Ian Murray Mackerras 1898-1980

Written by K.R. Norris.

Ian Murray Mackerras was born on 19 September 1898 at Balclutha in the south-east of the South Island of New Zealand where his father, James Murray Mackerras, and mother, Elizabeth Mary (née Creagh), were engaged in farming. His Scottish paternal grandparents had married in Melbourne, and his Irish maternal grandparents in Sydney. Shortly after Ian's birth, his parents moved back to Sydney, where they had been married. There Ian's brother, Alan Patrick, later to become Senior Lecturer in Engineering at the University of Sydney, was born on 28 August 1899. The family then returned to New Zealand until, in about 1902, the parents separated, and the mother returned to Sydney with the boys, to the comfort and support of her family. The grandparents, Patrick and Louisa Creagh, played a considerable part in raising the boys, who learned the use and care of tools in their grandfather's meticulously kept workshop. Patrick and his sons were solicitors, and, as with many other professional men living close to the Harbour, boats and sailing provided their principal recreation. The Mackerras brothers became expert yachtsmen.

Ian's birth certificate bears the given names Murray Ian Creagh, but quite early in his life, his mother's surname, Creagh, was dropped, and the order of his other two names reversed, so that he became Ian Murray Mackerras for the rest of his life.

Ian was educated at Sydney Grammar School, matriculating in 1915. Shortly afterwards (17 December 1915), with the consent of his mother, he enlisted in the Australian Imperial Force, advancing his age considerably to do so. He was allotted Number 5045 in the Army Medical Corps, and taken on strength as laboratory attendant in IHS Karoola on 19 December 1915. During the period of his service Karoola plied between Australia and the United Kingdom, either via the Cape or Suez, repatriating sick and wounded from the various battle zones (1). On one visit of the Karoola to Australia, Mackerras' service in her terminated (12 April 1917). The army records state that he 're-enlisted in New South Wales on 17 May 1917', but the term 're-enlisted' is probably loosely used here. More than likely he was given leave, during which he applied for a change that would make his participation in the war more adventurous. He was posted to the 8th Reinforcements, 31st Infantry Battalion, embarked at Sydney on 2 August 1917 in HMT A21 Miltiades, and disembarked at Glasgow on 2 October 1917. After being transferred to the Artillery and allotted the rank of gunner, he arrived in France on 13 December, and on 20 December was posted to the 51st battery, 13th Field Artillery Brigade, AFA (18 pounders). His army record has the laconic entry: 'Wounded in action "Gassed" on 28 May 1918'. This happened at Villers Bretonneux, in a sector where mustard gas shells were in free use by the Germans. Temporarily blinded, he was admitted to hospital in England on 9 June 1918. After recovery he returned to his unit in France on 17 November 1918, after the cessation of hostilities. He embarked for Australia on 28 February 1919, arrived in Melbourne on 13 April, and was discharged on 2 June, receiving the British War Medal and the Victory Medal. This was the end of his military activities until the 1939-45 War. After his discharge in 1919, there followed an intensive university career, marriage, parenthood and fifteen years of distinguished scientific research and administration in civil life.

Mackerras had early evinced an interest in medicine, when, as a schoolboy, a medical student friend draped him in a theatre gown and smuggled him into a theatre to watch an operation (2). His own biographical notes indicate that he was enrolled at the University of Sydney in April 1919. This was within a fortnight of his return to Australia, and even before his final discharge from the army. He recounted that he applied for and received an ex-serviceman's grant to do the medical course. As his studies progressed he realised that he wanted to be a zoologist, but if he switched to science he risked forfeiting, even having to refund, the grant which was so essential for his support. He thus proposed doing both courses simultaneously. His professors were apparently sympathetic, but one of the Deans foretold disaster in both courses, and withheld his approval. Mackerras then took his problem to the Vice-Chancellor, who agreed to the proposal. Anyone would have been justified in having reservations as to the likelihood of any ordinary man being able to perform this feat, but no ordinary man was involved, and in March 1924 Mackerras graduated MB, ChM, BSc, with First Class Honours in Zoology, the University Medal in Zoology, and the John Coutts Scholarship (shared). Mackerras wrote (1973) of his under-graduate years: 'The teaching at Sydney was good, some of it inspiring, and the teachers, especially in the clinical years, could still get to know their students as individuals'.

Mackerras states in his biographical notes that he found the influence of his Professor of Zoology, Launcelot Harrison, very stimulating. This is easily understood by comparing accounts of Harrison's career with the later development of Mackerras' interests. Harrison was a gifted field naturalist, active in Sydney scientific and natural history societies. He commenced a science course at the age of thirty. While still an undergraduate in 1911 he independently perceived the value of parasites in indicating the phylogenetic relationships of their hosts, a finding based on his studies of biting lice (Mallophaga) of birds. During the 1914-18 war Harrison served as Advisory Entomologist to the (British) Mesopotamian Expeditionary Force, and was credited with greatly reducing the mortality from malaria and typhus. A brilliant teacher, 'wide in his outlook and interests, he had the capacity of inspiring his students with the research spirit' (3). Mackerras stated that Zoology was 'probably the keenest and most friendly department in the faculty'. Harrison was deeply interested in the composition and origins of the Australian fauna, he supported the Antarctic radiation theory (4) before the appearance of Wegener's book (5), and further elaborated on it later (6) and he initiated informal discussions on these subjects in his department. His colleagues included a newly-appointed, full-time lecturer in Entomology, A.J. Nicholson, with whom Mackerras formed a life-long friendship. They became associated again later, when both joined CSIR.

In 1920 Mackerras met his future wife, Mabel Josephine Bancroft, who had enrolled for the second year of Medicine, after having completed a science course and having carried out post-graduate research at the University of Queensland. They seem to have paired off quite naturally. Quite probably Mackerras was already familiar with and deeply respectful of the achievements of Josephine's grandfather and father in medicine, zoology and other fields, and this would have given them a bond. During their student days they developed common interests in research and commenced the joint investigations that were such a notable feature of their life together.

The Senior Yearbook, 1923, of the University of Sydney Medical School quotes Mackerras as saying that 'sailing in an open boat is the best sport in the world'. Adding that all sailors are romantic, it went on to comment on his engagement to Josephine. During their university years Ian and Jo (as she was known to her friends) often fished from his skiff off North Head, making smears from the heart blood of their catches under what must have been very challenging conditions. They carried the fish home after sailing, and Josephine cooked them, while Ian stained the smears. After supper they would settle down at their microscopes, and search the smears for Haematozoa (7). This was the first of numerous joint research interests, and they still shared it many years later.

