The extant Stipeae are believed to be products <strong>of</strong> widespread hybridisation and subsequent stabilisation (Tsvelev 1977). There is “little doubt that the recombination <strong>of</strong> genetically divergent evolutionary lines in the Stipeae has been a factor in colonization <strong>of</strong> diverse habitat” (Johnson 1972 p. 26). Reconstruction <strong>of</strong> the eco-geological history <strong>of</strong> the Pampean stipoid areas <strong>of</strong> South America is at an early stage but some patterns are apparent. Since the mid Tertiary (c. 35 mybp) the Mesopotamian region <strong>of</strong> Argentina has been subjected to a marine transgression that flooded Pampasia, separating the Andean margin from the Brazilian-Uruguayan region, followed by a Pliocene (late Tertiary, 2-10 mybp) regression which allowed the development <strong>of</strong> fluvial plains with a diverse biota, then during the Quaternary (c. 2 mybp – 10 kybp), cooling and aridification alternated with warm to temperate humid periods, and finally in the Holocene (recent) there was another marine transgression (Aceñolaza 2004). In south-eastern Uruguay the Late Pleistocene (c 10-15 kybp) sediments show high concentrations <strong>of</strong> C 3 pooid Poaceae and Asteraceae indicating cool, dry <strong>grass</strong>lands, similar to those at the time in southern Brazil and on the Great Plains <strong>of</strong> North America; the Early Holocene (c. 7-10 kybp) shows a marked transistion to C 4 panicoid <strong>grass</strong>es with wetlands, indicating warmer and wetter conditions; the Mid Holocene (c. 4-6 kybp) showed dynamic climate oscillations from wet to dry with freshwater wetlands predominating; the Late Holocene (4 kybp to present) showed major increase in wetlands, and panicoid <strong>grass</strong>es, similar to southern Brazel but unlike conditions in the Argentine pampas which became more arid (Iriarte 2006). The cool arid periods in the Tertiary may have favoured stipoid speciation, while the cooler times throughout the Caionozoic have probably favoured the proliferation <strong>of</strong> stipoid dominated <strong>grass</strong>lands. The pattern <strong>of</strong> evolution <strong>of</strong> the <strong>Australia</strong>n Stipeae is largely unknown. The <strong>Australia</strong>n fossil record <strong>of</strong> all terrestrial plant taxa is absent or very poor over long periods <strong>of</strong> the Quaternary, although this period probably saw little speciation (McGowran et al. 2000). Poaceae taxa are difficult to distinguish palynologically, the fossil pollen record is generally poor in <strong>Australia</strong> (Kershaw et al. 2000) and <strong>grass</strong> macr<strong>of</strong>ossils are rare, although a potential Bambusites has been described from Tertiary stem impressions (Thomasson 1986). Phytolith analysis (see below) has potential for greater resolution, but has been little used in <strong>Australia</strong> (Kershaw et al. 2000). Morphology and anatomy A detailed description <strong>of</strong> N. neesiana extracted from published works has been compiled. Some descriptions <strong>of</strong> N. neesiana in monographs, floras and other publications are clearly stated to be <strong>of</strong> material found in the region <strong>of</strong> naturalisation (e.g. Burbidge and Gray 1970, Jacobs et al. 1989, Walsh 1994, Verloove 2005), while others are <strong>of</strong> material from the native range (e.g. Burkart 1969). Original or revisionary taxonomic papers generally provide details <strong>of</strong> the material described and examined. In some other descriptions, particularly non-taxonomic and informal publications, it is unclear what material is being described: these <strong>of</strong>ten cite dimensions, etc. that are clearly derivative <strong>of</strong> a single earlier author. Often one anatomical description clearly contradicts another (notably in dimensions) and it is not clear whether this is due to errors, inherent variation in the plant material or to selective assessment <strong>of</strong> character states within individual specimens and within populations. Where possible, errors have been noted. Form and habit: “Erect, strongly caespitose with shoots swollen and close-set at base” (Jacobs et al. 1989, Edgar and Connor 2000); lightly geniculate (Martín Osorio et al. 2000); but lacking the dense tussock form when growing in association with other <strong>grass</strong>es (Champion 1995). Branching intravaginal (Burkart 1969, Jacobs et al. 1989); short-leaved (Burbidge and Gray 1970); a robust tussock when established, “not as clumpy as Poa or Eragrostis” (Duncan 1993), consisting <strong>of</strong> “a number <strong>of</strong> independent tufts” (ACT <strong>Weeds</strong> Working Group 2002). Tillers