Literature review: Impact of Chilean needle grass ... - Weeds Australia

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Other grassland insects of conservation significance include Tropiderus childreni Gray (Phasmatodea: Bacteriidae), SA, Vic, NSW, savannah, mallee, grass, reportedly threatened by urbanisation in the Adelaide Hills (Hill and Michaelis 1988); Anisynta albovenata Waterhouse (Lepidoptera: Hesperiidae), SA, WA, NSW, associated with Austrostipa, reportedly threatened by roadworks, clearing and overgrazing (Hill and Michaelis 1988); Oriexenica kershawi kanunda Tindale (Lepidoptera: Satyriinae) of SA and Vic , associated with open grassland, reportedly threatened by fire at Canunda National Park (Hill and Michaelis 1988); Trapezites lutea (Tepper) (Lepidoptera: Hesperiidae) of SA and Vic found on Lomandra dura threatened by fire and drought in SA (Hill and Michaelis 1988). Xathorhoini, Larentiinae, Geometridae McQuillan (1999) reported on studies on the Xanthorhoini, a tribe of small, colourful, largely diurnal Geometridae (Lepidoptera) with high diversity and endemicity in Australian grasslands. Members of the subfamily Larentiinae are “numerous of the tablelands and mountains of sout-eastern Australia and Tasmania” (Common 1990 p. 375). The tribe is represented in Australia by about 20 spp. of Xanthorhoe Hűbner, mostly found in the south (Common 1990, but only 13 spp. listed by Nielsen et al. 1996), many species of Chrysolarentia Butler, three species of Acodia Rosenstock, one of Austrocidaria Dugdale, eight of Epyaxa Meyrick and two of Visiana Swinhoe (Nielsen et al. 1996). Adults of many Chrysolarentia spp. are “common in summer at the higher altitudes on the tablelands and mountains” of southern NSW, Victoria and Tasmania (Common 1990, as Euphyia Hűbner). Larval Xanthorhoini are sluggish and nocturnal and most species appear to be stenophagous on a small number of lowgrowing annual native herbs, including species of Hydrocotyle, Geranium and Acaena (McQuillan 1999). McFarland (1988) recorded details of the habitats and diets of X. actinipha (Lower) and X. vicissata (Guenée) which both consumed Medicago polymorpha L. in captivity. No Australian Geometridae are known to consume grasses or any other monocots (McFarland 1988). The studies of McQuillan (1999), in Tasmanian grasslands mainly dominated by Poa labillardieri or other Poa spp., Themeda triandra, exotic grasses, Gymnoschoenus sphaerocephalus (R.Br.) (Cyperaceae) or other Cyperaceae and Restionaceae, found that species richness increased with altitude, rainfall and greater sedge-content, and declined in areas with greater grass dominance, livestock grazing and weediness. Site species richness was lowest in highly modified grasslands and reached a maximum of 15 in unmodified grasslands and sedgelands, and 25 spp. were detected overall. Many species survived in small remnant grassland patches. Some species required grass tussocks for shelter in the adult stage and used tussocks as pheromonal ‘calling’ and mating sites. Synemon spp., Sun Moths (Lepidoptera: Castniidae) The Australian Castniidae or sun-moths consist of 20 (Douglas 2003b), 22 (Douglas and Marriott 2003) or 24 (Douglas 2000, Edwards 1996 1997a) named species and 22 (Douglas 2003b) or possibly 21 or more unnamed species of the endemic genus Synemon Doubleday (Edwards 1997a, Douglas and Marriott 2003). Douglas (2003a) provided a brief bibliography for the family. Adults normally fly only in bright sunshine (Edwards 1997a, Douglas 2003b), hence the common name, and resemble butterflies, having broad wings, clubbed antennae and usually bright colours (Douglas 2000). Females have very long, retractable ovipositors and deposit eggs underground between foodplant and soil or on the base of the tillers of the foodplant (Common and Edwards 1981, Edwards 1994, Dunn 1996, Douglas 2000). There is pronounced sexual dimorphism of adults, e.g. the the sexes of S. plana Walker were described as separate species in 1854 and 1874 and their identity was not recognised until 1926 (Edwards 1994). The larvae of different species feed inside the rhizomes of Lomandra spp. (Xanthorrhoaeaceae), Ecdeiocolea sp. (Ecdeiocoleaceae), in the tillers and then against the rhizomes of Lepidosperma viscidum R.Br. (Cyperaceae) or in tunnels entirely undergound, on the roots of grasses (Poaceae) or other sedges (Cyperaceae) (Common and Edwards 1981, Edwards 1994, Dunn 1996, Douglas 