One of their microscopes fell victim to their marriage plans, which matured after their graduation, as it had to be sold to finance their travel to Eidsvold in Queensland, Josephine's home town, where, on 5 April 1924, they had the local clergyman marry them under a large river-gum on the banks of the Burnett. Such a ceremony must have seemed quite unconventional to many people in those days. 'They always agreed that their best wedding present was a telegram from Sydney to tell them that they both had appointments, Jo as Junior Resident Medical Officer at Royal Prince Alfred Hospital, and Ian as Research Scholar in Zoology at the University of Sydney' (8).

In March 1925 Mackerras was awarded the Linnean Macleay Fellowship in Zoology, and continued to enjoy hugely being paid for indulging in a hobby. Some of the fruits of this period are a definitive paper on the Nemestrinidae, and the description of new Mydaidae, two families of flies that excite curiosity in most dipterists. In the first of several papers on the taxonomy of mosquitoes, he also continued to contribute to 'medical zoology', as he called it, 'that area of medicine which involves both man and other animals' (9). This field was to become a major part of his life activities, and one in which he made or contributed to the planning of, tremendous advances in knowledge.

In Australia in the early nineteen-twenties 'productive work in entomology...was predominantly taxonomic and predominantly in the hands of educated amateurs'. Mackerras developed a high regard for these men and came to know well the Sydney group, G.A. Waterhouse, H.J. Carter, E.W. Ferguson and G.M. Goldfinch. He learned much from them in informal discussions after meetings of the Linnean Society of New South Wales and in the Entomological Section of the Royal Zoological Society of New South Wales, as well as on collecting trips and on weekends at the Zoological Society's cottage in the Royal National Park. In his Sydney years, Mackerras also formed enduring friendships with Ian Clunies Ross, later to become Chairman of the Executive of CSIRO, and with P.D.F. Murray, later to become Professor of Zoology at the University of Sydney.

In 1926 Mackerras' Linnean Macleay Fellowship was renewed, but he was given leave of absence in January 1927 to carry on work at the Bureau of Microbiology of the New South Wales Department of Public Health that would have been neglected in the absence of Dr E.W. Ferguson on sick leave. Ferguson died in July 1927, and Mackerras was then offered a permanent appointment, which he accepted, resigning his Fellowship on 30 September. Ferguson had also graduated in medicine in Sydney and shared with Mackerras an intense interest in zoology in general, and in the tabanoid Diptera in particular. He had also had extensive experience in the Australian Army Medical Corps in England, France and what was then Palestine. From the frequency with which Mackerras referred to him throughout his life there is no doubt that, like Harrison, the older man was an important influence in his career. It it likely, therefore, that a sense of loyalty played a considerable part in Mackerras' ensuring that his friend's interests continued to receive attention. Josephine had given birth to David Mackerras, now Senior Lecturer in Electrical Engineering, University of Queensland, on 6 June 1926. It is therefore also possible that Mackerras was influenced by the need for a secure income, though some would stoutly contest any imputation that he ever let considerations of security affect the course of his career. Whatever his motives, he made the best possible use of the situation, and, in addition to performing his heavy load of routine bench-work, he was able to further his role in medical zoology by participating effectively in a study of the ectoparasites of rodents. In later years he actually stated that his routine duties had been very worthwhile because they provided him with valuable training and experience (10).

In 1928 Mackerras made a move that was crucial to the development of his career. The Council for Scientific and Industrial Research had been founded by the Commonwealth Government as a statutory authority to provide a better environment for creative research than was usually found in government departments. The function of CSIR was to undertake research for the benefit of Australia's primary and secondary industries, but the Executive concentrated the efforts of the Council at first principally on primary industry. The function of the Division of Economic Entomology, one of the four founding Divisions, was to investigate entomological problems of national importance for which remedies had not been readily forthcoming. While the first Chief of the Division, R.J. Tillyard, was overseas interviewing candidate entomologists (who for the most part proved to be too demanding regarding salaries) the Executive formed the opinion that two of the senior posts could be filled better by Australian applicants (letter from the Chief Executive Officer to Minister, 10 September 1928). Unlike the overseas applicants, Mackerras, one of the two Australians approached, was willing to accept a lower starting salary than he was receiving at the Bureau of Microbiology, so eager was he to join this young and promising research institution. The date of his appointment was 1 December 1928.

It is obvious that Mackerras' appreciation of the training value of routine work had not developed at this stage, as he was quick to side-step a suggestion by Tillyard that he move to Canberra to take over the mass-rearing of the wasp, Alysia manducator, for the biological control of the sheep blowfly. The Canberra laboratories were not to be ready until November 1929, and, at his suggestion, arrangements were made for him to work for a time at Sydney University. A student of A.J. Nicholson's, Mary E. Fuller, was appointed as a temporary assistant to do the work with Alysia. This arrangement had also been suggested by Mackerras to obviate having trivial tasks deflect him from his main objectives in the Division of Economic Entomology. His instructions were to develop and direct a program of research in veterinary entomology, including the problems of sheep blowfly, buffalo fly and arthropod-borne viral and protozoal diseases of cattle, a big enough task without technical burdens as well.

The Mackerras family became established in Canberra early in 1929. Wisely, the period before the laboratory became available was used in travelling extensively to make acquaintances among entomologists and primary producers, and to appraise the problems. In June 1929 Mackerras also attended a Pacific Science Congress at Bandoeng (now Bandung) in Java. During this visit he installed J.L. Windred at a laboratory in Buitenzorg (now Bogor) to carry out research on the buffalo fly. Mackerras returned via the Northern Territory and Western Australia, where he made enquiries into the buffalo fly and sheep blowfly problems.

Mackerras' scientific papers and those of his colleagues document the story of the next ten years of his career, as he excelled at writing up his own research, and vigorously encouraged those under his direction to do likewise. He was a highly effective critic, and if the subject matter warranted publication at all, he could assist an author to make the roughest of drafts into a clear and succinct contribution.

During that decade he undertook many arduous field trips throughout Australia, in the process establishing first-hand knowledge of the buffalo fly and sheep blowfly problems, and building up among graziers a tremendous fund of goodwill towards research. A happy touch of showmanship, which he possessed in those days, and the infectious nature of his enthusiasm played no small part in this exercise in public relations.

