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

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5°C warmer than under the canopy and much larger differences occurred in summer. Such contrasts will substantially affect the<br />

behaviour and activity levels <strong>of</strong> invertebrates.<br />

Farrow (1999) found no clear differences in the diversity <strong>of</strong> canopy-living insects in small burnt patches and unburnt areas in<br />

ACT <strong>grass</strong>lands and suggested that recolonisation occurred within 6 months <strong>of</strong> fire. Nearly twice as many individual insects<br />

were present in summer in areas burnt 6 months prior to sampling than in unburnt areas, and more than three times as many in<br />

the following spring. Species that benefit from more open ground, like many ants, probably commonly proliferate. Maintenance<br />

or periodic refreshment <strong>of</strong> plant diversity by fire will benefit a wide range <strong>of</strong> herbivorous taxa with particular food plants, and<br />

maintenance <strong>of</strong> the forb components will benefit nectar and pollen feeders. Maintenance or enhancement <strong>of</strong> animal diversity at<br />

the primary consumer level will flow on to higher tropic levels, benefitting the diverse array <strong>of</strong> predators and parasitoids.<br />

Greenslade (1994) found litter removal by fire was likely to markedly reduce Collembola diversity for at least two years. Other<br />

detritivores dependant on plant litter would be expected to be similarly disadvantaged.<br />

Any beneficial effects on fire on plant diversity may or may not flow on to invertebrates, depending, in part on whether source<br />

populations <strong>of</strong> new and recolonising species exist (Tscharntke and Greiler 1995). Driscoll (1994) concluded that some species<br />

may be completely intolerant <strong>of</strong> fire, while others may require it for survival. Any species that spend part <strong>of</strong> their lifecycle<br />

underground are probably relatively resistant to fire, while those that do not are likely to be more highly vagile and have an<br />

ability to recolonise from unburnt areas. Sedentary, non-subterranean species are probably most vulnerable to extinction, but<br />

there are unlikely to many <strong>of</strong> them in fire adapted vegetation. As with vascular plants (Stuwe 1994) and bryophytes (Morgan<br />

2004), the indigenous invertebrate flora <strong>of</strong> <strong>grass</strong>lands that have been subject to periodical fire over many thousands <strong>of</strong> years must<br />

consist largely <strong>of</strong> fire-adapted species, whether or not they are fire survivors or recolonisers.<br />

Fire regime management in Victoria now incorporates the goal <strong>of</strong> providing the conditions necessary for the persistence <strong>of</strong> the<br />

indigenous biota (Mansergh et al. 2006b) but implementation <strong>of</strong> this idealfor the various biotic elements, including animals, is<br />

difficult and is bound to produce contradictory results because different valuable elements <strong>of</strong> the biota may be advantaged or<br />

harmed by the same fire cycle. The difficulty for managers is to evaluate and juggle the <strong>of</strong>ten contrary needs and tolerances <strong>of</strong><br />

the various components in each <strong>grass</strong>land remnant.<br />

Nutrients and soil factors<br />

Lowland native <strong>grass</strong>land in <strong>Australia</strong> is found on relatively fertile, organic-rich, cracking clay soils on substrates <strong>of</strong> volcanic<br />

rocks (basalt and dolerite), fine-grained sedimentary rocks (including limestone) or recent alluvium (Kirkpatrick et al. 1995). In<br />

the past, temperate <strong>grass</strong>lands have erroneously been characterised as low in nutrients, based on analysis <strong>of</strong> soils. Mott and<br />

Groves (1994 p. 376) state that “<strong>Australia</strong>n <strong>grass</strong>land soils are <strong>of</strong> very low fertility, particularly <strong>of</strong> nitrogen and phosphorus, by<br />

comparison with those <strong>of</strong> <strong>grass</strong>lands elesewhere”. Sharp (1997) lists low soil nutrient content as one important factor<br />

determining the distribution <strong>of</strong> native <strong>grass</strong>land in the ACT. <strong>Australia</strong>n soils have in general been considered to be nutrient<br />

deficient (e.g. Roberts et al. 2006). However, like tropical rainforest, much <strong>of</strong> the system nutrients are held in the plants.<br />

Above- and below-ground biomass<br />

In temperate herbaceous communities 60-80% <strong>of</strong> vascular plant biomass is underground (see Reynolds 2006 p. 57). In <strong>grass</strong>lands<br />

up to 90% is below-ground (Wijesuriya and Hocking 1999) and it is usual for a high proportion <strong>of</strong> total plant biomass to be<br />

represented by roots and buried crowns. More <strong>of</strong> the energy captured in photosynthesis in <strong>grass</strong>lands is directed to below-ground<br />

parts than those above-ground and most <strong>of</strong> the nutrient circulation occurs below ground (Soriano et al. 1992). In <strong>Australia</strong>n<br />

<strong>grass</strong>lands most <strong>of</strong> the biomass is contained in the dominant <strong>grass</strong>es (Groves 1965). T. triandra- and Poa-dominated<br />

communities are more productive (i.e. produce more biomass) than the Austrodanthonia and Austrostipa <strong>grass</strong>lands that<br />

predominate in drier areas (Lunt and Morgan 2002). The mean root: shoot ratios in North American <strong>grass</strong>lands varies from 13 to<br />

2, being higher in cooler climates (Tscharntke and Greiler 1995). Cooler and drier <strong>grass</strong>lands generally have ratios between 13<br />

and 6, while warmer, more humid <strong>grass</strong>lands have ratios between 6 and 2 (Soriano et al. 1992). A ratio <strong>of</strong> 2.6 was calculated for<br />

one Argentinean pampas <strong>grass</strong>land (Soriano et al. 1992). Rodríguez et al. (1995) measured below-ground biomass down to 10<br />

cm depth in five grazed <strong>grass</strong>land communities in Spain and found that 61-68% <strong>of</strong> the total biomass was below ground, <strong>of</strong> which<br />

49-68% was in crowns and the remainder below 1 cm depth. Groves (1965) found that the root:shoot ratio <strong>of</strong> a T. triandra<br />

<strong>grass</strong>land had very high seasonable variability but that root biomass was generally 2 to 4 times that <strong>of</strong> above-ground parts, i.e.<br />

66-80% <strong>of</strong> the biomass <strong>of</strong> the overwhelming dominant <strong>grass</strong> was below ground. In one Pampas <strong>grass</strong>land 65% <strong>of</strong> the<br />

underground biomass was located above 10 cm, 85% above 30 cm and 100% above 70 cm, a similar distribution to most<br />

temperate, subhumid <strong>grass</strong>lands (Soriano et al. 1992).<br />

Underground net primary productivity in one Pampas <strong>grass</strong>land was estimated to be about 5000 kg ha -1 yr -1 , just 17% less than<br />

productivity above ground, with a below-ground: above-ground productivity ratio <strong>of</strong> 0.9 (Soriano et al. 1992).<br />

Soil nutrient levels<br />

The soils <strong>of</strong> <strong>Australia</strong>n native <strong>grass</strong>lands usually have low nutrient levels (Table 8), and a high organic matter content with<br />

organically bound N and P (Wijesuriya and Hocking 1999). However most <strong>of</strong> the nutrients in the system are locked up in the<br />

biomass <strong>of</strong> the <strong>grass</strong>es. In the Victorian basalt plains, soils derived from sediments have higher nutrient levels than those derived<br />

from basalt, and soils on stony rises have the highest fertility (Williams 2007).<br />

124

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