thomas stanley westoll. 3 july 1912−−19 september...

Elected F.R.S. 195219 September 1995:−−Thomas Stanley Westoll. 3 July 1912

Colin Patterson and R.A. Fortey

, 531-546, published 1 November 1999451999 Biogr. Mems Fell. R. Soc.

Email alerting serviceheretop right-hand corner of the article or click

Receive free email alerts when new articles cite this article - sign up in the box at the

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

on August 20, 2018http://rsbm.royalsocietypublishing.org/Downloaded from on August 20, 2018http://rsbm.royalsocietypublishing.org/Downloaded from

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

http://rsbm.royalsocietypublishing.org/subscriptions

http://rsbm.royalsocietypublishing.org/


THOMAS STANLEYWESTOLL3 July 1912—19 September 1995

Biog. Mems Fell. R. Soc. Lond. 45, 531–546 (1999)

on August 20, 2018http://rsbm.royalsocietypublishing.org/Downloaded from


THOMAS STANLEYWESTOLL

3 July 1912—19 September 1995

Elected F.R.S. 1952

B C P, F.R.S., R.A. F, F.R.S.

The Natural History Museum, Cromwell Road, London SW7 5BD, UK

Stanley Westoll was a vertebrate palaeontologist and anatomist, and a geologist. He is bestknown for important and innovative work on Palaeozoic fishes and their relationships withtetrapods, but also made substantial contributions in geology, especially on Devonian andCarboniferous stratigraphy in northern Britain.

F

Stanley was born in West Hartlepool, a seaport on the coast of County Durham. WestHartlepool was then almost a ‘new town’, developed around the harbour and dock that werebuilt in 1845–47 to ship the produce of the Yorkshire coalfield. Westolls had been in theSunderland area for at least 200 years, as mariners, shippers, shipowners and shipbuilders. Hisfather, Horace Stanley Westoll (1882–1952), was apprenticed to the sea, and in 1904–13travelled the world successively as Able Seaman, third mate, second mate and first mate infreighters out of Sunderland and West Hartlepool. He was one of the last to earn a Master’sCertificate in both sail and steam, but before he could command a ship he was invalided outof the Merchant Service with lung trouble. He then became assistant dockmaster in the TyneDock at Newcastle upon Tyne before returning to West Hartlepool as dockmaster. Stanley’smother was a schoolteacher who came from a farming family in the Yorkshire Dales; Stanleyhad fond memories of holidays spent there with his maternal grandparents. He was the oldestof three children, with a brother and a sister.

Although he was always fascinated by the sea and by shipping, Stanley recalled later ‘Myinterest in geology dates from my tenth birthday, for which a favourite uncle gave me W.Percival Westell’s splendid little book on rocks and fossils. Within a few weeks, on a familyholiday at Whitby, I had collected a better Dactylioceras than that figured in the book, andbecame a keen collector.’ The implement used for this fieldwork was the domestic

533 © 1999 The Royal Society



534 Biographical Memoirs

coalhammer; the book that Stanley recalled must have been Every boy’s book of geology byA.E. Trueman and W.P Westell, published in 1920 by the Religious Tract Society.

Stanley went to West Hartlepool Grammar School and recalled three teachers as beinginfluential: W.H. Bayliss (mathematics), W.H. Dowland (physics, who came to WestHartlepool with a First from Cambridge) and G.W. Young (chemistry). Stanley was anaccomplished athlete, and had trials for rugby with the county junior team. However, hisschool career was horribly interrupted at the age of 15. While larking with friends on the highboard at the swimming pool, he was pushed off and fell on the edge of the bath. He wasrescued from the bottom of the pool by the lifeguard, who revived him with artificialrespiration. Stanley was never sure whether the smashed pelvis that resulted came fromimpact with the edge of the bath or from the weight of the lifeguard. He spent months intraction in hospital, where, while the rest of him grew with adolescent vigour, the damagedright leg failed to keep pace. So he left hospital with one leg a couple of inches shorter thanthe other. It is hard to imagine the impact of such a tragedy on an extremely intelligent andactive boy. Certainly, Stanley never let his handicap deter him from physical activity, and onthe mental side he used his enforced idleness in hospital to teach himself Egyptianhieroglyphics.

Apart from his teachers, another early influence was Dr C.T. Trechmann (1885–1964)(26)*. Trechmann’s father owned a cement works at Hartlepool, and lived in some style atHudworth Towers, Castle Eden, Co. Durham. After graduating in geology and chemistry atNewcastle, Trechmann returned to work in his father’s business as an analytical chemist. Thefirm was later sold to ICI, and Trechmann became a man of leisure. He was a frequent visitorto Stanley’s school, arriving in the Rolls-Royce to take a few students out for an afternoon’sgeologizing. Stanley’s first publication (1), written as an undergraduate, was a petrologicalappendix to Trechmann’s paper on the geology of St Kitts, in the West Indies, whereTrechmann spent his winters. Trechmann was also interested in local archaeology andintroduced Stanley to worked flints, the subject of his second paper (2). Stanley kept in closetouch with Trechmann for many years.

While still at school, Stanley entered for the Hancock Junior Essay Prize of the NaturalHistory Society of Northumberland, Durham and Newcastle upon Tyne, submitting a pieceon ‘The geology of the district around West Hartlepool’. The Transactions of the Society for1928–29 record that six essays were sent in and that ‘the examiners … were unable torecommend any of these for the full prize, and consequently, by way of encouragement, minorprizes were awarded’. As his prize, Stanley chose Neaverson’s Stratigraphical palaeontology(1928), which remained in his library, with the Society’s Prize bookplate, until his death.

U

In October 1929, aged seventeen years and three months, Stanley entered Armstrong College,Newcastle upon Tyne (the college became King’s College, Durham University, in 1937, andthe University of Newcastle upon Tyne in 1963). He had won an open entrance scholarship,one of only four awarded. He graduated in 1933 with first class honours in geology, withmetallurgy and zoology as subsidiaries, and immediately began work for a Durham PhD on

* Numbers in this form refer to the bibliography at the end of the text.