People associated with Mackerras in the buffalo fly program were E. Handschin (a Swiss professor who studied the problem for two years in Java), H. Willings, J.L. Windred and T.G. Campbell. Substantial studies were made of the biology, phenology, ecology and ethology of the buffalo fly, and Handschin made attempts to bring about its biological control by liberating in North Australia Javanese parasitoids of muscoid flies and also their hybrids with related Australian forms ('hybrid vigour' was then enjoying a vogue). This biological control program had no detectable effect on the pest status of the insect, but other studies laid a valuable basis of knowledge for future investigations. In addition to planning and supervising the program, Mackerras studied the taxonomy of the buffalo fly, and the flies associated with it in dung.

In blowfly research Mackerras headed a team which comprised, for varying periods, A.J. Nicholson, M. Josephine Mackerras, M.R. Freney, Mary E. Fuller, C.R. Mulhearn, J.H. Riches (seconded from the Division of Animal Health and Nutrition), F.G. Holdaway, D.F. Waterhouse and D.J. Lee. Nicholson had been appointed to the Division as Deputy Chief, but, later, as Senior Entomologist, he transferred voluntarily to Mackerras' section, and spent some time on blowfly research. F.G. Lennox, working on insect toxicology, used the larvae of Lucilia cuprina as test insects and thereby contributed substantially to the blowfly program. Josephine Mackerras joined the staff in October 1930, as soon as a kindergarten had been started in Canberra, where their four-and-half-year-old David could be looked after during the day. In those times legislation forebade the employment by government of both man and wife. It is not known how Josephine survived a challenge which is in the records, but it is fortunate for Australian science that she did, as, for the next nine years her own papers and joint ones with Mackerras and others greatly expanded the knowledge on general and veterinary entomology.

Mackerras did much to elucidate the identity of the blowflies involved in sheep myiasis, and encouraged the extension of taxonomy into the larval stages (11) to short circuit the identification of flies from strikes. He assembled evidence that pointed to Lucilia cuprina being an introduced insect, and, with Josephine, performed some basic work on the attractiveness of living sheep for Lucilia cuprina. Under his direction, and often with a major personal input, the research program led to the discrediting of various methods of blowfly 'control' that had been advocated since early in the century, and which were now shown to involve much waste of effort. Central to this work was the demonstration of the crucial importance of Lucilia cuprina as an initiator of strike, the competitive disadvantages which this fly experienced in carrion in contest with other blowfly species (12), and the importance of living sheep as sources of Lucilia cuprina (13). The time-honoured burial of carcasses, primarily for sheep blowfly control, was thus demonstrated to be largely futile (14). A second, long-recommended approach to sheep blowfly control had been the destruction of adult flies by trapping. Mackerras participated in designing tests in which the incidence of strike was recorded in paddocks with and without traps, both in Western Australia and New South Wales. Statistical analysis showed that there had been a significant reduction in strike, but not marked enough to justify the considerable diversion of labour needed to carry out effective trapping. Poisoning of adult blowflies was another long-recommended method, but experiments carried out under Mackerras' direction suggested that this method also discriminated against secondary flies. He made positive contributions to the testing and adoption of the Mules operation, though, curiously, a tradition persisted for some years among sections of the veterinary fraternity that he had been opposed to the method. He also brought a new look to the development of sheep blowfly dressings, and the glyceroboric preparations developed by him in association with M.R. Freney and Josephine were magical compounds that were sorely missed when rises in the price of glycerine made their use uneconomic. His team did much to explore the conditions predisposing sheep to strike, the effects of strike on sheep, its cost to the industry, and the physiology, toxicology and behaviour of blowflies. Mackerras' thoughtful 'stock-taking' papers clarified the progress of the research, and preserved some realistic suggestions and concepts that never saw the light of day in research publications.

Other major studies during his period in Canberra were a joint investigation with Josephine and F.M. Burnet, then of the Walter and Eliza Hall Institute, Melbourne, on ephemeral fever of cattle, and another project, with Josephine and C.R. Mulhearn, on the transmission of anaplasmosis of cattle. These were no minor investigations, as they involved the use of 143 cattle lent by the Commonwealth Department of Health, and of 105 others lent by local owners. These cattle, and the divisional sheep flock of several hundred head, grazed on parts of the Black Mountain site and on sections of what is now the campus of the Australian National University, plus areas now occupied by parts of the suburb of Turner. Mackerras could remember the pet names of some of the cattle when their photographs were rediscovered a few years before his death. The paper on ephemeral fever is far more than a record of a piece of entomological research. In it they characterized the disease clinically, described the pathology, investigated the infectivity of blood from diseased animals, and studied the possibility of contact transmission and of mechanical transmission by interrupted feeding by the stablefly and several species of mosquitoes. Following their negative results with these insects they discussed the possibility of other insects being vectors, and, confirming earlier suggestions, ruled out all but the Ceratopogonidae, which, unfortunately, were not available for experimentation. Indeed, this family of blood-sucking midges was inculpated by subsequent workers. In Mackerras' own words (1973) the paper on ephemeral fever left 'no loose ends and recent studies have served to consolidate its findings'. The high quality of the work was confirmed by L.B. Bull in a foreword to the paper, which indicated that it had been undertaken in full consultation with his Division (Animal Health and Nutrition) and had made a 'definite contribution to knowledge'. Bull did not lightly hand out praise, and so this was praise indeed.

Thus, over the years 1928-1939, Mackerras' direction of research and his personal contributions brought a new perspective to the problems of buffalo fly and sheep blowfly control, and provided much new knowledge on two important cattle diseases. He also advanced the development of 'medical zoology', and made contributions to the advancement of zoology in general. With Mary E. Fuller he produced a classical study on the family Pelecorhynchidae, which they established. Outside of his own immediate interests he had a remarkable understanding of the work of colleagues. This was reflected years later, in his penetrating exposition of A.J. Nicholson's views on population dynamics, which showed an appreciation of the underlying themes that Nicholson's most vociferous critics never succeeded in achieving.

Mackerras acted as secretary to the Joint Blowfly Committee, comprised of representatives of the CSIR Divisions of Animal Health (later Animal Health and Nutrition) and Economic Entomology, and of the New South Wales Department of Agriculture. The clash of powerful personalities that occurred at these meetings must have helped to sharpen his innate awareness of the importance of human relations in science. In his biographical notes Mackerras states that during his early career with CSIR he 'came under the particular influence of R.J. Tillyard and J.A. Gilruth' (respectively first chiefs of the Divisions of Economic Entomology and Animal Health). Gilruth was a highly gifted and practical man. Tillyard, too was brilliant, and an unquenchable enthusiast, but with an outlook on science that was, at times, distinctly imaginative. Undoubtedly, however, Mackerras' eclectic approach would have ensured that he got the best out of both influences.

