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

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pyramidata (Benth.) P.G. Wilson and M. sedifolia (F. Muell.) P.G. Wilson that were originally dominant over most <strong>of</strong> the area,<br />

and resulted in their replacement by low-growing, spiny, unpalatable Sclerolaena spp., or by “degraded <strong>grass</strong>land vegetation ...<br />

<strong>of</strong> annual, and to a lesser extent perennial, <strong>grass</strong>es and herbs”. Kirkpatrick et al. (1995) thought that the original vegetation<br />

before livestock grazing might best be characterised as chenopod shrubland with a <strong>grass</strong>y ground layer, and suggested that<br />

drought, fire or rabbit grazing may have destroyed the original shrubland, rather than livestock grazing which merely prevented<br />

regeneration <strong>of</strong> woody species. According to Moore et al. (1973 p. 236) no stands <strong>of</strong> the original vegetation then existed, but<br />

observations suggested that the trees had been up to 9 m high, the shrub layer was well-developed but discontinuous, that<br />

Atriplex predominated on grey and brown clay soils and the Acacia was dominant on red-brown earths. The other main shrubs<br />

were the chenopods Rhagodia spinescens R.Br., Enchylaena tomentosa R.Br. and M. aphylla. The <strong>grass</strong>y ground layer varied in<br />

composition according to soil texture. Chloris truncata and Austrostipa “variabilis” (= A. scabra/A.nodosa) were then common<br />

on light-textured soils. Austrodanthonia caespitosa and C. truncata were the main species on clays, although Austrostipa<br />

artistiglumis was probably once more common or dominant. Leigh and Mulham (1965) considered the latter species to be found<br />

on most soil types and to <strong>of</strong>ten occur in dense, localised stands. Other common and widespread native <strong>grass</strong>es included Chloris<br />

ramosus B.K. Simon, Eragrostis spp., Eriochloa pseudoarcrotricha (Stapf ex Thell.) J.M. Black, Panicum spp., Sporobolus<br />

caroli Mez, (Leigh and Mulham 1965). Other common native species included Bulbine bulbosa (R.Br.) Haw., Lomandra effusa<br />

(Lindl.) Ewart, Hypoxis glabella R.Br., other Atriplex spp., Chenopodium spp., Disphyma crassifolium (L.) L.Bolus, Swainsona<br />

spp., Sida spp., Haloragis spp., Plantago varia R.Br., Asperula conferta Hook. f., Wahlehbergia spp., herbaceous Goodenia spp.<br />

and a large suite <strong>of</strong> small daisies (Leigh and Mulham 1965).<br />

Leigh and Mulham (1965) provided an illustrated compendium <strong>of</strong> the important pastoral plants <strong>of</strong> the Riverina <strong>grass</strong>lands.<br />

Grassland fauna<br />

The fauna <strong>of</strong> temperate <strong>Australia</strong>n native <strong>grass</strong>lands is very poorly known. Much <strong>of</strong> the vertebrate fauna, particularly the<br />

mammals, was eliminated during the early years <strong>of</strong> European occupation, and the invertebrate fauna has received limited<br />

attention, apart from a few agricultural pests and iconic native species, some exploratory inventory studies in Victoria and the<br />

ACT, and several detailed studies in the ACT. <strong>Australia</strong>n <strong>grass</strong>lands have been said to have relatively few specialist animal<br />

species and “a modest but distinctive array <strong>of</strong> animals that feed on on the foliage or the seeds <strong>of</strong> <strong>grass</strong>” (Keith 2004 p. 104).<br />

Grasslands are characteristed by high rates <strong>of</strong> herbivory compared to many other terrestrial ecosystems, with consumption<br />

efficiencies generally c. 25% compared to c. 5% in forests (Tscharntke and Greiler 1995). High root: shoot ratios in temperate<br />

<strong>grass</strong>lands means there is a large subterranean plant biomass to support soil fauna, and the subterranean standing crop consumed<br />

by insects is 2-10 times higher than the above-ground crop (Tscharntke and Greiler 1995). Thus an underground life stage is<br />

typical in <strong>grass</strong>land insect genera (McQuillan 1999).<br />

The seral stage <strong>of</strong> T. triandra <strong>grass</strong>land affects its suitability for animals. Biomass <strong>of</strong> the <strong>grass</strong> determines the floral composition<br />

