L.M. Hanlon and M.B. Mesgaran
Faculty of Science, Department of Resource Management and Geography, The University of Melbourne, Parkville, Victoria 3010, Australia.
Cite this article as:
Hanlon, L.M. and Mesgaran, M.B. (2014). The Biology of Australian Weeds 63. Thinopyrum junceiforme (Á. Löve & D. Löve) Á. Löve. Plant Protection Quarterly 29(4), 120-126.
The genus name Thinopyrum is derived from the Greek ‘thino; this’ for shore, and ‘pyros’, for wheat (Löve 1984), and the species name junceiforme is derived from the Latin ‘junceus’ for rush-like (Stearn 2004) and ‘formis; forme’ for resembling, shaped-like (Gledhill 2002).
The common name for Thinopyrum junceiforme (Figure 1) is sea wheatgrass. It is also referred to as sand couch-grass, sea couch (Hilton et al. 2006), and Russian wheatgrass (United States Department of Agriculture n.d.).
The genus Thinopyrum comprises tetraploid plants (2n = 4x = 28) (Löve 1980, Jauhar and Peterson 2001). It is a member of the tribe of Triticeae (family Poaceae), with a number of the species being resistant to drought, salinity and disease (Niento-Lopez et al. 2000). There are three species complexes within the genus Thinopyrum: T. elongatum (tall wheatgrass) (Host) D.R. Dewey, T. intermedium (intermediate wheatgrass) (Host) Barkworth & Dewey and T. junceum (Russian wheatgrass) (L.) Á. Löve. The taxa in T. junceum have been categorized variously as species or subspecies in different genera such as Triticum, Agropyron, Elymus, Elytrigia or Thinopyrum (Moustakas et al. 1986). There are many intergeneric hybrids which complicate the taxonomy of wheatgrasses (Refoufi and Esnault 2006).
Thinopyrum junceiforme falls within the complex of T. junceum, and is found around the Atlantic and Baltic coasts (Niento-Lopez et al. 2003). Nomenclature is complicated, with the complex T. junceum also being denominated as Agropyron junceum (L.) P. Beauv., A. junceum ssp. mediterraneum Simonet & Guin., Elytrigia juncea (L.) Nevski, and Elymus farctus (Viv.) Runemark ex Melderis (Niento-Lopez et al. 2003). Tetraploid Leymus spp. have also been confused with Thinopyrum, as both are perennial, wild relatives of wheat (Zhang and Dvorak 1991, Merker and Lantai 1997). Leymus occurs naturally in North America and Eurasia (Zhang and Dvorak 1991), and lacks the J genome of Thinopyrum (Wang and Jensen 1994). Thinopyrum junceiforme has also been denominated as Agropyron junceiforme (Á. Love & D. Love), A. junceum ssp. boreoatlanticum Simonet & Guin., Elytrigia junceiformis Á. Love & D. Love, and, like T. junceum, has also been known as Elymus farctus (Niento-Lopez et al. 2003). There is a clear morphological differentiation between the genera Thinopyrum and Elymus, with the former having rudimentary awns or lacking them altogether, whereas Elymus has wider laminae, and higher spike densities than Thinopyrum (Löve 1984, Niento-Lopez et al. 2000). The visual identification point for Thinopyrum junceiforme is a short, hirsute ligule that can be observed by pulling back the leaf blade at the collar region (Figure 2).
The currently accepted name for the species is Elytrigia juncea ssp. boreoatlantica (Simonet & Guin.) Hyl. (Valdes et al. 2009). However in this review, it is referred to by one of its heterotypic synonyms, Thinopyrum junceiforme, which is the name used in the most recent Australian and New Zealand literature, for example Heyligers (2006), Hilton et al. (2006) and Hilton et al. (2007).
