Welcome to the Worldwide Greathead family my One-Name Study - Dr David J Greathead
Dr David John Greathead # 1718 -
Author of "A Passage to the Cape of Good Hope"

David Greathead  1931–2006 

a Life in Biological Control  


The following article appeared in Biocontrol News and Information 28(1), 1N–28N pestscience.com http://www.pestscience.com/news.asp

Rebecca Murphy & Matthew J. W. Cock, CABI. the authors have kindly given me permission to include the article on my website

Dr David J Greathead

The founding editor of BNI, David Greathead, who died on 13 October 2006 at the age of 74, was an influential figure in biological control and also a world authority on Bombyliidae (bee flies). His career reflected many of the changes and developments in biological control over this period, which is hardly surprising since he was central to many of them. A naturally thoughtful demeanour coupled with an encyclopaedic knowledge of biological control endowed him with wisdom and foresight. His legacy includes successful and in some cases groundbreaking biological control initiatives, extensive publications in biological control and taxonomy, contributions to an international regulatory framework for biological 
control, and last but by no means least the many scientists whose early careers he fostered and some of whom are now well known names themselves.  His achievements were a direct result of his rare combination of broad perspective and attention to detail. As fellow-bombyliid expert Neal Evenhuis ( Bishop Museum , Hawaii ) says: “Everything was thoughtfully prepared and checked and rechecked before he would be satisfied,” but as Sean Murphy (a long-serving CABI scientist) says, “David always managed to see the bigger picture.” He inspired loyalty and affection in his staff, was excellent company and could spin a great story – his experiences gave him plenty of material to work with.
David was born on 12 December 1931 in London , UK , into a family with South African roots: his great great-grandfather led one of the first parties to be settled by the British Government in the Eastern Cape in 1820, and his great-grandfather was a member of the first South African colonial parliament in the late 1850s. David’s childhood was divided between the UK and South Africa , but he elected to stay in the UK for his university education when his family returned permanently to South Africa . He graduated from the University of London ’s Imperial College of Science and Technology (now Imperial College London) with a BSc in Zoology in June 1953, and was later awarded a PhD and a DSc.
In 1953 he was recruited by Dr (later Sir) Boris Uvarov to work at the Desert Locust Survey (DLS) and, in the words of Cliff Ashall (Officer-in-Charge of one of DLS's field research units) was “introduced to a varied and resourceful set of characters – men of a different breed.” While working at the All-Russian Plant Protection Institute in St Petersburg , Uvarov had famously discovered that solitary and gregarious desert locusts (Schistocerca gregaria) were different phases of the same species. Having left post-revolutionary Russia , he was recruited in 1920 to the staff of the Imperial Bureau of Entomology in London andwas subsequently directly involved in establishing the Anti-Locust Research Centre (ALRC). Uvarov, according to Elspeth Huxley in No Easy Way (ca. 1957), not only possessed single-minded drive and deep knowledge of locusts but, crucially, saw that countries would have to work together to solve the locust problem, and strove for this to happen. Over the next 10 years, he and another Russian émigré scientist, Zena Waloff, comprised the entire headquarters staff of the Commonwealth anti-locust effort (and ran this operation at a total cost of slightly over UK £7900). During WWII he advised the highly successful, largely military-based Middle Eastern Anti-Locust Unit that implemented desert locust campaigns in collaboration with the civilian East African Anti-Locust Directorate to protect vital food crops in eastern Africa . Post-war, Desert Locust Survey (DLS) and Desert Locust Control (DLC) took over this role. DLS, which David joined, is described by Cliff Ashall as “one of the more remarkable organisations ever to have operated in East Africa and the Middle East .”
Cliff tells how Uvarov chose David “to spearhead the research on locust natural enemies as part of the multi-disciplinary approach to a solution of the locust problem”, and this included work contributing to his PhD. He subsequently published a review of the natural enemies of Acridoidea1 (something he came back to much later in his career) while interest fostered by studies on bombyliid predators on egg pods of the desert locust developed into long-term involvement in biosystematic research on this group.  He also studied the effects of biotic factors on desert locust populations and looked at numerical changes in desert locust populations; a paper he wrote with Bill Stower2 remains one of the few published numerical population studies, and as such is proving important for locust control today.
For the next 8 years David worked at DLS in what Cliff refers to as “that great adventure which was locust research and control.” During this time he was involved in field work and research in Ethiopia , Somalia , Kenya and what was then the Aden Protectorate (now part of Yemen ). It was not a life for the faint hearted. David described the laborious work on choice of oviposition sites by locust swarms, studies he and Bill undertook with George Popov (another notable Russian émigré locust scientist, and traveller) in Somalia in 1953, continuing in Turkana in northern Kenya in 1954: “We used to delimit groups of egg-pods, scrape the surface and mark them all with matchsticks and then carefully excavate them and plot their position and condition on graph paper.” In later years, he expressed regret that the young scientists then involved in locust work had at best a few weeks at a time in the field. From tales he recounted, it is not hard to see why. He and George made a systematic survey of the locust recession areas, beginning with a journey in January–February 1954 from Cape Guardafui , the apex of the Horn of Africa, along the Gulf of Aden coast into Eritrea .  The journey was mostly not on tracks and probably not repeated since. They reached Lake Assal , in what was then French Somaliland (now Djibouti ).  This, the lowest point in Africa at 155 metres below sea level and the most saline body of water in the world, is set in a glistening white salt flat which they had to cross. They found they had to keep up a high speed because the salt crust, below which laythick sludge, began to break up if they slowed down.  Worse was to follow. Near the port of Assab in Eritrea (at the time federated with Ethiopia ) and close to the French Somaliland border they stopped to use the radio thinking they were out of sight. But when they set off again they were surrounded by the Ethiopian garrison and put under house arrest until their bona fides could be confirmed from Addis Ababa ; the soldiers thought they were the French army come to seize Assab. Moreover, they learnt afterwards, had the soldiers not been holding their topees on their heads with one hand as they came towards them at the double, someone might have been shot, for the soldiers had been told to shoot. Little surprise, then, that David later showed scant regard for, by comparison, minor privations experienced by his staff – and incredulity at the luxuries some biocontrol scientists regarded as essential in the field.
