AAS-Biographical memoirs-Ian Thornton

Ian Thornton was a fine zoologist, an accomplished academic acknowledged internationally as an authority in his field, and an admired leader and mentor to his colleagues and to generations of students. He came to Australia in early 1968 as Foundation Professor of Zoology at La Trobe University, Melbourne, and remained associated with that department, latterly as Emeritus Professor, for the rest of his life. He guided his department effectively until his retirement in 1991 and nurtured standards of excellence in research and teaching, whilst continuing to develop his research interests along two major lines: systematics and biogeography of Psocoptera (an insect order on which he was a recognized world authority) and Pacific-region island biogeography (becoming one of the leading regional biogeographers of his generation). Following his retirement, Thornton continued and diversified his academic activity, leading further physically strenuous expeditions to Indonesia and Papua New Guinea, publishing significant papers and a major book, and fostering international liaisons with universities in Indonesia and Laos.

Early Life

Ian Walter Boothroyd Thornton was a Yorkshireman through-and-through, born in Halifax and proud of his heritage and also of the fact that he was born (14 July 1926) on the anniversary of Bastille Day — a fact that somehow infused him in later life with a steadfast, forthright and in part revolutionary outlook, by which he always stood up strongly for what he believed to be just, was a staunch defender of his principles (and of his colleagues and staff), and imbued his dealings at times with an element of fun, and occasional risk. His father John, a Yorkshire dyer who had served in the First World War, died from peritonitis at the early age of 41, when Ian was only 10, but he remained close to his mother Alice Mary, née Crabtree, a Lancashire schoolteacher, until she died at the advanced age of 96. He had one sibling, his younger sister Mary Charlotte (later Mary Kitchen). The young Ian was an independent soul. He recollected his early schooldays as ‘rather tough’ and that for one period he was caned every morning for transgressions he was going to make that day! Although he was awarded a County Scholarship to Hipperholme Grammar School, he was later sent by his mother after his father’s death to a boarding establishment (Crossley and Porter’s Orphan Home and School), later returning to Hipperholme Grammar when war broke out in 1939. Ian excelled at school, both scholastically, with a number of school prizes to his name, and in sports, being Victor Ludorum in each of his last two years. His competitive nature, so well entrenched during his school years, persisted throughout his life. He also always remembered the full details of his paper round, and the hustling skills he acquired on the local church’s snooker table.

Around the end of the Second World War, Ian Thornton served (1944–1948) in the British army. After a short course in military engineering at Birmingham University he became an Officer Cadet Sapper in the Royal Engineers. He was later a commissioned officer in the King’s Own Yorkshire Light Infantry and parachutist in the 716 Parachute Brigade Company (6th Airborne Division). He served in the Middle East (Egypt, Palestine and Cyprus, with a trip to India to undertake a course in malaria biology and control), and was demobilized with the effective rank of Lieutenant. In 1948, Ian commenced his studies in Zoology at Leeds University, after marrying Jean (née Jean Frances Brown) at Hipperholme Methodist Church in August of that year. He remembered his undergraduate life fondly, and retained his meticulously transcribed undergraduate course notes throughout his life. Despite never having studied biology at school, Ian had decided in Palestine that he wished to become a zoologist, and the Professor of Zoology at Leeds, Eric Spaul, gave him a chance through a ‘trial year’ (supported also by the Professor of Botany, Irene Manton, whom Ian on their first encounter mistook for the cleaning lady!). He graduated with first class honours in Zoology and Botany, achieved the distinction of University Research Scholar (1951) and proceeded to a PhD, supported by a Nature Conservancy Research Studentship and supervised by Edward Broadhead, the authority on British Psocoptera and an ecologist of renown. Psocoptera, small insects that graze on algae and other microflora on the surfaces of bark and foliage, were little known at that time, and Ian was one of the first people to study their ecology in detail. The thesis involved a comparative study of the biology of three coexisting species of Elipsocus on a variety of tree species at Malham Tarn. Field surveys and laboratory experiments were combined elegantly in one of the pioneering studies on psocid ecology that was of much wider relevance in exploring how closely related species could coexist through niche differentiation. This work, completed successfully in 1953, resulted in his first two major papers on psocids (1, 2), and initiated his life-long interest in the twin strands of psocopteran systematics and ecology that would lead eventually to much wider considerations of evolution and biogeography.

Ian then moved from Yorkshire as a fledgling academic to work successively in three very different university environments, on different continents, in each of which his horizons and influence continued to expand. His long-time friend and colleague and fellow Leeds graduate, Alan Marshall, speculated in his eulogy at Ian’s funeral in Melbourne that throughout his life Ian vigorously pursued Charles Darwin’s advice (which Ian had himself quoted in his preface to Darwin’s Islands) that ‘nothing can be more improving to a young naturalist than a journey to distant countries’. Notwithstanding this, the standards and attitudes of a Yorkshire culture were to remain.

Sudan

Ian’s first appointment on leaving Leeds was as Lecturer in Zoology at the then Gordon Memorial College of Khartoum (later to become the University of Sudan [1956] but then affiliated with the University of London), for a three-year period, 1953–1956. This resulted in short papers on a variety of taxa and topics (scorpions, sea urchins, moths, succession in papyrus communities [3–6]), so diversifying his broad zoological interests and expertise. His major inspiration seems to have been the hydrologist Julian Rzoska, who was working on the Nile as a biological system and was perhaps instrumental in introducing Ian to ‘big picture ecology’, founded in the study of detail. Psocoptera took a temporary ‘back seat’.

Hong Kong

Thornton’s interests in Psocoptera re- established firmly when he moved to the University of Hong Kong as Senior Lecturer in Zoology (in a department then led by David Barker and later by John Phillips) in 1956. Here he was to remain for the next eleven years. His major initial research thrust was to collect and describe the local psocid fauna (7–11, 14), and to attempt to place them properly in the wider perspective of the fauna of south-east Asia and the western Pacific. He had met Lin Gressitt and other Pacific-region entomologists who were to become long-term friends at the Pacific Science Congress in Bangkok, soon after arriving in Hong Kong. Thus, in addition to descriptions of a substantial number of new taxa, this period also saw development of Ian’s interests in psocid dispersal and distribution, with studies on the wider fauna of the western Pacific, and dispersal mirrored by captures on ships and aircraft in the region (17, 18). Major studies, in part based on examination of the major regional accumulations of specimens at the Bishop Museum, Honolulu, resulted in conjunction with his postgraduate students, Wong Siu Kai (Peripsocidae and Ectopsocidae [26]), Lee Soo-Seong (Pseudocaeciliidae [23]) and Chui Wun Duen (Violet) (Hawaiian and Micronesian taxa); a fourth psocidological student of that era, Woo Kam Tien (Anita) moved with Ian to La Trobe, where she completed her analysis of the Galapagos psocid fauna (33). The results were a much fuller picture of the regional Psocoptera, with strong evolutionary and distributional underpinning to help explain the characteristics of the fauna. As later at La Trobe, Ian also supervised students working on a variety of non-psocid projects in Hong Kong; several of these were focused on the biology of rice pests, in relation to a collaborative project with Alan Marshall and Cliff Lewis of Imperial College.