The coastal dune flora of Australia changed rapidly from the late 1800s, following the introduction of South African, American and European plants such as Ammophila arenaria (L.) Link (marram grass), Cakile maritima Scop. (European sea rocket), Cakile edentula (Bigelow) Hook. (sea rocket), and Chrysanthemoides monilifera subsp. rotundata (L.) Norl. (bitou bush) (Hilton et al. 2006). Prior to that, Australia had only a few species that were capable of colonizing the area between the swash and foredune such as Atriplex billardierei (Moq.) Hook.f. (glistening saltbush) and Spinifex sericeus R.Br. (hairy spinifex) (Hilton et al. 2006). Sea wheatgrass now exerts a great influence over the establishment of both incipient dunes and foredunes, adversely affecting hairy spinifex colonies (Heyligers 1985, Hilton et al. 2006). Furthermore, in the foredunes, marram grass has displaced Austrofestuca littoralis (Labill.) E.B. Alexeev (beach fescue), and created steep, unstable dunes (Heyligers 1985, Hilton et al. 2006).
Sea wheatgrass was first recorded in Australian herbarium records in 1933, with a specimen collected from Ricketts Point, Victoria (MEL 0626849A). It may well have arrived much earlier, in ballast or cargo from the Windjammers, or sailing vessels, which plied between Europe and southern Australia from the 1830s through to 1950 (South Australia Maritime Museum n.d.). Australian herbarium records show that the first sea wheatgrass collections from other states were from Rocky Cape, Tasmania in 1948 (HO77017) and the Long Beach sand dunes, South Australia in 1983 (AD98409214).
Sea wheatgrass originates from the western European, Mediterranean, Atlantic and Baltic coasts (Figure 3) (Niento-Lopez et al. 2000, Heyligers 2006, Hilton et al. 2007), where it plays a major role in dune establishment (Heyligers 2006) as a pioneer species in embryo and incipient sand dunes (Harris and Davy 1986a, Hilton et al. 2007). In active foredunes of northwest Europe, sea wheatgrass is one of the few plants that can withstand periodic, temporary sand burial (Doody 2013), thereby creating dunes that are rarely higher than 1 m in elevation (Hilton et al. 2006). The plant has naturalized in Oregon and California on the west coast of the United States of America, where it is considered a native plant (United States Department of Agriculture n.d.). In Britain, sea wheatgrass is regarded as a primary dune colonizer, where it develops dense swards on pre-established foredunes (Harris and Davy 1986a). Sea wheatgrass is also found in southern New Zealand (Hilton et al. 2006).
In Australia, sea wheatgrass is found on the foredunes and incipient dunes of the southern coast of the mainland, from Henley Beach, Adelaide (34.92°S, 138.49°E) along the coast and inland to Naracoorte, on the South Australian border with Victoria (38.05°S, 140.94°E), and on the northern coast of Tasmania (Figure 3). In Victoria, it appears from the mouth of the Anglesea River (38.41°S, 144.18°E), around Port Phillip and Westernport Bays, including Phillip Island, and down to Wilsons Promontory National Park (39.06°S, 146.41°E) (Figure 3). Records do not indicate that it grows along the eastern coast of the mainland, except at Cape Conran Coastal Park, East Gippsland (37.79°S, 148.74°E). In Tasmania, sea wheatgrass is recorded from the mouth of Bottle Creek on the north west coast (41.1°S, 144.66°E) across to Cape Portland on the north east coast (40.75°S, 147.96°E) and it is also found on Flinders Island (40.00°S, 148.11°E) (Figure 3). The specimens recorded in the Australian Capital Territory are related to four samples collected from The Netherlands between 1962 and 1987, and grown in experimental plots (Australia’s Virtual Herbarium –b).
Sea wheatgrass is endemic across a wide latitudinal range in the Northern Hemisphere, extending from Finland (64°N, 26°E), to Spain’s Cadiz region (36.5°N, 6.28°W) (Hilton et al. 2006). As with many coastal species, the grass occupies a niche of high temperatures, high salinity, desiccation and abrasion from winds, and extremes of soil moisture content (Hesp 1991, Ievinsh 2006, Maun 2009). Additional factors in this dynamic niche include high light intensity and nutritional deficiencies (Hesp 1991, Martinez et al. 2001). Harris and Davy (1986a) argue that in their natural habitat, Elymus farctus (Thinopyrum junceiforme) tillers need vernalization in order to flower. Field observations from research in Australia note that the production of inflorescences is limited, and this may be because Australian coastal winter temperatures are not as low as those in Britain (M.B. Mesgaran unpublished data).