Cliff Ashall remembers David as a “pleasant, cooperative and industrious colleague of great integrity.” Nevertheless the locust days gave rise to one of the enduring legends about him, as Cliff recounts. David and Jerry Roffey, a fellow Imperial College graduate, joined DLS at the same time and “spent some time together in Somaliland and Eritrea . There was an Ethiopian locust officer with them who played his radio very loudly in the mornings, something that David did not agree with – and in spite of repeated pleas to turn down the volume it continued until David picked up a 303 rifle and put a bullet through the offending radio.” The story became embellished with time and, although apparently exasperated by this, David was known to put it to good use. In the 1980s, while Assistant Director of CIBC (the Commonwealth Institute of Biological Control; IIBC, the International Institute of Biological Control, as it became in September 1985), he was a popular visitor to Kenya and staff would vie with each other to put him up (something that afforded him wry amusement; as he once put it, he had to be careful to share himself around, and was never permitted the luxury of a hotel). These were the early days of personal computers and the dot matrix printer ruled the roost, or in this case the dining room. Eager to clear paperwork before setting off for a day in the field with David, Richard Markham (now Programme Director, Commodities for Livelihoods, Bioversity International) was printing out the results of his late-night labours at breakfast-time. This did nothing to allay David’s by-then renowned early-morning grouchiness, and it took but a single grumble about the noise and mess as the paper spewed out into the marmalade for the plug to be quickly pulled and the peace of a Kenya Highlands morning to be restored.
David married Annette in 1958. A graduate of the University of St Andrews in Scotland , Annette was recruited by ALRC in London and then temporarily seconded as a librarian to the International Red Locust Control Service in what was then Abercorn in Northern Rhodesia (now Mbale , Zambia ). After their marriage she was to become David’s professional colleague too, and her talents were an asset to CABI, both in Uganda where she worked with David on projects, and when they returned to the UK where she earned respect for her meticulous editing –notably for the Bulletin of Entomological Research.
David married Annette in 1958. A graduate of the University of St Andrews in Scotland , Annette was recruited by ALRC in London and then temporarily seconded as a librarian to the International Red Locust Control Service in what was then Abercorn in Northern Rhodesia (now Mbale , Zambia ). After their marriage she was to become David’s professional colleague too, and her talents were an asset to CABI, both in Uganda where she worked with David on projects, and when they returned to the UK where she earned respect for her meticulous editing –notably for the Bulletin of Entomological Research.
David joined the Commonwealth Agricultural Bureaux (CAB, now CABI) in 1962 to set up their first African base with the founding of the CIBC East African Station at Kawanda Research Station in Uganda , some eight miles west of Kampala . As such, James Ogwang (former head of the Biological Control Unit, National Agricultural Research Organization, Uganda ) describes him as “the grandfather of biocontrol in Uganda .”
The purpose of establishing CIBC stations in Africa was, David later wrote in the opening chapter of Biological control in IPM systems in Africa3, to assist African countries and to find natural enemies for export to other countries. This, as he outlined in the chapter, was during the aftermath of the era when synthetic pesticides had led many countries to abandon biological control, while remaining practitioners often tried to show that biological control was cheaper and permanent. The ease of shipment that air travel afforded had tempted many to economize on detailed ecological studies, and instead ship large numbers of species for release to see which would establish; lessons learnt from an earlier era were forgotten, inappropriate species were introduced, the success rate fell, and biological control came to be seen as something unlikely to succeed and to be used only as a last resort. Against this background, David’s emphasis on science-based biological control was invaluable.
Sean Murphy notes how: “From the start it was clear that David saw the need for creating centres of excellence – across the globe – to allow the science to flourish and for it to make a contribution to development”, and this was long before ‘development’ became the buzz word it is today. Gordon Tiley (then Pasture Agronomist at Kawanda, now at the Scottish Agricultural College ), describes how David “developed the East African station from practically nothing to a compact unit. A number of young expatriate and Ugandan scientists worked with and were trained by David, who placed particular emphasis on this particular aspect of the Unit’s work.” Encouragement of young scientists was to become one of David’s hallmarks. Gordon also says: “CIBC work frequently took David away on safari to all parts of East Africa in the Unit’s Land Rover. Being a small satellite, the Unit was constrained by limited laboratory and library facilities. Conditions for working were frequently technically and administratively challenging, though there was an excellent and comprehensive insect collection. However, with a quiet and organized approach, David sought to promote high standards of scientific professionalism.” David’s capacity to do excellent work under difficult circumstances could make him a hard act to follow. Ian Robertson (a lifelong friend from locust days) was thinking of David’s work in Uganda when, as Officer in-Charge, CIBC Kenya Station in the mid 1980s, he observed to junior colleagues that it was no point complaining to David about what he was expecting of them, because he had done far better work under far worse conditions and with far fewer resources. This tenacity, which David was to exhibit again and again during his career, was already matched by other traits Gordon Tiley describes that were to become familiar, and remain so, even as Director of IIBC: “He was approachable by staff at all levels and always willing to discuss a problem or to offer level headed advice, in characteristic measured tones and generally while lighting up or extinguishing his pipe!” Donald McNutt’s recollections from when he was posted to Kawanda as an Entomologist in 1967 highlight David’s open-mindedness: “My main work was testing the effectiveness of insecticides for pest control and the use of spraying machinery as opposed to biological methods but despite this David was always available to discuss problems with me. In particular he gave useful advice for a booklet I was writing on Insect collecting in the tropics.” He adds: “He was a realistic person who didn’t mind asking me to treat his house against possible cockroach breeding while he and his family went on vacation to South Africa .”