These studies on psocids thus laid a solid grounding as stimuli for development of later, more wide-ranging studies. Ian’s interests, eventually to become predominant, in island ecosystems and the processes of island biogeography were founded in psocids and during the Hong Kong phase of his career. A brief visit to Hawaii in 1961 indicated the explosive speciation of several psocid genera there — rivaling and to some extent paralleling the better documented case of Drosophila on the archipelago. He spent a year at the University of Hawaii’s Institute for Advanced Studies (1963) as a Visiting Senior Scholar, using the time to collect intensively in as many parts of the archipelago as he could reach. The resulting taxonomic monographs (57, 63, 93), mostly not published until some years afterward, are of lasting importance. They changed dramatically the earlier perspective of Hawaiian psocids given in the Insects of Hawaii monograph (Zimmerman 1948).

That year led to seminal changes in Ian Thornton’s thinking and his approach to research. Psocoptera became more firmly tools for exploration of wider evolutionary processes, rather than simply things to be described and enumerated in their own right, although the importance of doing this remained to ensure the reliability of the data he used. His maturing focus on processes of speciation on oceanic archipelagos was treated to what was effectively an independent replicate study in 1967, with a three-month stay on the Galapagos Archipelago, the biota of which had so inspired Charles Darwin more than a century before. This study revealed intriguing parallels with Hawaii and also some significant differences. The background information available was far less — simply, no psocids had been recorded previously from any of the Galapagos Islands. The major taxonomic outcome (published jointly with Anita Woo [33]) recorded 39 species, of which 18 were described as new. The similarities and differences between the faunas of these two archipelagos were important in the evolution of Thornton’s thinking. He regarded the Galapagos fauna as at a much earlier stage of evolution than that of Hawaii.

Hong Kong was not wholly about psocids! Ian’s interests encompassed other topics, such as the genetics of the white tigers of Rewa ([27] this resulting from a time he was marooned in Calcutta, thwarted from a planned visit to the Andaman Islands). He also collaborated in the classic studies of the genetics of the mimetic swallowtail butterfly Papilio memnon led by Cyril Clarke and Philip Sheppard, by collecting for them in Palawan, Hong Kong and parts of Indonesia (28). A book on Insects of Hong Kong he initiated with Phyllis Hore was later completed by Dennis Hill (book 2). The main impetus for this was Ian’s realization that there was considerable need for a locally focused entomology text, so that his students did not have to rely on those written more centrally for students in the northern hemisphere. Perhaps the greatest intellectual outcome from the Hong Kong years, though, was his widely read book Darwin’s Islands (book 1), mostly written in the year after he left the university and for a decade or more the standard account of the natural history of the Galapagos. The book was translated into Japanese following its initial publication in New York. The wide survey of the islands’ biota is interspersed with numerous personal observations interwoven with the established literature, as well as with ideas on evolution and conservation. As importantly, though, it is immensely readable, and the lucidity and insights that came to characterise Ian’s teaching and writing are already evident.

His academic progress was marked by promotion to Reader in Zoology (October 1966), but Ian had long also played a full part in the corporate life of the university. He was Acting Head or Head of the Department of Zoology on several occasions and Dean of the Faculty of Science, 1960–1963. He served on many boards and committees (including the University Senate, 1961–1965), and ‘Y. C. W.’ wrote in the University of Hong Kong Gazette (1967) on Ian’s departure, ‘Those of us who have at one time or another sat at the same conference table with him will remember his frankness, his keen observation, and his commonsense approach to problems’. These attitudes persisted, as did his concerns for students and colleagues. Again from Y. C. W.: ‘Dr Thornton enjoys a high reputation as a teacher and has a genuine concern for all his students…. Many of his students and junior colleagues will not forget the help and guidance they received from him on academic and other matters.’ Indeed, throughout his career, Ian was an inspirational teacher. In Hong Kong he was instrumental in introducing highly ‘urbanized’ Chinese students to rigorous field work on the then remote Lantoa Island. He consistently trusted his judgement of students, even to the extent of confronting eminent external examiners when he considered their opinions deficient. His tabletop duel (using toy swords, and ending with both protagonists falling off) with J. Z. Young (University College, London) resulting from one such defence of his students’ marks has passed into folklore.

Australia

Ian and Jean, with their children Angus and Jane, arrived in Australia on the Royal Interocean Lines ship ‘Tjiluwah’ on 6 January 1968, which he recalled as a ‘100 degree day’, to take up the Foundation Chair of Zoology at La Trobe University, Melbourne, Ian having selected this from amongst the three chairs he was offered around that time. The late 1960s was an exciting time in Australian universities, with an air of optimism brought about by the establishment of several new institutions, amongst which the promise of La Trobe was influential in Ian’s decision. Although interview (by a committee including Macfarlane Burnet) and selection were rigorous, Ian recorded that he was subsequently first offered the job by the then Vice-Chancellor, David Myers, whilst they occupied adjacent urinal stalls in the gentleman’s toilet! He formally accepted a few days later. Ian set about establishing a Zoology Department (initially as a non-departmentalized part of a wider School of Biological Sciences, following the educational philosophy of the university’s founders), based on his belief, from which he never deviated, that ‘Zoology is the study of animals, not just of books about animals’. He recognized the need to recruit colleagues, predominantly focused on ‘whole animal biology’, who were capable of communicating both knowledge and enthusiasm to their students. The major thrust of the department was to be ‘terrestrial zoology’; at that time Monash University (although also strong in terrestrial zoology) had firmly established regional leadership in freshwater biology (through Bill Williams and his colleagues), and the logistic difficulties of developing a strong marine programme were formidable. Ian also believed that the Professor should be the Head of Department, as both academic leader and mentor, and he fulfilled both roles for as long as he was allowed to do so (that is, until his retirement in 1991).

The scope of psocid studies initiated whilst in Hong Kong continued as the major research focus of Thornton’s first decade in Australia, but became conceptually expanded and geographically concentrated on the biogeography of the Pacific region, particularly the western side, and in particular on the psocid faunas of the Melanesian arcs of islands. Much of this work was based on Ian’s own field expeditions in a long-term ARGC-funded project (1971–1982), with Courtenay Smithers and (to a far lesser extent) Tim New as collaborators. Thus, over a period of some twenty years from the mid-1960s, Ian visited and collected psocids in Sri Lanka, the Himalayan foothills, Malaysia, Japan, many parts of Indonesia, Papua New Guinea, the Solomon islands, Palawan, many parts of the Melanesian arcs including Norfolk Island, New Zealand, the New Hebrides, Fiji, Tonga, the Society Islands, the Galapagos, Mexico, Colombia, Ecuador, Peru, Chile (including the Juan Fernandez archipelago), Argentina and Hawaii. He maintained detailed field journals for most of his field work, and the progressive list of places visited reads like a major gazeteer for this vast region. These studies gave him a unique personal perspective on an insect order and its evolution over a substantial part of the world.

Most of the field work was undertaken on shoestring budgets, and Ian’s Yorkshire upbringing and philosophy rendered him reluctant to operate on anything more than restricted personal financial input. Wherever possible, he would bargain hard to reduce costs of accommodation, car hire and so on, and he cared little what he ate — a packet of cornflakes was just as satisfying as a three-course spread. However, despite strenuous physical activity, he rarely succumbed to gastric or other upsets in the field. Much of the collecting was in remote areas, with a general tendency to move upward from lowlands to mountains on the basis that these would yield more ‘typical’ or endemic psocids than the more disturbed lower regions. In part, this was common sense in facilitating access to less-disturbed habitats. Ian frequently turned his eyes, and his body, toward the hills. Distributions along altitudinal gradients intrigued him, and he sometimes claimed that simply climbing up Mt Rinjani (Lombok) and seeing the changes along the way was a very fine lesson on tropical biology for any student to undertake.