pr<strong>of</strong>usely when grazed, forming dense (Bourdôt and Ryde 1986), wide clumps (Liebert 1996), that may “form a matt” (Slay 2002c p. 5); grazed tussocks are not large and resemble Festuca arundinacea Schreb. (Duncan 1993, Slay 2002c), Austrostipa spp. (Slay 2002c), or in New Zealand at any time <strong>of</strong> year Rytidosperma spp., in winter under hard grazing Sporobolus africanus (Poir.) Robyns and Tournay, and in the flowering and fruiting stage Bromus diandrus Roth (Slay 2002c). Areas <strong>of</strong> mature tussocks in New England Tablelands pastures have basal ground cover <strong>of</strong> c. 20% (Gardener et al. 2003b). Tussocks have an overall “yellowish-green” colour that contrasts with surrounding pasture (Bourdôt and Ryde 1986, Liebert 1996), although Snell et al. (2007 p. 10) stated that it “becomes yellow or straw like” in winter in colder regions, and “can be a darker green compared to most other pasture species” during early growth stages, and Slay (2002c) stated that in continuous pasture it may have narrower leaves and by lighter green in colour, possibly due to soil fertility and management practices. Mature tussocks typically have masses <strong>of</strong> dead leaves in the center and green leaves around the margin (Gaur et al. 2005). Heavily grazed tussocks in winter, according to Slay (2002c p. 11) “are ‘mushroom’ shaped ... with’flaggy’ 20 cms+ long leaves extending horizontally beyond the perimeter ... <strong>of</strong> the tussock”. The flowering heads are open with drooping branches (Weber 2003). Masses <strong>of</strong> maturing seed heads in dense infestations are “dark green-brown” in colour (Slay 2002c p. 12). After seed is shed, the stems change to “light green-silver” in colour (Slay 2002c p. 12), but remain green into late summer (p.22 and Slay 2001). The green slowly fades, and by autumn the stems are “light brown” (Slay 2002c) or straw-coloured and are easily dislodged from the plant (Slay 2002c). Like other <strong>grass</strong>es, the buds (apical meristems) are at or close to ground level and are protected under tightly enclosed leaf sheaths, so are more likely to survive fire and mammalian grazing (Wheeler et al. 1999). Young shoots are intravaginal (Watson and Dallwitz 2005), a condition that gives rise to the tussock form. In grazed situations in short pastures in the non-reproductive phase plants have a rather flat, almost rosette-like form. In the reproductive phase, plants are strongly upright, and consist almost totally <strong>of</strong> reproductive tillers, largely stalks with panicles, dead leaves and a few small living leaves. The contrast between these phases is extremely pronounced, and they can easily mistaken for different species. The flat, rosette-like growth stage, with smaller leaves, has been called the “sward form” and may be induced by slashing (Bedggood and Moerkerk 2002) and heavy grazing. Height: culm to 2 m (Jacobs et al. 1989, Edgar and Connor 2000, Slay 2002a), 30-100 cm (Hayward and Druce 1919), 60-200 (Weber 2003), 30-140 cm (Barkworth 2006), to 140 cm (Moraldo 1986), 40-90 (Baeza et al. 2007), 1 m or more in the absence <strong>of</strong> grazing (Bourdôt and Ryde 1986), up to 1 m (Martín Osorio et al. 2000, Germishuizen and Meyer 2003, Snell et al. 2007 ), to c. 26
1 m (Walsh 1994), ca. 90 cm (Verloove 2005); to 90 cm (Zanin 1998); 80 cm (Carolin and Tindale 1994), 60 cm (Stace 1997), 30- 100 cm (Burkart 1969, Barkworth and Torres 2001), 50-120 cm (Muyt 2001). Leaves: As with other <strong>grass</strong>es, the leaves mature and senesce progressively from the tip to the base (Wheeler et al. 1999). Lamina mid to dark green (Muyt 2001), flat or loosely inrolled (Jessop et al. 2006), inrolling occurring under stress including drought (Snell et al. 2007); flat to convolute (Barkworth 2006, Zanin 2008), or somewhat inrolled (Walsh 1994), plane or involute (Verloove 2005), rolled when the plant is under moisture stress (Bourdôt and Ryde 1986), sometimes tightly (Slay 2002a); sharp pointed (Martín Osorio et al. 2000); basal leaves up to 40 cm long cauline leaves 20 cm long and 3 mm wide (Martín Osorio et al. 2000); leaves in general to 30 cm long (Walsh 1994, Slay 2002c), mostly
- Page 1 and 2: Literature review: Impact of Chilea
- Page 3 and 4: Fire 49 Other disturbances 50 Shade
- Page 5 and 6: Conventions and standards Botanical
- Page 7 and 8: neesiana appears to be a habitat ge
- Page 9 and 10: population densities of existing sp
- Page 11 and 12: elated plants have similar defences
- Page 13 and 14: For invasion to occur there must be
- Page 15 and 16: As yet there appears to be no evide
- Page 17 and 18: experimental manipulation of specie
- Page 19 and 20: species tend to be those which tran
- Page 21 and 22: Taxonomy and nomenclature Stipeae N
- Page 23 and 24: Vernacular names ‘Needlegrass’