Professor Tecwyn Jones (Director, East African Agriculture and Forestry Research Organization [EAAFRO]; later Director, Commonwealth Instituteof Entomology [CIE]) says that David “was rightly widely commended for the success of CIBC’s biocontrol projects in East Africa .” This owed much to his wide-ranging abilities. One of the first projects he tackled concerned the Antestiopsis spp. complex, the main pests of Arabica coffee, which demonstrated how David went straight to the root of a problem and had the scientific skill to solve it; in this case, the identity of the pest was unclear. Donald McNutt, at the time working on Arabica coffee pests on Mt Elgon as Entomologist at Mbale, describes David as “a fine entomologist” and was “very impressed by the way he sorted out the Antestiopsis spp. complex”; David published a series of papers on this. Subsequent work on the sugarcane scale (Aulacaspis spp.) led to the establishment of a coccinellid introduced from Uganda to Mauritius but not control of the pest.  However, another coccinellid, Rhyzobius lophanthae (syn. Lindorus lophanthae) introduced from Mauritius to northern Tanzania was outstandingly successful in the continuous cropping system there and brought the pest under control within 18 months of being released. A (rare) well-funded project on cereal stemborers brought a second expatriate entomologist to the Station from about 1965 (Ed Milner, followed by Ikram Mohyuddin, then David Girling to 1973). David also worked on lantana biological control – a weed that continues to frustrate biocontrol scientists to this day. Insect agents (Teleonemia scrupulosa) achieved severe defoliation and dieback, but, David Girling says, “David came to realize that once the insects had knocked it down it just grew again if the land wasn’t cleared and used – early IPM?”
Professor Fred Legner from the University of California at Riverside (where he is now Emeritus Professor) spent time searching for natural enemies of the common housefly in Kenya and Uganda in 1966–67, under a joint project he and David were conducting for the US National Institutes of Health. This proved a significant partnership for Mauritius , where Stomoxys spp. stableflies were a severe constraint to dairy farming and cattle were kept in straw huts to protect them. First attempts to control flies by releasing New World parasitoids from dungfeeding flies had been unsuccessful in humid inland areas. Subsequently it was discovered the parasitoids controlled dung-breeding S. calcitrans, but not S. nigra which bred in the plentiful rotting vegetation (notably sugarcane trash) of the humid zone. Fred explains: “After I left, David supervised a study on breeding sites in Uganda . We came up with Tachinaephagus stomoxicida out of the work.” Between 1972 and 1975, sampling was carried out on banana trash and cut elephant grass. Pupae were shipped to Mauritius for study, and it became apparent that the natural enemy complex was markedly different from that in dung pits. Tachinaephagus stomoxicida was released and rapidly established in Mauritius , where it provided substantial control of S. nigra for most of the year – a case of careful ecological study reaping benefits.
In 1971 David published A review of biological control in the Ethiopian Region4, the fifth in CAB’s Technical Communications series, designed to review the development of biological control in the British Commonwealth . Tecwyn Jones pays tribute to David’s wider influence in East Africa: “As head of the CIBC East African Station, David, like all other entomologists in the region, was required to report annually on his work and plans for future research to the East African Specialist Committee for Agriculture and Entomology of the East African High Commission (EHC) – later the East African Community (EAC). I, as entomologist at EAAFRO, chaired that Committee (and submitted its findings and recommendations for approval to the EHC) and perhaps better than most came to know David’s value and standing on the committee.
“Throughout his time in East Africa , David was a major contributor to the deliberations of the Specialist Committee and was held in the highest esteem by all his fellow members. Thorough planning, meticulous attention to detail, and immaculate execution of every stage of his work, were the hallmarks of David’s own research, and of research projects under his authority. He also enjoyed the highest respect of his peers for his substantial and invaluable contribution to the delineation and work of the EA Plant Import and Export Committee and hence to the EA Plant Quarantine Service with its excellent record.”
If what has gone before suggests David was an overwhelmingly serious character, this was not the case.  Although he was dedicated to his profession and critical of what he considered poor scientific standards, Tecwyn Jones is keen to stress that there was another side to him: “David will long be remembered by his professional colleagues for his unique contribution to pest-management in East Africa – but no less so by all friends and acquaintances who knew him well for his personal attributes. He was a modest, unassuming, ever-helpful and kind person, whose advice and wise counsel was greatly valued by all. He was by nature rather shy and retiring but his studious thoughtful demeanour belied his keen sense of humour and a quick and healthy appreciation of the ridiculous – a combination which assured his welcome in professional and social gatherings of diverse composition and character.”