2000, Douglas and Marriott 2003). Douglas (1999) also mentions unnamed Juncaceae as larval food plants. Most species are found in areas with sandy or light textured soils (Edwards 1994). Final instar larvae pupate in their feeding gallery and the pupa works its way upwards to protrude from the ground or the foodplant to enable adult emergence (Edwards 1994, Douglas 2000). The whole life cycle occupies 1-3 years (Common and Edwards 1981, Edwards 1994, Douglas 2000, Endersby and Koehler 2006) so the adults present in any one year may represent only a small proportion of the population (Edwards 1994). All species are localised in their occurrence and the adult flight period is brief (Dunn 1996), “as short as a fortnight” (Edwards 1997b). Adults are usually present in only small areas (hundreds of square metres) (Dear 1996). Determination of what plant is actually eaten is a vexed question for some species. Edwards (1994) stated that digging the roots to find pupae “is the surest method … Finding protruding pupal shells within and between tussocks is the next most reliable”, female oviposition probing is unreliable as a foodplant indicator, although local distribution of adults, particularly females, is indicative. None of these methods establish actual feeding relationships. Nevertheless the appear to be the basis for the food plant relationships recorded, until recently with little criticism, in the literature. Determination of the actual food of the larvae is difficult and requires a clear protocol to indicate the various levels of uncertainty inolved (Edwards 1997a) The literature records species of Austrodanthonia as known or probable foodplants for five species found in Victoria including A. setacea (R.Br.) H.P. Linder and A. laevis (Vickery) H.P. Linder (Marriott 2004), while A. carphoides and A. auriculata are also probable food plants (Dear 1996) as well as Austrostipa sp. (Marriott 2004). Food plants of Queensland Synemon spp. include the grasses Chrysopogon sp. and Thellungia advena (Edwards 1997a). Recently Braby and Dunford (2006) identified N. neesiana and red-leg grass Bothrichloa macra as probable larval food plants for the Golden Sun Moth S. plana on the basis of the location and distribution of empty pupal cases (protruding from or beside tussocks) in ACT grasslands. Despite Edwards’ reliance on this method himself (e.g. Edwards 1994) he disputes this attribution (pers. comm. October 2006) believing it more likely that the moth has bred on undetected Austrodanthonia or in plants no longer visible above ground at the site, the juvenile stages being prolonged. More recently Gilmore et al. (2008) reported female oviposition on N. neesiana and pupal cases amongst a dense N. neesiana sward at Greenvale, Victoria. Gilmore et al. (2008) also reported oviposition on Austrostipa spp. and Microlaena stipoides. Edwards (1994) suggested that S. plana larvae may consume a mixture of roots, rhizomes and culm bases. S. plana is one of the few invertebrate taxa that have been used as flagship taxa for grassland invertebrate conservation in Australia (New 2000) and is ranked as critically endangered in Australia (Endersby and Koehler 2006, Gilmore et al. 2008). It 158
was once widespread in south- eastern Australia but in 1993 there were only three known populations recorded since 1950, one in the ACT and two in Victoria (Edwards 1994). Edwards (1994) investigated eight additional likely sites in Canberra and one in NSW and found populations at all except one in central Canberra. Recent surveys have found it at 43 sites in NSW and 12 in the ACT (Endersby and Koehler 2006). The South Australian distribution is based on a single individual from Bordertown (Edwards 1994) and it has not been found recently in that State (O’Dwyer 2004). Endersby and Koehler (2006 citing C. O’Dwyer pers. comm.) stated that: “Prior to 2003, the species had been reported from just six areas in Victoria – Broadford, Tallarook, Flowerdale, Dunkeld, Hamilton and near Nhill-Salisbury”. However that is a highly misleading statement that should have been qualified, perhaps as ‘probably extant populations’. Edwards’ (1994 pp. 32-33) map and list of historical records showed many more localities with much wider distribution. The map provided by O’Dwyer(1999) apparently leaves out historical records noted by Edwards (1994) including Ararat, Bendigo, Castlemaine, Gisbourne, Maryborough, Monbulk, Nagambie and Woodend. The map