Sand dune soils are generally poor in the macronutrients, nitrogen (N), phosphorus (P) and potassium (K) (Hawke and Maun 1988, Zhang 1996). In Victoria, the coastal sediments are carbonate-dominated west of Wilsons Promontory and silicon oxide-dominated eastward (Bird 1993). Literature on the nutritional status of Australian dune sands is lacking. However, in a coastal dune system on the west coast of the South Island of New Zealand, Sykes and Wilson (1991) found that the alkaline sand had low P levels, ranging between 4.8 µm mL-1 to 6.6 µm mL-1, but N levels were not included as they were <0.05%. As with other beach systems, there would be substantial variations in soil nutrients (Hawke and Maun 1988, Zhang 1996, Perumal and Maun 2006), where for example, the deposition of wrack, or organic matter, on beaches from wave action would periodically raise nutrient levels (Zhang 1996). Therefore, sea wheatgrass needs to tolerate both low, and fluctuating, levels of nutrients, and also needs to be tolerant of a wide range of conditions from inundation by sea water and mobile sand. In Australia, the plant grows lower on the beach, and closer to the swash than any native species (Hilton et al. 2006); this could be because no other plants can survive repeated seawater inundation unless they are halophytes.
Sea wheatgrass is one of three species in the genus Thinopyrum that are salt tolerant, along with tall wheatgrass and T. bessarabicum (Savul. & Rayss) Á. Love, and this tolerance is controlled by multiple genes on several chromosomes (Wang et al. 2003). Thinopyrum bessarabicum, for instance, has been found to withstand hydroponic solutions of 350 mM NaCl for prolonged periods (Gorham et al. 1985), whereas halophytes grow in concentrations of 400 mM NaCl, or higher (Flowers 2004).
Both native and exotic plants grow in association with sea wheatgrass on incipient dunes and foredunes in Victoria (L.M. Hanlon personal observations). On the incipient dunes, such plants may include native hairy spinifex, and the exotics Cakile spp. (sea rockets) and Euphorbia paralias L. (sea spurge). On the foredunes, the vegetation is more varied and includes the native plants Atriplex cinerea Poiret X A. paludosa (coast saltbush), and Ficinia nodosa (Rottb.) Goetgh., Muasya & D.A. Simpson (knobby club rush), with Lepidosperma gladiatum Labill. (coast sword-sedge) growing in the swale in the lee of incipient dunes. On open sites on the foredunes, hardy succulents such as native and naturalized Carpobrotus spp. (pigface), and native plants Rhagodia baccata (Labill.) Moq. (seaberry saltbush) and Tetragonia tetragonioides (Pall.) Kuntze (sea spinach), manage the inhospitable environment remarkably well, as does the native Geranium solanderi var. solanderi Carolin (native geranium). Exotic weed species such as Fumaria spp. (fumitory), Oxalis spp. (soursob) and Allium triquetrum L. (angled onion) are just a few of the smaller flowering plants, also found in association with sea wheatgrass on the foredunes.