Gordon Tiley saw similar qualities: “He was a close family man, with wife Annette and children Andrew, Sarah and Emma, plus their affectionate dog, Sheba .  They were all popular and active members of the Station community and did much to contribute to its social life. David himself possessed a somewhat dry reserved sense of humour but he was always congenial company. A valuable steadying influence in times of argument or the inevitable personality conflicts among the more boisterous elements in a compact community.” Charles Dewhurst, then working under the late Eric S. Brown with African armyworm (Spodoptera exempta), remembers visits to Kawanda, where they had one of their light traps, for the typical hospitality with which they were invited for supper by David and Annette. He also recalls David’s interest in bombyliids, adding: “David always requested us to bring any specimens of Bombyliidae that we might come across, and that has to be my main memory, as everywhere I have visited in the world, seeing and collecting bombyliids and knowing that David was always interested.” Hospitality is a thread that runs through many people’s memories of David and Annette. When they eventually returned to the UK and David was based at Imperial College at Silwood Park , they regularly entertained CIBC/IIBC staff and ‘Silwood’ students, and are particularly remembered for their hospitality to overseas students stranded in the UK at Christmas-time.
David and Annette remained in Uganda under increasingly difficult political circumstances under the regime of Idi Amin until 1973. By then permission to leave the country even temporarily was difficult to obtain, but David managed to extract a letter personally signed by the Minister of Internal Affairs allowing him to leave, with Annette and Emma, to conduct annual field work in neighbouring Kenya (where their two older children were at school). With the CIBC Station Landrover filled with laboratory equipment, and what possessions they could fit in once this was all stowed, they set off, arriving at the border after dark. It was, as David recounted in later years, a particularly tense moment when he handed over the letter. They watched the soldier read it, slowly. They were not sure what to expect next – but it was certainly not what happened. The soldier, clearly awed by the signature on the bottom, asked reverentially whether he could keep it. Bemused, David cordially replied that of course he could. And so they were waved through. But their troubles were not yet over. On reaching Nairobi , the Landrover was broken into and the microscopes stolen; CABI folklore has it that David was reprimanded for his carelessness.
When a new CIBC East African Station was subsequently established at the Kenya Agricultural Research Institute (KARI) at Muguga, the Landrover found a permanent home and the staff cherished it for many years, putting up with its increasingly cranky habits with varying degrees of forbearance until a new breed of four-wheel drive vehicles, of less character but also less likely to shed windscreen wipers in heavy rain, superseded it.  Even so it was not forgotten, and at David’s retirement dinner at Silwood Park in 1991, Garry Hill, then Director of the IIBC Kenya Station, presented him with the Landrover’s wing mirror as a memento.
From East Africa, David moved back to the UK , where he was based until his retirement. One of the first tasks he took on, with David Girling and colleagues at the CIBC European Station at Delémont in Switzerland , was editing a companion to his review of African biological control: A review of biological control in western and southern Europe5 was published in 1976. At first he had an office in the CIE headquarters at 56 Queen’s Gate, London, and later he moved to the old headquarters of CAB at Farnham Royal where he was, as Richard Hill (now with Hill & Associates, New Zealand, and on the BNI Editorial Board) recalls, “the sole CIBC staff member in the UK at that time and I would visit him at Farnham Royal to seek advice. The move to Silwood and the growing of CIBC UK soon followed, all driven by David.” During this period, David Girling adds, David continued with project work; for example surveying in Kenya and Ethiopia for parasitoids of olive pests for Mediterranean countries.
David, with David Girling’s assistance on the ground, was also involved with plans for the new CIBC East African Station in Kenya . The agreement for this was signed in January 1981 and it began business in facilities provided by KARI at Muguga, near Nairobi . However, the status of CIBC in the UK was far from assured at this time, but David, as David Girling puts it, “characteristically turned crisis into opportunity, persuading CAB that CIBC needed an information officer (me), a journal to promote biological control (BNI, later handing over the editorship to me), as well as an Assistant Director (David) and a new headquarters (Silwood).” Silwood Park formed part of Imperial College’s Department of Zoology and Applied Entomology (now Biology), and was home to world-renowned ecological research, and to one of the few MSc courses in Applied Entomology in the UK – a magnet for overseas students. David moved to Silwood in June 1981.  David Girling, who retired from CABI and as Editor of BNI in 1997 adds, “much of what you see today stems from that time.” David’s ability to think ahead of his time became familiar to a succession of scientific colleagues over the course of his career. What was less well known was that he came from a family of innovators (and he himself put a good deal of effort into tracing and writing up the family history6). To cite but one, David’s great-uncle, James Henry Greathead (1844–1896), is known as the ‘Father of the Tube’ and a statue of him stands outside Bank underground station in London; his improved design for a mechanical shield “made tunnelling deeper, cheaper and safer for the army of workers building the London Underground”7.
At first housed in a few rooms in the Victorian manor house that was the centre of Imperial College at Silwood Park – in Richard Hill’s words – “ensconced in the gallery rooms ‘through the looking glass’ as I always thought of them”, David oversaw the planning and construction of a new IIBC headquarters building in the developing science park, which was opened by Professor M. S. Swaminathan in 1989.  David became Director of IIBC in 1989, on the retirement of Fred Bennett, and continued to develop the UK Centre with strong links to Imperial College . His belief, as Sean Murphy puts it, “that good science was the way forward” lay behind the recruitment of Jeff Waage from Imperial College as Chief Research Officer (he became Director after David’s retirement).  David supported Jeff in the establishment of the Leverhulme Fellowship scheme, a joint CABI–Imperial College initiative that was to produce useful research with applications to biological control. He also took advantage of the proximity to Imperial College to apply his inclusive approach to integration of biological and non-biological control technologies –IPM in short – by developing links with the experts in pesticide application in tropical countries at the college’s International Pesticide Application Research Centre (IPARC). Both these were to later prove his wisdom in engaging widely with other disciplines.