Collecting trips with Ian tended not to be luxurious and relaxing, despite the envious comments made by colleagues who did not participate and thought of New Guinea and like places as ‘romantic’. He worked hard, remained focused on his objectives, and maintained a positive attitude under sometimes appalling and dangerous conditions. Ian’s competitive nature persisted on field trips so that visits to the local ‘expatriate club’ in (for example) remote parts of New Ireland or New Britain could become ‘interesting’. He prided himself on the skills at snooker obtained in his youth and commonly challenged the local champion to a game, which he resolved to win. If things were not going his way (a common occurrence, simply because most such local players sometimes seemed never to move away from the table!), a frequent gambit was to pause and casually ask his opponent whether the ball he was about to address was of a particular colour. After the usual surprised/annoyed retort, Ian would point out (correctly) that he was colour-blind and, having so disconcerted his opponent, commonly went on to win the game. Ian’s persistence, nevertheless, took him and his collecting companions to many remote areas that had never been explored before, some of which have now been changed dramatically by human pressures. The taxonomic treatment and faunal analysis of the accumulated Psocoptera added massively to knowledge of this complex and rapidly changing region. The succession of descriptive papers, many of them co-authored, and illustrated by Justine O’Regan, Jodie Kernutt, John Greer, Jenny Browning or Tracey Carpenter, are significant additions to the psocid literature. Altogether, Ian (alone or with his co-authors) described almost 750 new species of Psocoptera, a significant proportion of the documented world fauna. His work on psocids was recognized by the small global fraternity of psocidologists in dedicating eight species to him (as named ‘thorntoni’) and in the genus Thorntoniella whilst he was alive; a commemorative volume of papers on Psocoptera (Garcia Aldrete et al. 2005) augments these by a further two genera (Ianthorntonia, Thorntonodes) and four species.

However, this basic taxonomic work was simply a template for Ian’s increasing interests in island biogeography and patterns of distribution and speciation. Substantial papers on the distribution and origins both of taxa (e.g. Philotarsidae) and faunas (Hawai’i) are classics of much wider interest than to psocidologists alone. Ian’s DSc degree (Leeds, 1984) recognized the importance of this documentation and his developing syntheses, which later came to constitute some of his most significant work and to establish him among the forefront of modern Pacific-region biogeographers.

The second major theme, developed from the early 1980s on, was to lead to what many peers regard as Ian’s finest academic achievements. In 1982, with Ann (he had married Ann Juliana Patterson in 1980, following the dissolution of his first marriage in the mid-1970s) Ian had his first sight of the area that was to become his major scientific passion for the next decade and more — the Krakatau islands, nestled in the Sunda Strait between Sumatra and Java. Ian later noted his recurring feelings of excitement and awe each time the small fishing boats used for travel to the islands entered the caldera and moved along the base of the imposing towering sheer cliff face (800 m high) of Rakata. On that initial visit, Ian recognized the unique opportunity Krakatau provided for studying the colonization processes and development of tropical communities and ecosystems from a tabula rasa beginning. In contrast to Hawaii, where the emphasis of his studies had been on post- colonization radiations related to isolation, his perspective now broadened further to consider and study the initiation and development of tropical communities. The unique natural laboratory of the Krakataus comprised two distinct temporal sequences for studying the development of tropical systems. First, the cataclysmic 1883 eruption is widely believed to have obliterated all life from the islands, so that the condition of vegetation and animal assemblages on the three older islands (Rakata, Sertung, Panjang) in the 1980s represented the outcomes of a century of re-establishment from source areas of Java and Sumatra, each more than 40 km away. Second, and nested within this, the island of Anak Krakatau (‘Child of Krakatau’) emerged lastingly from the sea in the centre of the caldera in 1930, undoubtedly virgin land and providing a second, much younger sequence for study of colonization from the much closer source areas of the other three islands. Six expeditions to the Krakataus were organized and led by Ian, extending over almost a decade from 1984. They involved many colleagues and collaborators, and led to a series of papers of lasting interest and relevance in island biogeography. They culminated in Ian’s magnum opus Krakatau: The Destruction and Reassembly of an Island Ecosystem in 1996 (book 3), a book widely regarded as Ian’s finest academic achievement, and recognized by winning the 1996 Professional Scholarly Publications Award of the Association of American Publishers, Biological Sciences Category. These expeditions were hard work, at times frustrating, but probably all participants (including several honours and graduate students working in the tropics for the first time) viewed them as highlights in their academic careers. Camping on Anak Krakatau provided a remote but idyllic scenario in that harsh environment, and it came to be a place that Ian (and his companions) loved — despite the ever-present threat of volcanic activity, the fact that all food and water had to be carried to the island from Java, and the undoubted terrors of unpredictable sea crossings in rough weather. Tim New went on four of those expeditions and noted that Ian’s participation was enthusiastic and dynamic. Indonesian counterpart scientists became lasting friends, and sojourns in Bogor or elsewhere during the lengthy process of obtaining permits and other documentation allowed opportunity to collect in a variety of possible source areas for the Krakatau fauna. As in much of Ian’s earlier work, psocids were a focal group but now only one of numerous biota (even including bacteria [73–77] and soil nematodes) incorporated into the emerging picture. A highlight for Australian expeditioners was the participation of scientists from many other parts of the world. With the closest parallel study to the one on Anak Krakatau being based on the emergence of Surtsey (off Iceland, and which Ian had visited a year or so previously), a brief visit to Anak in 1990 by Sturla Fridriksson helped to foster insights from a broad and authoritative base, and provided an opportunity to discuss parallels more closely.

The major importance of the ‘Krakatau study’ was recognized by Ian being awarded the John Lewis Gold Medal by the Royal Geographical Society of Australasia (1992) and his election to the Fellowship of the Australian Academy of Science (1995). Later, Ian turned his attention to even wider aspects of the colonization of volcanic islands, appraising the role of Sebesi in the northern part of the Sunda Strait as a ‘stepping stone’ for colonization of the Krakataus (119), and (then in his 70s) making a further physically strenuous expedition to explore Mot Mot, a rare example of an island in a lake in the closed caldera of a volcanic island (Long Island) in Papua New Guinea. His major collaborator on this exploit was John Edwards, who had also visited Krakatau with Ian and who had worked extensively and innovatively on the colonization patterns following the eruption of Mt St Helens, Washington State, USA, in 1980. And, as for the Krakatau studies, Ian edited the series of papers to ensure that they appeared in co-ordinated and accessible form rather than being scattered widely. The Long Island papers constituted a special issue of the Journal of Biogeography, following the earlier Krakatau papers grouped in Philosophical Transactions of the Royal Society of London (1988, 1990) and GeoJournal (1992, this last being the proceedings of a two-day session organized by Ian at a Pacific Science Congress in Honolulu).

Ian’s zest for life and for his science was infectious. He revelled in academic discourse and persisted with argument until he got responses that (at least for the time being) satisfied his curiosity. He was a perceptive reviewer of grant applications and manuscripts and served on several editorial boards, as well as a term as Vice- President of the Australian Entomological Society. The example of enthusiasm and wonder he set to his students and colleagues is a lasting one, and generations of undergraduates had their scientific attitudes and perceptions honed and focused by his influence. Ian enjoyed teaching, both in the formal lecture-theatre context and on field courses, where his staying power was legendary. New was usually among the earliest risers on such trips, and recalls that it was common to find Ian still ‘instructing’ (not necessarily solely on scientific topics!) at around 5 a.m., with his youthful undergraduate audience ever more aware that they were due to start a strenuous day of field work within a couple of hours….

