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Biological Control of Serrated Tussock (Nassella Trichotoma): Is It Worth

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Biological Control of Serrated Tussock (Nassella Trichotoma): Is It Worth 94 Plant Protection Quarterly Vol.13(2) 1998 itch grass (Rottboellia cochinchinensis) as a Biological control of serrated tussock (Nassella case study, he found that highly specific fungal pathogens occur on this grassy trichotoma): Is it worth pursuing? weed in its centre of origin (Evans 1995). Indeed, both the rust and smut species D.T. BrieseA and H.C. EvansB under evaluation proved to be too specific A CSIRO Entomology, GPO Box 1700, Canberra, ACT 2601, Australia. with clearly defined pathotype-biotype associations. Thus, it is probable that a B IIBC, Silwood Park, Ascot, Berkshire, SL5 7TA, United Kingdom. suite of pathotypes will have to be selected in order to cover the range of weed bio- Summary measures, revolving around the use of the types present in the neotropical target area. The potential for biological control of the herbicide fluproponate, have not pre- Nonetheless, the view that natural en- weedy grass, Nassella trichotoma, in vented the continued spread of the weed emies of grasses are not sufficiently spe- Australia is discussed in the light of re- (Campbell and Vere 1995). It is clear that cific and that the risk to economically-im- cent surveys for pathogens in its native other control options need to be investi- portant species of grass is too great to per- range and an improved understanding of gated in order to develop a more effective mit implementation of classical biological the taxonomic relationships of Nassella management strategy for this weed. One control remains a common one, and has to other Australian and South American such option is classical biological control; discouraged such projects in Australia. genera of the tribe Stipeae. This informa- the introduction of a highly specific natu- This paper therefore re-examines the po- tion gives increased optimism that a suit- ral enemy(ies) from their native range that tential for biological control of N. tricho- able biological control agent or agents is capable of killing N. trichotoma plants or toma, in the light of an improved under- can be found. The type of research re- reducing their vigour, capacity for disper- standing of its taxonomic affinities and re- quired to do this is briefly discussed. sal and ability to compete with native and cent preliminary surveys for pathogen pasture grasses. natural enemies in its native range. Introduction The first survey of N. trichotoma was Serrated tussock (Nassella trichotoma undertaken by Wells (1977) in Argentina, Agent host-range and the taxonomic (Nees) Arech.) is a perennial grass of to assess the potential for its biological relationship of Nassella to other southern South American origin that was control in South Africa. He found that it grasses accidentally introduced into Australia in was not an aggressive or weedy plant in Given Wapshere’s (1990) contention that the early 1900s and has since become a se- its native range, due in part to the effects N. trichotoma is ‘probably too closely re- rious weed problem in south-eastern Aus- of natural enemies, including insects, lated to native Stipa spp. to allow the in- tralia (Campbell and Vere 1995). It has fungi and rodents. A follow-up survey for troduction of agents’, it is pertinent to re- also become a weed in New Zealand, insect natural enemies was made by Erb examine the taxonomic position of ser- South Africa and the United States. Ser- (1988), that identified 10 species of weevil rated tussock. Its relationship to other rated tussock is highly invasive, produc- and two moths. Unfortunately, while some grass taxa has long been problematic, at ing large quantities of seed that are widely species specialized on tussock grasses, all times being placed within the large genus dispersed by wind and capable of forming insects attacked a range of other grass spe- Stipa containing several hundred species dense infestations. cies. Evans (1991) reviewed the literature in Eurasia, South America and Australia. In Australia, the most seriously affected on arthropod natural enemies of grassy The one point of agreement has been that country, infestations occur mainly on pas- weeds and concluded that there is an ab- serrated tussock belongs to the tribe ture land, where the weed greatly reduces sence of records of host specific insects Stipeae, though this tribe has variously carrying capacity as it is unpalatable to which may be due to the simplistic and been placed in the subfamilies Pooidae stock. If forced to graze it, sheep lose uniform architecture of the Graminae and Arundinoidae in different morpho- weight and may die (Campbell and Vere which favours polyphagy and reduces the logical classifications of the Poaceae (Clay- 1995). Increasingly, serrated tussock is in- evolutionary pressures for monophagy. ton and Renvoize 1985, Watson et al. 1985). vading natural environments, in National While Erb (1988) confirmed that ‘the com- Recent molecular phylogenetic evi- Parks and water catchment areas, where it bination of insect and fungus attack can dence (Hsaio et