provided by O’Dwyer (2004) is more complete, showing c. 24 distribution localities prior to 1970 and c. 7 sites after that time. Gilmore et al. (2008) stated that there were only four records in the Melbourne area prior to 2003, near Keilor, Broadmeadows and Laverton. Recent records in Victoria include grasslands in the Deer Park, Craigieburn and Epping areas (Endersby and Koehler 2006, Gilmore et al. 2008), at Greenvale and Woodlands Historic Park (personal observations, Gilmore et al. 2008) and Moyhu (personal observations). Edwards (1994) considered S. plana to be restricted to Austrodanthonia patches in native grasslands, mostly on low hills or rises, sometimes in grazed and mown areas. He stated (p.34): “The foodplant is known to be Danthonia”. Driscoll (1994) stated that the moth requires Austrodanthonia carphoides and A. auriculata grassland for survival. According to O’Dwyer (1999) and O’Dwyer and Attiwill (1999) S. plana inhabits native grasslands and grassy woodlands with >40% Austrodanthonia cover with some sites having up to 75% cover, on soils ranging from sands, through loams to clays, with available P below 14μg/g. O’Dwyer (2004) states that its habitat is native grasslands dominated by Austrodanthonia spp. particularly including also A. eriantha and A. setacea. She noted that oviposition by S. plana has not been observed. At Craigieburn and Epping, Victoria, it has recently been found in habitat previously considered atypical, dominated by T. triandra (Endersby and Koehler 2006). The flight period of S. plana is variable from place to place and year to year (Driscoll 1994), possibly late October peaking in mid November and early December and through to at least the end of January in ACT (Edwards 1994), mainly 11 am – 2 pm (Edwards 1994). According to Driscoll (1994) the likely duration of the larval stage is 21-22 months. O’Dwyer (2004 p. 3) suggests “about 11 months”. Edwards (1994) mentioned the absence of parasite records, and recorded some bird predators of adults. O’Dwyer (2004) mentioned Willie Wagtail Rhipidura leucophrys and robber fly (Asilidae) predation of adults. Much of the literature on S. plana repeats unproved suppositions or speculation as facts, usually including Austrodanthonia spp. being food plants and native grassland habitat specificity, and fails even to provide full lists of known sites of occurrence. A large number of research projects on the species have failed to establish its most basic life history, including hosts plants and the duration of the larval stage. The most recent information (e.g. Braby and Dunford 2006, Endersby and Koehler 2006, Gilmore et al. 2008) indicate it is a widespread and common species that thrives in degraded pastures, including areas dominated by N. neesiana. The development of knowledge of this species is instructive. It has never been, as claimed by New (2000) a species “appraised reasonably fully” and its use as a flagship taxon has been of little value even for its own preservation, since so much of its basic life cycle remains poorly understood. S. selene has been considered a particularly significant taxon because it exists in five distinct parthenogenetic forms (Douglas 2000); however it has not been listed under Victoria’s Flora and Fauna Guarantee Act (Department of Sustainability and Environment 2009a) (Table 18). The Cryptic Sun Moth S. theresa Doubleday is morphologically similar to other Synemon species in the grass-feeding group and it has been suggested that it too feeds on Austrodanthonia (Douglas 2003b). No existing populations are known but locality data on old specimen labels indicate it was an inhabitant of open grassy woodland with Austrodanthonia and Austrostipa in the understorey (Douglas 2003b). It is listed as a threatened species under the Victorian Flora and Fauna Guarantee Act (Department of Sustainability and Environment 2009a). The Orange Sun Moth S. nais Klug occurs in the Mallee in Victoria and in South Australia and Western Australia. In the Mallee it is known from “a floristically diverse combination of open grassy areas interspersed with stands of trees and shrubs”, the grassy areas dominated by Austrodanthonia setacea and Austrostipa spp. (Douglas 2003b p. 7). Oviposition behaviour suggests that A. setacea and an unidentified Austrostipa sp. may be larval hosts (Douglas 2003b). It is listed as a threatened species under the Victorian Flora and Fauna Guarantee Act (Department of