In his early days at La Trobe, Ian was a key instigator in the formation of the School of Biological Sciences. He was a natural leader who inspired loyalty in his staff. His colleagues agreed with his strong belief that, as Professor, he should lead his department, and at the time when most departments at La Trobe were encouraged to elect their head Ian was endorsed as ‘permanent chairman’. He fought hard to defend the concept that a University should be a community of scholars free to pursue their research interests without interference from government. He strongly resented the rise of cohorts of ‘academistrators’ (his term, not entirely complimentary), and on occasion urged academic disobedience to resist externally imposed changes. He wrote formally to the University Council in 1988 under the heading ‘Take up the Mace!’ (a reference to the University’s ceremonial mace carried on formal academic occasions but — as far as we know — never used in anger), ‘asking Council for its support in the defence of my rights and responsibilities as a professor, and in defence of my discipline from outside interference’. Elsewhere, he argued his belief that ‘No-one realizes that universities cannot be run like businesses, because good universities are inherently inefficient operations — decisions are questioned, considered, mulled over, in a collegiate system’. The then recent changes to university priorities in Australia depressed him greatly, not least because he saw the opportunities for young people being eroded as funding and teaching capability declined, to the detriment of Australia’s future. He strongly resented his enforced retirement on grounds of age when he reached 65 and characteristically fought hard against this — even seeking professional advice as to whether he was subsequently able to apply for the job of his replacement, by which time the mandatory age retirement no longer existed! Ian served three periods as Dean of Biological Sciences (1970–1972, 1979–1981, 1985–1987) and was Acting Vice-Chancellor on two occasions. He sat on most of the University’s major boards and committees, where his determination, humour and abilities to think rapidly and laterally about many complex issues were useful counters to the tedium that some such bodies can adopt, and gained him the respect of colleagues throughout the institution. The University recognized his contributions, in conjunction with his scientific stature, by the posthumous award of the DSc degree, honoris causa, coincidentally presented to Ann on the first anniversary of his funeral.

Post-retirement, Ian lectured for many years in Natural Resource Management to Applied Sciences students at the Holmesglen College of TAFE, inspiring several of them to move on to university studies in related fields. In addition to his scientific achievements, Ian had a strong interest and involvement in fostering Australian/Indonesian academic co-operation and educational development. In the years after retirement, he was an academic adviser or guest lecturer at Udayana University (Bali), Mataram University (Lombok) and other Indonesian universities. As an Indonesian colleague recently expressed it, ‘Ian Thornton showed how Australian and Indonesian colleagues could work together’. His death occurred in Bangkok whilst he was returning from Laos, where he was advising the National University on the implementation of basic science courses. He is survived by Ann and three stepchildren, two adopted children from his earlier marriage to Jean, and six grandchildren.

Many zoologists have made notable contributions to different fields within their discipline, but Ian Thornton is memorable for the number of very different fields to which he made highly significant contributions. This stemmed from his immutable belief that a scholar should be allowed to follow his interests, and to his own acumen when interesting opportunities arose. He never took short cuts —a casual query from a student could engage him for several hours. Once he had decided a particular course, if the nature of the progress demanded laborious enterprise or even dangerous fieldwork he would not be deflected. He adhered firmly, and in our opinion correctly, to the ideal that zoogeography, ecology and indeed any aspect of the evolution of a group can be understood fully only after adequate systematic study. The considerable sacrifice and efforts needed to pursue fieldwork to collect Psocoptera in remote areas were simply ‘part of the game’ that he played so ably over much of his academic life.

Acknowledgments

We appreciate greatly the considerable help and advice given by Ann Thornton for this memoir. Longstanding colleagues and friends at La Trobe, particularly Pat Woolley, also generously shared their thoughts and reminiscences with us. The photograph shows Ian Thornton on Anak Krakatau, with the cliff of Rakata in the background.

References

Garcia Aldrete, A.N., Lienhard, C. and Mockford, E.L. (2005). Thorntonia. A Commemorative Volume for Ian W. B. Thornton. Publicaciones Especiales 20. Insituto de Biologia, Universidad Nacional Autonoma de Mexico. 205 pp.

Zimmerman, E.C. (1948). Insects of Hawaii. Vol. 2. University of Hawaii Press, Honolulu.

Publications

Books:

1. Thornton, Ian. (1971). Darwin’s Islands: a Natural History of the Galápagos. Doubleday, Natural History Press, New York. 322 pp.

2. Hill, D.S., Hore, P. and Thornton, I.W.B. (1982). Insects of Hong Kong. Hong Kong University Press, Hong Kong. 503 pp.

3. Thornton, Ian. (1996). Krakatau – The Destruction and Reassembly of an Island Ecosystem. Harvard University Press, Cambridge, Mass., USA. 346 pp.

Chapters of books:

1. Thornton, I.W.B. (1991). ‘Krakatau – studies on the origin and development of a fauna’, in The Unity of Evolutionary Biology (Proceedings of the Fourth International Congress of Systematic and Evolutionary Biology), E.C. Dudley (Ed.), pp. 396–408. (Dioscorides Press, Portland, USA.)

2. Thornton, I.W.B. (1996). ‘The origins and development of island biotas as illustrated by Krakatau’, in The Origin and Evolution of Island Biotas. New Guinea to Eastern Polynesia: Patterns and Processes, A. Keast and S.E. Miller (Eds), pp. 67–90. (SPB Academic Publishing, The Netherlands.)

3. Thornton, I.W.B. (1999). ‘The ecology of volcanoes: recovery and reassembly of living communities’, in Encyclopedia of Volcanoes, H. Sigurdsson (Ed.), 1057–1081. (Academic Press, San Diego, USA.)

Edited volume:

1. Thornton, I.W.B. (Scientific Ed.) (1992). Krakatau: a Century of Change. GeoJournal 28(2), 83–304 (editorial comments pp. 84–86, 129, 173–174, 232, 292).

Research papers in scientific journals:

1. Thornton, I.W.B. and Broadhead, E. (1954). The British species of Elipsocus Hagen (Corrodentia, Mesopsocidae). Journal of the Society for British Entomology 5(2), 47–64.

2. Broadhead, E. and Thornton, I.W.B. (1955). An ecological study of three closely related psocid species. Oikos 6(1), 1–50.

3. Thornton, I.W.B. (1956). Diurnal migrations of the echinoid Diadema setosum (Leske). British Journal of Animal Behaviour 4(4), 143–146.

4. Thornton, I.W.B. (1956). Notes on the biology of Leiurus quinquestriatus (H. & E. 1829) (Scorpiones, Buthidae). British Journal of Animal Behaviour 4(3), 92–93.

5. Thornton, I.W.B. (1957). Faunal succession in umbels of Cyperus papyrus L. on the Upper White Nile. Proceedings of the Royal Entomological Society of London A 32, 119–131.

6. Thornton, I.W.B. (1957). Notes on the ecology of the Acacia bagworm, Auchmophila kordofensis (Lepidoptera, Psychidae), in the environs of Khartoum. Sudan Notes and Records 38, 147–150.

