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

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<strong>grass</strong>es produced sigificantly more biomass when fertilsed with P than when fertilsed with N or not fertilised, and annual dicots<br />

were significantly more numerically dominant in plots receiving both nutrients. Annual <strong>grass</strong>es had significantly higher numbers<br />

when the dug soil was treated with both nutrients than when treated with N alone, but not P alone. Total biomass was<br />

significantly greater in plots receiving both nutrients. Addition <strong>of</strong> sucrose resulted in increased microbial activity and rapid, near<br />

complete exhaustion <strong>of</strong> soil nitrates. Biomass <strong>of</strong> dicot weeds and annual Poaceae was significantly lower in the sugar treatments<br />

than in disturbed, unfertilised plots.<br />

Small scale spatial and temporal N mineralisation fluxes across the C:N degradation threshold “are probably common in even<br />

strongly N-limited <strong>grass</strong>lands and may play a role in maintaining <strong>grass</strong>land diversity” (Wedin 1999) as well as <strong>of</strong>ferring limited<br />

opportunities for exotic plant invasion. However disturbances involving death <strong>of</strong> T. triandra produce major nutrient pulses in the<br />

soil, which strongly facilitate invasion <strong>of</strong> weeds. Elimination <strong>of</strong> T. triandra tips the competitive balance in favour <strong>of</strong> C 3 <strong>grass</strong>es,<br />

which produce litter with low C:N ratios, below the threshold that limits microbial breakdown, increase the mineralisation rate<br />

and perpetuate their own dominance. If N. neesiana happens to be one <strong>of</strong> those weeds, the cycling <strong>of</strong> N in the system is<br />

permanently altered. N. neesiana litter, produced largely in the summer after flowering, breaks down more rapidly than that <strong>of</strong> T.<br />

triandra so presumably gives rise to soil nutrient fluxes at a time <strong>of</strong> year more suitable to its own needs than that <strong>of</strong> T. triandra.<br />

Levels <strong>of</strong> plant-available nutrients will remain higher than in T. triandra swards, so conditions for the growth <strong>of</strong> other exotic<br />

weeds will be enhanced. Thus, once N. neesiana becomes established it too would appear to be able to perpetuate its own<br />

dominance. Grassland N cycling processes therefore explain both the original persistence and abundance <strong>of</strong> T. triandra and the<br />

permanent change from C 4 (T. triandra) to C 3 <strong>grass</strong> dominance (whether native or exotic) that have resulted from European<br />

grazing regimes and addition <strong>of</strong> chemical fertilisers (Groves and Whalley 2002, Groves et al. 2003).<br />

Nutrient enrichment, nutrient reduction and <strong>grass</strong>land restoration<br />

Species richness <strong>of</strong> native forbs declines with increasing P levels (McIntyre andLavorel 2007). Morgan (1998d) compared native<br />

and non-native species richness and cover in a T. triandra <strong>grass</strong>land and found a strong positive correlation between soil P<br />

(unstated method, unclear if this was Olsen ‘available P’) and the number and cover <strong>of</strong> non-native species, and a negative<br />

association for native species. There was more than twice the level <strong>of</strong> soil P at the edge <strong>of</strong> the <strong>grass</strong>land than in its centre (30-50<br />

m from the edge) and the edges were also enriched with ammonium and organic C. There were weak positive correlations<br />

between ammonium, soil pH, % organic C and exotic richness, and a weak negative correlation between soil pH and native<br />

richness, but no significant relationships for nitrate and sulfur. Non-native, perennial, high-biomass Poaceae were most<br />

dependent on high levels <strong>of</strong> soil nutrients and were suggested to be resource limited in undisturbed soils.<br />

Various endangered native plants are only known from areas where superphosphate has not been applied, but usually there are<br />

compounding disturbance factors such as soil cultivation that have probably played a role in reducing population sizes.<br />

Amphibromus pithogastrus is only known from unploughed sites to which superphosphate has not been applied (Ashton and<br />

Morcom 2004 p. 2). Cultivation destroys the deep-rooted native perennials, although “there are several instances <strong>of</strong> the reestablishment<br />

