OBITUARY

Ross Ewen Beever

MSc Auckland, PhD Leeds, FRSNZ, FNZIAHS, FAPPS 1946–2010

ROSS BEEVER was an exceptional natural scientist, and both mentor and friend to his many colleagues. He was New Zealand's most distinguished mycologist and plant pathologist. His main scientific contribution was in the study of fungi, their taxonomy, how they grow, how they interact with plants and how plant disease fungi causing plant disease may be controlled. He produced new and innovative work as well as combining previous work into new models of fungal biology. He also carried out significant studies on the biodiversity of both fungi and higher plants.

Ross was no tree-hugger; instead, he wanted to understand fungi and plants, he examined them critically in the field, he studied them in the laboratory, he analysed his results and he published his findings because he was keen that knowledge be used. He had great breadth of vision and was an insightful and objective observer, with research interests ranging from taxonomy through physiology and biochemistry to genetics and molecular biology. This is shown by his extensive list of publications (see list at the end of this obituary). The unusual breadth of his vision proved to be a major strength rather than a dilution of his individual efforts, as combined with this coherent vision was a depth of analysis that enabled him to make outstanding contributions to our knowledge. He did not avoid the difficult questions. His work was of basic importance to mycological and botanical science and to conservation, but he was also aware of commercial implications. He therefore undertook quality science relevant to the needs of New Zealand and he often chose deliberately to work on subjects that he knew were ultimately of economic importance. To do this, he drew on his solid background in chemistry, biochemistry and genetics.

Ross was born at Te Kuiti on 3 January 1946. His mother's family, the Brunskills, had a farm at Glorit, close to the Kaipara Harbour, and adjoining the native bush on Mt Auckland (Atuanui). Ross often visited the farm and he was always at ease in the country, an asset when so many of his scientific projects were in rural settings. His father, Jim Beever, taught in country schools at Piopio and Maungaturoto before shifting to Auckland as a foundation teacher and later Deputy Principal at Tamaki College (1957–1969), close by Landcare Research at Tamaki where Ross most recently worked. Jim Beever was an enthusiastic and knowledgeable amateur botanist, best known for his book A Dictionary of Maori Plant Names, and he instilled in Ross the love of our native flora that he was to retain for the rest of his life.

Ross was educated at Maungaturoto District High School, Remuera Intermediate and Auckland Grammar. He was largely self-taught in biology in his last years at school and it was therefore a special achievement that he was awarded in 1963 the Sinclair Scholarship for the top scholar nationally in biology. The same year he also won a University Junior Scholarship. He continued to excel at Auckland University, winning the Annual Prize in Botany, the Lancaster Memorial Prize, the Frances Briggs Junior and Senior Bursaries and in 1966 was a Senior Scholar in Botany and Chemistry. He then shifted to Mt Albert where he completed an MSc (1st class honours) under the supervision of Dr Ted Bollard FRSNZ, co-supervised by Professor Frank Newhook. His thesis topic was ‘Growth of fungi on potato extract’. ‘The Barn’, Bollard's laboratory at the Department of Scientific and Industrial Research (DSIR), was small and crowded with rather makeshift accommodation but it has proved to be extraordinarily influential in New Zealand biological science, producing six Fellows of the Royal Society of New Zealand and another who, on moving to Canberra, became a Fellow of the Australian Academy and an Honorary Fellow of the Royal Society of New Zealand. In 1968, Ross was appointed to the staff of Plant Diseases Division, DSIR and he remained with this division and its successor organizations, Plant Protection Division, DSIR and then Landcare Research, for the rest of his life.

In 1969 he was awarded a Sir Walter Mulholland Fellowship and he undertook a PhD at Leeds University, UK under Professor JRS Fincham FRS, his thesis title being ‘Genetic and biochemical studies of PEP carboxykinase in Neurospora crassa’. Ross did not travel alone to Leeds but was accompanied by his wife, Jessica, who did a PhD in plant physiology. Ross and Jessica (Spragg) had first met at Remuera Intermediate. They met again at university when both were doing botany and were keen members of the Auckland University Field Club. Both then did masterates based at the DSIR, Mt Albert. They married in 1969 and last year they celebrated 40 years of marriage and friendship together. They had two children, Rosemary and Graham, and in 2009 Graham and Melissa produced the first grandchild, Isabella. Jessica is an accomplished bryologist, working on New Zealand's moss flora, and has been co-located for many years with Ross at Landcare Research, first at Mt Albert and more recently at Tamaki, Auckland.