7. Thornton, I.W.B. (1959). A new genus of Philotarsidae (Corrodentia) and new species of this and related families from Hong Kong. Transactions of the Royal Entomological Society of London 111(2), 331–349.

8. Thornton, I.W.B. (1959). New species of Peripsocus Hagen 1866 (Corrodentia, Peripsocidae) from Hong Kong Island, with further descriptions of Peripsocus similis Enderlein (1903) and Peripsocus quercicola Enderlein (1906). Proceedings of the Royal Entomological Society of London B 28, 37–48.

9. Thornton, I.W.B. (1960). New Psocidae and an aberrant new myopsocid (Psocoptera) from Hong Kong. Transactions of the Royal Entomological Society of London 112(10), 239–261.

10. Thornton, I.W.B. (1961). Comments on the geographical distribution of Pseudocaecilius elutus Enderlein (Psocoptera), with descriptions of related new species from Hong Kong. Proceedings of the Royal Entomological Society of London B 30, 141–152.

11. Thornton, I.W.B. (1961). The Trichadenotecnum group (Psocoptera: Psocidae) in Hong Kong, with descriptions of new species. Transactions of the Royal Entomological Society of London 113(1), 1–24.

12. Thornton, I.W.B. (1962). Note on the genitalia of two New Zealand philotarsids (Insecta: Psocoptera). Journal of the New Zealand Institute of Science 5(2), 241–245.

13. Thornton, I.W.B. (1962). Psocids (Psocoptera) from the Batu Caves, Malaya. Pacific Insects 4(2), 441–455.

14. Thornton, I.W.B. (1962). The Peripsocidae (Psocoptera) of Hong Kong. Transactions of the Royal Entomological Society of London 114(9), 285–315.

15. Marshall, A.T. and Thornton, I.W.B. (1963). Micromalthus (Coleoptera: Micromalthidae) in Hong Kong. Pacific Insects 5(4), 715–720.

16. Thornton, I.W.B. (1963). The ecology of closely related species. Proceedings of the 9th Pacific Science Congress, Bangkok, 19–25. (Delivered in 1957.)

17. Thornton, I.W.B. (1964). Airborne Psocoptera trapped on ships and aircraft. Pacific Insects 6(2), 285–291.

18. Thornton, I.W.B. and Harrell, J.C. (1965). Air-borne Psocoptera trapped on ships and aircraft, 2 – Pacific ship trappings 1963–64. Pacific Insects 7(4), 700–702.

19. Thornton, I.W.B. (1965). Distribution patterns of endemic psocids (Psocoptera) in the Hawaiian Islands. Proceedings of the XIIth International Congress of Entomology, London 1964, 442–443.

20. Wong, S.K. and Thornton, I.W.B. (1966). Chromosome numbers of some psocid genera. Nature 211(45), 214–215.

21. Thornton, I.W.B. (1966). Isolation within archipelagos. Proceedings of the 11th Pacific Science Congress, Tokyo 5, 12.

22. Thornton, I.W.B. and Wong S.K. (1966). Some Psocoptera from West Bengal, India. Transactions of the Royal Entomological Society of London 118(1), 1–21.

23. Lee, S.S. and Thornton, I.W.B. (1967). The family Pseudocaeciliidae (Psocoptera) – a reappraisal based on the discovery of new Oriental and Pacific species. Pacific Insects Monographs 16, 1–116.

24. Thornton, I.W.B. (1967). The measurement of isolation on archipelagos, and its relation to insular faunal size and endemism. Evolution 21(4), 842–849.

25. Thornton, I.W.B. (1967). Wing reduction in endemic Hawaiian psocids. Journal of Natural History 1, 149–150.

26. Thornton, I.W.B. and Wong, S.K. (1967). A numerical taxonomic analysis of the Peripsocidae of the Oriental Region and the Pacific Basin. Systematic Zoology 16(3), 217–240.

27. Thornton, I.W.B., Yeung, K.K. and Sankhalz, K.S. (1967). The genetics of the white tigers of Rewa. Journal of Zoology 152, 127–135.

28. Clarke, C.A., Sheppard, P.M. and Thornton, I.W.B. (1968). The genetics of the mimetic butterfly Papilio memnon L. Philosophical Transactions of the Royal Society of London B 254, 37–89.

29. Thornton, I.W.B. and Wong, S.K. (1968). The peripsocid fauna (Psocoptera) of the Oriental Region and the Pacific. Pacific Insects Monographs 19, 1–158.

30. Wong, S.K. and Thornton, I.W.B. (1968). The internal morphology of the reproductive systems of some psocid species. Proceedings of the Royal Entomological Society of London A 43(1–3), 1–12.

31. Chui, W.D. and Thornton, I.W.B. (1972). A numerical taxonomic study of the endemic Ptycta species of the Hawaiian Islands (Psocoptera: Psocidae). Systematic Zoology 21(1), 7–22.

32. Thornton, I.W.B., Lee, S.S. and Chui, W.D. (1972). Psocoptera. Insects of Micronesia 8(4), 45–144.

33. Thornton, I.W.B. and Woo, A.K.T. (1973). Psocoptera of the Galapagos Islands. Pacific Insects 15(1), 1–58.

34. Smithers, C.N. and Thornton I.W.B. (1973). The Psilopsocidae (Psocoptera) of New Guinea. Proceedings of the Linnean Society of New South Wales 98(2), 98–103.

35. Smithers, C.N. and Thornton I.W.B. (1974). The Myopsocidae (Psocoptera) of New Guinea and New Caledonia. Transactions of the Royal Entomological Society of London 126(1), 91–127.

36. Smithers, C.N. and Thornton, I.W.B. (1974). The Psocoptera of Norfolk Island. Records of the Australian Museum 29(8), 209–234.

37. Thornton, I.W.B. and Smithers, C.N. (1974). The Philotarsidae (Psocoptera) of New Caledonia. Pacific Insects 16(2–3), 177–243.

38. New, T.R. and Thornton, I.W.B. (1975). Psocomorpha (Psocoptera) collected on recent expeditions to South America. Journal of Entomology B 44(1), 27–80.

39. Smithers, C.N. and Thornton, I.W.B. (1975). The first record of Stenopsocidae (Psocoptera) from New Guinea with descriptions of new species. Proceedings of the Linnean Society of New South Wales 100(2), 156–166.

40. Smithers, C.N. and Thornton, I.W.B. (1975). The Psocoptera of Lord Howe Island. Records of the Australian Museum 29(16), 453–471.

41. Thornton, I.W.B., Marshall, A.T., Kwan, W.H. and MA, Q. (1975). Studies on lepidopterous pests of rice crops in Hong Kong, with particular reference to the Yellow Stem- borer, Tryporyza incertulas (Walk.). Pest Articles and News Summaries 21, 239–252.

42. New, T.R. and Thornton, I.W.B. (1975). Psocomorpha (Psocoptera) from the Malayan Peninsula, including collections from forest canopy. Oriental Insects 9(4), 375–418.

43. Smithers, C.N. and Thornton, I.W.B. (1977). A new genus and some new species of Epipsocidae (Psocoptera) from the Melanesian Arc. Proceedings of the Linnean Society of New South Wales 102(2), 60–75.

44. Thornton, I.W.B. and New, T.R. (1977). Philotarsidae (Psocoptera) of the Bismarck Archipelago. Pacific Insects 17(4), 451–457.