Thomas Stanley Westoll 535

Late Permian fishes from the Marl Slate of Northumberland and Durham. His thesis, ‘ThePermian palaeoniscid fauna of Northumberland and Durham’, was submitted and the degreeawarded after just two years. Only the abstract was published (5), apparently because Stanleyfelt that the guts of his thesis had been appropriated by Hermann Aldinger, a German who in1933 went from Tübingen to Stockholm to work for a doctorate under E.A. Stensiö (1891–1984, For.Mem.R.S. 1946) at the Swedish Museum of Natural History. Aldinger’s subject wasLate Permian palaeoniscoid fishes from East Greenland, of the same age as Stanley’s MarlSlate fishes, and in December 1934 Stanley sent Aldinger a copy of his thesis. In the publishedwork (Aldinger 1937) Stanley is not mentioned in the acknowledgements, but his thesis iscited extensively in the text (as ‘Westoll 1934’, described in the bibliography as ‘Manuskr.’);Stanley’s reconstructions of the skulls of Palaeoniscus (fig. 25) and Acrolepis (fig. 74) arereproduced; and the new genus Reticulolepis is credited to ‘Westoll 1934’. Because the thesiswas unpublished, the new genus became Aldinger’s under the Code of ZoologicalNomenclature. One can understand Stanley’s displeasure, although in later life he seemed tobear no grudge against Aldinger.

U L

In October 1934, as Dr T.S. Westoll, the 22-year-old Stanley came to London to work underD.M.S. Watson, F.R.S. (1886–1973), Jodrell Professor of Zoology at University College andalready the doyen of British vertebrate palaeontology. Stanley had won a Senior ResearchAward from the Department of Scientific and Industrial Research (DSIR), worth £300 a year.A journal survives of his first year in London (30 September 1934 to 24 November 1935),filled with fascinating detail about his work, travel and entertainment. The first entry includes‘Digs quite comfortable—but will I be able to work at night?’. Stanley divided his timebetween University College and the British Museum (Natural History), where he renewed hisacquaintance with E.I. White (1901–85, F.R.S. 1956), in charge of fossil fishes. On winterSaturdays Stanley went to Twickenham to watch Harlequins play rugby, and on Sundayswalked around London. He soon made the first of many visits to Oxford (‘O! for a Leica!’,says his journal, after comments on the colleges; and ‘beer from silver tankards is much betterthan beer from anything else’, after dinner in hall at Oriel). He stayed with J.A. Moy-Thomas(1908–44), whom he had first met in Newcastle when Moy-Thomas came to study fossil fishesin the Hancock Museum. In Oxford, Stanley met E.S. Goodrich, F.R.S. (1868–1946), then theleading British comparative anatomist (whom Stanley did not like), G.R. de Beer (1899–1972,F.R.S. 1940; ‘a bouncing little man, absolutely bristling with intelligence’), W.J. Arkell (1904–58, F.R.S. 1941, ‘obviously very capable’) and J.Z. Young (1907–97, F.R.S. 1945, ‘a verybrilliant young man’), among others.

In December 1935, Gunnar Säve-Söderbergh (1910–48) came to London from Stockholm,where he was working under Stensiö on the Devonian amphibians (ichthyostegids) that hehad collected in East Greenland in 1931. Stanley and Säve-Söderbergh were invited to stay theweekend with Sir Arthur Smith Woodward, F.R.S. (1864–1944), who had retired in 1924 fromthe Keepership of Geology in the British Museum (Natural History) after a brilliant careeron fossil fishes. Lady Smith Woodward met the two at Haywards Heath station and took themto Hill Place, the Woodwards’ home. After dinner that night the ‘two aspiring youngsters wereallowed to sign the famous tablecloth on which Lady Woodward had embroidered in silk the




signatures of our precursors in the field’ (the tablecloth, with almost 350 signatures, is nowmounted in the Department of Palaeontology of the Natural History Museum). In thefollowing week, Stanley took Säve-Söderbergh to Newcastle and showed him the collectionsin the Hancock Museum.

Travels to Edinburgh, York and Manchester followed, and in the following JanuaryStanley crossed to Paris, to see the collections in the Musée National d’Histoire Naturelle, theÉcole des Mines, and the Sorbonne. There he met Jean Piveteau (1899–1991), Marcellin Boule(1861–1942) and others. In April 1935 Stanley made his first trip to Stockholm, meetingSäve-Söderbergh, Stensiö and his students Aldinger, Eigil Nielsen (1910–68), Erik Jarvik(1907–98) and Gustav Wängsjö. He was tremendously impressed by what he saw in Stensiö’sdepartment, and later (18), in reviewing Nielsen’s thesis, described the excellence of thetechnical support. Aldinger pleased him in particular: ‘I had no idea of his real brilliance; heis the most independently minded of the people now working with Stensiö’. From Stockholmhe went to Oslo, where he met Anatol Heinz (1898–1975) and Leif Støermer (1905–79), andsaw the enormous collections of fossil fishes from Spitsbergen. He returned to Newcastle viaBergen.

Back in London, Stanley began collaboration with Moy-Thomas on a Permian coelacanth(3), worked on Recent and Devonian lungfishes, and in September began collaboration withWilliam Graham-Smith (b. 1911) on the new Devonian lungfish Fleurantia, using specimensthat Graham-Smith had collected in 1934 at Scaumenac Bay (now known as Miguasha),Quebec, Canada (8). In July and August, Stanley made a tour of Scotland, studying museumcollections and visiting the classic Old Red Sandstone fish localities, several of which wereshown to him by Graham-Smith’s parents. For part of this trip he was accompanied by ErikJarvik from Stockholm (travelling by motor cycle), and at Thurso they met the Cambridgezoologists C. Forster-Cooper, F.R.S. (1880–1947) and F.R. Parrington (1905–81, F.R.S. 1962)(‘had a skinful of port and played billiards’, wrote Stanley in his journal). Forster-Cooper(who had been working the quarry for seven summers) took them next day to AchanarrasQuarry, the first of Stanley’s many visits there.