Sustainability and Environment 2009a). Only a single population of the Striated Sun Moth Synemon sp. aff. collecta is known in Victoria, from near Shelley. Its habitat may have originally been open grassy woodland and grassland dominated by Austrodanthonia spp. (Douglas 2003b). Edwards (1996) noted that the “grass feeding species in particular have sufferred drastic reductions in distribution from the use of grasslands for agriculture”; urban development threatens populations of some species and invasion by introduced weeds has been identified as a threat to S. plana: “Without grazing or mowing the low-growing natives can become shaded and eventually choked out by taller exotic plants” (Edwards 1993 p. 17, Edwards 1994). In western Victoria this species survives in sheepgrazed Austrodanthonia pastures in which fertilisers and pesticides are not used (Dear 1996). According to Douglas and Marriott (2003 p. 90): “Any disturbance, through ploughing and other types of cultivation and/or excessive invasion of exotic grasses and forbs” leads to the disappearance of Synemon spp. Exotic perennial and annual grasses, not including Nassella spp., are listed as threats by O’Dwyer (2004). Cultivation in Two Wells area of South Australia was blamed for the extinction of S. selene in South Australia (Douglas 2000). These opinions should be viewed with some scepticism since they are based almost solely on observational correlations. S. plana continues to exist in areas of the ACT subject to light grazing or mowing, and the underground larval stage probably confers resistance to fire (Driscoll 1994). Douglas (2000) suggested that moderate disturbance from grazing, slashing or possibly fire appears to be necessary for S. selene to flourish (Douglas 2000). Douglas and Marriott (2003) advocated weed control and regular mowing, burning or grazing to remove accumulated dead grass “that provides cover for predators and reduces the extent of acceptable sites for oviposition”. 159
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Literature review: Impact of Chilea
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Fire 49 Other disturbances 50 Shade
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Conventions and standards Botanical
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neesiana appears to be a habitat ge
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population densities of existing sp
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elated plants have similar defences
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For invasion to occur there must be
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As yet there appears to be no evide
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experimental manipulation of specie
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species tend to be those which tran
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Taxonomy and nomenclature Stipeae N
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Vernacular names ‘Needlegrass’
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to Bouchenak-Khelladi et al. 2009).
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1 m (Walsh 1994), ca. 90 cm (Verloo
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Figure 2. Anatomy of the seed of N.
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are the seeds larger/smaller, longe
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also based on a misunderstanding of
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Table 2. Modified Feekes Scale for
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Argentina, in the provinces of Chac
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Figure 3. Recorded distribution of
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1994). Only 3 of 186 exotic grasses
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According to Morfe et al. (2003) th
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populations have been found in the
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(Honaine et al. 2006). The flechill
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In Australia the altitudinal range
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Proximity to urban development appe
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In the southern Brazilian campos of
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arundinacea (Gardener et al. 2005).