45. Thornton, I.W.B. and New, T.R. (1977). The Philotarsidae (Psocoptera) of Australia. Australian Journal of Zoology, Supplementary Series 54, 1–62.

46. Thornton, I.W.B. and Smithers, C.N. (1977). Philotarsidae (Psocoptera) of New Guinea. Pacific Insects 17(4), 419–450.

47. Thornton, I.W.B., Wong, S.K. and Smithers, C.N. (1977). The Philotarsidae (Psocoptera) of New Zealand and islands of the New Zealand plateau. Pacific Insects 17(2–3), 197–228.

48. Thornton, I.W.B. (1978). White tiger genetics – further evidence. Journal of Zoology, London 185, 389–394. (Review in Science Report, The Times, 1.ix.78, p.14.)

49. Thornton, I.W.B. and Lyall, I. (1978). Psocoptera from Chilean Nothofagus. Pacific Insects 19(1–2), 1–16.

50. Thornton, I.W.B. and Smithers, C.N. (1978). Philotarsidae (Psocoptera) of the Solomon Archipelago. Pacific Insects 18(3–4), 227–233.

51. Smithers, C.N. and Thornton, I.W.B. (1979). Psilopsocidae and Myopsocidae (Psocoptera) of the Bismarck, Solomon and New Hebrides archipelagos. Records of the Australian Museum 32(16), 513–545.

52. Thornton, I.W.B. (1980). Plate tectonics and the distribution of the insect family Philotarsidae (Order Psocoptera) in the Southwest Pacific. Palaeogeography, Palaeoclimatology, Palaeoecology 31, 251–266.

53. New, T.R. and Thornton, I.W.B. (1981). Psocoptera from central and southern Chile. Pacific Insects Monographs 37, 136–178.

54. Thornton, I.W.B. (1981). Psocoptera of the Fiji Islands. Pacific Insects Monographs 37, 1–105.

55. Thornton, I.W.B. (1981). Psocoptera of the Tongan Archipelago. Pacific Insects Monographs 37, 106–135.

56. Thornton, I.W.B. and New, T.R. (1981). Psocoptera from Robinson Crusoe Island, Juan Fernandez Archipelago. Pacific Insects Monographs 37, 179–191.

57. Thornton, I.W.B. (1981). Psocoptera of the Hawaiian Islands. Parts I and II. Introduction and the nonendemic fauna. Pacific Insects 23(1–2), 1–49.

58. Thornton, I.W.B. (1981). The systematics, phylogeny and biogeography of the psocopteran family Philotarsidae. Systematic Entomology 6(4), 413–452.

59. Smithers, C.N. and Thornton, I.W.B. (1981). The Psocidae (Insecta: Psocoptera) of New Guinea, including a new coleopteriform genus from high on Mt Wilhelm. Australian Journal of Zoology 29(6), 921–969.

60. Smithers, C.N. and Thornton, I.W.B. (1981). ‘The role of New Guinea in the evolution and biogeography of some families of psocopteran insects’, in J.L. Gressitt (Ed.) Biogeography and Ecology of New Guinea Vol. 2, 621–638 (W. Junk, Publishers, The Hague).

61. Thornton, I.W.B. (Scientific Ed.). (1983). Proceedings of Symposium on Biogeography and Plate Tectonics in the SW Pacific, 15th Pacific Science Congress, Dunedin, (1983). GeoJournal 7(6), 479–564.

62. Thornton, I.W.B. (1983). ‘Vicariance and dispersal: confrontation or compatibility?’, in I.W.B. Thornton (Ed.) Symposium on Biogeography and Plate Tectonics in the Pacific. GeoJournal 7(6), 557–564.

63. Thornton, I.W.B. (1984). Psocoptera of the Hawaiian Islands. Part III. The endemic Ptycta complex of species: systematics, distribution and possible phylogeny. International Journal of Entomology 26(1–2), 1–128.

64. Thornton, I.W.B. (1984). Review of ‘Taxonomy, phylogeny and biogeography of the genus Cosmopsaltria, with remarks on the historic biogeography of the subtribe Cosmopsaltriaria (Homoptera: Cicadidae)’ by J.P. Duffels. International Journal of Entomology 26(1–2), 171–173.

65. Thornton, I.W.B. and Smithers, C.N. (1984). The systematics of the Calopsocidae, an Oriental and Melanesian family of Psocoptera. Systematic Entomology 9(2), 183–244.

66. Thornton, I.W.B. (1984). Krakatau – the development and repair of a tropical ecosystem. Ambio 13(4), 216–225.

67. Thornton, I.W.B. (1984). Psocoptera and Wallace’s Line: collections from the islands of Bali and Lombok. Treubia 29(2), 83–177.

68. Thornton, I.W.B. (1984). An unusual psocopteran from New Guinea, and its relationships within the Philotarsidae. International Journal of Entomology 26(4), 378–385.

69. Thornton, I.W.B. (1985). The geographical and ecological distribution of arboreal Psocoptera. Annual Review of Entomology 30, 175–196.

70. Thornton, I.W.B. (1985). A preliminary survey of the psocopteran fauna of the Krakatau Islands. Proceedings of the Symposium on 100 years development of Krakatau and surroundings. Jakarta, L.I.P.I. 466–470.

71. Thornton, I.W.B., Zann, R.A., Rawlinson, P.A., Tidemann, C.R., Adikerana, A.S. and Widjoya, A.H.T. (1988). Colonization of the Krakatau Islands by vertebrates: equilibrium, succession and possible delayed extinction. Proceedings of the National Academy of Science of the USA (Ecology) 85, 515–518.

72. Thornton, I.W.B. and Rosengren, N.J. (1988). Zoological Expeditions to the Krakatau islands, 1984–1985: General Introduction. Philosophical Transactions of the Royal Society of London B 322, 273–316.

73. Graves, S.R., Plummer, D.C., Hives, N., Harvey, K.J. and Thornton I.W.B. (1988). Antibiotic resistance patterns of soil bacteria (Gram-negative rods) from the Krakatau Is (Rakata) and W. Java. Philosophical Transactions of the Royal Society of London B 322, 317–326.

74. Graves, S.R., Rosengren, N.J., Kennelly- Merrit, S.A., Harvey, K.J. and Thornton, I.W.B. (1988). Antibiotic resistance patterns and relative concentrations of bacteria (Gram-negative rods) from ash deposits of various ages on the Krakatau Is. Philosophical Transactions of the Royal Society of London B 322, 327–338.

75. Graves, S.R., Kennelly-Merrit, S.A., Tidemann, C.R., Rawlinson, P.A., Harvey, K.J. and Thornton, I.W.B. (1988). Antibiotic- resistance patterns of enteric bacteria of wild mammals on the Krakatau Is and W. Java. Philosophical Transactions of the Royal Society of London B 322, 339–354.

76. Graves, S.R., Rawlinson, P.A., Kennelly- Merrit, S.A., McLaren, D.A., Harvey, K.J. and Thornton, I.W.B. (1988). Enteric bacteria of reptiles on Java and the Krakatau Islands. Philosophical Transactions of the Royal Society of London B 322, 355–362.

77. Thornton, I.W.B. and Graves, S.R. (1988). Colonization of the Krakataus by bacteria and the development of antibiotic resistance. Philosophical Transactions of the Royal Society of London B 322, 363–368.

