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

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8.4 and 9.3 respectively (Slay 2001, Slay 2002a). The minimum maintenance level <strong>of</strong> crude protein for livestock is 7% (Slay<br />

2001). Gardener et al. (1999 p. 10) compared N. neesiana productivity and feed analysis with Festuca arundinacea, and Grech<br />

(2007a) compared it with Dactylis glomerata.<br />

Flowering and fruiting<br />

According to Barkworth and Torres (2001 p. 462) and Barkworth (2006) <strong>Australia</strong>n populations “do not appear to set seed in the<br />

exposed inflorescences”. These erroneous statements are possibly based on a few herbarium specimens collected during drought<br />

conditions.<br />

Plants in their first year <strong>of</strong> growth do not flower (Sethu Ramasamy pers. comm. 13 June 2007). Benson and McDougall (2005)<br />

provided the primary juvenile periods <strong>of</strong> many other <strong>grass</strong>es but failed to provide one for N. neesiana. No definitive published<br />

information appears to be available on the juvenile period.<br />

The factors that trigger a shift from vegetative to reproductive growth, are partially known. Low winter temperatures are<br />

necessary for vernalization (Gardener et al. 1996b). (See Sinclair 2002 for the variety <strong>of</strong> factors involved in floral inducation in<br />

<strong>grass</strong>es). Rainfall appears to play a role. In southern Victoria bolting generally commences in mid October (D. McLaren, 26<br />

October 2006).<br />

According to Gardener et al. (2003a) the main ‘flowering’ period is November to February in <strong>Australia</strong> and October to January<br />

in South America (based on herbrium specimens) with some flowering as late as May. Near Guyra on the Northern Tablelands <strong>of</strong><br />

NSW ‘flowering’ commenced between early October and early November and finished between early March and late April<br />

(1994 and 1995, Gardener et al. 2003a). Two separate flowering periods were observed near Guyra in 1995, but not in 1996 or<br />

1997. However Gardener et al. (2003a, etc.) had a very broad, inaccurate definition <strong>of</strong> ‘flowering time’ covering the period from<br />

the first emergence <strong>of</strong> the panicle from the leaf sheath to the time when the last mature seeds were dropped.<br />

Burkart (1969) may be more accurate, giving the flowering period in Argentina as October and November. In south western<br />

Europe, flowering occurs from May to July (Verloove 2005). Slay (2002c) is pehaps most specific, stating that anthesis<br />

(flowering) occurs around mid-November in New Zealand, while Slay (2001) observed flowering over a period <strong>of</strong> approximately<br />

17 days from mid to late November in an infestation at Waipawa, New Zealand. In <strong>Australia</strong>, according to Cook (1999), panicle<br />

production occurs from late spring to early summer, sometimes with a secondary, much less prolific, flush in autumn. Snell et al.<br />

(2007 p. 8) noted that there is “a second panicle seeding period during autumn” in northern NSW. Bedggood and Moerkerk<br />

(2002) state that flowering occurs from early October through to March or April and that panicle seed are produced in autumn in<br />

northern areas. In the ACT N. neesiana usually flowers from November to February (ACT <strong>Weeds</strong> Working Group 2002). In<br />

South <strong>Australia</strong> flowering or fruiting inflorescences are present in November and December (Jessop et al. 2006). Flowering in<br />

Victoria is mostly October to February (Walsh 1994) and in <strong>Australia</strong> generally from spring to early summer with occasional<br />

plants to April (McLaren, Stajsic and Iaconis. 2004).<br />

According to the ACT <strong>Weeds</strong> Working Group (2002) and Ens (2005 citing Ens 2002a) N. neesiana “ has the ability to flower all<br />

year round” given favourable environmental conditions. Bedggood and Moerkerk (2002) recorded that it will flower in response<br />

to summer rain. High density <strong>of</strong> tiller production (and thus seed production) appears to require consistent rainfall during the<br />

reproductive period (Gardener et al. 2003a). Slay (2002c) noted that a second flush <strong>of</strong> panicles can be produced if the normal<br />

late spring panicles are removed by mowing or grazing, and that isolated plants are more likely to have a secondary flowering<br />

period than plants in dense populations.<br />

The flowering periods recorded mostly appear to apply to large populations in broad areas. No information seems to be available<br />

on site specific aspects <strong>of</strong> flowering, the proportion <strong>of</strong> the population that has a particular floral phenology, or the individual<br />

variation <strong>of</strong> a single plant. There is no indication <strong>of</strong> the duration <strong>of</strong> flowering on a particular panicle or the sequence and timing<br />

<strong>of</strong> individual flower opening and closing on a panicle. The time <strong>of</strong> day that the flowers open and their period <strong>of</strong> opening do not<br />

appear to have been recorded. Information about the timing <strong>of</strong> fertilisation, the period <strong>of</strong> receptivity <strong>of</strong> the stigma, the period<br />

between pollination and fertilisation and between fertilisation and embro development have rarely been determined for any<br />

plants (de Tirquell 1986) and are unknown for N. neesiana.<br />

The period from flowering to seed maturation is about 7 weeks (Liebert 1996). According to Slay (2002c), seed became viable<br />

within 8 days <strong>of</strong> flowering, 76% <strong>of</strong> seed was still ‘s<strong>of</strong>t’ after 14 days, with 60% viable after 22 days. In New Zealand most seed<br />

reaches full maturity between late December and early January and is quickly shed, but seed produced on sub-panicles that<br />

emerge from stem nodes matures later (Slay 2002c).<br />

The seed is a velate caryopsis, i.e. the caryopsis falls free <strong>of</strong> other spikelet parts (Sendulsky et al. 1986). Panicle seed matures<br />

and is shed from mid to late summer (Gaur et al. 2005), generally in mid December in southern Victoria (D. McLaren, 26<br />

October 2006). Near Guyra on the Northern Tablelands <strong>of</strong> NSW most seeds matured between late December and mid-February<br />

(1994 and 1995, Gardener et al. 2003a), and was generally shed in December-January (Gardener et al. 2003ba). Slay (2001)<br />

noted that panicle seed at one site was almost all shed simultaneously in early January 2001. Most panicle seed has been shed by<br />

late February in NSW (Duncan 1993) and this is the case in Victoria (Liebert 1996). Stem cleistogene seed from ensheathed<br />

inflorescences matures later than panicle seed, in February (Slay 2002a). In the Melbourne region in 2006 under drought<br />

conditions, most panicle seed was shed in November-December, largely in late November-early December (pers. obs.). Puhar<br />

(1996) noted that seed began to drop at St Albans, Victoria, in early January 2006.<br />

Slay (2002c) modified the Feekes Scale used to define the growth stages <strong>of</strong> cereals to classify the reproductive growth stages <strong>of</strong><br />

N. neesiana (Table 2).<br />

34

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