<strong>of</strong> native <strong>grass</strong>land after ploughing” (Kirkpatrick et al. 1995 p. 80). Weed invasion is enhanced more by nutrient<br />

addition than by cultivation alone (Kirkpatrick et al. 1995).<br />

There are potentially large nutrient additions to small vegetation remnants from farmland via windblown fertiliser, soil and plant<br />

material (Hobbs 1989) and ‘run-on’ <strong>of</strong> nutrient enriched water (Sharp 1997) that could significantly enhance weed invasion<br />

(Hobbs 1989). Proximity to urban development presumably has similar risks, particularly in regard to nutrient enrichment by<br />

atmospheric nutrient deposition. N enrichment is occurring globally in the atmosphere, water and soils. As much as 70% <strong>of</strong> the<br />

reactive nitrogen in the global system is the result <strong>of</strong> human activity and deposition rates in industrialised countries increased<br />

500% in the last 100 years (Hooper 2006) and are expected to double globally by 2050 (Mooney et al. 2006). The increases have<br />

been largest in agricultural lands, where native <strong>grass</strong>lands are mostly found (Aguiar 2005). The N comes from various sources:<br />

fossil fuel combustion, agricultural fertilisers, use <strong>of</strong> agricultural legumes, mobilisation <strong>of</strong> soil nitrogen by soil disturbance, and<br />

ammonia from livestock and human sewage being the most important. Much reactive N enters the soil from acid rain and<br />

particulate deposition and this is largely ammonium or ammonia (Heil et al. 1988, Hooper 2006). Wind-blown dust contains high<br />

amounts <strong>of</strong> N (Eldridge and Mensinga 2007) and increased dust deposition has likely been a feature <strong>of</strong> the landscape associated<br />

with increased agricultural development. Heil et al. (1988) measured bulk and throughfall ammonium deposition in a<br />

Netherlands <strong>grass</strong>land and found that a significant proportion is captured by the canopy and assimilated by the plants. Total<br />

deposition was estimated to be 5.9 kg ha -1 over the 3.5 month growing period in undisturbed <strong>grass</strong>land with a leaf area index <strong>of</strong><br />

2.0-5.7 m 2 m -2 , and 2.5 kg ha -1 over the same period in mown <strong>grass</strong>land with a leaf area index <strong>of</strong> 1.8-3.2 m 2 m -2 . Assimilation in<br />

the canopy was calculated to be 4.7 kg ha -1 in the undisturbed <strong>grass</strong>land and 1.2 kg ha -1 in the mown <strong>grass</strong>land. Such increases in<br />

ammonium availability are more than sufficient to alter competitive relationships between plants and enable fast-growing species<br />

to outcompete slow-growing ones. Grasslands as a whole are not greatly N-limited and are expected to be in the intermediate<br />

range <strong>of</strong> affected ecosystems, but N addition has been demonstrated to have pr<strong>of</strong>ound effects on <strong>grass</strong>land biodiversity,<br />

promoting dominance <strong>of</strong> a few species, generally fast-growing with high shoot-root ratios, and the suppression <strong>of</strong> many (see<br />

Tilman 1987, Aguiar 2005).<br />

Long term N fertiliser application is also associated with increased soil acidity (Aguiar 2005), which in turn is likely to have<br />

differential effects on <strong>grass</strong>land species, Austrodanthonia spp. for example having low tolerance to acidic conditions than<br />

Microlaena stipoides (Sharp 1997). Fertilisation <strong>of</strong> <strong>grass</strong>lands also decreases species diversity <strong>of</strong> native invertebrates across a<br />

wide range <strong>of</strong> taxa and simplifies the trophic web, although normally a few species benefit (Driscoll 1994).<br />

Increased atmospheric CO 2 improves N use efficiency <strong>of</strong> C 3 <strong>grass</strong>es (Milton 2004 see her ref.). A meta-analysis indicates that<br />

under CO 2 enrichment, C 3 species increased their biomass by 44% while C 4 species increased theirs by 33%, so elevated levels<br />

are expected to have a relatively high impact in mixed C 3 /C 4 <strong>grass</strong> systems, although experimental findings have so far been<br />

mixed (Aguiar 2005). Higher water use efficiency by plants with increased CO 2 levels should decrease transpiration and increase<br />

soil water content (Aguiar 2005). Increasing concentration <strong>of</strong> CO 2 is also likely to increase rates <strong>of</strong> microbial denitrification in<br />

the soil (Hooper 2006).<br />

127

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