By October, Stanley had worked out ‘the cosmine-resorption story’ and found de Beer andGoodrich ‘very intrigued’ when he told them it in Oxford. This became his paper (4) on thesnout of Osteolepis, a remarkably original work, using specimens gathered that summer in thefield and in the museums of Aberdeen, Cambridge, Edinburgh, Elgin, Inverness, London andYork. His idea was that the shiny cosmine covering the dermal bones in Osteolepis and manyother Devonian crossopterygians is periodically resorbed, so permitting growth of theseparate underlying bones. The idea had profound effects on systematics, because in manycases specimens with and without cosmine had been assigned to different genera or evenfamilies. The paper also includes significant proposals about the homologies of dermal bones,and introduces the term ‘anamestic’ for bones ‘that merely exist as space fillers’; Stanley’sword is now widely used. One of his arguments for periodic resorption of cosmine was theexistence of specimens showing cosmine with concentric grooves, which he took to be ‘thestadia of past resorptions’. His theory was immediately adopted by Forster-Cooper (1937),and these grooves are now called Westoll-lines (following Bystrow (1942)).

Beyond its originality, Stanley’s Osteolepis paper (4) is an early example of anothercharacteristic of his work, frequent reference to forthcoming publications that will expand orilluminate one point or another. As time went by, these promises were fulfilled less frequently,if only because Stanley had too many irons in the fire.




After cosmine resorption, Stanley turned to the homologies of the cheek bones,particularly the squamosal, in the different groups of fishes. The published paper (7) includesa postscript added after a second visit to Stockholm in April 1936 (accompanied byMoy-Thomas and Errol White).

In August 1936 the Geologists’ Association held a field-meeting in Orkney and Shetland,led by H.H. Read (1889–1970, F.R.S. 1939). Stanley acted as secretary to the excursion,having written himself a memo ‘go to Orkneys next summer’ after hearing Sir John Flett,F.R.S. (1869–1947), Director of the Geological Survey and a native of Dingwall, lecture at theGeological Society in November 1934. By the end of 1936 he had completed a substantialpaper on the Devonian fishes of the region (6).

In the spring and summer of 1937, Stanley made his first visit to North America, with theaid of a grant of £100 from the Royal Society and the J.B. Tyrrell Fund (£40) from theGeological Society. He visited all the major museums on the east coast, and travelled west asfar as the Field Museum in Chicago. He met many American palaeontologists who were tobecome friends, notably G.G. Simpson (1902–84, For.Mem.R.S. 1958) in New York, and A.S.Romer (1894–1973, For.Mem.R.S. 1969) at Harvard. He was accompanied on part of themuseum tour by D.M.S. Watson. Early in May, Stanley met W. Graham-Smith at St John,New Brunswick (Graham-Smith had been in North America for a month or so, and hadbought a second-hand car for the trip to Canada). The two began fieldwork in the famousUpper Devonian exposures of Scaumenac Bay (Miguasha), where Graham-Smith hadworked alone in 1934. Graham-Smith expected their work to continue until September, but inJune Stanley ‘dropped a bombshell’ (letter from W.G.-S., January 1998) by announcing thathe must leave to attend the International Geological Congress in Moscow (21–29 July 1937).So the two packed up their fossils and left, stopping only to discover the Devonian vertebratefossil locality at D’Aiguillon, Quebec. When that locality was ‘discovered’ by Canadians in1946 (Russell 1947), the first fish to be described from it was named Cephalaspis westolli

Russell (1954), in recognition of Graham-Smith and Stanley’s priority. There is no record ofStanley’s having attended the Moscow Geological Congress.

The proceeds of Graham-Smith and Westoll’s fieldwork at Miguasha were extraordinarilyvaluable. Among the specimens that they brought home was the holotype of Elpistostege

watsoni, a partial skull roof described by Stanley in 1938 (9) as the ‘missing link’ (‘a perfecttransition’) between fishes and tetrapods. Stanley used Elpistostege to elaborate his theory ofhomologies between the skull roofing bones of tetrapods and rhipidistians. He argued that thenames of the principal roofing bones (frontals, parietals) had ‘largely been misapplied infishes’. Taking the pineal foramen as a marker, he showed that the differences in patternbetween rhipidistians and early amphibians could be seen as due to changes in proportions, sothat the ‘frontal’ of fishes is homologous with the tetrapod parietal, the fish ‘parietal’ is thetetrapod postparietal, and the tetrapod frontal is represented in most rhipidistians by nasals.Stanley was noncommittal about whether his new animal was fish or tetrapod, but otherssoon treated it as the first tetrapod. Schultze (1996) gives a full account of Elpistostege and itssubsequent history. Stanley’s new theory of bone homologies was enthusiastically accepted bywhat A.S. Romer called the ‘Anglo-Saxon school’, but was rejected by Stensiö’s ‘Stockholmschool’. Controversy continued for many years, but most would feel that Stanley’s view is nowgenerally accepted. Apart from Elpistostege, the Graham-Smith and Westoll expedition toScaumenac Bay furnished material later used by several of Stanley’s PhD students.




U A

In 1937, when his three-year DSIR award expired, Stanley won an 1851 Exhibition ResearchFellowship, but relinquished it almost immediately when appointed to a Lectureship inGeology at the University of Aberdeen. In 1937 its Department of Geology and Mineralogyconsisted of a newly appointed Professor, T.C. Phemister (1902–82), Stanley and one assistant(Scott Simpson, 1915–81, later Professor of Geology at Exeter). Among the undergraduatesStanley found F.H. (later Sir Frederick) Stewart (F.R.S. 1964), who went on to becomeProfessor of Geology at Edinburgh and a lifelong friend. Stanley also found a department ‘inneed of several spring-cleans; they had a fire in 1906 & things are just as they were left, withscorched and scattered labels’ (letter to E.I. White, October 1937). He was soon teaching 26hours a week and wrote ‘there is enough work here for a galley-slave, & I’m the one’.

