Dr Ross E Beever

Dr Ross Beever died prematurely on 3 June, 2010, after a short, severe illness. His death brings sadness to all who knew and worked with him, and marks a severe loss for New Zealand science. Nevertheless, his life as a highly intelligent and productive scientist, as a personable and collaborative colleague and mentor, and as a good and loyal friend, will be remembered with great admiration and respect by all with whom he was associated. A recent comment succinctly epitomises Ross Beever: “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.” (GJ Samuels, personal communication).

Ross Beever began university education at the University of Auckland, where he completed a BSc in 1966, as Senior Scholar in botany and chemistry. He then graduated MSc (Hons I) in 1969 after completing thesis research in fungal physiology. He joined Plant Diseases Division of New Zealand’s Department of Scientific & Industrial Research in 1968, and then attended Leeds University, United Kingdom, after receipt of a Sir Walter Mulholland Fellowship for postgraduate studies in fungal genetics. He was awarded a PhD in 1972, and returned to continue his career in plant pathology at Plant Diseases Division in Auckland. With the re-organisation of science in New Zealand in 1992 he joined Landcare Research, and continued research as a plant pathologist and biologist in that institution.

Dr Beever’s achievements made him New Zealand’s most renowned plant pathologist/mycologist. His research spanned a wide range of topics and sub-disciplines related to plant pathology, mycology, botany and plant biology, and was distinguished both by excellence in creation of basic knowledge and for development of solutions for practical plant disease problems. A particular characteristic of his work has been the productive collaboration he has fostered across several plant science disciplines. His achievements have been recognised by numerous distinctions and prestigious awards, and he maintained very high levels of research output throughout his 42-year career, publishing over 170 refereed research papers.

Awards and distinctions

• 2009 Fellow of the Australasian Plant Pathology Society

• 2007 Honorary Life Member, Auckland Botanical Society

• 2007 Science Excellence Award (Landcare Research)

• 2005 Fellow of the New Zealand Institute of Agricultural and Horticultural Science

• 2004 Fellow of the Royal Society of New Zealand

• 2004 Keynote speaker, 13th International Botrytis Symposium, Turkey

• 2003 Marsden Fund award ‘Why fungi like sex’

• 2003–2008 Senior Editor Australasian Plant Pathology

• 2003-International Board Member, The Botrytis Genome Project (INRA, Paris)

• 2003 Invited speaker at 8th Int. Congress of Plant Pathology, Christchurch

• 2001 New Zealand Science and Technology Bronze Medal

• 2000 Tennant Lecturer (University of Otago)

• 2000 Marsden Fund award ‘Phytoplasma genome’

• 1997 Marsden Fund award ‘Fungal hydrophobins’

• 1994 USNZ Cooperative Science Program (USDA, Corvallis, USA)

• 1989 USNZ Cooperative Science Program (USDA, Corvallis, USA)

• 1987–94 Member, Fungicide Task Group (New Zealand Committee on Pesticide Resistance)

• 1983 Trimble Fellowship

• 1980–81 Department of Scientific & Industrial Research Study Award

• 1980 Trimble Fellowship

• 1972–73 Department of Scientific & Industrial Research Study Award

• 1969–72 Sir Walter Mulholland Fellowship

• 1966 Senior Scholar Botany and Chemistry (Auckland University)

• 1963 University Junior Scholarship

Services to the Australasian Plant Pathology Society

Dr Beever became a member of Australasian Plant Pathology Society in 1985. At that time he and other Auckland members of the Society were heavily involved with organisation of the 5^th Biennial Australasian Plant Pathology Conference, held at Auckland University. Since then he continued membership of the Society, making particular contribution as a Senior Editor of Australasian Plant Pathology from 2003 to 2008.

