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

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Caro (1966) illustrated a plant, ligule, glumes and seed of var. hirsuta. Burkart (1969) illustrated a whole plant and seed of var. neesiana, seed of var. longiaristata and the caryopsis of var. neesiana. Torres (1993) illustrated seeds of vars. neesiana, longiaristata and gracilior. Hocking (2002) suggested that more than one “type” of N. neesiana may be present in Australia. All the Australian specimens examined by Vickery et al. (1986) ‘seemed’ to be N. neesiana var. neesiana. Walsh (1994), who examined Victorian material, did not contradict this. Britt (2001) and Britt et al. (2002) stated that the varieties present in Australia were “unknown”. Gardener et al. (2005) considered the plants they studied on the Northern Tablelands of NSW were var. neesiana and stated that it appeared to be the only variety present in Australia. Jessop et al. (2006) classed all South Australian material as subsp. neesiana. Nevertheless, the large number of varieties and forms recognised by American taxonomists indicates the existence of considerable infraspecific phenotypic variation. Information at hand is inusufficient to indicate whether this variation is merely phenotypic plasticity in response to environmental variation or represents distinct genotypes. Misapplied names In Europe the name Stipa setigera (= Nassella mucronta) has been widely misapplied to N. neesiana and all verifiable records of these taxa in Europe are N. neesiana (Verloove 2005). N. neesiana and N. mucronata are easily confused (Vàzquez and Devesa 1996). Zanin et al. (1992, cited by Gardener 1998) used the name Nassella setigera var. setigera for N. neesiana vars. gracilior, virescens and hirsuta in Brazil. Overbeck et al. (2007) also used the name S. setigera for what is apparently N. neesiana. Longhi- Wagner and Zanin (1998) used the name Stipa setigera for N. neesiana and recorded it from Paraguay. Hybridisation etc. Many groups of grasses arose by intergeneric hybidisation (Tsvelev 1984) and interspecific hybridisation is very common in Poaceae (Wheeler et al. 1990) and in Stipeae. According to Tsvelev (1977) all extant Stipeae are of hybrid origin and the tribe itself may have arisen this way. Grasses in general are highly dispersable and fecund, a characteristic of higher taxa with high speciation rates (Levin 2006). Wind pollination and simultaneous flowering of grass species provides much opportunity for cross-fertilsation (Groves and Whalley 2002). Sterile intergeneric stipoid hybrids are very common (Johnson 1972). Oryzopsis caduca Beal (?name: not in Barkworth 2006) is a hybrid between Achnatherum hymenoides (Roem. and Schult.) Barkworth and Nassella viridula (Trin.) Barkworth, and A. hymenoides crosses spontaneously with 11 stipoid species in the USA, including Nassella pulchra (Hitchc.) Barkworth, N. cernua (Stebbins and Love) Barkworth and species of Stipa, Achnatherum, and Heterostipa, producing plants that have all been classified as Oryzopsis bloomeri (Boland) Ricker (Johnson 1972). “In favorable years and under advantageous conditions of habitat disurbance hundres [sic] of individuals have been observed in some hybrid swarms” (Johnson 1972 p. 25). Barkworth (1990 2006) and Barkworth and Torres (2001) considered N. viridula itself to be most probably an alloploid, with Achnatherum and Nassella progenitors, or possibly an autoploid derivative of a common ancestor of Achnatherum and Nassella. According to USDA FEIS (2006), the occasional hybrids of N. viridula with A. hymenoides produce the sterile Achnella caduca (Beal) Barkworth. Watson and Dallwitz (2005) listed such hybrids as X Stiporyzopsis B.L. Johnson and Rogler and X Achnella Barkworth. Tsvelev (1984 p. 903) considered intra-sectional hybrids in Stipa to be “probably ... not so rare”, but inter-sectional hybrids to be “rarer”. Examples of the latter include S. gregarkunii