At the outbreak of war in 1939, Stanley tried to enlist in the army. He passed the medicaland was not found out until it came to drawing uniform, when his request for a special bootdrew attention to his disability. He joined ARP (Air Raid Precautions) instead. TomPhemister did join up, leaving Stanley as acting head of department, and from 1942 to 1945he also had to stand in for the Professor of Metallurgy. Stanley served as geological adviser inconstruction of the anti-submarine barriers at Scapa Flow (after the battleship Royal Oak

was torpedoed there in October 1939), a job that involved precarious flights in military lightaircraft between Aberdeen and the Orkneys. In 1943, Stanley wrote to A.S. Romer that he was‘in anti-chemical warfare’. This involved training people in anti-gas precautions. Over-exposure to noxious gases in these exercises gave him sinus problems that requiredintervention in 1945–47 and persisted into old age.

Although the early war years must have been extraordinarily busy, they did not impedeStanley’s ambitious scientific programme. He published on coelacanths (10) (a study based onmaterial gathered during his 1937 tour of North America), rhipidistians (11) and on moregeneral problems such as early amphibian phylogeny (14) and the relation between dermalbones and sensory canals (13). In 1939 he collaborated with Rex Parrington, in Cambridge,on the bones of the palate in reptiles and mammals (12), reconciling the conflict betweenembryology and palaeontology on the homologies of the vomer, parasphenoid andpterygoids. At this time, Stanley kept a note of the dates of submission, proofs andpublication of his papers: the speed with which Nature, in war-torn London, processed papers(13, 14, 20) now seems amazing: paper (13) was completed on 16 July 1941 and published on 9August; paper (14) was completed on 19 May 1942 and published on 13 June.

Stanley’s first major synthetic work came out in 1943 (16), on tetrapod origins, inBiological Reviews, and is still frequently cited (he wrote that ‘it was cut tremendously afteracceptance owing to paper-shortage’). In it he gives a thorough review of the problem ofskull-bone homologies, disagreeing extensively with Säve-Söderbergh. While working on thatpaper, Stanley made the mistake of studying a draft of fig. 3 on the London Underground.Only with difficulty did he avoid arrest for espionage after a fellow-passenger called the police.The drawings are remarkably like plans of engines of war.

Stanley’s first monograph followed in 1944 (19), on the Late Carboniferous haplolepidfishes. Most of the specimens on which it is based were from American collections, borrowedby Stanley during his 1937 tour. He used this work as a thesis for an Aberdeen DSc, awardedin July 1941. The work was published (after a long delay) by the American Museum ofNatural History, where on receiving the manuscript, W.K. Gregory submitted it to the New




York Academy of Sciences as a candidate for the 1942 A. Cressy Morrison Prize (two prizesper year, each of $200, for ‘the most acceptable papers in any field of natural science’).Stanley did not win but had to make do with an Honorable Mention and publication of hisabstract (15).

In 1943, Stanley had turned his attention to one of the more remarkable events invertebrate history—the development of the mammalian ear ossicles and the transformation ofthe fish hyomandibular—which is used to support the jaw—into the stapes, a middle-ear boneused to transmit sound. He was prompted to do this by his own collecting of specimens ofEusthenopteron, the putative tetrapod ancestor from the Upper Devonian of Quebec, whichdisplayed the hyomandibular. He recognized various projections and processes on thehyomandibular that could be homologized with processes on the tetrapod stapes. He alsosuggested by means of the reconstruction of hypothetical intermediate stages that, despite thevery different appearances in frogs, lizards and mammals, the typanum (eardrum) and thetypanic recess (middle-ear cavity) were homologous throughout the tetrapods and not derivedindependently (Gaup 1913). Stanley argued that variation among living taxa is due todifferent orientation of the stapes and to the development of ‘outpushings’ of the originaltympanic recess cavity.

In December 1939, Stanley married Dorothy Wood, daughter of a family of Aberdoniantrawler owners, whom he had met shortly after arriving in Aberdeen. Their son Neil, born in1941, went on to complete a first degree and a doctorate in geology at Edinburgh and is nowin mineral exploitation in Ontario, Canada.

On Christmas Eve 1946 Stanley crossed the Atlantic, by plane, to attend two meetings, thesecond of which would make his name as an evolutionist. The first meeting, in Chicago, wasthe sixth annual meeting of the Society of Vertebrate Paleontology (SVP), to which Stanleywas elected in 1943. The second, at Princeton on 2–4 January 1947, was the Conference onGenetics, Paleontology and Evolution. This Princeton conference was the result of severalyears’ planning, detailed in the foreword to the published volume (Jepsen et al. 1949). It isnow seen as the proof or crowning achievement of the ‘modern synthesis’ of evolutionarybiology that occurred during the preceding decade (see, for example, Mayr (1980)). Therewere 23 participants in the published conference proceedings, all except four from the USA.The other four were from Britain: E.B. Ford, F.R.S. (1901–88), J.B.S. Haldane, F.R.S. (1892–1963), whom Stanley knew from University College London in the 1930s, David Lack (1910–73, F.R.S. 1951), and Stanley. The topic that Stanley chose for his contribution was lungfishevolution. His published paper (21), the longest in the book, after a full descriptive section,quantified evolutionary changes in about 100 characters of the lungfish skull and postcranialskeleton, plotted (fig. 11) against time as ‘rate of loss of characters of the ancestral type’. Theresulting curves, one for the skull, one for the body, and one for the total, show very rapidchange in the Devonian and Early Carboniferous, followed by much slower change from theLate Carboniferous to Recent. Stanley used this example in a commentary on some ofSimpson’s conclusions in his pioneering Tempo and mode in evolution (Simpson 1944), a bookthat Stanley did not see until after the Princeton conference. His paper is remarkably original,and is still frequently cited. In Simpson’s next book (Simpson 1953, p. 22) he wrote, ‘Westoll’sexample is so instructive and he has based on it so important a discussion … that his data aresummarized here’.

Stanley’s presentation at the Princeton conference must have impressed both his Americanand British colleagues greatly. Early in 1949 he was elected a Corresponding Member of the




American Museum of Natural History (where Simpson was), a high honour that was renewedevery five years until his death. In 1948 he was proposed for Fellowship of the Royal Societyby D.M.S. Watson, seconded by Sir Edward Bailey, F.R.S. (1881–1965, then Director of theGeological Survey) and supported by E.B. Ford and J.B.S. Haldane, among others.

