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

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Nevertheless, based largely on Northern Hemisphere knowledge, the basic insect phytophage assemblage, for the smallest grasses, consists of one species each of Eurytomidae (Hymenoptera), Cecidomyiidae (Diptera) and Pseudococcidae (Hemiptera) (Tscharntke and Greiler 1995). The large, cosmopolitan Common Reed, Phragmites australis (Cav.) Trin. ex Steud., is the most speciose host known, attacked by c. 100 insect species (Tscharntke and Greiler 1995) and has a total of over 160 associated arthropods, approximately half of which are endophages (Witt and McConnachie 2004). Its diverse flora may be largely a result of a long evolutionary history, it being a “Palaeogenic relict” belonging to the”most ancient of modern grasses” the Arundineae (Tsvelev 1984 p. 59) and its broad geographical distribution. The phytophage complement is dependent on many factors, but larger faunas are associated with wider geographical range, large size, predictable occurrence and perenniality (Lawton and Schroder 1977). More than 24 arthropod species have been identified as potential biological control agents for the intertidal grass Spartina alternifolia Lois. in the USA, while Calamagrostis epigejos (L.) has 10 endophagous arthropod species (Witt and McConnachie 2004). Isoptera (Termites) Drepanotermes grass-harvesting termites are a conspicuous feature of arid Australia but are not found in the grasslands of temperate south-eastern Australia (Watson 1982), where most grasses appear to escape termite attack. Coleoptera Unlike on dicotyledonous groups, beetles (Coleoptera) are generally relatively depauperate on grasses. Only 8 spp. are known from Phragmites australis and only 2 of 24 apparent specialists on Spartina alternifolia are beetles (Witt and McConnachie 2004). Coleoptera are “relatively abundant” on smaller grasses including Sporobolus spp. and Nassella trichotoma, but are mostly pollen feeders (Witt and McConnachie 2004). Thysanoptera Thrips (Thysanoptera) are common inhabitants of grasses. Chirothrips species breed only in the flowers of grasses and can reduce seed production and limit grass regeneration (Mound and Palmer 1972). Six species are recorded in Australia, all introductions. Native thrips of grass flowers are species of Odontothripella. Other species associated with Poeaceae flowers include Caliothrips striatopterus (Kobus), common in grass flowers in Queensland, Haplothrips froggatti Hood common in subtropical Australia, Desmothrips reedi Mound and D. tenicornis (Bagnall). Many other species are associated with grasses including Phibalothripis longiceps (Karny), leaf tissue-feeding species related to Anaphothrips, usually found in leaf funnels, Podothrips species predatory on coccoids at the base of grasses in the tropics, and species found in the bases of grass tussocks that feed on fungi (Mound and Palmer 1972). Desmothrips reedi (Aelothripidae)lives at the base of grasses, Moundothrips and Phibalothrips (Thripidae) live on grasses and Odontothripella are found on grass flowers (Mound and Heming 1991) including O. compta Pitkin, O. reedi Pitkin and O. unidentata Pitkin (Pitkin 1972). Introduced Limothrips is common on grasses in cooler areas of southern Australia often living on leaves, the introduced Aptinothrips rufus is abundant on grasses and Haplothrips (Phlaeothripidae) live in grass flowers (Mound and Heming 1991), five species (H. anceps Hood, H. angustus Hood, H. gowdeyi (Franklin), H. froggatti Hood and H. pallescens (Hood)) apparently breeding there, one species living in the base of tussocks, and other species including Antillothrips cingulatus (Hood), Apterygothrips australis Pitkin and Podothrips xanthopus Hood living on or amongst grasses, the latter probably predatory on small arthropods such as coccids, mites or other thrips (Pitkin 1973). Diptera True flies are relatively more common on Poaceae than on dicotyledons, notably gall midges Cecidomyiidae, leafminer flies Agromyzidae and grass flies Chloropidae. On a world basis, many stem-boring and stem-galling Diptera on grasses have narrow host ranges (Witt and McConnachie 2004). Many cecidomyiid species appear to be monophagous at species level (Witt and McConnachie 2004). The Australian cecidomyiid fauna is extremely poorly known (Harris 1979, Gagné 2007, R. Adair and R. Gagné pers. comms.) but several species belonging to three genera, Contarinia, Lasioptera and Geromyia have been recorded from grasses, largely in northern Australia (Harris 1979). Larval Cecidomyiidae feed in the inflorescence, inside the culms, or under leaf sheaths often near the base of the plant. The Contarinia spp. inhabit inflorescences and seed heads, and include C. brevipalpis Harris which attacks Eragrostis brownii (Nees) Kunth, C. dichanthii Harris which attacks Dichanthium spp. including D. sericeum (R.Br.) A. Camus, and undescribed