David’s outward-looking approach is endorsed by many people he worked with over his career. Peter Kenmore (Food and Agriculture Organization of the United Nations, FAO) describes him as: “A real pioneer, and a stalwart for biocontrol, who encouraged diverse approaches so long as they had been fieldtested.  FAO's first field biocontrol training course in rice, hosted by CIBC in India 25 years ago [in 1982], was made possible because David agreed to our nearly exclusive emphasis on conservation-oriented, rather than ‘classical’, biocontrol.” Harry Evans, a plant pathologist recruited by David from CIBC’ssibling institute, the Commonwealth Mycological Institute, CMI (later the International Mycological Institute, IMI), says, “David had the vision to realise that CIBC needed a more holistic approach to biological control, and, despite serious internal reservations, he managed to persuade CABI to invest in a pathology capability. Subsequently, a pathologist was appointed in 1984 to develop projects against both invasive weeds and arthropod pests.  More investments followed as specialist facilities were included in the plans for the new building and greenhouse infrastructure in order to handle both low- and high-risk pathogens. This also gave CIBC the opportunity to further enhance its role as a third country quarantine centre. Thanks to David’s legacy, high-profile pathology projects could be undertaken; including the highly successful one against the desert locust – a subject, of course, close to his heart.” The IIBC UK Centre grew to have a substantial pathology staff who also became involved in classical biological control of weeds, including the successful control of rubbervine (Cryptostegia grandiflora) in northern Australia by a rust fungus; David had first looked for natural enemies of this plant on the Kenya coast in 1973.
The inclusion of quarantine facilities at the Silwood Park site also meant that staff and students based at a UK university were able to study tropical pests.  David took advantage of the stability of a managerial role and association with Imperial College to get involved in supervision of research students in the 1980s, including PhD students Aristóbulo LópezÁvila from Colombia working on parasitoids of Bemisia tabaci and ‘Ravi’ Raveendranath from Sri Lanka working on Telenomus spp. egg parasitoids of Spodoptera spp.
David’s easy engagement with people created a network of international linkages that CABI benefits from to this day. Dr S. P. Singh (formerly Director, Project Directorate of Biological Control [PDBC], Bangalore , India ) describes David’s role in the evolution of CABI’s links with India . From when he was a post-graduate student in Russia , S. P. Singh had harboured a desire to meet the “stalwarts of biological control from CIBC.” His opportunity came in 1984 when, working in Bangalore as a Project Coordinator of the All India Coordinated Research Project on Biological of Crop Pests and Weeds, “I got the news that Dr David Greathead is visiting” and of their meeting he says, “when I met Dr Greathead, then Assistant Director, CIBC, I eagerly explained the activities and the progress of work and also put forth the expansion plan of the project to co-ordinate research, transfer viable technology on biological control of important crop pests and weeds and to serve as a nodal agency for introduction, exchange and conservation of biological control agents at national level. He listened carefully and offered several suggestions, and told me that such a type of expansion requires a lot of public funding.” Of this first meeting, S. P. Singh sums David up as “a very pleasant, modest, unassuming and helpful person with depth of knowledge and breadth of vision.” Although S. P. Singh was to meet David only once more, they continued to correspond, and he drew on David’s published material – notably, he comments, the BIOCAT database. In the years that followed, “collaboration and interactions with CABI improved” and continued to flourish after the formation in 1993 of PDBC with its 16 co-ordinating centres and laboratories.  The association has led to joint CABI–ICAR workshops and many other meetings and seminars involving CABI staff – indeed some have become regular visitors and collaborators. Thus, S. P. Singh concludes, “The seeds of collaboration sown by Dr Greathead seem to have germinated and flourished.”
Sean Murphy recognizes the importance of David’s influence and impact at a personal level: “I met with David in the early 1980s (when I was a student in the UK ) when he was already a leading light in biological control. I (and others) quickly learnt that this was a man who had real-life practical experience of trying to get science working for mankind – and who was succeeding – but also (somehow!) managed to keep the ‘romance’ of the science alive by being a practising scientist and a teacher. This was so important to younger scientists who at that age need to be shown how what they have learnt can make a difference.” Sean stresses that alongside all his ‘political’ and technical achievements was, “David the teacher and mentor. David always had time to discuss and share experiences, and most of all to help.” And also, “David the scientist. Once one of my staff in Kenya showed David some (we thought) beautifully prepared insect specimens (from coffee plants) for identification. David sucked on his pipe and after a long pause said, ‘Mmm, they need proper labels.’ But this was not a critical David – it was just David the professional.” Richard Hill remembers with gratitude David's guidance in his early research, when he was a student at Imperial College at Silwood Park but also responsible for the New Zealand Department of Scientific and Industrial Research (DSIR) gorse biocontrol project. He recollects how “a green, early 20-something New Zealand scientist first visited David at Farnham Royal to talk about the gorse project. He was always helpful and full of ideas, and of course I soaked up the stories about biological control history.” Richard’s memories of those days include “David's good company and droll sense of humour.” James Ogwang tells how, also as a student at Silwood Park , “Dr Greathead was a reference for me, perhaps one of the pillars that influenced me to develop interest in biocontrol. I remember him as a simple easy-to-approach fellow who was always smoking his pipe.” The significance to biological control of David’s encouragement of young scientists is well-illustrated by this, for James went on to be a driving force in the biocontrol effort against water hyacinth in East Africa, and the instigator of the community-based mass releases of Neochetina weevils that famously led to the weed’s biological control on Lake Victoria in the late 1990s.