Mackerras' period in Canberra was marked by many a confrontation with the clerical side of the administration. The grudging treatment of expenses for the extensive travel he was obliged to undertake was particularly distasteful to him, with his generous and honest nature. His abiding enthusiasm for getting on with the job also made him chafe at any delays involved in working through official channels. Although, in these respects, CSIR was by no means the worst of the depression-acerbate government authorities in those days, it nevertheless provided him with sufficient challenge to augment his innate desire to cut red tape, and to side-step 'working by the book'. Many people profited in later years, particularly in the army, by his ability to sweep aside encumbrances imposed by bureaucracy. However, his relationships with the scientific personnel of the administration were always most cordial, and he enjoyed the very warm friendship and mutual respect of Professor A.C.D. Rivett, first as Chief Executive Officer of CSIR, and later, as Sir David, Chairman of the Executive.

Mackerras and Josephine worked long hours and often a six-day week. Mackerras did not expect such dedication from his team, though the loyalty his leadership elicited ensured that all did a fair day's work, and more when the program really required it. He did take an impish delight in recounting how he once recalled an unfortunate colleague from his honeymoon to send him on a field trip, but he was generally very considerate, and, with the touch of a born administrator, he knew full well the value of a judicious word of praise, bestowed where it was due.

The Mackerrases showed a warm kindness to single colleagues, obliged to live in boarding houses in what was then a very humdrum town. They frequently entertained young people, who found in their home a chance to join in scientific discussions and to relax in a stimulating and cheerful atmosphere. For recreation in their Canberra days the Mackerrases fished, swam, went on collecting expeditions and listened to recorded music. Both also learned to fly light aircraft, and participated in the activities of the Canberra Aero Club. Their licenses permitted them to carry passengers, and Mackerras also secured an Air Navigator's License. His examiners were astounded at the excellence of his pass, which was unprecedented in a self-taught candidate.

The outbreak of the 1939-45 War on 3 September 1939 triggered another important phase in the lives of the couple, as both immediately offered to serve. Mackerras was appointed a Provisional Captain in the Australian Army Medical Corps on 12 October 1939, and the following day was appointed Major in the Australian Imperial Force, with number NX18. He was instructed to report for duty at the 2/1 Australian General Hospital on 1 January 1940, attended brief refresher courses, and embarked for overseas service at Sydney on 10 January 1940. He and his fellow-participants in this first convoy out of Australia nicknamed themselves 'The Mayflowers'.

Mackerras disembarked at Kantara on the Suez Canal on 13 February 1940. He was posted as Pathologist at the 2/1 Australian General Hospital at Gaza Ridge near Gaza, as it then was. All reports of his activities in the Mediterranean Theatre are warm with praise. Though he was kept very busy at Gaza Ridge, he still found time to let his natural enthusiasms express themselves, and a friend remarked how his efficiently-functioning Pathology Department otherwise became like a small zoo. One visitor reported that while he was in the laboratory a snake slithered across the floor, and Mackerras did not completely allay his alarm by telling him that it was harmless (15). In January 1941 he was despatched, with special transport, to investigate flies and fly-breeding in the Western Desert, where diarrhoea was expected to take its toll of the troops as the weather warmed up. In recaptured Tobruk his recommendations for sanitary precautions, and for the quartering of troops in relation to the disposition of endemic peoples went far towards solving the problem of enteric infections. His services in sanitary entomology were also most valuable during the advance to Benghazi, when, 'after the experience of rapid movement in the desert the corporate and individual conscience in the lowly but necessary matters of hygiene and sanitation had relaxed vigilance'. Characteristically, his recommendations emphasised the need for research into the types of flies involved in the problems, and also the importance of selecting personnel well qualified for this approach. During the assault on Tobruk, Mackerras demonstrated his great versatility by participating in organising a highly effective blood transfusion service to meet the needs of the wounded who had to receive surgery.

No scientific publications resulted from his activities in the Mediterranean and the Middle East, and relatively few from his other activities in the 1939-45 War. His importance as a director of the research of other men precluded the achievement of an extensive publication list. Those papers that he did publish as 'sole' author, are generous in their acknowledgement of the contributions of others.

With the entry of Japan into the war (8 December 1941), Australian attention was, of necessity, focussed on the Pacific. Mackerras was recalled, reaching Melbourne on 23 May 1942. He was again Mentioned in Despatches (30 June 1942). In August he rapidly settled the nature of a problem that had arisen among troops in the Northern Territory, identifying the disease as infectious hepatitis, rather than leptospirosis, as local investigators had suggested. His summary disposal of this problem reflected his extensive experience of hepatitis in the Middle East, and re-affirmed his long-established mastery of microscopy and histology.

Mackerras was now about to begin his activities in relation to organising the study and control of malaria, dengue and scrub typhus, which stand out as a great wartime achievement, and a remarkable contribution to human welfare in the long term. His first involvement with malaria was in relation to an outbreak among troops and civilians in Cairns in 1942. However, this was of relatively minor concern, and, with the Japanese becoming established in New Guinea (Salamaua) from 8 March 1942 onwards, it was apparent that there would be extensive operations in the Territory, and that malaria would be a major factor in the success or failure of campaigning there. Thus, during the period 15-24 June 1942, he visited New Guinea in company with Colonel N.H. Fairley. 'They found a high spleen rate in native children in widely-separated areas on the south and north coasts, and confirmed that the areas likely to be occupied by troops were hyperendemic. What was more disturbing was the high rate of infection among troops stationed in and near the Port Moresby area, which before the war had been well controlled and relatively safe' (16). Shortly after their return to Melbourne they submitted a report to the Director General of Medical Services which detailed the measures necessary for the protection of the troops, and emphasised the dire consequences that would result from neglecting to implement them.