Research achievements

Fungal physiology

Dr Beever made fundamental contributions to knowledge of fungal physiology, particularly including phosphorus nutrition, culminating in a major review of this field relevant to plant pathology and mycorrhizal research (Beever & Burns 1980). He also completed a pioneering study of hydrophobins, proteins which polymerise into sheets (rodlets) on fungal surfaces facilitating many fungal processes including dispersal of dry-spored species (Beever & Dempsey 1978). These rodlet layers, which are amphipathic with the hydrophilic inner surface facing the fungal cell wall and the hydrophobic outer surface facing the exterior (fungal GORE-TEX®), are one of the hallmark characteristics of filamentous fungi, facilitating hyphal growth and spore dispersal.

Fungicide resistance: field studies, genetics and mode of action

Dr Beever developed understanding and effective management of fungicide resistance in plant pathogens, particularly in Botrytis cinerea. He documented levels of resistance in New Zealand, developing an hypothesis explaining field behaviour of pathogens. This emphasised the balance between fitness gain associated with resistance acquisition and fitness loss associated with the genetic changes leading to resistance (Beever et al. 1991). He helped establish industry guidelines for reducing the development of fungicide resistance in plant pathogens in New Zealand. He also initiated studies to understand the genetics and physiology of fungicide resistance in plant pathogens, to explain field observations. He discovered that acquisition of resistance to dicarboximide fungicides results in a loss of fitness linked with increased sensitivity to high osmotic stress (Beever 1983). As well as discovering and characterising the gene associated with dicarboximide resistance (Cui et al. 2002), these studies resulted in a major paper demonstrating, at the genetic level, how pathogen strains in the field have continue to evolve in the face of ongoing fungicide usage (Cui et al. 2004).

Genetics of fungal pathogens

Dr Beever initiated studies of the genetics of fungal plant pathogens to help understand their field behaviour. Emphasis of this research was on Botrytis cinerea, a major pathogen notorious for its variability. He developed protocols to define vegetative compatibility groups in this species (Beever & Parkes 2003), and provided a definitive review of the taxonomy and genetics of Botrytis species (Beever & Weeds 2004). Coupled with this work, he explored the potential of ‘low pathogenesis’ or ‘mild’ strains of pathogens as novel biocontrol agents (Weeds et al. 2000, Beever et al. 2005). He also recognised that nitrogen non-utilising mutants retain high fitness and thus can be used as ‘marked’ strains for ecological studies such as those helping to define infection periods in the field.

Mycoviruses

Dr Beever initiated (with plant virologist colleagues) a programme exploring the potential of mycoviruses as biocontrol agents against fungal plant pathogens. This ‘over the horizon’ approach was initiated in light of the increasing limitations on chemical control of plant pathogens. Two remarkable viruses in Botrytis cinerea have been discovered which show sequence similarity to plant viruses infecting Allium spp., hinting at possible common ancestry (Howitt et al. 2006). A major impediment in the practical application of mycoviruses for biocontrol is the restriction of virus transmission by vegetative incompatibility, and this phenomenon was explored in parallel studies, especially in Botrytis cinerea.

Phytoplasma diseases

The pathology of plants native to New Zealand has traditionally received little attention, except for a phase of intensive work immediately after World War II on a Phormium yellows disease of New Zealand flax (Phormium tenax), a plant cultivated at that time for fibre. This situation changed in the late 1980s, when the highly distinctive New Zealand cabbage trees Cordyline australis (ti kouka) began dying in large numbers throughout northern New Zealand. This was of much concern to the New Zealand public, who regard cabbage tree as a national icon, and a special Cabinet grant was provided for research to understand the cause of “sudden decline” of cabbage trees. Dr Beever led a team charged with finding the cause of the disease. After much investigative plant pathology (Beever et al. 1996), the cause was identified as the phytoplasma “Ca. Phytoplasma australiense” (Andersen et al. 2001). Identification of this pathogen was hampered by rapid host death and the low titre of the pathogen in most parts of affected plants, except for some apices of underground rhizomes. Genetic studies have indicated a number of different lineages in this pathogen, which is also the cause of Phormium yellows disease and diseases of strawberry and other crops, both in Australia and New Zealand. Phylogeographic analysis, a technique seldom used in plant pathology, suggested that there are both ancient (pre-human) and modern populations of the pathogen in New Zealand (Andersen et al. 2006). Phytoplasmas are insect transmitted, and symptomatology suggested that cabbage tree and strawberry were both terminal hosts. Ecological studies indicated that an unrelated wild plant, Coprosma robusta (karamu), acts as a reservoir in the environment of the pathogen (Beever et al. 2004), and led to the proposal that one of the triggers of the disease epidemic in cabbage trees may have been the widespread planting of karamu in restoration plantings. Recently, it has been found that a native plant hopper Zeoliarus (Oliarus) oppositus is a (perhaps the major) vector of the phytoplasma to strawberry and cabbage trees (Beever et al. 2008). In complementary research the team sequenced the genome of a New Zealand isolate of “Ca. Phytoplasma australiense” and its associated plasmids (Liefting et al. 2006). Sustained research over more than a decade, combining field and laboratory research, has resulted in detailed understanding of the ecology of phytoplasmas in a seminatural situation.