Ross was to become primarily a plant pathologist and mycologist but he also did innovative work on fungal physiology. His work with Don Burns in the 1970s on phosphate uptake by the fungus Neurospora as a model system was a novel and significant contribution to our understanding of fungal nutrition with implications for both our native plant communities and for pastoral and horticultural cropping and forestry. Many soils in New Zealand are low in phosphorus and plants rely on fungal symbionts (mycorrhizas) for its uptake. The work established for the first time how plants achieve a constant phosphorus uptake (homeostasis) in the face of a 1000-fold concentration range in the external environment, through relatively small modulations of two coexisting transport systems. These studies in phosphorus uptake culminated in a major review.

Fungi have to respond to stress. Ross demonstrated for the first time that polyols are used by filamentous fungi as compatible solutes, increasing in concentration internally in response to external osmotic stress. This work later linked to a major project Ross developed on the dicarboximide class of fungicides that target the osmo-sensing histidine kinase signalling pathway.

Another contribution of particular note to fungal physiology was his study with Graeme Dempsey of hydrophobins, unique proteins that polymerize into microscopic rodlets on hyphal and spore surfaces, making them water-repellent. Such rodlets play diverse but essential roles in hyphal growth, spore dispersal and the interactions between fungi and plants and animals. The importance and novelty of this work was demonstrated by the appearance of an electron micrograph of the rodlets on the cover of Nature. Subsequently, Ross was part of a team with Matt Templeton that established the structure of the hydrophobin from Neurospora.

Botrytis cinerea is one of the world's most destructive fungi. Fungicide resistance poses a particular problem in controlling this disease of grapes, kiwifruit, berryfruit and many ornamentals. Ross, together with a number of colleagues, made a long-term study of fungicide resistance from the cellular or genetic level to studies of epidemiology and field behaviour. This work focused in particular on resistance to the dicarboximide fungicides first released about 30 years ago for Botrytis control. He and his colleagues surveyed the occurrence of fungicide resistance in New Zealand and developed a model to account for competition between resistant and sensitive strains in the field. He emphasized the balance between the fitness gain associated with the acquisition of resistance and the fitness loss associated with the genetic changes leading to resistance. As well as discovering and characterizing the gene associated with dicarboximide resistance (the osmo-sensing histidine kinase), a major paper demonstrated how, at the genetic level, fungal strains in the field continued to evolve in the face of continuing fungicide usage. This applied work was combined with more fundamental studies on the basis of fungicide resistance, showing that resistance to dicarboximides is due to a change in the amino acid sequence of histidine kinase, the first enzyme in a major osmo-sensing signalling pathway in fungi. This established the biochemical basis for fungal resistance to dicarboximides. Other work on the genetics of Botrytis showed that populations of the fungus in the wild consist of a series of ‘vegetative compatibility groups’, a most significant finding which has important implications for the development of biocontrol agents. All these studies underpinned practical management advice to the New Zealand agrichemical industry and to growers on managing fungicide resistance in general.

Chemical control of fungi has obvious limitations. The potential of low-pathogenic strains of Botrytis as novel biocontrol agents was therefore investigated. Another programme initiated with Robyn Howitt and the plant virologists Richard Forster and Mike Pearson explored the potential of mycoviruses as biocontrol agents. Two remarkable viruses were discovered in Botrytis cinerea and characterized. These have subsequently been designated as type species for two new genera within the family Flexivirideae.