In May 1942 a conference on mosquito-borne diseases had been held in Brisbane. It was attended by Australian and Allied military representatives of preventive medical services, and resulted in an appeal to the Prime Minister for the establishment of army entomological units. This appeal was successful, though it was not until 25 October that Mackerras was given the title of Director of Entomology at Land Headquarters and instructed to get on with the job. He was promoted to Lieutenant Colonel on 26 May 1943. He and his Assistant director, Major F.N. Ratcliffe encountered tremendous problems in getting well-equipped and well-trained malaria control units into the field, and in solving these, Mackerras had ample opportunity to demonstrate his prowess for overcoming administrative inertia (17).

Similar problems were encountered in convincing the establishment that research on vector-borne diseases in New Guinea was necessary. Even when, by June 1943, there were some 25 000 servicemen already with malaria in the south-west Pacific theatre, it was difficult to convince non-scientific people in the army of the need for research, although a provisional plan for a research unit had been formulated as a result of discussions between Mackerras, H.K. Ward (Professor of Bacteriology in the University of Sydney) and Colonel E.V. Keogh (Director of Pathology and Hygiene). Eventually the Director General of Medical Services, Major General S.R. Burston, convinced General Sir Thomas Blamey of the need to establish the unit, and the Land Headquarters Medical Research Unit was set up at Cairns. Mackerras was responsible for the detailed planning of the entomological aspects of the program. The unique medical research output, under the direction of N.H. Fairley (then Brigadier) has been described elsewhere. The entomological aspects of the work were described by Ford (18) in his obituary of Josephine Mackerras, who had been called up in 1942. In Cairns she was, at first, put in charge of the mosquito culturing, but later she also participated in tests of anti-malarial drugs.

Under Mackerras' direction of the entomological services, the armed forces of Australia were relieved of much of the threat of mosquito-borne diseases and scrub typhus. By contrast the Japanese neglected prophylactic measures, and they paid dearly for it. Out of the many notable personal contributions that Mackerras made, perhaps one of the most outstanding was his organization of malaria-control measures in the Milne Bay area, January-March 1943. Here the Japanese had been defeated by 6 September 1942, but, in the aftermath, malaria was a severe problem to the Australian servicemen. Mackerras saw that a number of anti-malaria measures were implemented, and a dramatic fall occurred in the incidence of infections. '...In a report on the entomological and epidemiological aspects of Milne Bay, (he) found that the prompt reduction in malaria was undoubtedly directly related to the control of the vectors' (19).

In February and March 1944 Mackerras planned and participated in experiments to establish the vector status for dengue of a number of species of New Guinea mosquitoes that occurred in infected areas where the classical vector, Aedes aegypti, was scarce or absent. In the final, critical experiments, mosquitoes were collected at Lae and Finschhafen, fed on dengue patients, and flown to Sydney, where they were later fed on healthy volunteers. The experiments showed conclusively that Aedes scutellaris was an important vector, and so the way was cleared for counteractive measures to be planned.

From the beginning of his activities in the Pacific theatre Mackerras also directed and personally contributed to research to establish the vector status for malaria of Australian and New Guinea anophelines, which pointed to the over-whelming importance of mosquitoes of the Anopheles punctulatus complex. This information was crucial in the orientation of control measures.

By March 1944 the Japanese were being pushed back in New Guinea, malaria control measures, for the time being at least, were on a firm footing, and active participation by Mackerras seemed no longer so essential. He was therefore assigned to liaison work overseas, and his title altered to Advisor in Parasitology (23 March). He embarked at Sydney on 22 March, arriving in Washington on 17 April for a tour of the eastern United States 'to present the results of entomological research in malaria and (scrub) typhus in the South-West Pacific Area (SWPA), to give information regarding malaria control in the SWPA, to enquire into recent developments in insecticides and repellents, and to emphasise the value of plasmoquine in the treatment of malaria and in the limitation of its spread' (20). This he also did in England, arriving there on 28 June, and returning to the United States on 12 September, after a diversion to the west coast of Africa, from 19 to 31 August, to advise on malaria control. The visits were well received in the U.S.A. and in England, and they drove home the important part that scientific research had played in a war in the tropics and emphasised the importance of personal contact.

After returning to Australia in November 1944, Mackerras assumed the direction of experiments on aerial spraying with DDT that F. N. Ratcliffe had commenced during his absence overseas. His contribution was the more valuable through his having amassed information on relevant techniques during his period overseas as Advisor in Parasitology. Following the early experiments in Victoria and subsequent tests in the Cairns area, experiments were carried out in Papua New Guinea, December 1944-January 1945. Basic knowledge of the behaviour and biology of the malaria vectors was exploited in the course of these experiments, which demonstrated conclusively that aerial spraying with DDT-pyrethrin mixtures could eliminate malaria vectors. F.N. Ratcliffe carried out operational experiments in Bougainville and elsewhere in the Solomon Islands, and the techniques developed were then widely used in New Guinea, Bougainville and Borneo. Thus another effective weapon became available in the battle against malaria.

In December 1944 Mackerras moved to the north coast of New Guinea, his title having now been altered to that of Malariologist, 1st Australian Army. There he became involved in studies of malaria incidence which ultimately led to the demonstration of atebrin-resistance in the disease organisms (21). To provide an epidemiological background for these studies, Mackerras participated in a malaria survey at Wewak. The survey showed that the Japanese were the most important source of gametocytes infecting the mosquitoes that conveyed malaria to the Australian troops, and that particular wind patterns contributed to the problem.

Japan surrendered on 15 August 1945, but needless to say, this did not immediately free Mackerras of his military obligations. His transfer to the Reserve of Officers was effected on 21 February 1946, and on the same day he recommenced duty with CSIR Division of Economic Entomology. Thus ended more than six years of intensive and inspired activity on behalf of the Australian war effort, during which his personal efforts and direction of the research of others contributed to outstanding advances in the knowledge and prophylaxis of malaria, dengue and scrub typhus. In addition to being twice Mentioned in Despatches, as noted above, he was awarded the 1939-45 Star, Africa Star, Pacific Star, Defence Medal, War Medal and Australia Service Medal. His military career did not cease on 21 February, however, as on 2 September 1953 he was appointed Lieutenant Colonel in the Citizen Military Forces and Commander of the 1st Mobile Malaria Control Company, posts which he held until 19 September 1956, when he was transferred to the Reserve of Officers (Retired List).

