing the relationships of the fauna or flora of several different

localities. Proc. Linn. Soc. Lond. 158, 99-103.The logarithmic series and the comparison of island floras. Proc. Linn. Soc. Lond. 158, 104-108.

T he generic relationships of species in small ecological communities. J . Anim. Ecol. 16, 11-18. The logarithmic series and its application to biological problems. J . Ecol. 34, 255-272.

1949 Jaccard’s General Coefficient and Coefficient of Floral community in relation to thelogarithmic series and the index of diversity. Ann. Bot. N .S . 13, 53-58.

1950 The application of the logarithmic series to the frequency of occurrence of plant species inquadrats.^. Ecol. 38, 107-138.

1951 A note on the relative sizes of genera in the classification of animals and plants. Proc. Linn.Soc. Lond. 162, 171-175.

Diversity as a measurable character of an animal or plant population. Annee biol. (3), 27, 129-141.

Intra-generic competition as illustrated by M oreau’s records of East African bird com m unities. J. Anim. Ecol. 20, 246-253.

1952 Sequences of wet and fine days considered in relation to the logarithmic series. Q. J l R. met.Soc. 78, 91-96.

1953 The relative abundance of different species in wild animal populations. J. Anim. Ecol. 22,14-31.

1960 The range and pattern of insect abundance. Am. N at. 94, 137-151.1964 Some experiences of a Biologist with R. A. Fisher and Statistics. Biometrics 20, 301-307.

Black flies (Sim uliidae) in Scotland1964 (With L. D avies, A. E. R. D ow ne & B. W eitz) Studies on Black flies (Simuliidae) taken in a

light trap in Scotland. Trans. R. ent. Soc. Lond. 114, 1-47.Nocturnal activity of Black flies (Simuliidae) Nature, Lond. 201, 105.

1965 Black flies (Diptera, Simuliidae) in a suction trap in the central highlands of Scotland. Proc.R. ent. Soc. Lond. 40 92-95.

Notes on swarms of male Black flies (Simuliidae) in South east Scotland. Entomologist 103, 135-138.

Miscellaneous entomology and general biology1913 Some biological notes on Raphidia maculicollis, Step. Entomologist 46, 6-8.

A summary of the present knowledge of the Protura. Entomologist 46, 225-232.




1914 Phytodecta viminalis, a viviparous British beetle. Entomologist 47, 249 250.Notes on Podagrion pachymerum, a chalcid parasite of mantis eggs. Entomologist 47, 262-266.

1916 (With F. W. E dwards) Sciara tritici, a fly injurious to seedlings. Ann. appl. Biol. 2 258- 262. (With P. A. B uxton) On the biology of Sphodromantis guttata. Trans. R. ent. Soc. Lond. 1916,

86- 100.1917 Some problems of sex-ratios and parthenogenesis. J . Genet. 6, 255-267.1923 Notes on the food and habits of some Trinidad birds. Bull. Dep. Agric. Tnn. Tobago 20,

1929

1931193919401941 194519501951 1971

123-185.The duties of an administrative entomologist. Scient. Wkr 5, 36- 39. Reprinted 1960.^. Inst.

A new Nemoptera larva and a Mantis parasite from Egypt. Bull. Soc. ent. Egypt 1928, 40-42. A possible racial difference in pupating habit in napi and its variety bryomae.

Entomologist 62, 202-204.Some entomological experiences in the tropics. Trop. Agric., Tnn. 8, 119-123.A tent-building W hite-fly (Aleurodidae) in Trinidad Entomologist 73, 225-227.On ‘T ype’ specimens. Ann. ent. Soc. Am. 33, 621-624.The sense of hearing in fishes. Nature, Lond. 147, 543.An insect-catching grass. Entomologist 76, 37-38.The way by which I have come. (Autobiographical.) Countryman 41, 293-298.On educating an entomologist. Emp. Cott. Grow. Rev. 28, 97-102.Notes on a fungus eating fly in North Britain and in North America. Entomologist 104, 1-3.

Statistics and literary style1940 A note on the statistical analysis of sentence length as a criterion of literary style. Biometnka

31 , 356-361. Reprinted in 1969 in Statistics and Style (ed. Dolezel & Bailey). New York: Elsevier.

1 9 4 4 The number of publications written by biologists. Ann. Eugen. 12(2), 143-146.1956 A note on an early statistical study of literary style. Biometrika 43, 249-256.1967 Writers, readers and arithmetics. New Scient. 35, 88-91.

Books1930 The migration of butterflies. Edinburgh: Oliver & Boyd.1958 Insect migration. London: Collins.1964 Patterns in the balance of nature. London and New York: Academic Press. 1970 Style and vocabulary: numerical studies. London: Griffin and Sons.



Carrington Bonsor Williams, 7 October 1889 - 12 July...

Documents

Transcript of Carrington Bonsor Williams, 7 October 1889 - 12 July...