One bizarre episode was involved in Stanley’s lungfish paper (22). In November 1946, amonth before Stanley was to leave for Princeton, Dr W.M. Lehmann of the University ofBonn sent to Errol White photographs and X-rays of a lungfish skull that he had discoveredin the Hunsrückschiefer, an Early Devonian slate famous for superb preservation of fossils.White, knowing that Stanley was working on lungfishes and that no other Early Devonianlungfish had been reported, immediately sent the material to Stanley. Within a couple of daysStanley made labelled tracings from the photographs, identified the animal as Dipnorhynchus,and wrote White a long letter on his interpretation. When Stanley’s Princeton paper appearedin 1949 it included drawings and a description of Lehmann’s specimen (as Dipnorhynchus

lehmanni), with references to a manuscript cited as ‘Lehmann and Westoll, 1949’ andacknowledgement of Lehmann’s ‘great generosity in permitting the writer to use excellentphotographs of the fossil’. ‘Lehmann and Westoll’ appeared in 1952 (23), with only thesection on ‘Discovery and provenance’ credited to Lehmann, quoted from a 1947 letter fromhim to Stanley. Remarkably, Stanley had never seen the specimen, only the photos and X-rayssent to him by White, and plaster casts that Stanley had obtained from Krantz, the dealer inBonn. Even more remarkably, four years later, Lehmann (1956) published his own descriptionof the fossil. In an introduction that hardly conceals his irritation, he ran through the eventsdescribed above, save the last. Lehmann was unaware of the 1952 Royal Society publication ofwhich he is first author.

In April 1948 Stanley travelled to Paris to take part in a colloquium with much the sameaim as the Princeton conference: to bring together a select group of geneticists and palaeon-tologists to discuss the evolutionary process. Stanley was by far the youngest of the invitedparticipants, who included Haldane, Simpson, Stensiö, C.H. Waddington, F.R.S. (1905–72)and Watson. Stanley did not present a paper, but contributed to the published discussion.

In August 1948 the 18th International Geological Congress (postponed from 1940) washeld in London. After the formal congress, D.M.S. Watson, Stanley and Errol White led atwo-week excursion (2–26 September) to Palaeozoic vertebrate fossil sites. Stanley wasresponsible for the Scottish half of this trip; planning it had occupied much of his time overthe preceding year. The excursion was a great success, acknowledged by the dozenparticipants (Gregory & Colbert 1949). Immediately after the excursion, Stanley packed up inAberdeen and moved south.

U N T (I)

In October 1948, Stanley took up his appointment as J.B. Simpson Professor of Geology atNewcastle, succeeding H.G.A. Hickling, F.R.S. (1883–1954), who had taught Stanley as anundergraduate and supervised his PhD. The department that he took over was staffed by G.S.Mockler, A. Raistrick and S.I. Tomkieff (appointed Reader in 1948), all of whom had beenthere since the 1920s and had taught him in the 1930s. He found a ‘cramped andunderequipped’ department, struggling to cope with the postwar influx of students. Ingeology, student numbers had increased from 90 to over 400. Stanley soon appointed two new




lecturers and began working towards acquiring extra space, if not a new building. At the endof his second autumn term (December 1949) he wrote, ‘I’ve tried ten weeks of 9 a.m.–2 a.m.,and merely keep stationary’ (letter to E.H. Colbert). In 1950 he took on the first of a longseries of PhD students; by 1952 there were five such students.

Stanley’s reputation as an evolutionist at this time is exemplified by his correspondence in1950–51 with Julian Huxley, F.R.S. (1887–1975), discussed by Swetlitz (1995). Huxley hadcome to believe that large-scale evolution (beyond the species level) had come to a stop, andsought empirical support for this view from those palaeontologists that he thought best ableto supply it, G.G. Simpson and Stanley.

When Stanley moved from Aberdeen to Newcastle, his wife and son did not go with him,and he and Dorothy were divorced in 1951. In July 1952 he married Barbara SwansonMacadie. They had first met, auspiciously, in Achanarras Quarry in the summer of 1948,when Stanley and two students were preparing it for the excursionists from the InternationalGeological Congress. Barbara, who had just graduated in geology from Edinburgh, hadcycled to Achanarras from her home in northern Caithness in search of fossil fishes for theEdinburgh department. Before the marriage she worked for four years in the GeologicalSurvey in London, assisting James (later Sir James) Stubblefield, F.R.S. (b. 1901), then ChiefPalaeontologist. In September 1952 the newly married couple sailed to New York on theQueen Elizabeth and travelled north to Harvard, where Stanley was to spend a semester asAgassiz Visiting Professor of Vertebrate Paleontology at the Museum of ComparativeZoology (MCZ). These visiting appointments at Harvard were often followed by offers ofjobs. So it was with Stanley, and in the SVP News Bulletin of June 1953 Al Romer, Director ofMCZ, announced that Stanley was appointed Alexander Agassiz Research Professor and wasexpected to arrive soon. The same source reported in February 1954 that Stanley’s arrival waspostponed until July, and in June 1954 that he was to remain in England. The Westolls’decision not to emigrate was taken in the Cunard building in Liverpool, after they had visitedthe US consul to acquire visas. The consul asked Stanley whether he knew any communists(this was the time of McCarthyism). To the consul’s obvious distress, Stanley replied that hedid. ‘How?’ was the next question; Stanley answered ‘in the same way as your President’ (i.e.in the course of duty). This exchange swung the balance in Stanley’s and Barbara’s decision toremain at Newcastle. Al Romer bore no grudge, and was thereafter a frequent visitor toNewcastle; in 1955 the Westolls took the Romers round Scotland for two weeks.

In 1958, after a long and painful gestation, a Festschrift for D.M.S. Watson was publishedunder Stanley’s editorship (some of the included papers were ‘in press’ for four years, andStanley mislaid the typescript of at least one) (24). Stanley’s own contribution was anembellishment of a nineteenth-century idea, most elegantly stated by Goodrich (1906), thatthe pectoral and pelvic fins of jawed vertebrates were modifications of an original paired andmuscled fin fold that ran continuously down each side of the trunk. Through his observationson Silurian and Devonian cephalaspid, anaspid, acanthodian and placoderm fishes, hesuggested that, although the basic idea was correct, the pectoral fins were so differentlydeveloped within each of these groups that they must have arisen independently from arudimentary paired ancestral fin fold, neither supported by endoskeleton nor equipped withmuscle. His style of argument is typical of much of his work in following sequences ofdevelopment of pectoral fins within each of these groups of fishes upwards through thegeological succession to show that the early forms had poorly developed fins, whereas thelater ones bore increasingly specialized fins.