Soon after demobilisation, the Mackerrases moved to Brisbane, where at the CSIR Veterinary Parasitology Laboratory at Yeerongpilly, Mackerras assumed the leadership of a team investigating problems in veterinary entomology. From the outset of his short period there (22 March 1946-2 June 1947) he displayed all his old vigour and enthusiasm. Within a short time he had prepared for publication a paper based on work carried out by the unit before his return, plus experiments in which he had since participated. His co-author wanted to delay while the inevitable few loose ends were tied up, but Mackerras firmly drew his attention to CSIR's obligation to make its results available to cattle owners, who, in some places, were in dire straits because of arsenic resistance in their tick populations. There was no detectable lag in his readjustment to research and administration in civil life!

On 2 June 1947 Mackerras took up an appointment as the first Director of the newly established Queensland Institute of Medical Research (QIMR) in Brisbane. It seems likely that his exemplary performance and unfailing willingness to be of service while in the Mediterranean theatre were remembered by people influential in the selection, and his activities in the Pacific theatre were a further demonstration of his suitability for the post. His achievements in the QIMR have been well documented (22), and need not be covered in detail here. He defined the special area of work for the Institute as 'that group of infections the study of which lies on the borderline between medicine and zoology', and later produced several papers on this topic. How closely he followed this aim is evidenced by the reputation that the Institute achieved in its work on the typhus group of organisms, including Q fever, murine typhus, scrub typhus and tick typhus, and the virus diseases, Australian arbo-encephalitis and dengue fever. Many other fields of 'medical zoology' and parasitology were advanced, and some straight-out medical problems explored.

At the outset Mackerras took measures to ensure that he had a free hand. He wrote an article in 1947 entitled Ways and Means of Research, which was strategically placed in the Institute's Second Annual Report. In it he expressed his views on what a research institution should aim to do, and how scientists should be trained and handled so as to ensure a sustained output of high-class research. Mackerras well recognised the undesirability of outside (ie, government) interference, and successfully exerted himself to keep this from affecting the activities of his staff. He also shielded them from routine tasks and from the burden of extension work, and took full responsiblity for all that happened in the research programs.

Josephine was appointed as the Institute's Senior Parasitologist on 1 September 1947, and again Mackerras enjoyed fruitful collaboration with her on a number of projects. In his busy fourteen years as Director he found time to carry out research on the taxonomy of the blood-sucking flies, Simuliidae, and Tabanidae. During his extensive research on the taxonomy, evolution and zoogeography of the Tabanidae he maintained warm, brotherly relations with the other world masters on the group, G.B. Fairchild, C.B. Philip and H. Oldroyd.

Despite a heavy burden of research and administration, Mackerras was also able to accomplish some major 'outside' activities, for which his enthusiasms or special abilities fitted him. On becoming established in Brisbane he was invited to become a member of the Great Barrier Reef Committee, which sponsored research on the Reef. He became Chairman of this Committee for the years 1955-56, and, with characteristic vigour, worked successfully to raise funds, both government and private, to implement the Committee's plans to erect a research station on Heron Island. Visits to the island, which he and Josephine made during this period, would have delighted them, as general zoologists, and provided them with a change from their arduous regular labours, which they probably did not realise they needed. One of the fruits of Mackerras' visits there was a thoughtful paper on the peculiarities of marine insects, a topic in which he remained interested for the rest of his life. His contribution to the advancement of the objectives of the Committee were recognised by his election as an Honorary Life Member in 1969.

Another major 'extra-mural' task performed by Mackerras while Director of the QIMR was to chair a Committee set up by the Commonwealth Government and the Government of New South Wales to investigate the reasons for the failure of the Cattle Tick Eradication Campaign conducted in New South Wales in 1956-57. The Committee's Report is a penetrating and highly constructive document, which clearly bears the stamp of Mackerras' deep knowledge of parasitology and epidemiology.

Mackerras remained Director of the Queensland Institute of Medical Research until 6 August 1961, when Josephine, whose 65th birthday was on the 7th, retired. He then retired also, leaving the Institute with a well-established world reputation for research of the first rank.

Both Ian and Josephine Mackerras were appointed in August 1961 as Research Fellows in the Division of Entomology, in the Commonwealth Scientific and Industrial Research Organization, both bodies having been rechristened on 19 May 1949. Mackerras was to edit a text-book on the insects of Australia to succeed Tillyard's Insects of Australia and New Zealand, which was outdated and long out of print, and Josephine was to carry out research on the taxonomy of Australian cockroaches, the need for which had become evident during her studies on the vector status of domestic cockroaches in the dissemination of salmonellosis in Brisbane.

The Insects of Australia or The Insects, as it became known to those involved in its production, was six and a half years in preparation. During this period the editor was served by a sequence of dedicated secretaries who slaved cheerfully on the mountains of typescript, and to meet his somewhat finical requirement for clean copies of even slightly altered pages.

There was a succession of artists, sometimes numbering as many as seven at once. The temperaments of several of them, who lacked scientific training, and sometimes proved insensitive to briefing, aroused the editor's ire on occasions, but, to offset this, several others had scientific training or experience, and needed little supervision. Mackerras himself, in places, assumed the role of artist, notably in Chapter 9: Composition and Distribution of the Fauna. The uniformity of the art work was due largely to the integrative role that he performed.

The uniformity of the text was also largely due to his skilful editing. Most of the authors conceded that, with his literary abilities and vast knowledge of taxonomy, biology and morphology, it was to their advantage to allow him a free rein in arranging the content and presentation of their chapters, and few objected to their final form. The overall results commend editorship of this type, but the uniqueness of Mackerras strictly limits the frequency with which the process can be repeated. Really serious problems arose in several cases over matters of opinion, in which, in the end, the chapter authors had to have their own way. It was because of such occurrences that Mackerras refused to have his name on the book as editor. He was firm in his belief that the book would be unreasonably blemished by the inclusion of some views that he considered to be misleading, inappropriate to the book, or else based on outmoded interpretations in anatomy.

In the mechanics of the production of the work Mackerras revealed astonishing abilities in fields in which he had no previous experience, such as the layout of pages, appropriate placement and grouping of illustrations, and the preparation of a highly compact and effective index.

Josephine's death on 8 October 1971 brought to an end forty-six years of happy married life, during which they had worked shoulder to shoulder for a total of about four decades. Their shared projects resulted in twenty-four joint papers. The philosophical way in which Mackerras bore Josephine's death reflected his realisation that she had been delivered from further great suffering.