P. Smirnov, reportedly a result of crossing between S. pulcherrima C. Koch (section Stipa) and S. caucasica Schmalh. (Section Smirnovia), and S. kopetdaghensis Czopan. (section Smirnovia), possibly a cross between S. caucasica Schmalh. (section Smirnovia) and S. zalesskii Walensky subsp. turcomania (P. Smirn.) Tzvel. (section Stipa). The presence of hybrids in S. aggr. capillata L. and in section Stipa was undoubted, and in section Barbatae a hybrid S. arabica Trin. and Rupr. subsp. arabica X S. hohenacheriana Trin. and Rupr. subsp. nachiczevanica Tzvel. ( S. hohenackerana according to Barkworth 2006) had been recorded (Tsvelev 1984). Hybridisation of N. neesiana does not appear to have been reported. However Verloove (2005) examined specimens from South America and France with characters intermediate between N. neesiana, N. mucronata (H.B.Kunth) R.W.Pohl and/or N. poeppigiana (Trin. and Rupr.) Barkworth. Barkworth and Torres (2001) considered N. mucronata, N. mexicana (Hitchock) R.W. Pohl and N. leucotricha (Trin. and Rupr.) R.W. Pohl to comprise a species complex, with intergrades of the former two species in northern South America and of N. mucronata with N. leucotricha in northern Mexico. Data in Britt et al. (2002) suggest the possibility of some gene flow between N. neesiana and N. leucotricha at Melton, Victoria. The new combinations of exotic and native stipoids now occurring in south-eastern Australia would appear to create new possibilities for hybridisation. It should be kept in mind that hybrid individuals might be found in Australia, and any suspected examples should be collected. Hybrids between N. neesiana races or regional populations may occur as a result of pollen flow, as may hybrids with other Nassella spp., or possibly with other introduced Stipeae and native Austrodanthonia spp. Evolutionary origin The evolutionary origin of the Poaceae is obscure. Grasses are presumed to have existed since the Mid-Cretaceous (c. 120 mybp, Mesozoic Era) on the basis of fossil leaves (Tsvelev 1984). Stebbins (1986) suggested their first appearance in the Late- Cretaceous, but molecular clock estimates suggest an origin about 83 mybp (Prasad et al. 2005). A minimum age of 90 mybp (Cretaceous) for the crown group of Poaceae has recently been suggested (Bouchenak-Khelladi et al. 2009). Thomasson (1986) found that the oldest definite grass fossils were from the Oligocene (c. 36-24 mybp, Tertiary Era) of North America, that fossils of probable grasses were also known from the Oligocene of Germany, and that Mesozoic leaf impression fossils from Europe, North America and Mongolia were only “possible” grasses. Jones (1999a) stated that grass fossils occur in the Eocene (45 mybp) in the Americas and Africa. Currently the oldest unequivocal grass macrofossils are recorded from the Paleocene-Eocene boundary c. 56 mybp (Piperno and Sues 2005). Bouchenak-Khelladi et al. (2009) considered a spikelet with a minimum age of 55 mybp to represent a crown node for almost all recognised grass genera. However grass phytoliths representative of a diverse range of taxa have been found in 70 mybp (late Cretaceous) fossilised dinosaur dung (Prasad et al. 2005) (65-67 mybp according 24
to Bouchenak-Khelladi et al. 2009). The earliest fossil grass pollen has been found in the early Tertiary, with doubtful earlier records from the Cretaceous (Thomasson 1986), and presumed records (Monoporites) from 70-60 mybp (Prasad et al. 2005). Grass pollen first appears in the Australian record in the Paleocene (Macphail et al. 1994), at the end of the Paleocene (c. 58 mybp) (Keith 2004), or about 50 mybp (Keith 2004). Undoubted fossils, with seeds very similar to modern Stipa, Piptochaetium and Phalaris, occur in rocks dated to the mid-Tertiary (c. 35 mybp) long after the family first evolved (Stebbins 1972). Bouchenak-Khelladi et al. (2009) indicate that the BEP clade (Bambusoideae, Ehrhartoideae and Pooideae), which includes