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Dynamics of plant populations in Heteropogon contortus - Tropical ...

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78 D.M. Orr<br />

native pasture although the 3 earlier papers<br />

reported data from a subset <strong>in</strong>clud<strong>in</strong>g 2 land<br />

classes (narrow-leaved ironbark and silver-leaved<br />

ironbark) at 3 nom<strong>in</strong>al stock<strong>in</strong>g rates (0.3, 0.6 and<br />

0.9 beasts/ha) <strong>in</strong> both native pasture and legumeoversown<br />

native pasture. Further details <strong>of</strong> this<br />

graz<strong>in</strong>g study are provided <strong>in</strong> Orr et al. (2004a).<br />

Site and burn<strong>in</strong>g treatment<br />

Two additional paddocks, each 6.0 ha <strong>in</strong> size, were<br />

located <strong>in</strong> a mixed narrow-leaved ironbark/silverleaved<br />

ironbark landscape and were fenced dur<strong>in</strong>g<br />

w<strong>in</strong>ter 1992. Prior to the study reported here, the<br />

areas conta<strong>in</strong>ed <strong>in</strong> these 2 paddocks were parts <strong>of</strong><br />

additional areas with<strong>in</strong> the orig<strong>in</strong>al study<br />

described <strong>in</strong> Orr et al. (2004a) and, generally, had<br />

been grazed <strong>in</strong>termittently between 1989 and<br />

1992.<br />

One beast grazed each <strong>of</strong> these 2 additional<br />

paddocks (stock<strong>in</strong>g rate <strong>of</strong> 0.15 beasts/ha)<br />

between autumn 1992 and spr<strong>in</strong>g 1995 and the<br />

pastures were burnt each spr<strong>in</strong>g (1992–1995)<br />

follow<strong>in</strong>g the first ra<strong>in</strong>fall event greater than<br />

25 mm between August and November. After<br />

burn<strong>in</strong>g <strong>in</strong> spr<strong>in</strong>g 1995, the paddocks were stocked<br />

with 2 beasts (stock<strong>in</strong>g rate <strong>of</strong> 0.3 beasts/ha) until<br />

the study concluded <strong>in</strong> autumn 1996.<br />

Measurements<br />

Population changes. In each <strong>of</strong> the paddocks, 20<br />

permanent quadrats, each 0.5 × 0.5 m, were<br />

established <strong>in</strong> W<strong>in</strong>ter 1992 <strong>in</strong> 2 nests each <strong>of</strong> 10<br />

quadrats. Measurements <strong>in</strong> these quadrats were<br />

made <strong>in</strong>itially <strong>in</strong> W<strong>in</strong>ter 1992 and subsequently<br />

each autumn until 1996 us<strong>in</strong>g procedures outl<strong>in</strong>ed<br />

<strong>in</strong> Orr et al. (2004a). Briefly, the position <strong>of</strong><br />

<strong>in</strong>dividual H. <strong>contortus</strong> and Aristida spp. <strong>plant</strong>s<br />

<strong>in</strong> each quadrat was charted us<strong>in</strong>g a pantograph<br />

(Williams 1970) and the diameter measured<br />

firstly along the widest diameter and secondly the<br />

diameter <strong>in</strong> the perpendicular direction to this<br />

widest diameter.<br />

For the other 2 treatments, population data for<br />

both H. <strong>contortus</strong> and Aristida spp. were available<br />

for each year between 1990 and 1996. However,<br />

for the purposes <strong>of</strong> the comparisons<br />

reported here, data for the 1990, 1991 and 1992<br />

<strong>populations</strong> <strong>in</strong> each <strong>of</strong> these 4 paddocks were<br />

merged and these merged data were regarded as<br />

the orig<strong>in</strong>al population. By do<strong>in</strong>g this, all <strong>populations</strong><br />

had a common start<strong>in</strong>g time <strong>in</strong> 1992.<br />

Basal area <strong>of</strong> H. <strong>contortus</strong> and Aristida spp.<br />

Basal area <strong>of</strong> H. <strong>contortus</strong> and Aristida spp. was<br />

calculated on an <strong>in</strong>dividual quadrat basis as the<br />

sum <strong>of</strong> the areas occupied by <strong>in</strong>dividual <strong>plant</strong>s <strong>in</strong><br />

the quadrat while mean <strong>plant</strong> size was determ<strong>in</strong>ed<br />

as the total area divided by the number <strong>of</strong> <strong>plant</strong>s.<br />

(Further details are provided <strong>in</strong> Orr et al. 2004a;<br />

2004c.)<br />

Seed production <strong>of</strong> H. <strong>contortus</strong>. Seed production<br />

<strong>of</strong> H. <strong>contortus</strong> was calculated as the product <strong>of</strong><br />

<strong>in</strong>florescence density and seeds per <strong>in</strong>florescence.<br />

Briefly, <strong>in</strong>florescence density was measured annually<br />

<strong>in</strong> the permanent quadrats dur<strong>in</strong>g late March<br />

between 1993 and 1996. The number <strong>of</strong> seeds per<br />

<strong>in</strong>florescence was determ<strong>in</strong>ed annually from 5<br />

<strong>in</strong>florescences collected at random from each<br />

paddock. (Further details are provided <strong>in</strong> Orr et al<br />

2004b.)<br />

Soil seed bank <strong>of</strong> H. <strong>contortus</strong> and Aristida spp.<br />

The soil seed banks <strong>of</strong> H. <strong>contortus</strong> and Aristida<br />

spp. were estimated <strong>in</strong> spr<strong>in</strong>g each year between<br />

1992 and 1995 by germ<strong>in</strong>at<strong>in</strong>g seed conta<strong>in</strong>ed <strong>in</strong><br />

soil cores collected from the areas surround<strong>in</strong>g<br />

the permanent quadrats. Briefly, 4 cores, each<br />

5 cm diameter and 5 cm deep, were bulked to produce<br />

each sample and there were 15 samples (i.e.<br />

60 cores) from each paddock. In the subsequent<br />

summer, each sample was spread as a th<strong>in</strong> layer<br />

on top <strong>of</strong> sand <strong>in</strong> a 15 cm diameter dra<strong>in</strong>ed plastic<br />

pot and seed <strong>in</strong> these samples was germ<strong>in</strong>ated by<br />

water<strong>in</strong>g with an overhead spr<strong>in</strong>kler for 30 m<strong>in</strong>utes<br />

daily <strong>in</strong> a glasshouse (Orr et al. 1996). Seedl<strong>in</strong>gs<br />

were identified and counted after 6 weeks.<br />

(Further details are provided <strong>in</strong> Orr et al. 2004b).<br />

Data analysis<br />

Data were analysed by standard analysis <strong>of</strong><br />

variance as a completely randomised design with<br />

3 treatments and 2 replications. Plant survival<br />

was analysed us<strong>in</strong>g a proportional hazards survival<br />

model (Cox 1972).<br />

Results<br />

Seasonal conditions<br />

The overrid<strong>in</strong>g climatic condition throughout the<br />

study at “Glenwood” was drought with 6 consecutive<br />

years <strong>of</strong> below average ra<strong>in</strong>fall start<strong>in</strong>g <strong>in</strong>

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