He had published some papers on tabanid taxonomy while the textbook was in production, but after the book was finished, he had the freedom to intensify his studies, and a further series of papers on the Australasian tabanid fauna resulted. As a taxonomist he was a master. Any tabanid submitted would be taken to the microscope forthwith, and confidently classified after a few seconds of study, unless a dissection was required. Loss of his capacity to make dissections helped to put an end to his career in taxonomy. To his fellow dipterists he remarked with wry humour, 'When the amplitude of one's shake exceeds the size of the specimen, it is time to give up'. By about 1972 he could no longer dissect genitalia from tabanids without jeopardising the entire specimen. Deteriorating eyesight also contributed to terminating his cherished studies.

Mackerras retired on 31 December 1974, not long after the appearance of The Insects of Australia: Supplement 1974, which he had processed as meticulously as he had done the parent volume. His colleagues will long remember, with affection, his image over the years of his Research Fellowship in Canberra: at his desk, wearing a home-made cardboard eyeshield, writing deliberately and perspicuously in his impeccable script, or poring in total absorption over draft or proof. When a knotty problem arose, his behaviour was quite characteristic. He would tip back his chair, settle his heels on the table, and while cogitating, alternate between gnawing his finger-nails and drawing on one of the fat cigarettes that he rolled from 'the makings'. In his later years his cigarette-rolling left a generous carpet of tobacco around his chair, and one could easily tell where he had dropped in for a chat during the day. If his hands had become shaky, however, his intellect and memory were unimpaired, and he could discuss lucidly an enormous range of topics. For one who had never practised, he also had a remarkable command of medical science. He was always ready for sympathetic discussion of personal medical problems with friends. Though comfort could not always be drawn from the statistics that he could quote with astonishing facility, he was judicious as to whether he revealed them or not. A superb conversationalist, he had a rich fund of stories about interesting episodes and colourful people in his life.

Regrettably, the later years of Mackerras' life were clouded by the promulgation of Australian Customs Regulation 13A, to control the despatch of insect specimens from Australia to other countries. This legislation invited criticism because it was very clumsily framed, but Mackerras maintained that, in any form, it was an unwarrantable affront to the international spirit of science, and he regarded it as his duty to the scientific community to exert his every effort to bring about its repeal. He was unsuccessful, and he did not survive to see the introduction of long-promised guidelines to streamline the application of the regulation. In any case, his view was that 'No fiddling with guidelines can sustain a regulation that is fundamentally bad'. It saddened him that the confrontations over this regulation damaged long-standing friendships.

Except for an occasional excursion to Queensland, Mackerras lived quietly and frugally in his unit in Lyneham after retirement, watching cricket on television when it was available, until commercial elements intruded into the game, when he disowned it in disgust, and concentrated on listening to orchestral music from among the vast store of treasured selections he had on tape. His only other luxury was an evening dram or two of whisky, which, he maintained, in deference to an ulcer, had to be only of the finest quality, and bottled in Scotland. If friends dropped in, it added to his pleasure to pour nips for them also.

After surviving a mild cerebral haemorrhage earlier in the year, Mackerras succumbed to a massive one on 21 March 1980. Like Josephine, he had left instructions for a secular funeral ceremony.

To the academic and military honours and medals mentioned earlier can be added a most impressive list of other distinctions and activities. He was elected a Fellow of the Australian Academy of Science at its first election in 1954, and served on its Council in 1955-57. He was elected a Fellow of the Royal Australasian College of Physicians in 1950, and was also a member of the College of Pathologists of Australia. He served on the Faculty of Medicine of the University of Queensland, 1947-61, the Research Advisory Committee of the National Health and Medical Research Council, and the Advisory Council of the School of Public Health and Tropical Medicine, University of Sydney (now the Commonwealth Institute of Health). In 1961 Mackerras delivered the 35th Bancroft Oration to the Queensland Branch of the Australian Medical Association (unpublished). He had a long involvement with the Australian and New Zealand Association for the Advancement of Science. He was President of the Zoology Section at the Sydney meeting in 1952, he was elected a Fellow in 1957, and was awarded the Association's Mueller Medal in 1961. Mackerras was awarded the Clarke Medal of the Royal Society of New South Wales in 1950, and an Honorary Degree of Doctor of Science by the University of Sydney in 1971, a distinction which he cherished very much.

Mackerras was also at various times a Member of Council of the Linnean Society of New South Wales, President of the Naturalists' Society of New South Wales, and President of the Royal Society of Queensland. Having been a Foundation Member, he was also elected an Honorary Life Member of the ephemeral, Canberra-based, Royal Society of Australia. Mackerras was the patriarch and prime mover of major entomological associations in Australia. He was a member of the Entomological Society of Queensland for many years, and twice its President, and he will always be remembered for his potent influence in the establishment of the Australian Entomological Society, and as its first President, a story which has been told by himself in co-authorship with Marks and by Marks (23). He was President of the Society over the years 1965-67, and was elected its first Honorary Member in August 1969. Mackerras was also very happy to receive news of his election, in 1970, as an Honorary Fellow of the Royal Entomological Society of London.

As mentioned earlier, during his Directorship of the Queensland Institute of Medical Research, he acted as Chairman of a Committee of Enquiry into the Cattle Tick Problem in New South Wales (and into the reasons for the failure of the 1955-57 eradication campaign there). Mackerras then served for some years as Chairman of the Research Advisory Committee for the Research Station that was set up at Wollongbar in New South Wales, in response to his Committee's recommendations. In 1957 he was also Chairman of a CSIRO Committee of Review of the Cattle Tick Research carried out by the Division of Entomology, and, in 1959, of a Committee of Review of the entire program of the Division of Entomology. Both Committees submitted highly constructive reports.

It will always remain a puzzle why no civil honour was ever added to Mackerras' impressive list of degrees and awards.

His own, regrettably telegraphic, biographic notes, written during the preparation of The Insects, state: 'Research has oscillated between zoology (mostly entomology) and medicine (microbiology) throughout life, with a special interest always in problems of evolution and zoogeography; spent a lot of time laying foundations for the evolutionary studies'. Naturally he says nothing about the quality of the research, which is more important than the field, but undoubtedly it was uniformly of the highest, and the volume of it enormous. In taxonomic fields it bridged wide gaps that existed half a century ago. His own summary virtually sweeps under the carpet his war-time contributions to the control of vector-borne diseases, and effectively disguises his contributions in veterinary entomology. Immodesty was certainly not one of his attributes!

A feature of his life that illustrates Mackerras' great loyalty is the way in which he cherished, almost worshipped, the Bancroft history in science and medicine. This is epitomised in his description of the Tabanid fly, Cydistomyia bancroftae, where he drew attention to the fact that 'so begins the fourth generation of specific epithets based on the family name', and he must have been very pleased when Doherty quite naturally drew him into his account of The Bancroft Tradition in Infectious Disease Research in Queensland.