Phytophthora diseases

Dr Beever recognised Phytophthora as a genus of major biosecurity importance and of potential significance to the health of native ecosystems. He helped to clarify the role of Phytophthora cinnamomi in native forests (Johnston et al. 2003) and restoration planting. He led a Ministry of Agriculture and Forestry-funded project clarifying the species of Phytophthora occurring in New Zealand using DNA sequencing techniques (Beever et al. 2006). This recognised the presence of P. kernoviae in New Zealand, a pathogen of significant biosecurity importance previously known only from Cornwall in England (Ramsfield et al. 2009). More recently he recognised that species of Phytophthora, including an undescribed species tag named Phytophthora Taxon Agathis, pose a threat to kauri (Agathis australis), an iconic tree of northern New Zealand (Beever et al. 2009).

Taxonomic mycology and plant biology

Dr Beever had continuing interests in the taxonomy and biology of New Zealand native truffles and puffballs (Castellano & Beever 1994; Bridge et al. 2008), and he identified novel plant pathogens affecting native New Zealand flora (Johnston & Beever 1994; Beever 2007). He also studied the botany of native New Zealand plants in collaboration with botanists in Landcare Research. This included research on the genetics of Cordyline (Beever 1981), and study of adaptation in Cordyline australis (Harris et al. 2001, 2003, 2004), work which demonstrated the scientific basis for eco-sourcing of plants in restoration.

Key publications

Fungal physiology

Beever RE, Bollard EG (1970) The nature of the stimulation of fungal growth by potato extract. Journal of General Microbiology60: 273–279.

Beever RE (1975) Regulation of 2-phosphoenolpyruvate carboxykinase and isocitrate lyase syntheses in Neurospora crassa. Journal of General Microbiology86: 197–200.

Beever RE, Dempsey GP (1978) Function of rodlets on the surface of fungal spores. Nature272: 608–610.

Burns DJW, Beever RE (1979) Mechanisms controlling the two phosphate uptake systems in the fungus Neurospora crassa. Journal of Bacteriology 139: 195–204.

Dempsey GP, Beever RE (1979) Electron microscopy of the rodlet layer of Neurospora crassa conidia. Journal of Bacteriology 140: 1050–1062.

Beever RE, Redgwell RJ, Dempsey GP (1979) Purification and chemical characterization of the rodlet layer of Neurospora crassa conidia. Journal of Bacteriology140: 1063–1076.

Beever RE, Burns DJW (1980) Phosphorus uptake, storage and utilization by fungi. Advances in Botanical Research8: 127–219.

Templeton M.D, Rikkerink EHA, Beever RE (1994) Small, cysteine-rich proteins and recognition in fungal-plant interactions. Molecular Plant-Microbe Interactions 7: 320–325.

Sunde M, Kwan AHY, Templeton MD, Beever RE, Mackay JP (2008) Structural analysis of hydrophobins. Micron 39: 773–784.

Fungicide resistance

Beever RE, Byrde RJW (1982) Resistance to the dicarboximide fungicides. In Dekker J, Georgopoulos SG eds. Fungicide Resistance in Crop Protection. Wageningen, Pudoc. pp. 101–117.