Growth and development
Morphology and physiology
Sea wheatgrass is a rhizomatous, perennial grass, growing to approximately 50 cm in height, but under favourable conditions it can grow as tall as 80 cm. Plants can grow from a single node and produce as many as 20–100 tillers. The blue-green leaves of sea wheatgrass are glabrous below and finely pubescent above, usually 30 cm in length, but they can grow up to 50 cm, with the widest part of the blade varying from 3 mm to 8 mm (average 5 mm) (M.B. Mesgaran unpublished data). Most populations from South Australia have wider leaves than those of Victoria or Tasmanian populations (M.B. Mesgaran unpublished data). Spike lengths are about 15 cm, with each spike bearing 10 spikelets. Sea wheatgrass is a C3 cool-season grass, but other data describing its physiology or biochemistry are lacking. Notwithstanding this, temporary burial is a common occurrence in plants of sandy environments, and many plants such as sea wheatgrass survive such burial, with the short-term suspension of physiological activity such as photosynthetic capacity, which is quickly reinstated once uncovered (Harris and Davy 1988, Perumal and Maun 2006). This is due to newly-emerged leaves from previously buried plants having an increased chlorophyll content (mg g-1 fresh weight), and a higher energy content in subterranean organs (Yuan et al. 1993, Perumal and Maun 2006).
Young plants of sea wheatgrass lack the axillary meristems and energy reserves required to grow new shoots when buried, and they re-allocate their resources from non-photosynthetic organs to maintain the photosynthetic ones, until the plant is uncovered by winds or storms (Harris and Davy 1988). It has been shown that multi-node rhizome fragments have more success in emergence, and from greater depths, than do single-node fragments of sea wheatgrass (Harris and Davy 1986b).
In Britain, sea wheatgrass requires vernalization in order to flower; however flowering usually occurs in the second year of growth, and Harris and Davy (1986a) found that such flowering was limited due to the proximity of plants to wave disturbance along the swash, as well as grazing by rabbits. It is hypothesized that as the winters in Australia are not as cold as those in Europe, flowering is limited, appearing in December, January, and occasionally in February (Figure 4) (M.B. Mesgaran unpublished data), although the flowers do not persist for long on the stems (Rudman 2003).
The seasonal variation in the ability of sea wheatgrass rhizome buds to produce adventitious shoots and roots under glasshouse conditions was found to be strongly correlated to nitrogen as a limiting factor, with carbohydrate reserves also implicated (Harris and Davy 1986b). Additionally, dormancy was found to show a peak in late winter and early spring, with a sharp decline in late spring-early summer (Harris and Davy 1986b). This variation was inversely related to the growth rate of the parent plants at time of harvesting (Harris and Davy 1986b).
Approximately half of the 150 described species of arbuscular mycorrhizal (AM) fungi are found in sand dunes (Koske et al. 2004). In particular, Glomus intraradices N.C. Schenck & G.S. Sm. a species of AM fungi, was shown to be highly tolerant of harsh conditions such as aridity and salinity, reducing the concentration of sodium in the shoots of plants in saline environments (Yamato et al. 2012). However, there is no literature specifically on the involvement of AM fungi with sea wheatgrass, although grasses in general tend to be facultatively mycorrhizal (Brundrett et al. 1996), or weakly mycorrhizal (Ramos-Zapata et al. 2011) in their association with AM fungi. The early work of Forster (1979) which focussed on aggregation of sand by plants and microbes, asserted that microbial abundance associated with sea wheatgrass (under the heterotypic synonym of Agropyron junceiforme) was less in winter when annual species were dying, and many perennial species were dying back.
Sea wheatgrass can reproduce both sexually and asexually (Löve 1984), but production of flowering tillers in Britain (Harris and Davy 1986a) and mature seeds in Australia (M.B. Mesgaran unpublished data) is low. In Australia, the main route of propagation is by rhizome growth and fragmentation. The plant has cells with four sets of chromosomes (Merker and Lantai 1997, Jauhar and Peterson 2001), with the genome of J1J1J2J2 (Colmer et al. 2006), and is a self-crosser, although it is capable of cross-fertilization (Moustakas et al. 1986). Refoufi and Esnault (2006) contend that it has little genetic diversity, as assessed by isozymes and RAPD markers.