David’s return to Europe did nothing to dim his enthusiasm for helping developing countries conduct safe and effective biological control. It was one of his motives in championing the need for international guidelines. Increasing environmental awareness had had a double-edged impact on biological control: potential environmental as well as economic nontarget effects of introduced biocontrol agents were starting to be seen to be significant; meanwhile, the emergence of IPM, in response to overuse of pesticides, was leading to increased adoption of biological control as its cornerstone. Thus countries with little or no previous experience of biological control were starting to make introductions of biological control agents, both for classical biological control and formulated as biological pesticides. Around 1989, David, on behalf of IIBC, together with the International Organization for Biological Control (IOBC), approached FAO to propose an international code of conduct. As David later wrote8, “FAO commissioned Professor Michael Way from Imperial College of Science, Technology and Medicine, London , an advisor to FAO on IPM, to prepare a review and discussion document on the need for a code, in association with IIBC and in collaboration with the FAO Integrated Pest Management Programme.” With this as the starting point, a worldwide consultative process over the ensuing years led to the development of the code as an International Standard for Phytosanitary Measures (ISPM) of the International Plant Protection Convention (IPPC: an international treaty for protection of plant resources), under the guidance of Dr Gerard Schulten of FAO and with support from David, culminating in its endorsement by FAO member countries at the end of 1995 and formal publication in 1996 as ISPM No. 39.
An assessment of ISPM No. 3, conducted by Moses Kairo, Matthew Cock and Megan Quinlan in 200310, described its publication as timely: in many developing countries the economic and social factors influencing biological control decisions tended to be more concerned with economic and food security issues than impact on indigenous species. They comment:  “It is those mostly developing countries recently starting to use biological control or with an opportunity to use biological control, which benefited most from ISPM No. 3. Until ISPM No. 3 was prepared, there was little guidance available to these countries and none with the international authority that is embodied in ISPM No. 3. [It] gave them increased confidence to proceed, based on the assurance that they were following international standards and procedures.” The authors also note that it “has provided a good basis for facilitation of regional projects and dialogue between countries facing similar problems.” The authors end by acknowledging that although many people were involved in the process, the efforts of Gerard Schulten and David Greathead were particularly important in seeing ISPM No. 3 through to finalization and ratification. David’s thoroughness and patience were key to the ultimate success of an initiative.  He himself used to observe ruefully that he was often labelled as pessimistic when he pointed out difficulties with other people’s bright ideas; his strength was that he not just foresaw problems, but persevered until ways had been found to overcome them. [ISPM No. 3 was revised and republished in April 2005.]
The quantity and quality of David’s publications were recognized in 1977 when London University awarded him a DSc. He continued to contribute significantly to biological control literature, including, with Jeff Waage, Opportunities for biological control of agricultural pests in developing countries published in 198311, and he edited with Jeff the Royal Entomological Society of London’s symposium volume Insect parasitoids in 199412. However, a major contribution during this phase, and still used today, was the BIOCAT database. This, according to David Girling, was initially a card database, kept by David, of all introductions of insect natural enemies (parasitoids and predators) for biological control of insect pests worldwide; his wife Annette took over running it when it was put on computer. David recognized, and he and Annette say in their 1992 review of BIOCAT in BNI13, “the results of introductions of agents of classical biological control are of great interest, not only to biological control practitioners, but also to ecologists interested in biogeography, and the process of colonization by invading species, to taxonomists who may encounter unfamiliar species and to conservationists concerned with their impact on native biota.”
Notwithstanding his progressive approach to biological control, David had definite ideas about what made a good biological control scientist, and among things he instilled into his recruits was the importance of taxonomy, encouraging them to develop a specific interest. His own interest in Bombyliidae and other Diptera, especially in the Afrotropical Region, was how Neal Evenhuis came across him: “I first became acquainted with David 23 years ago when I was compiling for a book all the published scientific literature of bee flies and was adding to it a short history of its workers.” Neal wrote to coworkers he was including, requesting a photo. “Everyone sent me portraits very quickly and without much fanfare, wishing me well in my endeavor.  Except David. I had never corresponded with him previously and he said he would get back to me, but only after finding just the right photo. I was baffled by the response. What could he have in mind? I was just asking for a simple portrait. All the photos of the other workers sent to me were the run-of-the-mill portraits or the typical pose by the microscope.  Except David. David had a photo done especially for my book. It was of him smartly dressed, smoking his pipe, and his head slightly tilted as though finding something of interest while examining flowers on a shrub. It was the best photo of the bunch and it typified David's method of work.” Neal acknowledges David’s influence in a way that many will recognize:  “He generously took me ‘under his wing’ as it were and – in addition to letting me in on his incredible knowledge of African bee flies – he also taught me about the necessities of scientific work: patience, thoroughness, and even diplomacy in dealing with co-workers.”
Jeff Waage (now Professor of Applied Ecology, Imperial College London) says: “My favourite memory of David was during his time as Director of IIBC, while I was his Deputy, perhaps because he had such an influence on me, when I finally took that role.” He realized that, “David was not one of those people who likes management for its own sake. For him, management seemed more of a duty or a service, undertaken in order to support his team, to help us to develop our programmes and to protect us from the whims of the organization above. He was approachable, sympathetic, and supportive, as such a manager would be. He could be a powerful calming force to a fretful scientist. He did this with the aid of a pipe, the filling, lighting and smoking of which created those frequent, thought pauses that turned the crisis into a process of solution.”