<strong>of</strong> the <strong>grass</strong>land, presence <strong>of</strong> food plants, amount <strong>of</strong> shade and the structure <strong>of</strong> habitat. Watson (1995 cited by Lunt and Morgan<br />

2002) “found substantial seed predation beneath a dense <strong>grass</strong> cover, but little in burnt open areas”.<br />

In this <strong>review</strong>, the vertebrates are first examined, with a particular emphasis on the extinct mammalian fauna, then invertebrates,<br />

with a particular emphasis on insects, including an examination <strong>of</strong> several rare and threatened taxa, and a detailed <strong>review</strong> <strong>of</strong><br />

what is known about the fauna <strong>of</strong> <strong>grass</strong>es.<br />

Vertebrates<br />

Many <strong>of</strong> the mammals and birds that forage in <strong>grass</strong>lands require structural habitat features for shelter, nesting, etc. that<br />

<strong>grass</strong>lands do not provide (Keith 2004), this being the case for example with insectivorous bats. But today, and in geologically<br />

recent times, the <strong>of</strong>ten complex vegetation mosaics around or within south-eastern <strong>Australia</strong>n <strong>grass</strong>lands have meant that<br />

vertebrates dependent on woodland and shrubland for shelter etc. have been widely able to use <strong>grass</strong>lands to meet some <strong>of</strong> their<br />

requirements.<br />

Vertebrate species can be detrimentally or beneficially impacted by the floristic and structural vegetation changes caused by<br />

weeds, including alterations to food supply and foraging potential, nest sites, cover, predator protection, etc. (Brown et al.<br />

1991). Conversely, the activities <strong>of</strong> vertebrates may hinder or facilitate weed populations by bioturbation, defecation and<br />

urination, feeding and other activities.<br />

Mammals<br />

Grassland mammal assemblages are characterised by the lack <strong>of</strong> arboreal species and <strong>of</strong> browsers, and restricted diversity <strong>of</strong><br />

grazers (Webb 1977). <strong>Australia</strong>n <strong>grass</strong>lands were once occupied by a marsupial megafauna, which disappeared, like similar<br />

faunas in North and South America, during the Miocene and the end <strong>of</strong> the Pleistocene (Webb 1978). The major extinction phase<br />

in <strong>Australia</strong> occurred in the late Pleistocene, possibly mostly c. 26 kbp and continuing to c. 18-15 kbp, and has been associated<br />

with general continental drying (Kershaw et al. 2000). The marsupial megafauna occurred in 250-750 mm rainfall zones, and one<br />

<strong>of</strong> three groups was specifically adapted to southern <strong>grass</strong>lands (Kershaw et al. 2000). Aboriginal <strong>Australia</strong>ns may have played<br />

little part in their disappearance, since fossil evidence suggests coexistence for at least 20,000 years, although recent redating<br />

appears to have markedly reduced this overlap period (Kershaw et al. 2000) and consensus now favors hunting as the major<br />

cause <strong>of</strong> extinctions (Flannery 1994, Johnson 2009). Subsequent to the extinction <strong>of</strong> the megafauna, another set <strong>of</strong> medium sized<br />

animals largely disappeared from south-eastern <strong>Australia</strong>n <strong>grass</strong>lands, mainly during early historical times (Table 15).<br />

According to Lunt et al. (1998), 8 <strong>of</strong> 26 mammal species (excluding bats), mostly rodents and small to medium sized marsupials,<br />

once found in the <strong>grass</strong>y plains <strong>of</strong> south-eastern <strong>Australia</strong> are now no longer present. In the South East <strong>of</strong> South <strong>Australia</strong> the<br />

mammals that have disappeared since European occupation have been those that “lived mainly in woodlands and <strong>grass</strong>lands”<br />

(Aitken 1983 p. 127). Numerous species have been lost in Victoria including Potoroidae (Bettongs Bettongia spp., Rufous<br />

Bettong Aepyprymnus rufescens Gray), Macropodidae (e.g. Eastern Hare-wallaby Lagorchestes leporides (Gould), Bridled<br />

Nailtail Wallaby Onychogalea fraenata (Gould), Toolache Wallaby Macropus greyi Waterhouse, Tasmanian Pademelon<br />

142

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