Beever RE (1983) Osmotic sensitivity of fungal variants resistant to dicarboximide fungicides. Transactions of the British Mycological Society80: 327–331.

Beever RE, Brien HMR (1983) A survey of resistance to the dicarboximide fungicides in Botrytis cinerea. New Zealand Journal of Agricultural Research26: 391–400.

Hoksbergen KA, Beever RE (1984) Control of low-level dicarboximide-resistant strains of Botrytis cinerea by dicarboximide fungicides. New Zealand Journal of Agricultural Research 27: 107–111.

Beever RE, O’Flaherty BF (1985) Low-level benzimidazole resistance in Botrytis cinerea in New Zealand. New Zealand Journal of Agricultural Research28: 289–292.

Beever RE, Elvidge J (1986) Green fruit rot of apricot caused by Botrytis cinerea resistant to benzimidazole fungicides. New Zealand Journal of Agricultural Research26: 299–304.

Beever RE, Laracy EP (1986) Osmotic adjustment in the filamentous fungus Aspergillus nidulans. Journal of Bacteriology168: 1358–1365.

Beever RE, Laracy EP, Pak HA (1989) Strains of Botrytis cinerea resistant to dicarboximide and benzimidazole fungicides in New Zealand vineyards. Plant Pathology38: 427–437.

Pak HA, Beever RE, Laracy EP (1990) Population dynamics of dicarboximide-resistant strains of Botrytis cinerea on grapevine in New Zealand. Plant Pathology 39: 501–509.

Beever RE, Pak HA, Laracy EP (1991) An hypothesis to account for the behaviour of dicarboximide-resistant strains of Botrytis cinerea in vineyards. Plant Pathology40: 342–346.

Beever RE, Parkes SL (1993) Mating behaviour and genetics of fungicide resistance of Botrytis cinerea in New Zealand. New Zealand Journal of Crop and Horticultural Science21: 303–310.

Cui W, Beever RE, Parkes SL, Weeds PL, Templeton MD (2002) An osmosensing histidine kinase mediates dicarboximide fungicide resistance in Botrytotinia fuckeliana (Botrytis cinerea). Fungal Genetics and Biology 36: 187–198.

Cui W, Beever RE, Parkes SL, Templeton MD (2004) Evolution of an osmosensing histidine kinase in field strains of Botryotinia fuckeliana (Botrytis cinerea) in response to dicarboximide fungicide usage. Phytopathology 94: 1129–1135.

Genetics of fungal pathogens

Beever RE, Olsen TL, Parkes SL (1995) Vegetative compatibility groups in Colletotrichum gloeosporioides (Glomerella cingulata) from apple and other fruits. Australasian Plant Pathology24: 126–132.

Weeds PL, Beever RE, Long PG (1998) New genetic markers for Botrytis cinerea (Botryotinia fuckeliana). Mycological Research 102: 791–800.

Weeds PL, Beever RE, Sharrock KR, Long PG (1999) A major gene controlling pathogenicity in Botryotinia fuckeliana (Botrytis cinerea). Physiological and Molecular Plant Pathology 54: 13–35.

Weeds PL, Beever RE, Long PG (2000) Competition between aggressive and non-aggressive strains of Botrytis cinerea (Botryotinia fuckeliana) on French bean leaves. Australasian Plant Pathology 29: 200–204.

Beever RE, Parkes SL (2003) Use of nitrate non-utilising (Nit) mutants to determine vegetative compatibility in Botryotinia fuckeliana (Botrytis cinerea). European Journal of Plant Pathology109: 607–613.

Beever RE, Weeds PL (2004). Taxonomy and genetic variation of Botrytis and Botryotinia. In: Elad Y, Williamson B, Tudzynski P, Delen N eds. Botrytis: Biology, Pathology, and Control. Dordrecht, Kluwer. Pp 29–52.