As part of the worldwide celebration of the centenary of Darwin’s On the origin of species,in 1959 Stanley was invited to Melbourne to take part in a symposium. He chose to make thisa round-the-world trip (he boasted that it took 79 days, one less than that of Phileas Fogg andPassepartout). He went to Australia via Harvard, San Francisco, Hawaii, Fiji and NewZealand, and returned via Indonesia. In San Francisco, Sam Welles, vertebratepalaeontologist at Berkeley, took Stanley to meet one of his heroes, C.S. Forester, whoinscribed a Hornblower novel for him. In Hawaii he was lucky enough to see an eruption ofKilawea.

A

In August 1966 the Royal Society became aware of plans by the Ministry of Defence (MoD)and the BBC to ‘develop’ Aldabra, the atoll north of Madagascar, then part of British IndianOcean Territory. Anxious to avoid the destruction of a unique habitat that would inevitablyfollow establishment of the proposed staging-post, Council appointed Dr D.R. Stoddart, ofthe Department of Geography at Cambridge, to accompany an MoD/BBC expedition toAldabra in September 1966. After Stoddart’s report, the Society mounted a scientificexpedition to Aldabra from August 1967, and in December 1967 set up the Aldabra ResearchCommittee ‘to co-ordinate and extend the existing proposals for research work andconservation there’. Stanley was appointed Chairman of that committee, a job that was tolast ten years and give him a good deal of work. One of his reasons for accepting was the factthat Aldabra is the nearest neighbour of the Comores, home of Latimeria, and he hoped(wrongly) that Britain might acquire its own source of coelacanths. During the next ten yearsStanley did not miss a single meeting of the committee or its offspring, the numeroussub-committees and working groups to which he was appointed.

The Aldabra Research Committee met first in February 1968, and in September 1968Stanley (with David Stoddart and Dr M.E.D. Poore, Director of the Nature Conservancy)visited Aldabra and the Seychelles aboard Manihine, research vessel of the East AfricanMarine Fisheries Organization. A converted Hull trawler, Manihine had no reputation forcomfort; Stoddart recalls Stanley ‘with his gammy leg heroically struggling below decks with aglass carboy of formaldehyde that had broken loose in a storm, while the rest of us were tiedto our bunks’ (letter, February 1996). The MoD plans for a staging-post on Aldabra weresoon dropped because of defence cuts. However, the Royal Society’s engagement thereflourished. In 1969–71 a research station was built to house a permanent staff and visitingscientists, and in 1973 the Society acquired the lease of the island (the previous lessee at firstasked £100 000 for it, but eventually settled for £7500 and 50 tortoises per year). Stanley andStoddart organized a Discussion Meeting at the Society in March 1969 on the results of thefirst expedition (30), and a second meeting in March 1977 on longer-term results (32).

In the mid-1970s the future of the Royal Society Research Station on Aldabra becameprecarious. The Seychelles were to become independent in June 1976, and would havesovereignty over Aldabra. In 1974–75 funding difficulties necessitated cuts in permanent staffon the station, which together with irregularities in supply (by ship) led to problems withmorale. In March and April 1976, Stanley, together with Stoddart and D.J.H. Griffin of theSociety’s staff, flew to the Seychelles. They did not get to Aldabra, but had discussions withministers and officials in Mahé. Meanwhile Council set up a group to review the future of thestation; it proposed that research should terminate in March 1980. With estimated




expenditure of £80 000 for 1977, the station was a heavy drain on the Society’s grant-in-aid,and Council agreed to closure. The President of the new Republic of Seychelles attended the1977 Discussion Meeting at the Society, and after it Stanley asked to retire from the AldabraResearch Committee. He did so in June 1977, and was replaced by Richard (later Sir Richard)Southwood, F.R.S. At Stanley’s last meeting the Executive Secretary said ‘Officers were awarethat it was largely thanks to Professor Westoll’s leadership of the Committee that the researchprogramme had been so successful’. By March 1979 the station held only a skeleton staff andno visiting scientists, but in 1977–78 David Stoddart had made several visits to the Seychellesand succeeded in setting up the Seychelles Island Foundation, which took over the station inMarch 1980, when the Aldabra Research Committee was dissolved.

During its existence, the Royal Society Research Station housed over 100 staff and visitingscientists (listed in Stoddart (1978)), and Aldabra became ‘among the best known oceanicislands and coral atolls on Earth’ (Stoddart 1978). The station still exists, having been rebuiltin 1997–78 with a grant from the World Bank. David Stoddart (letter, February 1998) reportsthat the Aldabra bibliography now contains over 1200 items.

U N T (II)

The ten years that Stanley gave to Aldabra were also among the busiest of his tenure atNewcastle. With the general expansion of British universities (mainly in the 1960s), theacademic staff of Stanley’s department had grown from the three whom he found in 1948 totwelve (including two professors) in the mid-1960s and to sixteen (with three professors) at hisretirement in 1977. Much of his time during this period was inevitably taken up byadministrative duties, and the palaeontological and anatomical studies on which he had madehis name were carried forward by his students during this time (25, 27–29, 33). However, hefound opportunities to contribute to a number of compendia of Upper Palaeozoicstratigraphy, of which his contribution to the Correlation Chart of Devonian rocks of theBritish Isles (31) summarized some of his previous work. Stanley had made periodic, short,published contributions to stratigraphy since 1939, and in his later years he published moreextensively on Devonian and Carboniferous stratigraphy and sedimentology. Discussions onthe boundary between the Silurian and Devonian Systems were influenced by the views ofJack Shirley in the Newcastle department, and favoured a horizon that was not, ultimately,selected. However, the debate that preceded the fixing of the boundary at a rock section in theCzech Republic remains a type example of modern stratigraphic procedure. Stanley made adeliberate effort to make Newcastle a centre of Carboniferous (and coal) studies, appropriateto its location, notably by the appointment of Duncan Murchison, with whom he edited avolume on coal-bearing strata published in 1968. Murchison it was who wrote the firstobituaries after Stanley’s death in 1995.