al. 1995, Barker et al. 1995) threatens native grasslands and modifies provide a control’ of serrated tussock, suggests that Stipeae does not sit comfort- natural habitats. The most recent survey South Africa did not pursue this work. ably in either sub-family, though only of its distribution in New South Wales Wapshere (1990) examined the possi- Simon (1993) has erected a new subfamily (Jones and Vere 1998) found that serrated bilities of controlling Australian grassy (Stipoideae) for it. Simon (1993) listed only tussock occupies 880 000 ha, indicating weeds biologically. He reported that, al- four genera within the tribe Stipeae that continued spread in the past 20 years. though no fungus had been recorded from are present in Australia: Nassella contain- Moreover, an additional 2 million ha have N. trichotoma, several rust and smut fungi ing six introduced species (McLaren et al. occasional plants present and can be con- infected both members of the genus 1998), Stipa containing over 60 native spe- sidered at risk. In Victoria, infestations Nassella and the closely related Stipa in cies (Vickery et al. 1986), Piptatherum rep- have increased to 130 000 ha with a poten- South America. This led him to conclude resented by a single species, P. miliacea, tial range of 4 million ha (D. McLaren, per- that biological control was unlikely for introduced as a pasture grass but not cur- sonal communication). In Tasmania, the serrated tussock, because Stipa contains rently utilized widely, and Piptochaetium, area infested is smaller (c. 1500 ha) but several native Australian species that are represented by a single weedy species in- spreading (Goninon 1998). The economic useful pasture grasses and the risk to them troduced from South America (Simon losses attributed to serrated tussock are would preclude the introduction of any 1993). currently of the order of $40m per year in pathogen. In contrast, Evans (1991) as- Nassella has at various times been con- New South Wales (Jones and Vere 1998) sessed the potential of fungal pathogens sidered a subgenus of Stipa or a genus in and $5m per year in Victoria (Nicholson as biological control agents of tropical its own right (Barkworth 1990), while N. 1997). grassy weeds and considered that, al- trichotoma has been shifted between the Serrated tussock is a declared noxious though there was still an urgent require- two genera. Nassella was first revised by weed in New South Wales, Victoria and ment for more basic research, the pros- Parodi (1947) who excluded N. trichotoma, Tasmania. Despite this, current control pects appeared to be promising. Using while Corti (1951) assigned it as the Plant Protection Quarterly Vol.13(2) 1998 95 unique member of the section Nassellopsis Eurasian lines within Stipa. This is the classification Piptatherum maintained today by some South Ameri- 1 2 3 can botanists (Zuloaga et al. 1994), Stipa Trikeraia whereas Australian taxonomists included it within the genus Nassella (Vickery and Ptilagrostis Achnatherum * Jacobs 1980). In the most recent revision, Obtusae * Barkworth (1990) redefined the circum- scription of the genus to include all spe- Boreobtusae cies of Stipeae with strongly overlapping lemma margins and lemma apices that are fused into a crown. This expanded genus Nassella comprises 79 species including N. tricho- 5 Stipa (sect. Stephanostipa) toma, N. neesiana and 60 South American 4 Austrostipa * ArchaeostipaArchaeostip a species formerly assigned to Stipa. 3 Piptochaetium 2 Barkworth and Everett’s (1987) exami- Podopogon nation of the relationships of taxa within 1 Hesperostipa * the Stipeae, including for the first time a comparison of Australian and non-Aus- Australian lines American lines tralian Stipa species, sheds most light on Figure. 1. The phylogenetic relationships of monophyletic taxa in the tribe taxonomic position vis a vis biological con- Stipeae. Numbers indicate where several monophyletic groups occur within trol. They identified a number of mono- phyletic species-groups, including five en- one genus, and asterisks indicate those genera or monophyletic groups demic to Australia, covering the previ- containing species previously assigned to the genus Stipa. There are several ously described genera (Figure 1). While other groups within the American lines of Stipa. Figure modified from South American species within Stipa were Barkworth and Everett (1987). close to species of Nassella, both these groups were quite distinct from the Aus- Table 1. Genera of the tribe Stipeae present in Australia that would need to tralian groups of Stipa (Figure 1). In fact, be considered in any host-testing program for biological control agents of these differences are considered sufficient serrated tussock. to warrant erecting a new genus, Austro- stipa comprising 13 subgenera, to describe Genus No. of species Origin Status the Australian native taxa (Jacobs and Austrostipa 62 Australia native taxon with several useful Everett 1996). In addition, all three exotic pasture and several weedy species species present in Australia that were as- Nassella 6 South America declared noxious weeds or signed to Stipa have recently been placed potentially weedy in other genera; S.
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