Beever RE, Plummer KM, Wurms KV (2005) Novel approaches to controlling fruit pathogens. New Zealand Plant Protection58: 68–73.

Beever RE, Parkes SL (2007) Vegetative compatibility groups in Cryptosporiopsis actinidiae. New Zealand Journal of Crop and Horticultural Science35: 67–72.

Mycoviruses

Howitt RLJ, Beever RE, Pearson MN, Forster RLS (1995) Presence of double-stranded RNA and virus-like particles in Botrytis cinerea. Mycological Research 99: 1472–1478.

Howitt RLJ, Beever RE, Pearson MN, Forster RLS (2001) Genome characterization of a Botrytis virus F, a flexuous rod-shaped mycovirus resembling plant “potex-like” viruses. Journal of General Virology 82: 67–78.

Tsai P-F, Pearson MN, Beever RE (2004) Mycoviruses in Monilinia fructicola. Mycological Research 108: 907–912.

Howitt RLJ, Beever RE, Pearson MN, Forster RLS (2006) Genome characterization of a flexuous rod-shaped mycovirus, Botrytis virus X, reveals high amino acid identity to genes from plant ‘potex-like’ viruses. Archives of Virology 151: 563–579.

Pearson MN, Beever RE, Boine B, Arthur K. 2009 Review: Mycoviruses of filamentous fungi and their relevance to plant pathology. Molecular Plant Pathology 10: 115–128.

Phytoplasma diseases

Beever RE, Forster RLS, Rees-George J, Robertson GI, Wood GA, Winks CJ (1996) Sudden decline of cabbage tree (Cordyline australis): search for the cause. New Zealand Journal of Ecology20: 53–68.

Liefting LW, Andersen MT, Beever RE, Gardner RC, Forster RLS (1996) Sequence heterogeneity in the two 16S rRNA genes of Phormium yellow leaf phytoplasma. Applied and Environmental Microbiology 62: 3133–3139.

Liefting LW, Beever RE, Winks CJ, Pearson MN, Forster RLS (1997) Planthopper transmission of Phormium yellow leaf phytoplasma. Australasian Plant Pathology 26:148–154.

Liefting LW, Padovan AC, Gibb KS, Beever RE, Andersen MT, Newcombe RD, Beck DL, Forster RLS (1998) “Candidatus Phytoplasma australiense” is the phytoplasma associated with Australian grapevine yellows, papaya dieback, and Phormium yellow leaf diseases. European Journal of Plant Pathology 104: 619–623.

Beever RE (2001) Phytoplasmas, a new threat to New Zealand plants? Auckland Botanical Society Journal 56: 24–26.

Andersen MT, Beever RE, Sutherland PW, Forster RLS (2001) Association of ‘Candidatus Phytoplasma australiense’ with sudden decline of cabbage tree in New Zealand. Plant Disease 85: 462–469.

Beever RE, Wood GA, Andersen MT, Pennycook SR, Sutherland PW, Forster RLS (2004) “Candidatus Phytoplasma australiense” in Coprosma robusta in New Zealand. New Zealand Journal of Botany42: 663–675.

Andersen MT, Newcomb RD, Liefting LW, Beever RE (2006) Phylogenetic analysis of “Candidatus Phytoplasma australiense” reveals distinct populations in New Zealand. Phytopathology 96: 838–845.

Pearson MN, Clover GRG, Guy PL, Fletcher JD, Beever RE (2006) A review of the plant virus, viroid and mollicute records for New Zealand. Australasian Plant Pathology 35: 217–252.

Liefting LW, Andersen MT, Lough TJ, Beever RE (2006) Comparative analysis of the plasmids from two isolates of “Candidatus Phytoplasma australiense”. Plasmid 56, 138–144.

Beever RE, Andersen MT, Winks CJ (2008) Transmission of “Candidatus Phytoplasma australiense” to Cordyline australis and Coprosma robusta. Journal of Plant Pathology90 (2, Supplement): S2.459. [Abstracts of ICPP 2008, 9th International Congress of Plant Pathology, Torino, Italy 24–29 August 2008].