P

Stanley Westoll was a clubbable and gregarious man. He was a born raconteur, with aseemingly bottomless fund of anecdotes and stories. He was musical, although he played noinstrument. In London in 1935 he joined the English Madrigal Choir, and complained in hisjournal of the conductor ‘using tonic sol-fa for teaching modal music’. That journal also




includes knowledgeable comment and criticism on concerts he attended. In the 1960s and1970s the ‘top half’ of the Aeolian Quartet were regular house guests of the Westolls whenthey came annually to Newcastle for a week of chamber music performances. He championedthe independence of palaeontology from the geological sciences. When the PaleontologicalAssociation was founded in 1957—as a breakaway from the Geological Society of London—Stanley Westoll was the first member of the new group, a fact of which he remained proud.

I

Stanley was an excellent and willing teacher and lecturer. He usually spoke without notes,apparently extempore, with fluent authority. Throughout his time at Newcastle he taught allthe first-year introductory geology except crystallography. Murchison (1996) wrote ‘Hisphenomenal memory enabled him to cover wide blackboards with world stratigraphicsequences in immaculate handwriting to the intense frustration of students who had no sheetsof paper sufficiently wide or long… nor had they the speed of writing to reproduce thesequences before they were removed and the next set begun’. Roger Miles, a PhD student withStanley in the early 1960s, attended Stanley’s third-year lectures on vertebrate palaeontologyand recalls his astonishment when each Friday afternoon Stanley talked without notes for twohours, covering the board with accurate drawings, from memory, of specimens of Cephalaspis,or a series of fish skulls, and so on.

Stanley took on over twenty doctoral students, the majority of them working onstratigraphic problems in northern Britain. He felt a special responsibility for his students invertebrate palaeontology, of whom there were: A.D. Walker, on Triassic reptiles from Elgin(PhD 1957), who stayed on as a lecturer in Stanley’s department; R.S. Miles, on theplacoderm Coccosteus (PhD 1963); S. Mahala Andrews, on the axial skeleton of rhipidistians(PhD 1968); Peter Zaborski, on the ‘protocoelacanth’ from Miguasha (PhD 1977); Paul Roseon the cervical region of fishes and tetrapods (PhD 1980); D.M. Pearson, on the Devonianactinopterygian Cheirolepis (PhD 1977); Susan Turner, on Palaeozoic microvertebrates (PhD1984). Several of these worked mainly on specimens in Stanley’s collection, and he felt entitledto put his name on the resulting publications.

The lasting influence of Stanley’s research comes mainly from his early work, in the 1930sand 1940s, on cosmine (4), skull-roofing bone homologies (9), tetrapod origins (16–18), andlungfish evolution (21). All this is still commonly cited, as it deserves to be, for it was brilliant,original, and communicated in lucid, economical prose. Once Stanley had his chair and hadbeen elected F.R.S. (both before the age of forty), his subsequent work was not, perhaps, asprolific or original, and his influence was mostly through the direction of the Department ofGeology at the University of Newcastle upon Tyne. For forty years, Stanley Westoll was oneof the well-known figures in vertebrate palaeontology, and his memory survives in the workof several of the leading specialists who received their training at this hands.

A

Colin Patterson died before this memoir was entirely complete [a memoir of Colin Patterson is included in thisvolume]. In his notes he gratefully acknowledged the receipt of information from W. Graham-Smith, J.G.




Maisey, Alick Walker, Alec Panchen, Roger Miles, Peter Zaborski, Malcolm Stoddart, Duncan Murchison andBarbara Westoll. Richard Fortey would like to acknowledge the help of Peter L. Forey, Brian Gardiner and L.Robin Cocks in completing the work for publication.

The frontispiece photograph was taken in 1952 by the Godfrey Argent Studio, and is reproduced withpermission.

R

Aldinger, H. 1937 Permische Ganoidfische aus Ostgrînland.Meddr Groenland 102 (3), 1–392.Bystrow, A.P. 1942 Deckknochen und Zähne der Osteolepis und Dipterus. Acta Zool. 23, 263–289.Forster-Cooper, C. 1937 The Middle Devonian fish fauna of Achanarras. Trans. R. Soc. Edinb. 59, 223–239.Gaup, E.S. 1913 Die Reichertsche Theorie. Archiv Anat. Physiol. Suppl. 1912, 1–146.Goodrich, E.S. 1906 Notes on the development, structure and origin of the median and paired fins of fish.

Q. J. Microsc. Sci (N.S.) 50, 333–376.Gregory, J.T. & Colbert, E.H. 1949 The C-16 (Vertebrate Paleontology) excursion of the XVIII International

Geological Congress, 1948. News Bull. Soc. Vertebr. Paleont. 25, 6–9.Jepsen, G.L., Simpson, G.G. & Mayr, E. (eds) 1949 Genetics, paleontology and evolution. Princeton University

Press.Lehmann, W.M. 1956 Dipnorhynchus lehmanni Westoll, ein primitiver Lungenfisch aur dem rheinischen

Unterdevon. Paläont. Z. 30, 21–25.Mayr, E. 1980 Prologue: some thoughts on the history of the evolutionary synthesis. In The evolutionary

synthesis (ed. E. Mayr & W.B. Provine), pp. 1–48. Cambridge, MA: Harvard University Press.Murchison, D. 1996 Thomas Stanley Westoll (1912–1995). In Annual Report, The Geological Society of

London, 1996, pp. 22–23. London: Geological Society.Russell, L.S. 1947 A new locality for fossil fishes and eurypterids in the Middle Devonian of Gaspé, Quebec.

Contr. R. Ont. Mus. Palaeont. 12, 1–6.Russell, L.S. 1954 A new species of Cephalaspis from the Devonian Gaspé Sandstone at D’Aiguillon.