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al. 2008). Cues for masting may be
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Approximately 200 alien grass speci
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Dispersal of seed in contaminated s
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In New Zealand, Hurrell et al. (199
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No emergence was observed in undist
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and high impact (“ability to caus
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also noted that despite a wide rang
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a small reduction in seedhead produ
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Slashing and mowing Slashing can re
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Themeda re-establishment McDougall
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species (Lawton and Schroder 1977 p
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y increased importance of ant grani
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BIODIVERSITY “Biodiversity ... on
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According to Woods (1997 p. 61) “
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2004, Richardson and van Wilgen 200
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negative depending on the particula
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Competition with native plants Comp
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asexual seed production, so local f
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GRASSLANDS Grasses: “... the most
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susceptible to N. neesiana invasion
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Floristic composition, vegetation s
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proportion of the flora then presen
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tussock space (Stuwe and Parsons 19
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Like vascular plant diversity, comm
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Opinions differ on the nature and i
Page 107 and 108: Species Common Name Family Aust ACT
Page 109 and 110: in shifting the distribution, exten
Page 111 and 112: of these systems is largely explain
Page 113 and 114: pasture. The least understood trans
Page 115 and 116: tend to benefit more from relaxed c
Page 117 and 118: Bovids crop their food between the
Page 119 and 120: are therefore less likely to distur
Page 121 and 122: esult in a “short-term flush” o
Page 123 and 124: Fire effects on weeds Moore (1993 p
Page 125 and 126: Table 8. Typical nutrient levels in
Page 127 and 128: grasses produced sigificantly more
Page 129 and 130: Fossorial vertebrates are or were o
Page 131 and 132: (Rosengren 1999). Approximately one
Page 133 and 134: Table 12. Areal extent and conserva
Page 135 and 136: Foreman (1997) investigated the eff
Page 137 and 138: Willis (1964) considered that the f
Page 139 and 140: Austrostipa-Enneapogon) from around
Page 141 and 142: Woodlands and New England Grassy Wo
Page 143 and 144: Thylogale billardierii), Peramelida
Page 145 and 146: Its original habitat on the mainlan
Page 147 and 148: Table 17. Endangered reptile specie
Page 149 and 150: equirement, but unlike plants and v
Page 151 and 152: e assigned the same biodiversity sc
Page 153 and 154: Nematodes are mostly minute animals
Page 155 and 156: found in all mainland states, O. co
Page 157: Keyacris scurra, Melbourne (1993) o
Page 161 and 162: eing sluggish and wingless, and exi
Page 163 and 164: estoration and, if Australia follow
Page 165 and 166: close to the plant are able to bury
Page 167 and 168: Species *Chirothrips mexicanus Craw
Page 169 and 170: Table A2.1 (continued) Species Life
Page 171 and 172: Table A2.1 (continued) Species *Het
Page 177 and 178: Nematodes of grasses and grasslands
Page 179 and 180: REFERENCES Aceñolaza, F.G. (2004)
Page 181 and 182: Benson, D. and McDougall, L. (2005)
Page 183 and 184: Chan, C.W. (1980) Natural grassland
Page 185 and 186: DNRE (Department of Natural Resourc
Page 187 and 188: Fuhrer, B. (1993) A Field Companion
Page 189 and 190: Groves, R.H. and Whalley, R.D.B. (2
Page 191 and 192: Iaconis, L. (2004) Chilean needle g
Page 193 and 194: Levine, J.M., Adler, P.B. and Yelen
Page 195 and 196: McDougall, K.L. (1987) Sites of Bot
Page 197 and 198: Morfe, T.A., McLaren, D.A. and Weis
Page 199 and 200: Perelman, S.B., León, R.J.C. and O
Page 201 and 202: Saunders, D.A. (1999) Biodiversity
Page 203 and 204: Thellung, A. (1912) La flore advent
Page 205 and 206: Weiss, J. and McLaren, D. (2002) Vi
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