Jeff also saw characteristics many others have recognized: “The other feature of David’s management that left a permanent stamp on the persona of IIBC was his continuing interest in research and the day to-day business of biocontrol. In so many organizations, you find staff and management tend to differentiate, taking on different interests and priorities.  In IIBC, we were all, like David, just curious scientists. He set the example, and that enabled us to all remain one team of colleagues, whatever our secondary management role might be. And he let us all be our own managers – under David, IIBC was a place where you could chase any good idea you wanted, as long as you could find the money. In gentle and supportive ways, David would get involved with many projects.”
This was to prove crucial in a ground-breaking project that, not entirely by coincidence, brought David’s career full circle (although not in a literal sense) to where it started in Eritrea when, as Eileen Stower described to Cliff Ashall, David first joined DLS and he and fellow recruit Jerry Roffey could often be seen in Asmara “full of the joys of spring careering round the town in the back of a 15-hundredweight truck hanging onto the canopy irons.” Jeff Waage picks up the story more than 30 years later: “I remember clearly coffee time conversations [at Silwood] with him and Chris Prior [now Head of Horticultural Science at the Royal Horticultural Society, Wisley], batting about the challenges of locust control, the effect of oils on fungal spores and insect infection in Papua New Guinea, and my rantings about getting enough resources to do successful tropical biopesticide development. And then, eureka, an idea was born that grew and grew and much later became LUBILOSA (Lutte Biologique Contre les Locustes et Sauteriaux). Around other cups of coffee, he would challenge us about classical biological control, drawing from his vast knowledge and his BIOCAT project. Again and again, David contributed while manager to so many of IIBC’s most creative moments in his modest way. Gentleman manager and gentleman scientist, he was a very singular person.”
The LUBILOSA programme14, which went on to develop Green Muscle® as a biopesticide for acridids, grew out of concerns about the use of chemical pesticides during the locust plagues of the late 1980s which fuelled a demand for an alternative. A short concept paper by Chris Prior and David in the FAO Plant Protection Bulletin in 198915 identified deuteromycete fungi as promising candidate pathogens for locust control. From this initial idea, CABI went on to lead, with the International Institute of Tropical Agriculture, a multi-national, multi-institutional team which confirmed that an isolate of the fungus Metarhizium anisopliae var. acridum (IMI 330189) was the most effective biological control agent available, and developed robust formulation and application technology to allow it to be deployed as an effective biopesticide, Green Muscle®, which has subsequently proved its credentials in many field trials against locusts and grasshoppers in Africa.  Commercially produced for the first time in South Africa in 1999, it is registered throughout West and East Africa, and is recommended by FAO for use in environmentally sensitive areas; most recently FAO organized a trial of Green Muscle® against local hopper outbreaks in Mauritania in October 2006.  David’s early population studies with Bill Stower2 are achieving new significance: understanding multiplication rates and population numbers is becoming important in deploying Green Muscle® to manage population size in pre- and early post swarming locust populations.
The significance of David’s contribution to LUBILOSA went beyond the belief that biological control could work as part of locust control. The groundbreaking work in the programme was not based solely on the recognition of a suitable pathogen, but also relied on advances in formulation and application technology, so the fungal spores could be formulated as an oil suspension with a long shelf life, and sprayed using standard ultra low volume spinning disk spray equipment. His conviction that biological control should be based on science meant there was support for the recruitment of postdoctoral researchers, such as Matt Thomas (now with CSIRO Entomology, Australia ), under the Leverhulme Fellowship scheme to investigate critical features of locust biology and ecology. Roy Bateman (then CABI, now returned to IPARC), who was involved in the application aspects of the biopesticide development, says that “what marked David out was his breadth of view. He was inclusive in his thinking; for example, welcoming of the pesticide scientists and recognizing their value, despite being a world authority on biological control – and this was ultimately to the benefit of the locust programme.” Although much of this took place after David retired, it was his knowledge and foresight that allowed it to germinate, and his encouragement of scientists from diverse disciplines that laid the foundations for its ultimate success.
At 60, David came up against CABI’s obligatory management retirement age, and against his wishes stepped down as Director of IIBC. However, he was awarded an Honorary Senior Research Fellowship at the Centre for Population Biology, Imperial College London at Silwood Park , and remained professionally very active in biological control and bombyliid taxonomy. He continued to maintain BIOCAT, for example, and kept up a regular flow of information and ideas to BNI. Although saddened by some of the changes at CABI, he remained a stalwart support to staff, and as ready as ever to discuss ideas and problems and dispense advice based on his unparalleled knowledge.
Sean Murphy reflects on David’s influence: “I think most would agree that as one moves through life, one crosses paths with a few people who end up deeply influencing one’s thoughts and even how one approaches a significant part of one’s life. David was a leader, a great thinker, and a visionary and path maker and (as the messages I have seen from across the globe clearly show) he had a ‘guiding’ impact on many people.” Speaking for CABI, he adds: “He commanded respect because of who he was and what he stood for. David will not leave us – there is too much of him in what we now do.” S. P. Singh echoes these sentiments on behalf of the wider biocontrol community in saying, “the community will continue to traverse the path shown by him.” But, in the words of Gordon Tiley, News of his most untimely death will have been received with shock and sorrow by all who knew him.”
We extend our deepest sympathy to Annette and her family at the loss of this most singular scientist, man, husband, father and grandfather.