The pathology of plants native to New Zealand has traditionally received little attention. This situation changed in the late 1980s when the New Zealand cabbage tree, Co rdyline australis, started dying in large numbers. Ross had long had an interest in the botany of the cabbage tree and this, combined with his knowledge of plant diseases, made him the ideal person to lead a team to investigate the ‘Sudden Decline’ of this national icon. There was much public speculation, not always well-informed, as to the possible causes and that the pathological studies were continued to a successful conclusion a decade later was in large part due to Ross's quiet determination. The real cause was found to be a phytoplasma, ‘Candidatus Phytoplasma australiense’. Phytoplasmas are poorly known bacteria that are very difficult to study because they cannot be grown in culture. The work was made even more difficult by the rapid death of the host and the relatively low titre of the pathogen in most parts of affected plants, except for some apices of underground rhizomes. This particular phytoplasma is also responsible for Phormium yellows disease (the disease that was partly responsible for flax growing in New Zealand becoming non-commercial), and diseases of strawberries and grapes. There are therefore serious implications for some commercial crops. Coprosma species, particularly C. robusta, act as reservoirs of the pathogen and native plant hoppers Zeoliarus (Oliarus) spp. are perhaps major vectors of the phytoplasma. Sudden Decline of cabbage trees was a very difficult and contentious problem and the resolution brought great satisfaction to those responsible.

The successful elucidation of the cause of cabbage tree Sudden Decline was also due to teamwork. This was typical of Ross, his ability to develop productive collaboration within his discipline and across disciplines both within Landcare Research, and before that, within DSIR, and also across New Zealand institutional and international boundaries, bringing together specialists of different skills. This collaboration is demonstrated by the many co-authors with whom he published.

Another national icon that is currently being threatened by disease is the kauri, Agathis australis. Ahead of others, Ross recognized the major biosecurity importance of the fungus-like oomycete genus Phytophthora and its potential risk to our native ecosystems. He helped to clarify the destructive role of P. cinnamomi in native forests and restoration plantings. He led a MAF-funded project clarifying the species of Phytophthora in New Zealand using DNA sequencing techniques and this revealed the presence in the country of P. kernoviae, a potential biosecurity risk. More recently he recognized that species of Phytophthora, especially the as-yet undescribed species that he named Phytophthora taxon Agathis (PTA), were involved in death of kauri from seedlings to mature trees. Right to the end, he was leading a team of researchers from Landcare Research, Scion, Plant & Food Research, and Auckland Regional Council studying this major threat to kauri, and engaging closely with MAF, DOC and iwi. He also co-organized an international conference for over 100 researchers on Phytophthora in forests and natural ecosystems, held in New Zealand in March 2010 and highlighting the seriousness of the kauri dieback disease.

The biodiversity of fungi has received comparatively little attention compared with other components of New Zealand's biota. Ross was particularly interested in the truffle-like fungi that form their fruiting bodies underground or hidden in litter. In conjunction with Mike Castellano (Oregon), Ross described nine new species in the genera Gallacea (G. dingleyae, G. eburnea), Hysterangium (H. neotunicatum, H. rugisporum, H. rupticutis, H. youngii), and Protubera (P. hautuensis, P. nothofagi, P. parvispora). Ross developed the hypothesis that in New Zealand, unlike the rest of the world, truffle-like fungi have evolved to be dispersed by (mainly extinct) ground-feeding birds. A particular interest was the mycorrhizal puffball Pisolithus, restricted in New Zealand to thermal areas. Completion of a taxonomic monograph of New Zealand's truffle-like fungi was a planned undertaking to follow intended partial retirement. He also described with Peter Johnston the new species Elsinoe dracophylli; and with UK mycologists Paul Bridge and Brian Spooner and Landcare Research's Duckchul Park clarified the taxonomy of the mushroom genus Leratiomyces along with two new combinations L. erythrocephalus and L. cucullatus.

Ross was a strong advocate for others to learn about fungi, and readily shared his knowledge and expertise with all both personally and through publications. In 1986, he co-led establishment of the annual New Zealand Fungal Foray, a national event open to amateurs and professionals that is now in its 24th year. In addition to scientific papers about fungi, Ross also prepared chapters on fungi in popular books about the natural history of Rangitoto Island, Chatham Islands and the Waitakere Ranges. He also co-authored a richly illustrated popular guide to common macrofungi of New Zealand. His many identified fungal collections and cultures were meticulously annotated and curated for deposition in the Landcare Research collections – New Zealand Fungal and Plant Disease Herbarium (PDD) and International Collection of Micro-organisms from Plants (ICMP).