Mackerras was fascinated with evolution and continental drift. In contemplating these stupendous processes he found something of a religion, if he needed one, to replace the traditional ones that he eschewed. His philosophy was to enjoy this life to the full, and to leave to the timorous any thought of a shadowy after-life.

Like Harrison and other zoologists of the first half of this century, Mackerras had contributed to zoological studies which showed that continental drift had occurred, long before any geologists accepted the theory. In his paper on the Nemestrinidae, Mackerras commended the theory that the study of these insects 'showed that South America was joined to Australia by Antarctica'. At this time he had not read Wegener's book on continental drifts. In his paper on the Pelecorhynchidae, he stated that 'There is apparently no geological or bathymetrical evidence for the existence of past land bridges'...(which were at one time freely invoked to account for the distribution of animal groups)...'and the only remaining hypothesis which meets the conditions is that of Wegener (1924)...'It satisfies at any rate many of the geophysical and geological requirements, and it meets the biological facts we have discussed,...as well as those considered by Harrison. We may accept it provisionally as a sound working hypothesis from the biological point of view, leaving it to the geologists to substantiate it or modify it, or to propose a satisfactory alternative'. With what pleasure, therefore, must Mackerras have watched the reality of continental drift established over the last few decades by the developing science of plate tectonics.

Mackerras was a man of great physical courage, as his wartime activities testify. It is a sobering thought that, with a little less luck, his combatant role in the 1914-18 War might well have robbed us of the fruits of his activities in the South-West Pacific in the 1939-45 War. In the latter theatre he seemed to think nothing of flying into places near Japanese-occupied areas, and this at a time when hazards of air travel were high enough, even without the enemy. He used to advocate 'the old Mackerras principle of having a "look-see" for oneself'. He was equally fearless in any necessary confrontations in civil life. He never abandoned his dedication to truth around a conference table, or in arguments with scientific colleagues. Even the warm loyalty he accorded his friends in other matters never extended to his breaching this principle, which is also reflected in obituary notices he wrote: he gave his full measure of praise, but if there were critical things to say, his principles obliged him to say them, but he found gentle ways of doing so. Not for one of his mettle to add to the world's store of insipid stained glass saints!

Mackerras' humanitarian and intellectual achivements rate him as one of the really great Australians of the century.

About this memoir

This memoir was originally published in Historical Records of Australian Science, vol.5, no.2, 1981. It was written by K.R. Norris, a graduate of the University of Western Australia, who worked in the CSIRO Division of Entomology from 1937 to 1979, chiefly on pests of pastures and livestock. He first met Mackerras in Perth, Western Australia in 1935, and they were later associated in research in veterinary entomology at Yeerongpilly, Queensland and in the preparation of The Insects of Australia in Canberra.

Acknowledgments

The author wishes to acknowledge valuable assistance given by Dr David Mackerras, Dr Elizabeth N. Marks, Mrs S.W. Bailey, Mrs P.H. Nicholson, Dr Dorothea F. Sandars, Sir Frederick White, Dr D.F. Waterhouse, Dr R.H. Wharton, Dr R.N. McCulloch, Dr J.H. Calaby and Mr A.L. Dyce.

Notes

  • (1) José, A.W. (1943), The Royal Australian Navy. The Official History of Australia in the War of 1914-1918. 9. Angus and Robertson, Sydney.
  • (2) Marks, Elizabeth N. (1980), Obituary. Ian Murray Mackerras. Aust. Ent. Soc. News Bull., 16: 50-6.
  • (3) 'H.', J.P. (1928), Obituary. Professor Launcelot Harrison, Nature, 122: 65-6
  • (4) Harrison, L. (1924), The migration route of the Australian marsupial fauna, Aust. Zoologist, 3: 347-63.
  • (5) Wegener, A. (1924), The origin of continents and oceans, Methuen, London.
  • (6) Harrison, L. (1928), The composition and origins of the Australian fauna, with special reference to the Wegener hypothesis. Rept. 18th Meeting Australasian Ass. for the Advancement of Science. 18: 322-96.
  • (7) Doherty, R.L. (1978), The Bancroft tradition in infectious disease research in Queensland, Med. J. Aust., 1978, 2: 560-3, 591-4.
  • (8) Ford, E. (1972), Obituary, Mabel Josephine Mackerras, Med. J. Aust., 1972, 1: 604-8.
  • (9) Ibid. loc.sit.
  • (10) MacPherson, R.K., Personal communication.
  • (11) Fuller, Mary E. (1932), The larvae of the Australian sheep blowflies. Proc. Linn. Soc, N.S.W., 57: 77-91
  • (12) Fuller, Mary E. (1934), The insect inhabitants of carrion: A study in animal ecology, Coun. Sci. Ind. Res. (Aust.), Bull., 82.
  • (13) Waterhouse, D.F. (1947), The relative importance of live sheep and of carrion as breeding grounds for the Australian sheep blowfly, Lucilia cuprina. Coun. Sci. Ind. Res. (Aust.), Bull., 217.
  • (14) Fuller, Mary E. (1932), The blowfly problem. Notes on the effect of carcass burial. J. Coun. Sci. Ind. Res. (Aust.), 5: 162-4.
  • (15) MacPherson, R.K., Personal communication.
  • (16) Walker, A.S. (1952), Australia in the War of 1939-1945. Series Five. Medical. 1. Clinical Problems of War, Australian War Memorial, Canberra.
  • (17) Ibid. Medical. 2. Middle East and Far East.
  • (18) Ford, E. (1972), Obituary. Mabel Josephine Mackerras. Med. J. Aust., 1972, 1: 604-8
  • (19) Walker, A.S. (1952), Medical. 3. The Island Campaigns.
  • (20) Ibid.loc.sit.
  • (21) Fairley, N.H. (1946), Malaria in the South-West Pacific, with special reference to its chemo-therapeutic control, Med. J. Aust., 1946, 2: 145-62.
  • (22) Doherty, R.L. (1978), The Bancroft tradition in infectious disease research in Queensland. Med. J. Aust., 1978, 2: 560-3, 591-4.
  • (23) Marks, Elizabeth N. (1980), Obituary, Ian Murray Mackerras, Aust. Ent. Soc. News Bull., 16: 50-6.

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