1.   Greathead, D.J. (1963) A review of the insect enemies of Acridoidea (Orthoptera). Transactions of the Royal Entomological Society of London 114: 437–517.

2.   Stower, W.J. & Greathead, D.J. (1969) Numerical changes in populations of the desert locust with special reference to factors responsible for mortality. Journal of Applied Ecology 6: 203–235.

3.   Greathead, D.J. (2003) Historical overview of biological control in Africa . In: Neuenschwander, P, Borgemeister, C. & Langewald, J. (eds) Biological control in IPM systems in Africa . CABI Publishing, Wallingford , UK , pp. 1–26.

4.   Greathead, D.J. (1971) A review of biological control in the Ethiopian Region. Technical Communication No. 5. Commonwealth Institute of Biological Control. CAB, Farnham Royal , UK , 162 pp.

5.   Greathead, D.J. (ed) (1976) A review of biological control in western and southern Europe . Technical Communication No. 7. Commonwealth Institute of Biological Control. CAB, Farnham Royal , UK , 182 pp.

6.   Greathead, D.J. (1997) A passage to the Cape of Good Hope . D.J. Greathead, Wargrave , UK , 132 pp.

7.   Cooper, J. The Greathead family one name study:www.greathead.org/

8.   Greathead, D.J. (1997) An introduction to the FAO code of conduct for the import and release of exotic biological control agents. Biocontrol News and Information 18: 117N–118N.

9.  IPPC (1996) ISPM No. 3. Code of conduct for the import and release of exotic biological control agents. [Revised as: ISPM No. 3 (2005) Guidelines for the export, shipment, import and release of biological News 9N control agents and other beneficial organisms. www.ippc.int/IPP/En/default.jsp]

10. Kairo, M.T.K., Cock, M.J.W. & Quinlan, M.M. (2003) An assessment of the use of the code of conduct for the import and release of exotic biological control agents (ISPM No. 3) since its endorsement as an international standard. Biocontrol News and Information 24: 15N–27N.

11. Greathead, D.J. & Waage, J.K. (1983) Opportunities for biological control of agricultural pests in developing countries. World Bank Technical Paper No. 11, pp. 1–44.

12. Waage, J.K. & Greathead, D.J. (eds) (1986) Insect parasitoids. 13th Symposium of the Royal Entomological Society of London . Academic Press, 406 pp.

13. Greathead, D.J. & Greathead, A.H. (1992) Biological control of insect pests by insect parasitoids and predators: the BIOCAT database. Biocontrol News and Information 13: 61N–68N.

14. LUBILOSA website: www.lubilosa.org/

15. Prior, C. & Greathead, D.J. (1989) Biological control of locusts: the potential for exploitation of pathogens. FAO Plant Protection Bulletin 37: 37–48.

We are grateful to Cliff Ashall, Roy Bateman, Keith Cressman, Charles Dewhurst, Harry Evans, Neal Evenhuis, David Girling, Keith Harris, Jocelyn Hemming, Richard Hill, Tecwyn Jones, Peter Kenmore, Fred Legner, Joyce Magor, Donald McNutt, Sean Murphy, James Ogwang, Mark Ritchie, Ian Robertson, S. P. Singh, Gordon Tiley and Jeff Waage for help and contributions. We owe a particular debt to Keith Harris for sources, contact details and advice.