Stipeae, originated in the Paleocene (57 mypb), while the other major clade, including most C 4 species, is younger, originating in the late Eocene (40 mybp). C 4 physiology was probably well developed by the Miocene (Thomasson 1986), the oldest origin in grasses being approximately 30.9 mybp (Christin et al. 2009, Bouchenak-Khelladi et al. 2009). Tsvelev (1977 1984) convincingly argued that Poaceae first evolved in response to colder and drier conditions in mountainous areas with an increasingly continental climate. Stebbins (1986) argued a probable origin from Joinvillea-like ancestors (Joinvilleaceae) and differentiation into major families in lowland, open tropical savannahs with seasonal droughts; the non sequitur of a grass-less savannah going unremarked. A South American (Gondwanan) origin for the family has been suggested on the basis of the distribution of extant taxa, with diversification of subclades in Gondwana by the late Cretaceous (Prasad et al. 2005). Although considerable diversification within the family took place in the mid-Miocene (Piperno and Sues 2005), most modern tribes probably existed by the Paleocene (early Tertiary, c. 50 mybp), probably along with many modern genera (Tsvelev 1984, Jones 1999a). The Stipeae probably arose from primitive Pooideae (Stebbins 1986) which existed at least as early as 70 mybp (Prasad et al. 2005). Jones (1999a) cited authors who argued the tropical origin of grasslands in areas that were cooling and developing seasonal aridity, in forest-savannah ecotone. Poaceae are uncommon in the fossil record until the mid-Miocene (16-11 mybp) (Piperno and Sues 2005) and “only became widespread 25 to 15 million years ago when cool, dry conditions kicked in” (O’Donoghue 2008 p. 39). The unimportance of mammals, except for South American gonwanatherians, with typical grazing adaptations such as hypsodont teeth until the Oligocene and Miocene also indicates that grasses were a minor component of vegetation before this time (Prasad et al. 2005). Barkworth and Everett (1986 p. 261) noted that the character sets that define non-Australian supra-specific stipoid taxa do not occur in Australian taxa, even though Australian stipoids in aggregate possess almost all of these characters. They believed that Stipeae originated in Gondwana, suggesting a late Jurassic or Cretaceous (c. 135 mybp) origin for the tribe (Jones 1999a). Tsvelev (1977) more or less agreed, noting that the impoverished stipoid flora of Africa (excluding the Mediterranean areas) probably resulted from repeated long dry periods and the absence of high mountain refugia. Africa began to split from the South American section of Gondwana in the mid to late Mesozoic (c. 165-70 mybp) leaving Australia and South America still linked through Antarctica by the early Tertiary (c. 65 mybp) (Barlow 1981). Separation of Antarctica and Australia occurred in the Paleocene (53-50 mybp) and accelerated in the middle-late Eocene (c. 43-36 mybp) (McGowran et al. 2000). Several authors have favoured the Gondwanan centre of origin of Stipa (sens. lat.) including Moraldo (1986 p. 205) who placed it in the ‘suture zones between South American, Antarctica and Australia’. South America has a very rich stipoid flora, paralleled only by that of Eurasia (Tsvelev 1977). The current centre of diversity of Nassella is Argentina with c. 72 species, with greatest diversity in the north west, and 26 indigenous species. Uruguay has 23+ spp., and the greatest diversity in Bolivia and Chile is in the central Andes, adjacent to Argentina (Reyna and Barkworth 1994, Barkworth and Torres 2001, Barkworth 2006). The pampas or Rio de la Plata grasslands have 25 Nassella species (Gardener et al. 1996b). Whether the genus evolved in the Pampas region is not known. There are no macrofossil Stipeae known from South America (Barkworth 2006). Tsvelev (1977) considered Oligocene (36-25 mybp) fossil panicles from Colorado to be Stipa florissanti (Knowlt.) MacGinitie, and