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

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(Barkworth 2006), 10-25 cm (Barkworth and Torres 2001), 5-30 cm (Burkart 1969), 10-25 cm (Moraldo 1986), 13-28 cm (Baeza<br />

et al. 2007), 14-35 cm (Zanin 2008) or 20-35 cm long (Martín Osorio et al. 2000) or up to 30 cm long, open, branches drooping,<br />

flexuous (Jacobs et al. 1989, Edgar and Connor 2000) or erect to nodding (Barkworth 2006), up to 40 cm long, “the branches<br />

breaking up at maturity” (Carolin and Tindale 1994 p. 772); loose (Walsh 1994, Muyt 2001), lax, nodding (Hayward and Druce<br />

1919), drooping (Muyt 2001), sometimes interrupted, to 40 cm long (Walsh 1994); rachis smooth to slightly scabrous, branches and<br />

pedicels with stiff hairs (Jacobs et al. 1989); pedicels 1.5-12 mm (Baeza et al. 2007); branches thin, angular, scabrid (Hayward and<br />

Druce 1919) or pubsecent (Martín Osorio et al. 2000); overall “very distinctive purplish colour” (Duncan 1993) or “initially ...<br />

striking violet ... with ....green awns” (Slay 2002c); branches 2.5-8.5 cm with 2-5 spikelets (Barkworth 2006), 3-5 spikelets (Hayward<br />

and Druce 1919), average <strong>of</strong> c. 15 spikelets in the aerial section and a progressive reduction in length <strong>of</strong> infloresence sections and<br />

number <strong>of</strong> spikelets at lower nodes (Connor et al. 1993), averages <strong>of</strong> 16 to 27.4 seeds per panicle (Gardener et al. 2003a).<br />

Disarticulating above the glumes (Verloove 2005). Published descriptions <strong>of</strong>ten lack precise descriptions <strong>of</strong> the subsidiary<br />

clandestine panicle sections. According to one description (Jacobs et al. 1989, Edgar and Connor 2000): “Occasionally an aerialtype<br />

branch may be produced at the uppermost culm node, and so far as can be judged, remains unsheathed”. This may be<br />

equivalent to what Slay (2002c p. 21) refers to as “another ‘emerged seed head’ [that] may develop from the top node underneath<br />

the terminal leaf sheath” on larger plants (see his Fig. 26). Slay (2002a) noted that such secondary panicles develop in the leaf<br />

sheath <strong>of</strong> all culm nodes, are progressively smaller towards the base <strong>of</strong> the plant and may occasionally produce exposed,<br />

strongly-awned seed. The ‘overt’ panicle is purplish in the flowering stage, becoming more silver as the seeds ripen (Bourdôt<br />

and Ryde 1986). Plants grown from single tillers and from seed produced a mean <strong>of</strong> 18 and 16 panicles per plant respectively<br />

after 6-9 months (Hartley 1994). The potential seed yield per panicle <strong>of</strong> plants in a dense sward in New Zealand was 38 (Slay<br />

2001). As in other paniculate <strong>grass</strong>es, the panicle matures basipetally, the uppermost spikelet developing first (Stebbins 1972).<br />

Flowers:<br />

hermaphrodite; pedicels 1-8 mm long, angled, scabrous, pubescent (Barkworth 2006), drooping (Slay 2002c); terminal panicle<br />

spikelets – glumes unequal (Hayward and Druce 1919) or subequal (Walsh 1994, Barkworth 2006); 10-22 mm (Barkworth 2006),<br />

16-20 mm (Walsh 1994), 15-25 (Moraldo 1986), 16-25 (Walsh 1998), 14-21 mm (Barkworth and Torres 2001), up to c. 2 cm (Weber<br />

2003), 15-20 mm long (Verloove 2005), shorter than the awn column, the upper to 15 mm, the lower to 20 mm (Jacobs et al. 1989)<br />

or the lower 17-20 mm long and the upper 2-3 mm shorter (Martín Osorio et al. 2000) or the lower 13-15 mm and the upper 12-<br />

14 mm (Baeza et al. 2007); the lower 15-17 x 0.4-1 mm, the upper 13-15 x 0.5-1 mm (Zanin 2008); 1.8-2.3 mm wide (Barkworth<br />

2006), narrowly lanceolate (Martín Osorio et al. 2000, Barkworth 2006), linear lanceolate (Hayward and Druce 1919), acuminate<br />

(Hayward and Druce 1919, Walsh 1994), produced into awn-like processes to 3 mm (Jacobs et al. 1989); 3-5 veined (Barkworth<br />

2006), 3-nerved (Hayward and Druce 1919, Jacobs et al. 1989, Zanin 2008), the lower 3-nerved (Martín Osorio et al. 2000), or 5-<br />

nerved, the upper 3-nerved, scabrous-pubescent on the nerves and/or margins (Verloove 2005), nerves scabrous (Moraldo 1986,<br />