Phytophthora diseases

Beever RE, Ramsfield TD, Dick MA, Park D-C, Fletcher MJ, Horner IJ (2006) Molecular characterisation of New Zealand isolates of the fungus Phytophthora [MBS305]. Landcare Research Contract Report LC0506/155 to MAF Policy, Wellington. 43 pp.

Johnston PR, Horner IJ, Beever RE (2003) Phytophthora cinnamomi in New Zealand’s indigenous forests. ‘Phytophthora in Forests and Natural Ecosystems’. 2nd International IUFRO Working Party Meeting, 30 Sept–5 Oct 2002, Albany, Western Australia. Eds JA McComb. GE StJ Hardy, IC Tommerup. Murdoch University Print, Perth.

Ramsfield TD, Dick MA, Beever RE, Horner IJ, McAlonan MJ, Hill CF (2009) Phytophthora kernoviae in New Zealand. Proceedings of the 4 ^th meeting of the International Union of Forest Research Organizations (IUFRO) Working Party S07.02.09: Phytophthoras in Forests and Native Ecosystems. Gen Tech. Rep. PSW-GTR-221. Albany, Ca, USA: US Departments of Agriculture, Forst Service Pacific Southwest Research Station: 47–53.

Beever RE, Waipara NW, Ramsfield TD, Dick MA, Horner IJ (2009) Kauri (Agathis australis) under threat from Phytophthora? Proceedings of the 4^thmeeting of the International Union of Forest Research Organizations (IUFRO) Working Party S07.02.09: Phytophthoras in Forests and Native Ecosystems. Gen Tech. Rep. PSW-GTR-221. Albany, Ca, USA: US Departments of Agriculture, Forst Service Pacific Southwest Research Station: 74–85.

Taxonomic mycology and plant biology

Beever RE (1981) Self-incompatibility in Cordyline kaspar (Agavaceae). New Zealand Journal of Botany19: 13–16.

Castellano MA, Beever RE (1994) Truffle-like basidiomycotina of New Zealand: Gallacea, Hysterangium, Phallobata and Protubera. New Zealand Journal of Botany 32: 305–328.

Johnston PR, Beever RE (1994) Elsinoë dracophylli sp. nov. New Zealand Journal of Botany 32 : 519–520

Harris W, Beever RE, Smallfield B (2001) Variation in response to cold damage by populations of Cordyline australis and some other species of Cordyline (Lomandraceae). New Zealand Journal of Botany 39:147–159.

Harris W, Beever RE, Parkes SL, Webster R, Scheele S (2003) Genotypic variation of height growth and trunk diameter of Cordyline australis (Lomandraceae) grown at three locations in New Zealand. New Zealand Journal of Botany 41: 637–653.

Moyersoen B, Beever RE, Martin F (2003) Genetic diversity of Pisolithus in New Zealand indicates multiple long-distance dispersal from Australia. New Phytologist 160: 569–579.

Harris W, Beever RE, McSeveny T, Parkes SL (2004) Genotypic variation of dead leaf retention by Cordyline australis (Lomandraceae) populations and influence on trunk surface temperatures. New Zealand Journal of Botany 42: 833–845.

Hosaka K, Bates ST, Beever RE, Castellano MA, Colgan III W, Domínguez LS, Geml J, Giachini AJ, Kenney SR, Nouhra ER, Simpson NB, Trappe JM. (2006) Molecular phylogenetics of the gomphoid-phalloid fungi with an establishment of the new subclass Phallomycetidae and two new orders. Mycologia 98: 949–959.

Beever RE (2007) Notes on Chatham Island fungi and some plant pests. Journal of the Auckland Botanical Society62: 28–31.

Bridge PD, Spooner BM, Beever RE, Park D-P (2008) Taxonomy of the fungus commonly known as Stropharia aurantiaca, with new combinations in Leratiomyces. Mycotaxon 103: 109–121.

Prof Richard E Falloon

FNZIAHS, FAPPS

Dr A Ross Ferguson

FRSNZ, FNZIAHS