Hydrochory, or the passive dispersal of propagules by water, is the most likely vector for the dispersal of sea wheatgrass caryopses along the coastline of southern Australia. Drift bottle programmes by Olsen and Shepherd (2006) demonstrated that surface water flows along the South Australian coast, east through Bass Strait, then south east past the west coast of Tasmania in winter, reversing the direction in summer. Thus, currents could be responsible for the transportation of plant fragments to the shores of Tasmania from the mainland. Heyligers (2007) noted that dispersal patterns of four other introduced species to Australian beaches follow the circulation of ocean currents around Australia, and such currents may have been responsible for the dispersal of propagules to the South Australian coast. Although no research has been conducted on the dispersal mechanisms of sea wheatgrass, it is likely to be via such fragments being scarped, or torn, from eroding sand dunes during storms (Hilton et al. 2006, Hilton et al. 2007). For example, following a severe storm on the Norfolk (UK) coast in 1978, sea wheatgrass (under the heterotypic synonym Elymus farctus), rapidly re-colonized the swash, and both seeds and fragments of rhizomes were of equal importance in establishing new clumps, and in producing similar tiller densities (Harris and Davy 1986a). Similar recruitment was observed in the 1960s at Shallow Inlet, Wilsons Promontory where a sand spit developed after the previous spit was washed out in 1901. The new spit initially lacked vegetation but in the 1960s, sea wheatgrass was found to have colonized it (Heyligers 2006). Clumps of sea wheatgrass rhizomes have also been observed at the mouth of the Glenelg River at Nelson in Victoria, where there was no actively growing parent plant on the beach, suggesting that the rhizomes might have been dispersed by hydrochory (M.B. Mesgaran unpublished data).
Physiology of seeds and germination
Woodell (1985) suggested that seed germination patterns of coastal plants can be separated into three categories and that their response to germination in saline conditions correlated with their habitat. He found that the greatest germination response of sea wheatgrass seeds (n=180) was in freshwater (53%), followed by 18% in half-strength sea water, 5% in full-strength seawater, and the complete inhibition (0%) in one and a half strength sea water (sodium chloride (NaCl) concentration 600 mM) (Woodell 1985). Seeds from all treatments recovered sufficiently when transferred to distilled water for some germination to occur (Woodell 1985). Even though the seeds germinated under controlled conditions, it is likely that under natural conditions, sea wheatgrass seeds in the swash may be stimulated to germinate following precipitation. That sea wheatgrass seeds do not germinate in full-strength sea water may also aid in its dispersal by keeping seeds dormant and the embryo surviving on the endosperm within the seed coat until the seed reaches land.
In Australia, reproduction is largely vegetative from new shoots off nodes along the highly meristematic rhizomes, which are produced in great lengths (Figure 5). Under glasshouse conditions, each 5 cm node of sea wheatgrass produced up to 30 m rhizome length in the course of one season (M.B. Mesgaran unpublished data). The length of most internodes was approximately 7 cm, and skewed toward lengthier internodes (Figure 6).
On land, single-node sea wheatgrass rhizome fragments can emerge from depths of up to 17 cm. Multi-node fragments can emerge from greater depths (>17 cm), producing more emergent shoots and more quickly than single-node shoots, especially in late winter to early spring (Harris and Davy 1986b).
Population dynamics data for sea wheatgrass in Australia are not available. The maximum life-span of the species is unknown, as are mortality rates between life stages, apart from the rate of regeneration after dispersal by water, or burial by sand, discussed above. Research is required on population dynamics to enable the development of weed management strategies, rather than ad hoc herbicide applications. Such strategies could be used by all beach management authorities.
In Victoria, anecdotal evidence suggests that sea wheatgrass is impacting upon the rookery of Eudyptula minor J.R. Forster (fairy penguin) on Phillip Island’s Summerlands beach, by creating steep-fronted incipient dunes that are too high for the birds to climb in order to access their burrows (P. Dann personal communication 2013). Furthermore, on Phillip Island and beaches in Victoria such as those on the Barwon Coast and Geelong, the endangered Thinornis rubricollis Gmelin (hooded plover), which prefers a nest scrape with little vegetation on which to lay its eggs, is likely to have its nesting sites encroached upon by sea wheatgrass (Cousens et al. 2012). Steep-fronted incipient dunes that are thickly vegetated with sea wheatgrass are also found along the South Australian coast, such as at Normanville (Figure 7).