Jacobs et al. 1989), central nerve prominent with rigid hairs, lateral nerves with rudimenary hairs and lightly rough (Martín<br />

Osorio et al. 2000); glabrous (Jacobs et al. 1989, Watson and Dallwitz 2005, Barkworth 2006); overall maroon (Cook 1999) or<br />

purple (Bourdôt and Ryde 1986), the lower violet, the upper clear to violet at anthesis (Slay 2002c); violet below, hyaline above<br />

(Jacobs et al. 1989), strongly (Walsh 1994) purplish with hyaline apex and margins (Verloove 2005), hyaline or violet (Moraldo<br />

1986), hyaline and dyed purple in the upper half (Martín Osorio et al. 2000), green-violet (Baeza et al. 2007), brownish with white<br />

margins (Hayward and Druce 1919), losing colour but retained on plant for some time after seed fall (Muyt 2001), retaining a<br />

pinkish tinge even after the seeds have dropped (Liebert 1996); florets 6-13 mm long, 1-1.5 mm wide, terete, widest just below<br />

the crown (Barkworth 2006); anthoecium (including callus, lemma body and crown, plus the concealed palea) cylindrical, 7.5-9<br />

mm but up to 10 mm long, diameter c. 1.2 mm (Verloove 2005) or ±cylindrical 7-11.5 mm long (Barkworth and Torres 2001) or<br />

fusiform 6.3-11.5 mm long, c. 1-1.5 mm wide, white or violet in colour (Burkart 1969), purple (Muyt 2001) or corona purple<br />

(Bourdôt and Ryde 1986), lemma white, corona violet, awn light green at anthesis (Slay 2002c); basal spikelets more or less<br />

cleistogamous, enclosed by uppermost leaf sheath (Burbidge and Gray 1970), other spikelets chasmogamous. [“In the <strong>Australia</strong>n<br />

species [<strong>of</strong> Stipa sens. lat.] there are no externally visible differences between the two, but the cleistogamous spikelets usually<br />

have shorter stamens. Both types may be found scattered through a panicle” (Vickery at al. 1986 p. 11)]; lemma and palea – see<br />

description below under ‘Fruit’; ovary glabrous (Jacobs et al. 1989); lodicules (scales below the stamens and ovary, regarded as<br />

a reduced perianth): 2, c. 1mm (Jacobs et al. 1989), membranous, glabrous (Watson and Dallwitz 2005), hyaline, transparent<br />

(Baeza et al. 2007), nerveless; styles 2, plumose (Jacobs et al. 1989); ovary 1-1.5 mm, glabrous (Baeza et al. 2007); stigmas 2<br />

(Watson and Dallwitz 2005), white (Slay 2002c), 2-2.5 mm (Baeza et al. 2007); anthers 3, penicillate,3-3.5 (Barkworth 2006) or<br />

4-4.5 mm long (Baeza et al. 2007), or up to 3.2 mm long in chasmogamous flowers and in cleistogamous flowers reduced to one<br />

fertile 0.5-0.7 mm long and 2 sterile, 0.1-0.2 mm long (Edgar and Connor 2000), yellow (Slay 2002c); pollen grains smaller than<br />

in Aveneae, almost spherical (tribal characters, Tsvelev 1977).<br />

axillary cleistogamous spikelets - extremely variable (Gardener and Sindel 1998); the various floral parts are progressively<br />

reduced in number and size from the upper to the basal spikelets (Connor et al. 1993) i.e. loss <strong>of</strong> glumes and reduction <strong>of</strong> awn.<br />

See description under “Cleistogenes”, below.<br />

Fruit = ‘seed’ (Fig. 2). Caryopsis “a dry monospermic indehiscent fruit” (Sendulsky et al. 1986);( = grain, includes the hilum and<br />

embryo). 3.5-5 x 0.6-1 mm (Zanin 1998), 4.8-5.5 x 0.9-1.2 mm (Baeza et al. 2007), 3-5 (Barkworth 2006), 4-5 mm (Verloove<br />

2005) or 6-8 mm (Martín Osorio et al. 2000) long; cylindrical (Burkart 1969), obovoid (Baeza et al. 2007); tightly enclosed by<br />

the lemma (Bourdôt and Ryde 1986); clear-c<strong>of</strong>fee coloured (Baeza et al. 2007); hilum (the scar left on the caryopsis at the point<br />

<strong>of</strong> attachment) linear (Jacobs et al. 1989), 2 mm long (Baeza et al. 2007); embryo small (Jacobs et al. 1989), 1.5-1.6 mm (Baeza et<br />

al. 2007), festucoid, 1 / 6 - 1 / 3 <strong>of</strong> the grain (for Stipeae- Tsvelev 1984), with an epiblast and a neglibible mesocotyl internode,<br />

without a scuteller tail (Watson and Dallwitz 2005); endosperm hard, without lipid, containing compound starch grains (Watson<br />

and Dallwitz 2005).<br />

28

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