Naturaliste Can. 81, 245–254.Schultze, H.-P. 1996 The elpistostegid fish Elpistostege, the closest the Miguasha fauna comes to a tetrapod. In

Devonian fishes and plants of Miguasha, Quebec (ed. H.-P. Schultze & R. Cloutier), pp. 316–337.Munich: Verlag Dr Friedrich Pfeil.

Simpson, G.G. 1944 Tempo and mode in evolution. New York: Columbia University Press.Simpson, G.G. 1953 The major features of evolution. New York: Columbia University Press.Stoddart, D.R. 1978 Aldabra and the Aldabra Research Station. Phil. Trans. R. Soc. Lond. B 286, 3–10.Swetlitz, M. 1995 Julian Huxley and the end of evolution. J. Hist. Biol. 28, 181–217.

B

The following publications are those referred to directly in the text. A full bibliographyappears on the accompanying microfiche, numbered as in the second column. A photocopy isavailable from the Royal Society Library at cost.

(1) (1) 1932 Description of rock specimens from Brimstone Hill and three other localities in St. KittsB.W.I. Geol. Mag. 69, 259–264.

(2) (2) 1933 (With A. Raistrick) A prehistoric site on the South Durham coast. Vasculum 19, 139–144.

(3) (3) 1935 (With J.A. Moy-Thomas) On the Permian coelacanth, Coelacanthus granulatus, Ag.Geol. Mag. 72, 446–457.

(4) (5) 1936 On the structures of the dermal ethmoid shield of Osteolepis. Geol. Mag. 73, 157–171.(5) (6) The Permian palaeoniscid fauna of Northumberland and Durham. Abstr. Thes. Doct.

Univ. Durham 1935–1936, pp. 14–15.




(6) (7) 1937 The Old Red Sandstone fishes of the north of Scotland, particularly of Orkney andShetland. Proc. Geol. Ass. 48, 13–45.

(7) (8) On the cheek-bones in teleostome fishes. J. Anat. 71, 362–382.(8) (10) (With W. Graham-Smith) On a new long-headed dipnoan fish from the Upper Devonian

of Scaumenac Bay, P.Q., Canada. Trans. R. Soc. Edinb. 59, 241–266.(9) (12) 1938 Ancestry of the tetrapods. Nature 141, 127–128.(10) (17) 1939 On Spermatodus pustulosus Cope, a coelacanth from the ‘Permian’ of Texas. Am. Mus.

Novit. no. 1017, 1–23.(11) (22) 1940 New Scottish material of Eusthenopteron. Geol. Mag. 77, 65–73.(12) (23) (With F.R. Parrington) On the evolution of the mammalian palate. Phil. Trans. R. Soc.

Lond. B 230, 305–355.(13) (25) 1941 Latero-sensory canals and dermal bones. Nature 148, 168.(14) (28) 1942 Ancestry of captorhinomorph reptiles.Nature 149, 667–668.(15) (31) 1943 The Haplolepidae, a new family of late Carboniferous bony fishes [Abstract]. Trans.

N.Y. Acad. Sci. (2) 5, 60–62.(16) (32) The origin of the tetrapods. Biol. Rev. 18, 78–98.(17) (34) The origin of the primitive tetrapod limb. Proc. R. Soc. Lond. B 131, 373–393.(18) (35) The hyomandibular of Eusthenopteron and the tetrapod middle ear. Proc. R. Soc. Lond.

B 131, 393–414.(19) (36) 1944 The Haplolepidae, a new family of Late Carboniferous bony fishes: a study in taxonomy

and evolution. Bull. Am. Mus. Nat. Hist. 83, 1–122.(20) (42) 1946 Triassic fishes from East Greenland [Book review]. Nature 158, 75–76. 20/7.(21) (46) 1949 On the evolution of the Dipnoi. In Genetics, paleontology and evolution (ed. G.L. Jepsen,

G.G. Simpson & E. Mayr), pp. 121–184. Princeton University Press.(22) (48) Evolutionary trends in the Dipnoi. Int. Congr. Zool. 13, 513–516.(23) (53) 1952 (With W. Lehmann) A primitive dipnoan fish from the Lower Devonian of Germany.

Proc. R. Soc. Lond. B 140, 403–421.(24) (57) 1958 (Ed.) Studies on fossil vertebrates. London: Athlone Press.(25) (67) 1963 (With R.S. Miles) Two new genera of coccosteid Arthrodira from the Middle Old Red

Sandstone of Scotland, and their stratigraphical distribution. Trans. R. Soc. Edinb. 65,179–210.

(26) (72) 1965 (With C.A. Fleming) Charles Taylor Trechmann [Obituary]. Proc. Geol. Soc. Lond. no.1628, 207–208.

(27) (78) 1968 (With R.S. Miles) The placoderm fish Coccosteus cuspidatus Miller ex Agassiz from theMiddle Old Red Sandstone of Scotland. Part I. Descriptive morphology. Trans. R. Soc.Edinb. 67, 373–476.

(28) (79) 1970 (With S.M. Andrews) The postcranial skeleton of Eusthenopteron foordi Whiteaves.Trans. R. Soc. Edinb. 68, 207–329.

(29) (80) (With S.M. Andrews) The postcranial skeleton of rhipidistian fishes excludingEusthenopteron. Trans. R. Soc. Edinb. 68, 391–489.

(30) (81) 1971 (Ed., with D.R. Stoddart) A Discussion on the results of the Royal Society expedition toAldabra 1967–68. Phil. Trans. R. Soc. Lond. B 280, 1–654.

(31) (94) 1977 Northern Britain. In A correlation of the Devonian rocks in the British Isles (GeologicalSociety Special Report no. 8) (ed. M.R. House), pp. 66–93. Edinburgh: ScottishAcademic Press.

(32) (95) 1978 (Ed., with D.R. Stoddart) The terrestrial ecology of Aldabra. Phil. Trans. R. Soc. Lond.B 286, 1–263.

(33) (97) 1979 (With D.M. Pearson) The Devonian actinopterygian Cheirolepis Agassiz. Trans. R. Soc.Edinb. 70, 337–399.



thomas stanley westoll. 3 july 1912−−19 september...

Documents