78. Thornton, I.W.B., New, T.R. and Vaughan, P.J. (1988). Colonization of the Krakatau Islands by Psocoptera. Philosophical Transactions of the Royal Society of London B 322, 427–444.

79. New, T.R., Bush, M., Thornton, I.W.B. and Sudarman, H.K. (1988). The butterfly fauna of the Krakatau Islands after a century of colonization. Philosophical Transactions of the Royal Society of London B 322, 445–458.

80. Compton, S.G., Thornton, I.W.B., New, T.R., and Underhill, L. (1988). The colonization of the Krakatau Islands by fig wasps and other chalcids (Hymenoptera, Chalcidoidea). Philosophical Transactions of the Royal Society of London B 322, 459–470.

81. Thornton, I.W.B., New, T.R., McLaren, D.A., Sudarman, H.K. and Vaughan, P.J. (1988). Airborne arthropod fall-out on Anak Krakatau and a possible pre-vegetation pioneer community. Philosophical Transactions of the Royal Society of London B 322, 471–480.

82. New, T.R. and Thornton, I.W.B. (1988). A pre-vegetation population of crickets subsisting on allochthonous aeolian debris on Anak Krakatau. Philosophical Transactions of the Royal Society of London B 322, 481–486.

83. Thornton, I.W.B. and New, T.R. (1988). Freshwater communities of the Krakatau islands. Philosophical Transactions of the Royal Society of London B 322, 487–492.

84. Thornton, I.W.B. and New, T.R. (1988). Krakatau invertebrates: the 1980s fauna in the context of a century of colonization. Philosophical Transactions of the Royal Society of London B 322, 493–522.

85. New, T.R. and Thornton, I.W.B. (1988). Epipsocetae (Psocoptera) from Peru. Studies on the Neotropical Fauna and Environment 23(4), 225–250.

86. Cole, P.J., New, T.R. and Thornton, I.W.B. (1989). Psocoptera of Flinders, King and Deal Islands, Bass Strait. Journal of the Australian Entomological Society 28, 31–38.

87. Vaughan, P.J., Thornton, I.W.B. and New, T.R. (1989). The Psocoptera of the Krakatau Islands, Indonesia. Treubia 30(1), 1–93.

88. Thornton, I.W.B. (1990). Psocoptera (Insecta) of the island of Moorea, French Polynesia, and comparisons with other Pacific island faunas. Bulletin du Muséum National d’Histoire Naturelle, Paris 4 ser., 11, A(4), 783–828.

89. Endersby, N.M., New, T.R. and Thornton, I.W.B. (1990). Psocoptera from the Grampians and Mt. Arapiles, Western Victoria – a biogeographic analysis. Journal of the Australian Entomological Society 29, 215–224.

90. Thornton, I.W.B., Zann, R.A. and Stephenson, D.G. (1990). Colonisation of the Krakatau islands by land birds and the approach to an equilibrium number of species. Philosophical Transactions of the Royal Society of London B 328, 55–93.

91. Tidemann, C.R., Kitchener, D.J., Zann, R.A. and Thornton, I.W.B. (1990). Recolonization of the Krakatau Islands and adjacent areas of West Java, Indonesia, by bats (Chiroptera) 1883–1986. Philosophical Transactions of the Royal Society of London B 328, 121–130.

92. Thornton, I.W.B., New, T.R., Zann, R.A. and Rawlinson, P.A. (1990). Colonization of the Krakatau Islands by animals: a perspective from the 1980s. Philosophical Transactions of the Royal Society of London B 328, 131–165.

93. Thornton, I.W.B. (1990). Psocoptera of the Hawaiian Islands. Part IV. The endemic genus Palistreptus (Elipsocidae): systematics, distribution and evolution. Bishop Museum Bulletin of Entomology 4, 1–57.

94. Smithers, C.N. and Thornton, I.W.B. (1990). Systematics and distribution of the Melanesian Psocidae (Psocoptera). Invertebrate Taxonomy 3, 431–468.

95. Vaughan, P.J., Thornton, I.W.B. and New, T.R. (1991). Psocoptera from Southern Sumatra and West Java, Indonesia: source faunas for colonization of the Krakatau Islands. Treubia 30(2), 103–164.

96. Thornton, I.W.B. (1991). Replacement name for Aaroniella badonelli Thornton (Psocoptera, Philotarsidae). Bulletin du Muséum National d’Histoire Naturelle, Paris 4 ser.13, A (3–4), 483.

97. New, T.R. and Thornton, I.W.B. (1992). The butterflies (Insecta, Lepidoptera) of Anak Krakatau, Indonesia: faunal development in early succession. Journal of the Lepidopterists’ Society 46(2), 83–96.

98. Thornton, I.W.B. and Browning, J.A. (1992). Myopsocidae (Insecta: Psocoptera) from Java, including a discussion of the known Indonesian species. Revue Suisse de Zoologie 99(2), 343–367.

99. New, T.R. and Thornton, I.W.B. (1992). Colonization of the Krakatau Islands by invertebrates. GeoJournal 28(2), 219–224.

100. Rawlinson, P.A., Zann, R.A., Van Balen, S. and Thornton, I.W.B. (1992). Colonization of the Krakatau Islands by vertebrates. GeoJournal 28(2), 225–231.

101. Thornton, I.W.B. and Walsh, D. (1992). Photographic evidence of rate of development of plant cover on the emergent island Anak Krakatau from 1971 to 1991 and implications for the effect of volcanism. GeoJournal 28(2), 249–259.

102. Thornton, I.W.B., Ward, S.A., Zann, R.A. and New, T.R. (1992). Anak Krakatau – a colonization model within a colonization model? GeoJournal 28(2), 271–286.

103. Thornton, I.W.B. (1992). K.W. Dammerman – fore-runner of island equilibrium theory? Global Ecology and Biogeography Letters 2, 145–148.

104. Endang, S.K. and Thornton, I.W.B. (1992). Psocidae (Insecta: Psocoptera) from the islands of Bali and Lombok, Indonesia. Treubia 30(3), 319–379.

105. Thornton, I.W.B., Ward, S.A., Zann, R.A. and New, T.R. (1993). The Anak Krakatau Question. GeoJournal 29(4), 421–425.

106. Thornton, I.W.B., Zann, R.A. and Van Balen, S. (1993). Colonization of Rakata (Krakatau Is) by non-migrant land birds from 1883–1992 and implications for the value of island equilibrium theory. Journal of Biogeography 20, 441–452.

107. Maeto, K. and Thornton, I.W.B. (1993). A preliminary appraisal of the braconid (Hymenoptera) fauna of the Krakatau Islands (Indonesia) in 1984–1986, with comments on the colonizing abilities of parasitic modes. Japanese Journal of Entomology 61(4), 787–801.

108. Schmidt, E.R. and Thornton, I.W.B. (1993). The Psocoptera (Insecta) of Wilsons Promontory National Park, Victoria, Australia. Memoirs of the Museum of Victoria 53(2) (1992), 137–220.

109. Schmidt, E.R., Thornton, I.W.B. and Hancock, D. (1994). Tropical fruitflies (Diptera: Tephritidae) of the Krakatau Archipelago in 1990 and comments on faunistic changes since 1982. Ecological Research 9, 317–324.

110. Compton, S.G., Ross, S.J. and Thornton, I.W.B. (1994). Pollinator limitation of fig tree reproduction on the island of Anak Krakatau (Indonesia). Biotropica 26(2), 180–186.

