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

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population densities <strong>of</strong> existing species. K-strategists have low mobility and are adapted to late successional stages and stable<br />

environments where the carrying capacity is approached and competition is high (Matthews 1976, Rejmánek and Richardson<br />

1996). For perennial plants, the ability to propagate vegetatively has <strong>of</strong>ten been considered important (Newsome and Noble<br />

1986).<br />

Table 1. General characteristics <strong>of</strong> successful vs. unsuccessful invasive organisms. Sources: Newsome and Noble 1986, New<br />

1994, Adair 1995, Rejmánek and Richardson 1996, Williamson and Fitter 1996, Cox 2004, Whitney and Gabler 2008.<br />

Successful invaders<br />

large native range<br />

wide climatic tolerance<br />

abundant in native range<br />

high vagility<br />

high reproductive rate<br />

short generation time<br />

reproduction requiring a single parent<br />

small propagule size<br />

high ecological flexibility<br />

wide physical tolerance<br />

wide genetic variability<br />

larger than related taxa<br />

rapid growth<br />

absence <strong>of</strong> specialised requirements<br />

low susceptibility to attack by other organisms<br />

special competitive mechanisms<br />

r strategists<br />

association with humans (commensal)<br />

Unsuccessful invaders<br />

small native range<br />

narrow climate tolerance<br />

rare in native range<br />

low vagility<br />

low reproductive rate<br />

long generation time<br />

reproduction requiring two parents<br />

large propagule size<br />

low ecological flexibility<br />

narrow physical tolerance<br />

narrow genetic variability<br />

smaller than related taxa<br />

slow growth<br />

specialised requirements<br />

high susceptibility to attack by other organisms<br />

no special competitive mechanisms<br />

K strategists<br />

not associated with humans (not commensal)<br />

Subsets <strong>of</strong> inasive characteristics can be combined to define particular ‘strategies’ possessed by invasive plants. Newsome and<br />

Noble (1986) identified four such strategies, based on the suites <strong>of</strong> ecophysiological characters possessed by different types <strong>of</strong><br />

weeds:<br />

1. Gap-grabbers – early germinators with fast initial growth enabling preoccupation <strong>of</strong> ecological space.<br />

2. Competitors - taller growing (light) or with deeper or more extensive roots (water and nutrients).<br />

3. Survivors – longevity due to resistance to mortality factors or clonal growth.<br />

4. Swampers – mass germinators.<br />

Rejmánek (2000) found that Eurasian and North African <strong>grass</strong> species adventive in eastern and western North America are more<br />

<strong>of</strong>ten those with large rather than small native latitudinal ranges and that the latitudinal size <strong>of</strong> the native range is highly<br />

correlated with the size <strong>of</strong> the introduced range. Species with larger ranges may be more successful because <strong>of</strong> the larger<br />

absolute size <strong>of</strong> the propagule pool and because they are more likely to interact with long distance dispersers (Levin 2006).<br />

Schmidt et al. (2008) highlighted a particular character suite for invasive <strong>grass</strong>es: 1. smaller seed size than the native species; 2.<br />

plastic morphological traits that enable the invader to adjust to water and N deficiencies; 3. faster growth to sexual maturity than<br />

the native species; and 4. ready stem dehiscence at the lower node (i.e. stoloniferous growth form). However Lonsdale (1994)<br />

found no relationship between weediness and height growth, relative growth rate, time to maturity (annual or perennial) or seed<br />

weight <strong>of</strong> exotic <strong>grass</strong>es introduced into northern <strong>Australia</strong>. Instead the successful weeds were most likely to be species judged<br />

to be useful, high performing or persistent in experimental agronomic field trials.<br />

Numerous studies have investigated particular biological traits and their relationships with the success <strong>of</strong> invasive plants.<br />

Hamilton et al. (2005) investigated specific leaf area (the ratio <strong>of</strong> the light-capturing area per unit dry mass), plant height and<br />

seed mass and found significant correlations between invasion success and small seed mass at regional and continental scales,<br />

and between high specific leaf area at the continental scale. Greater environmental heterogeneity at regional levels, with<br />

consequent increased biotic resistance, was invoked as one cause <strong>of</strong> the differences across spatial scales. Rejmánek (1995) found<br />

that invasiveness <strong>of</strong> Pinus species correlated with small seed weight, short juvenile period, short mean intervals between large<br />

seed crops and vertebrate dispersal.<br />

Various attempts have been made to assess the most important invasiveness characters by analysing subsets <strong>of</strong> regional weed<br />

floras. For example, Gassó et al. (2009) assessed invasive success on the basis <strong>of</strong> area occupied in mainland Spain and found that<br />

wind dispersed species were the most invasive, followed by animal dispersed species, and that residence time, when

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