species on Heteropogon contortus (L.) P. Beauv. ex Roem.and Schult. and Themeda triandra (Harris 1979). There are “probably hundreds” of undescribed Australian species (Harris 1979 p. 168). Lasioptera species are found on inflorescences of grasses including Panicum, Setaria, Bothriochloa and Heteropogon, while the single Geromyia species is “probably restricted to Setaria” (Harris 1979 p. 164). No Cecidomyiidae species are known from Austrostipa or Austrodanthonia spp. It is highly likely that exploration of the temperate Australian fauna will reveal many species that inhabit native grasslands and attack grasses (Adair and Gagné pers. comms.). The biology of a New Zealand species that feeds on developing seeds of New Zealand ‘snow-tussocks’ Chionochloa spp. has been described by Kolesik et al. (2007). The larvae do not form galls but overwinter in the floret after feeding on developing seeds. The species appears to be the main seed predator driving mast seeding of these grasses, destroying up to 60% of florets (Kolesik et al. 2007, Kelly et al. 2008). Agromyzidae has relatively high host plant specificity with most species specific to a single plant genus or family, Asteraceae hosting the greatest number of species, and polyphagy (across plant families) being rare (Spencer 1977). Larvae may be leaf miners, internal feeders throughout the plant or selective feeders in roots, stems or flowers (Ferrar 1987, Spencer 1989). Adults feed on plant sap via punctures in the tissue made by the female (Ferrar 1987). No Australian Agromyzidae are known from Stipeae (Spencer 1977), but members of the family are recorded from other grasses in Australia. Spencer (1977) listed Pseudonapomyza ?spinosa Spencer from Urochloa subquadripara (Trin.) R.D. Webster at Darwin, NT (as Agromyza sp. on Brachiaria miliformis (Presl.) Chase in Kleinschmidt 1965), P. spinosa from the introduced Eleusine indica (L.) Gaertn. and Agromyza sp. from Oplismenus compositus (L.) Beauv., while Kleinschmidt (1965) also listed Cerodontha australis Malloch from Poa annua L. and Pseudonapomyza spicata (Malloch), a “common grass-mining species”, probably of exotic origin, from Eleusine indica. Elsewhere in the world P. spinosa attacks wheat and barley and “other wild grasses certainly serve as hosts” (Spencer 1977). U. subquadripara is widespread in NSW, but not in areas of temperate grassland, the common weed E. indica is 154
found in all mainland states, O. compositus is not known from Victoria or NSW. Other Pseudonampomyza spp. with angulate third antennal segments have host plants restricted to Poaceae, but no known Australian hosts: P. probata Spencer, P. rara Spencer, P. salubris Spencer and P. pudica Spencer. Agromyza mellita Spencer and A. venusta, both known from northern Queensland have male genitalia with “the characteristic form of grass-feeders” (Spencer 1977 p. 122). Some Anthomyiidae (Diptera) also attack grasses including the wheat bulb fly Delia coarctata (Fallén), D. extreminata (Malloch) in Bromus in the USA, and Phorbia spp. mining stems and shoots (Ferrar 1987). Others are known to breed on Epichloe fungi on grasses (Ferrar 1987). Larvae of numerous Chloropidae species live in young shoots and stems of grasses and may feed largely on bacteria (Colless and McAlpine 1991) or be saprophagous rather than phytophagous (Ferrar 1987), however little is known of the biology of Australian species (Sabrosky 1989). Ferrar (1987) provided summary information on some pests of cereals and pasture grasses elsewhere in the world, plus a tabulation of the genera known from Poaceae. Most Chloropinae attack Poaceae or Cyperaceae, as do some Oscinellinae (Ferrar 1987). Some Oscinella species appear to be monophagous on particular grass species, while others have multiple known grass hosts (Ferrar 1987). Thirty-two Chloropidae spp., mostly endophages, have been collected from Phragmites, over 20 of which are monophagous (Witt and McConnachie 2004). Diplotaxa similis Spencer has been commonly found in the florets of Chionochloa spp. in New Zealand and has been viewed as the major seed predator, although they may actually often destroy the flower before seed formation, and there has been considerable confusion about whether the damage is caused by the chlropopid or Cecidomyiidae (Kelly et al. 1992). The wingless D. moorei (Salmon) Spencer is a predator of Festuca novae-zelandiae author (Kelly et al. 1992). Some Ephydridae, mostly Hydrellini, attack grasses, and some are cereal pests, but most breed in aquatic situations or wet areas and feed on decaying matter, algae etc. (Ferrar 1987). Hydrellia spp. are recorded from Lolium, Poa, Pennisetum, Holcus, Panicum, Echinochloa and other grasses outside Australia (Ferrar 1987) Hemiptera Hemiptera also appear to have good representation on grasses. Specialist insects on Spartina alternifolia are mainly Hemiptera (Witt and McConnachie 2004). Among the Heteroptera, the Leptocorisini (Alydidae: Leptocorisinae) is closely associated with grasses and includes a number of rice pests, while the Stenodernini (Miridae) and Blissinae (Lygaeidae) are also grass inhabitants (Ahmad 1965). Lygaeids are mostly seed eaters but the Blissinae (Chinch Bugs) feed on grass sap (Slater 1991). Subfamily Cyminae of Lygaeidae contains “small, brown bugs that live in seed heads and resemble seeds of various sedges and rushes” (Slater 1991 p. 502) and possibly includes some grass feeders. The pachygronthine Stenophyella macreta Horváth “is often common in seed heads of grasses” (Slater 1991 p. 502) and “appears to feed on a number of grasses even when they are completely dry” (Slater 1976 p. 135). Nabidae are predacious but oviposit in grass stems (Gross and Cassis 1991). Many grassfeeding Homoptera have a narrow host range, but mealybugs Pseudococcidae and soft scales Coccidae are generally polyphagous (Witt and McConnachie 2004). Farrow (1999) found that Cicadellidae (leafhoppers) were very numerous in grasslands in the Australian Capital Territory and this family along with other planthoppers (Delphacidae, Eurymelidae, Flatidae, Membracidae, Ricanidae), froghoppers (Cercopidae), cicadas (Cicadidae) and allies, collectively classified in the suborder Auchenorryhncha, has been identified as a highly appropriate higher taxon for evaluation of the conservation significance of grasslands and monitoring environmental and habitat change, at least in Europe (Biedermann et al. 2005). They are diverse, with many rare species, wholy herbivorous, their numerical abundance supports higher trophic levels, and perhaps most importantly, rhey have high and rapid sensitivity to a range of disturbances (Biedermann et al. 2005). The European grassland fauna however is much better described than that of Australia, baseline studies of whole faunas have been undertaken and some effects of grassland management on the faunas have been determined. Poaceae probably supports a higher diversity of Australian cicadas (Cicadidae) than any other plant family, but few species inhabit the grasslands of the south-east (Moulds 1990). Nymphs of the grass-feeding species feed on grass roots, and the adults of some species oviposit in grass stems or leaves (Moulds 1990). Evans (1966) considered the cicadelloid fauna of Australian grasses and herbs to be depauperate and Australian grasslands to carry “a very small leafhopper population and ... even fewer cercopids” (p. 20) in comparison with other parts of the world. However Day and Fletcher (1994 p. 1119) noted that there had been “little systematic collecting” of Cicadelloidea in Australia and that distribution data was “very inadequate”. According to Evans (1966 p. 21) both Cicadelloidea and Cercopoidea were “not usually abundant on grasses” in Australia, and most grassfrequenting species were introduced, but he acknowledged that records of food plants were very scanty. However Evans (1966 p. 133) thought it “probable” that all species of Hecalini (Cicadellidae) are grass-feeders. Hecaleus arcuatus (Motschulsky) known from Queensland and outside Australia is recorded from T. triandra and Heteropogon (Day and Fletcher 1994). Day and Fletcher (1994) listed numerous species with known grass hosts in Australia but their designation of “host” is not rigorous; e.g. Evans (1966) stated that Mircrolopa minuta Evans was collected on grasses, which became in Day and Fletcher (1994 p. 1123): “Known hosts: Poaceae (Evans 1966: 87)”. Their host information “frequently reflects the plant species on which specimens were colleced and do not necessarily reflect the true hosts” (Day and Fletcher 1994 p. 1121). Hymenoptera Many ‘seed chalcids’ Eurytomidae (Hymenoptera) are host specific and different species develop in different positions within the culm (Witt and McConnachie 2004). They can cause major reductions in forage yield and seed weight, including a reduction of 60% in Heterostipa comata (Trin. and Rupr.) Barkworth, with consequent reduction in germinability (Witt and McConnachie 2004). A Tetramesa sp. in Africa that bores in the stems of Sporobolus spp. was found to infest 33% of culms at one site. Infested culms were significantly shorter than uninfested and 60% had deformed infloresences (Witt and McConnachie 2004). The tribe Cephini of the sawfly family Cephidae consists of grass-mining specialists, a number of which are cereal pests, but is restricted to the Nearctic and Holarctic (Ivie 2001). 155
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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
Page 103 and 104: Like vascular plant diversity, comm
Page 105 and 106: 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: Nematodes are mostly minute animals
Page 157 and 158: Keyacris scurra, Melbourne (1993) o
Page 159 and 160: was once widespread in south- easte
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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