noted that the American grass specialist Agnes Chase considered them identical with the extant species Stipa mucronata (now Nassella mucronata (Kunth) R.W. Pohl. Barkworth). Everett (1986) accept the Miocene North American Berriochloa primaeva Thomasson to be the earliest stipoid fossil. According to Barkworth (1990) “nasselloid” fossils are present in Late Miocene-Early Pliocene (c. 13-5 mybp) deposits in the USA. These have Nassella-like lemma epidermal patterns, and were considered to be Nassella by Thomasson (1986). Other described fossil stipoids from the USA include Oligocene and Miocene Stipidium and Stipa and Oligocene Piptochaetium (Thomasson 1986). Barkworth and Torres (2001) appeared to accept four North American fossil Nassella species, but point out that only a single Nassella sp. is today present in the areas of Colorado, Kansas and Nebraska where the fossils were found. Tsvelev (1977) accepted lower Miocene fossils as Nassella and Piptochaetium spp. along with other grasses, indicating that prairie grasslands existed at that time. Johnson (1972) prematurely considered this fossil record provided geological evidence of a stipoid evolutionary ‘hot-spot’ in the high plains of Nebraska during the Tertiary and argued for a North American focal point of polyploidisation. Barkworth and Everett (1986) considered that American stipoids consisted of groups derived directly from Gondwana, such as Nassella (in the narrow sense) and Piptochaetium, and from a separate independent group that initially occupied Eurasia, notably Achnatherum. According to Barkworth (1990) citing Tsvelev (1977), the North American Stipa sens. lat. evolved from South American taxa, and European taxa from North American, however this is not evident in my reading of Tselev (1977), who argued that stipoids in South America evolved in parallel with those in North America and Eurasia over a very long period prior to the Pliocene, and that they first evolved around the Tethys sea between Africa and Eurasia when all the continental masses were joined in Pangaea (i.e. in the Triassic period c. 200 mybp). Tsvelev (1977) thought that evolution of the Stipeae in all areas simultaneously involved elongation of the spikelet and all its parts, and the lengthening of the awns, usually correlated with elongation of the glume apices to prevent premature shedding of the floret. Based on anatomical and distributional data, he had no doubt that Stipa sens. lat. evolved before the formation of lowland grasslands and considered Achnatherum to be the most primitive stipoid genus. 25
Page 1 and 2: Literature review: Impact of Chilea
Page 3 and 4: Fire 49 Other disturbances 50 Shade
Page 5 and 6: Conventions and standards Botanical
Page 7 and 8: neesiana appears to be a habitat ge
Page 9 and 10: population densities of existing sp
Page 11 and 12: elated plants have similar defences
Page 13 and 14: For invasion to occur there must be
Page 15 and 16: As yet there appears to be no evide
Page 17 and 18: experimental manipulation of specie
Page 19 and 20: species tend to be those which tran
Page 21 and 22: Taxonomy and nomenclature Stipeae N
Page 23: Vernacular names ‘Needlegrass’
Page 27 and 28: 1 m (Walsh 1994), ca. 90 cm (Verloo
Page 29 and 30: Figure 2. Anatomy of the seed of N.
Page 31 and 32: are the seeds larger/smaller, longe
Page 33 and 34: also based on a misunderstanding of
Page 35 and 36: Table 2. Modified Feekes Scale for
Page 37 and 38: Argentina, in the provinces of Chac
Page 39 and 40: Figure 3. Recorded distribution of
Page 41 and 42: 1994). Only 3 of 186 exotic grasses
Page 43 and 44: According to Morfe et al. (2003) th
Page 45 and 46: populations have been found in the
Page 47 and 48: (Honaine et al. 2006). The flechill
Page 49 and 50: In Australia the altitudinal range
Page 51 and 52: Proximity to urban development appe
Page 53 and 54: In the southern Brazilian campos of
Page 55 and 56: arundinacea (Gardener et al. 2005).