Hilton et al. (2006) proposed that the sand-binding ability of sea wheatgrass makes it more resilient to erosive processes in comparison to native flora. Indeed, sea wheatgrass is one of four exotic species that were noted by Heyligers (1985) as being more efficient than native species at trapping sand and building dunes where otherwise dunes would not have formed. Such dunes have the potential to limit sediment movement, thereby changing the ecosystems and geomorphology of the coastlines on which they appear. Thus it is of concern that sea wheatgrass can rapidly colonize the swash and incipient dunes after propagules are washed ashore following storms. One example where sea wheatgrass has spread with great rapidity is along the Younghusband Peninsula in South Australia, where James (2012) reports that the plant has spread at approximately 18 ha per year, outcompeting native species and altering ecosystems.
Sea wheatgrass is a potential gene source for salt tolerance in wheat (Wang et al. 2003), which has been investigated in Triticeae in general, as well as in T. bessarabicum (Gorham et al. 1985, Gorham et al. 1986). Sea wheatgrass has also been investigated as a potential gene source for scab resistance in wheat (Jauhar and Peterson 2001).
Groves et al. (2003) list sea wheatgrass as a weed species, albeit not specifically a coastal weed, as there is no formal listing or category for coastal weed species. Cousens et al. (2012) note that sea wheatgrass is one of the dominant exotic species on the coast displacing native vegetation. However, many coastal managers, including state government co-ordinators, are unaware of the existence of sea wheatgrass, let alone its potentially adverse impacts (R. Cousens personal communication 2014), which may explain the lack of specific legislation related to this species.
Phillip Island Nature Parks (PINP) in Victoria, has used Verdict™ 520, Fusilade™, Starane™, glyphosate and metsulphuron-methyl at the recommended rate for similar plants, as well as lower rates (J. Fallow personal communication 2014). Staff at PINP report that the lower rates of herbicides were more efficient than the recommended rate, as it is believed that lower rates allow for translocation of the herbicide throughout the plant, rather than killing the aerial parts alone via one larger application.
The Barwon Coast Foreshore Committee of Management, Victoria, has also used Verdict™ 520. The application rate was 50 mL of Verdict 520 per 100 L of water, plus 500 mL of Uptake™ spray oil added to the product, as an adjuvant to improve the spreading and wetting qualities of the herbicide (W. Chapman personal communication 2014). It has been reported that this treatment is making an impact on sea wheatgrass, with minor off-target damage to native grasses (W. Chapman personal communication 2014). Additionally, it has been found that two applications, 2–4 weeks apart, as suggested by the manufacturer of Verdict™ 520 for use on other invasive grasses, were not required for the suppression of sea wheatgrass (W. Chapman personal communication 2014).
On Phillip Island, and around the Barwon Coast, Theba pisana (Müller) (sandhill snail) has been observed by the authors to graze on sea wheatgrass, causing damage to leaves. However this invertebrate is not specific to sea wheatgrass or coastal areas, and is also found in land crops in Australia (Fox 2011).
We are grateful for the Elizabeth Ann Crespin Scholarship (2013), which partially funded the work by Lynda M. Hanlon. We thank Roger Cousens, The University of Melbourne, for discussions about coastal weeds. We also thank PINP for access to beaches to study the plant and to Jon Fallaw for information on herbicide usage on Summerlands Beach, Phillip Island. We are appreciative of discussions with Warren Chapman, Barwon Coast Foreshore Committee of Management, regarding detailed application rates of herbicides used on the Barwon Coast, and for access to beaches to study the plant.
Australia’s Virtual Herbarium (n.d. –a). Map of the distribution of Thinopyrum junceiforme in Australia. http://avh.ala.org.au/occurrences/search?taxa=Thinopyrum+junceiforme#tab_mapView (accessed 22 January 2014).