Although Ross was primarily a mycologist and plant pathologist, he also made a number of important botanical contributions. His first botanical discovery of note came at the age of 16 when he re-found and recognized the ‘Waipoua orchid’, previously known from a single, rather tattered specimen. This parasitic orchid, associated mainly with taraire trees, was subsequently named Danhatchia australis. Ross continued to investigate many botanical questions and published numerous notes, especially in the Auckland Botanical Society Journal and in Tane, the publication of the Auckland University Field Club. He studied the variants such as the large-leafed forms on offshore islands, he unravelled the self-incompatibility breeding systems of cabbage trees and, in collaboration with Warwick Harris, demonstrated that cabbage trees are adapted to their local environment, thereby providing a scientific basis for the policy of eco-sourcing. Using physiological manipulation, he obtained viable seed of the world's then rarest plant, a single female of Pennantia baylisiana, thus enabling hundreds of offspring to be grown and hence ensuring its survival. He was a very active member of the Auckland Botanical Society and served as its very effective president from 1983 to 1987.

He was also active in the Offshore Islands Research Group that developed among former members of the Auckland University Field Club, and co-organized a symposium on the northern offshore islands of New Zealand in 1983, co-editing the proceedings in 1986. Recalling his association with Atuanui–Mt Auckland as a child, Ross participated in the Atuanui Restoration Project. His selection of plants for landscaping the grounds surrounding Landcare Research at Tamaki, Auckland, is one of his many tangible legacies. Botany was a hobby as well as a profession.

During his career, Ross received many awards that recognized the excellence of his research. He received DSIR study awards in 1972 and 1980, Trimble Fellowships in 1980 and 1983, and USNZ Cooperative Science Program Awards in 1989 and 1994. He was Principal Investigator on one Marsden Award and Associate Principal Investigator on two others. He was invited speaker at numerous conferences both in New Zealand and overseas. He was the Tennant Lecturer at the University of Otago in 2000. He received the Science and Technology Bronze Medal in 2001 ‘for significant contributions to mycology and plant pathology’. He was elected a Fellow of the Royal Society of New Zealand in 2004, a Fellow of the New Zealand Institute of Agricultural and Horticultural Science in 2005 and a Fellow of the Australasian Plant Pathology Society in 2009. In 2007 he was, like his father before him, elected an Honorary Member of the Auckland Botanical Society in recognition of all he had done for the Society.

We individually knew Ross for more than 40 and 32 years, respectively and shared a laboratory with him for many years. We quickly learnt to value him as a scientist, as a colleague, as a mentor and as a friend. He was committed to his research, both for its own sake and for the good of the community. He was quietly determined – even stubborn – and at times this was very necessary, especially in the discussions on Sudden Decline of cabbage trees. However, he never allowed disagreements to become personalized. At worst, he would become very quiet or mutter, fortunately in private, ‘lightweight’ – a term he usually restricted to selected managers or administrators. He did himself take on administrative duties including team leadership, laboratory directorship, and was a strong and successful advocate for science-based decision-making in management issues. Despite his many management responsibilities and his frequent involvement in strategizing and applying for research funds, his productivity remained remarkably constant throughout his scientific career. His insistence on quality and his willingness to collaborate made him an excellent mentor of younger scientists and research technicians.

Ross died prematurely at 64 when he was still actively initiating new research ideas, and had much work left to finish and manu scripts to write. A scholarship will be established by Landcare Research in his honour. It is natural to regret the things left undone and the things that might yet have been achieved. Instead we prefer to remember his many remarkable achievements and what he brought to our lives. His career and his personality are well summarized in an email we received from an old friend and mycological colleague, Gary Samuels: ‘Ross was among the nicest and smartest people I ever knew. He was an amazing empirical scientist and naturalist who represented the best of New Zealand.’

Ross Beever—Publication List

(not including most unpublished reports or abstracts)

Acknowledgements

We thank Dr Jessica Beever for assistance and for allowing ARF to read autobiographic notes prepared by Ross. We also thank Mark Andersen, Mike Pearson, Erik Rikkerink and Matt Templeton for helpful comments.

Notes

Based on an obituary available from The Royal Society of New Zealand: http://www.royalsociety.org.nz/organisation/academy/fellowship/obituaries/ross-ewen-beever