Page 57 and 58: al. 2008). Cues for masting may be
Page 59 and 60: Approximately 200 alien grass speci
Page 61 and 62: Dispersal of seed in contaminated s
Page 63 and 64: In New Zealand, Hurrell et al. (199
Page 65 and 66: No emergence was observed in undist
Page 67 and 68: and high impact (“ability to caus
Page 69 and 70: also noted that despite a wide rang
Page 71 and 72: a small reduction in seedhead produ
Page 73 and 74: Slashing and mowing Slashing can re
Page 75 and 76:
Themeda re-establishment McDougall
Page 77 and 78:
species (Lawton and Schroder 1977 p
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y increased importance of ant grani
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BIODIVERSITY “Biodiversity ... on
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According to Woods (1997 p. 61) “
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2004, Richardson and van Wilgen 200
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negative depending on the particula
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Competition with native plants Comp
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asexual seed production, so local f
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GRASSLANDS Grasses: “... the most
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susceptible to N. neesiana invasion
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Floristic composition, vegetation s
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proportion of the flora then presen
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tussock space (Stuwe and Parsons 19
Page 103 and 104:
Like vascular plant diversity, comm
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Opinions differ on the nature and i
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Species Common Name Family Aust ACT
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in shifting the distribution, exten
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of these systems is largely explain
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pasture. The least understood trans
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tend to benefit more from relaxed c
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Bovids crop their food between the
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are therefore less likely to distur
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esult in a “short-term flush” o
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Fire effects on weeds Moore (1993 p
Page 125 and 126:
Table 8. Typical nutrient levels in
Page 127 and 128:
grasses produced sigificantly more
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Fossorial vertebrates are or were o
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(Rosengren 1999). Approximately one
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Table 12. Areal extent and conserva
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Foreman (1997) investigated the eff
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Willis (1964) considered that the f
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Austrostipa-Enneapogon) from around
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Woodlands and New England Grassy Wo
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Thylogale billardierii), Peramelida
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Its original habitat on the mainlan
Page 147 and 148:
Table 17. Endangered reptile specie
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equirement, but unlike plants and v
Page 151 and 152:
e assigned the same biodiversity sc
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Nematodes are mostly minute animals
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found in all mainland states, O. co
Page 157 and 158:
Keyacris scurra, Melbourne (1993) o
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was once widespread in south- easte
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eing sluggish and wingless, and exi
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estoration and, if Australia follow
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close to the plant are able to bury
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Species *Chirothrips mexicanus Craw
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Table A2.1 (continued) Species Life
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Table A2.1 (continued) Species *Het
Page 173 and 174:
Page 175 and 176:
Page 177 and 178:
Nematodes of grasses and grasslands
Page 179 and 180:
REFERENCES Aceñolaza, F.G. (2004)
Page 181 and 182:
Benson, D. and McDougall, L. (2005)
Page 183 and 184:
Chan, C.W. (1980) Natural grassland
Page 185 and 186:
DNRE (Department of Natural Resourc
Page 187 and 188:
Fuhrer, B. (1993) A Field Companion
Page 189 and 190:
Groves, R.H. and Whalley, R.D.B. (2
Page 191 and 192:
Iaconis, L. (2004) Chilean needle g
Page 193 and 194:
Levine, J.M., Adler, P.B. and Yelen
Page 195 and 196:
McDougall, K.L. (1987) Sites of Bot
Page 197 and 198:
Morfe, T.A., McLaren, D.A. and Weis
Page 199 and 200:
Perelman, S.B., León, R.J.C. and O
Page 201 and 202:
Saunders, D.A. (1999) Biodiversity
Page 203 and 204:
Thellung, A. (1912) La flore advent
Page 205 and 206:
Weiss, J. and McLaren, D. (2002) Vi
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