111. Thornton, I.W.B., Ward, S.A., Zann, R.A. and New, T.R. (1994). Further comments on the Anak Krakatau Question. GeoJournal 33(4), 493.

112. Thornton, I.W.B. (1994). Figs, frugivores and falcons: an aspect of the assembly of mixed tropical forest on the emergent volcanic island, Anak Krakatau. South Australian Geographical Journal 93, 3–21. (Based in part on the Brock Memorial Lecture, given to the Royal Geographical Society of Australasia [South Australia Branch] in August 1993.)

113. Wang, Q., Thornton, I.W.B. and New, T.R. (1994). Systematics and biogeography of the Australian-New Guinean genus Thoris Pascoe (Coleoptera: Cerambycidae: Phoracanthini). Invertebrate Taxonomy 8, 839–860.

114. Thornton, I.W.B., Partomihardjo, T. and Yukawa, J. (1994). Observations on the effects, up to July 1993, of the current eruptive episode of Anak Krakatau. Global Ecology and Biogeography Letters 4, 88–94.

115. Schedvin, N.S., Cook, S. and Thornton, I.W.B. (1994). The diversity of bats on the Krakatau Islands in the early 1990s. Biodiversity Letters 2, 87–92.

116. Wang, Q., New, T.R. and Thornton, I.W.B. (1995). Phylogeny and distribution of the phoracanthine genus Atesta (Coleoptera: Cerambycidae) from Australia. Systematic Entomology 20, 229–238.

117. Wang, Q., Thornton, I.W.B. and New, T.R. (1996). Biogeography of the phoracanthine beetles (Coleoptera: Cerambycidae). Journal of Biogeography 23, 75–94.

118. Thornton, I.W.B., Compton, S.G. and Wilson, C.N. (1996). The role of animals in the colonization of the Krakatau Islands by Ficus species. Journal of Biogeography 23, 577–592.

119. Ward, S.A. and Thornton, I.W.B. (1999). Guest Editorial. Equilibrium theory and alternative stable equilibria. Journal of Biogeography 25, 615–622.

120. Runciman, D., Cook, S., Riley, J., Wardill, J. and Thornton, I.W.B. (1999). The avifauna of Sebesi, a possible stepping-stone to the Krakatau Islands. Tropical Biodiversity 5(2), 1–9.

121. Wang, Q., Thornton, I.W.B. and New, T.R. (1999). A cladistic analysis of the phoracanthine genus Phoracantha Newman (Coleoptera: Cerambycidae: Cerambycinae), with discussion of biogeographic distribution and pest status. Annals of the Entomological Society of America 92(5), 631–638.

122. Smithers, C.N., Peters, J.V. and Thornton, I.W.B. (2000). The Psocoptera (Insecta) of Norfolk and Philip Islands: occurrence, status and zoogeography. Proceedings of the Linnean Society of New South Wales 121, 101–111.

123. Ward, S.A. and Thornton, I.W.B. (2000). Chance and determinism in the development of isolated communities. Global Ecology and Biogeography 9, 7–18.

124. Thornton, I.W.B., Mawdsley, N.A. and Partomihardjo, T. (2000). Persistence of biota on Anak Krakatau after a three-year period of volcanic activity. Tropical Biodiversity 7, 25–43.

125. Thornton, I.W.B. (2001). Colonization of an island volcano, Long Island, Papua New Guinea, and an emergent island, Motmot, in its caldera lake. I. General introduction. Journal of Biogeography 28, 1299–1310.

126. Harrison, R.D., Banka, R., Thornton, I.W.B., Shanahan, M. and Yamuna, R. (2001). Colonization of an island volcano, Long Island, Papua New Guinea, and an emergent island, Motmot, in its caldera lake. II. The vascular flora. Journal of Biogeography 28, 1311–1338.

127. Schipper, C., Shanahan, M., Cook, S, and Thornton, I.W.B. (2001). Colonization of an island volcano, Long Island, Papua New Guinea, and an emergent island, Motmot, in its caldera lake. III. Colonization by birds. Journal of Biogeography 28, 1339–1352.

128. Cook, S., Singadan, R. and Thornton, I.W.B. (2001). Colonization of an island volcano, Long Island, Papua New Guinea, and an emergent island, Motmot, in its caldera lake. IV. Colonization by non-avian vertebrates. Journal of Biogeography 28, 1353–1364.

129. Shanahan, M., Harrison, R.D., Yamuna, R.Y., Boen, W. and Thornton, I.W.B. (2001). Colonization of an island volcano, Long Island, Papua New Guinea, and an emergent island, Motmot, in its caldera lake. V. Colonization by figs (Ficus spp.), their dispersers and pollinators. Journal of Biogeography 28, 1365–1378.

130. Edwards, J.S. and Thornton, I.W.B. (2001). Colonization of an island volcano, Long Island, Papua New Guinea, and an emergent island, Motmot, in its caldera lake. VI. The pioneer arthropod community of Motmot. Journal of Biogeography 28, 1379–1388.

131. Thornton, I.W.B., Cook, S., Edwards, J.S., Harrison, R.D., Schipper, C., Shanahan, M, Singadan, R. and Yamuna, R. (2001). Colonization of an island volcano, Long Island, Papua New Guinea, and an emergent island, Motmot, in its caldera lake. VII. Overview and discussion. Journal of Biogeography 28, 1389–1408.

132. Endang, S.K., New, T.R. and Thornton, I.W.B. (2002). The Psocidae (Psocoptera) of Java and the eastern islands of Indonesia. Invertebrate Systematics 16, 200–276.

133. Thornton, I.W.B., Runciman, D., Cook, S., Lumsden, L., Partomihardjo, T., Schedvin, N., Yukawa, J. and Ward, S.A. (2002). How important were stepping stones in the colonisation of Krakatau? Biological Journal of the Linnean Society 77, 275–317.

Miscellaneous publications:

1. Thornton, I.W.B. (1983). J. Linsey Gressitt (1914)–(1982). GeoJournal 7(6), 481–482.

2. Thornton, I.W.B. (Ed.) (1985). 1984 Zoological Expedition to the Krakataus. Preliminary Report. La Trobe University Department of Zoology. Miscellaneous Series No. 1, 57 pp.

3. Thornton, I.W.B. (Ed.) (1986). 1985 Zoological Expedition to the Krakataus. Preliminary Report. La Trobe University Department of Zoology. Miscellaneous Series No. 2, 63 pp.

4. Thornton, I.W.B. (1986). Krakatau Rebirth. Australian Geographic 1(2), 40–54.

5. Thornton, I.W.B. (1987). A Guide to Krakatau. Sponsored by the Krakatau Foundation and P.H.P.A. (Forest Protection and Nature Conservation, Indonesia). 21 pp.

6. Thornton, I.W.B. (Ed.) (1987). 1986 Zoological Expedition to the Krakataus. Preliminary Report. La Trobe University Department of Zoology. Miscellaneous Series No. 3, 59 pp.

7. Thornton, I.W.B. (1989). The recolonisation of Krakatoa by animals. Pacific Science Association Information Bulletin 41(3), 13–23.

8. Thornton, I.W.B. (1990). Message from Melbourne. Bio News 28, 4–6.

T. R. New, Department of Zoology, La Trobe University, Victoria.
C. N. Smithers, The Australian Museum, Sydney.

A. T. Marshall, Department of Zoology, La Trobe University, Victoria.

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