- Page 25: to Bouchenak-Khelladi et al. 2009).
- Page 29 and 30: Figure 2. Anatomy of the seed of N.
- Page 31 and 32: are the seeds larger/smaller, longe
- Page 33 and 34: also based on a misunderstanding of
- Page 35 and 36: Table 2. Modified Feekes Scale for
- Page 37 and 38: Argentina, in the provinces of Chac
- Page 39 and 40: Figure 3. Recorded distribution of
- Page 41 and 42: 1994). Only 3 of 186 exotic grasses
- Page 43 and 44: According to Morfe et al. (2003) th
- Page 45 and 46: populations have been found in the
- Page 47 and 48: (Honaine et al. 2006). The flechill
- Page 49 and 50: In Australia the altitudinal range
- Page 51 and 52: Proximity to urban development appe
- Page 53 and 54: In the southern Brazilian campos of
- Page 55 and 56: arundinacea (Gardener et al. 2005).
- Page 57 and 58: al. 2008). Cues for masting may be
- Page 59 and 60: Approximately 200 alien grass speci
- Page 61 and 62: Dispersal of seed in contaminated s
- Page 63 and 64: In New Zealand, Hurrell et al. (199
- Page 65 and 66: No emergence was observed in undist
- Page 67 and 68: and high impact (“ability to caus
- Page 69 and 70: also noted that despite a wide rang
- Page 71 and 72: a small reduction in seedhead produ
- Page 73 and 74: Slashing and mowing Slashing can re
- Page 75 and 76: 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
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Species Common Name Family Aust ACT
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in shifting the distribution, exten
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of these systems is largely explain
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pasture. The least understood trans
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tend to benefit more from relaxed c
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Bovids crop their food between the
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are therefore less likely to distur
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esult in a “short-term flush” o
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Fire effects on weeds Moore (1993 p
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Table 8. Typical nutrient levels in
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grasses produced sigificantly more
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Fossorial vertebrates are or were o
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(Rosengren 1999). Approximately one
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Table 12. Areal extent and conserva
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Foreman (1997) investigated the eff
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Willis (1964) considered that the f
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Austrostipa-Enneapogon) from around
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Woodlands and New England Grassy Wo
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Thylogale billardierii), Peramelida
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Its original habitat on the mainlan
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Table 17. Endangered reptile specie
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equirement, but unlike plants and v
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e assigned the same biodiversity sc
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Nematodes are mostly minute animals
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found in all mainland states, O. co
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Keyacris scurra, Melbourne (1993) o
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was once widespread in south- easte
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eing sluggish and wingless, and exi
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estoration and, if Australia follow
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close to the plant are able to bury
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Species *Chirothrips mexicanus Craw
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Table A2.1 (continued) Species Life
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Table A2.1 (continued) Species *Het
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Table A2.1 (continued) Species Life
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Table A2.1 (continued) Species Life
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Nematodes of grasses and grasslands
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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
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Thellung, A. (1912) La flore advent
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Weiss, J. and McLaren, D. (2002) Vi