Australia’s Virtual Herbarium (n.d. –b). Thinopyrum junceiforme. http://avh.ala.org.au/occurrences/search?taxa=Thinopyrum+junceiforme#map (accessed 15 October 2014).
Bird, E.C.F. (1993). ‘The coast of Victoria: the shaping of scenery’. (Melbourne University Press, Melbourne). 324 pp.
Brundrett, M., Bougher, N., Dell, B., Grove, T. and Malajczuk, N. (eds) (1996). ‘Working with mycorrhizas in forestry and agriculture’. (Australian Centre for International Agricultural Research, Canberra).
Colmer, T.D., Flowers, T.J. and Munns, R. (2006). Use of wild relatives to improve salt tolerance in wheat. Journal of Experimental Botany 57, 1059-78.
Cousens, R., Kennedy, D., Maguire, G. and Williams, K. (2012). ‘Just how bad are coastal weeds? Assessing the geo-eco-psycho-socio-economic impacts’. (Australian Government, Rural Industries Research and Development Corporation, Canberra).
Doody, J.P. (2013). ‘Sand dune conservation, management and restoration’. (Springer, Dordrecht, Netherlands). 303 pp.
Flowers, T.J. (2004). Improving crop salt tolerance. Journal of Experimental Botany 55, 307-19.
Forster, S.M. (1979). Microbial aggregation of sand in an embryo dune system. Soil Biology and Biochemistry 11, 537-43.
Fox, E. (2011). Mediterranean snails Theba pisana. White snails, Italian snails, coastal snails, sand hill snail. http://corangamite.landcarevic.net.au/lismore-land-protn/publications/mediterranean-snails-information-and-control (accessed 18 December 2014).
GBIF, Global Biodiversity Information Facility, (n.d.) Global distribution of Thinopyrum juncieforme. http://data.gbif.org/ (accessed 3 February 2014).
Gledhill, D. (2002). ‘The names of plants.’ Third edition. (Cambridge University Press, Cambridge). 326 pp.
Gorham, J., Budrewicz, E., McDonnell, E. and Jones, R.G.W. (1986). Salt tolerance in the Triticeae: Salinity-induced changes in the leaf solute composition of some perennial Triticeae. Journal of Experimental Botany 37, 1114-28.
Gorham, J., McDonnell, E., Budrewicz, E. and Jones, R.G.W. (1985). Salt tolerance in the Triticeae: growth and solute accumulation in leaves of Thinopyrum bessarabicum. Journal of Experimental Botany 36, 1021-31.
Groves, R.H., Hosking, J.R., Batianoff, G.N., Cooke, D.A., Cowie, I.D., Johnson, R.W., Keighery, G.J., Lepschi, B.J., Mitchell, A.A., Moerkerk, M., Randall, R.P., Rozefelds, A.C., Walsh, N.G. and Waterhouse, B.M. (2003). ‘Weed categories for natural and agricultural ecosystem management’. (Bureau of Rural Sciences, Australian Government Department of Agriculture, Fisheries and Foresty, Canberra). 194 pp.
Harris, D. and Davy, A.J. (1986a). Strandline colonization by Elymus farctus in relation to sand mobility and rabbit grazing. Journal of Ecology 74, 1045-56.
Harris, D. and Davy, A.J. (1986b). Regenerative potential of Elymus farctus from rhizome fragments and seed. Journal of Ecology 74, 1057-67.
Harris, D. and Davy, A.J. (1987). Seedling growth in Elymus farctus after episodes of burial with sand. Annals of Botany 60, 587-93.
Harris, D. and Davy, A.J. (1988). Carbon and nutrient allocation in Elymus farctus seedlings after burial with sand. Annals of Botany 61, 147-57.
Hawke, M.A. and Maun, M.A. (1988). Some aspects of nitrogen, phosphorus, and potassium nutrition of three colonizing beach species. Canadian Journal of Botany 66, 1490-6.
Hesp, P.A. (1991). Ecological processes and plant adaptations on coastal dunes. Journal of Arid Environments 21, 165-91.
Heyligers, P.C. (1985). The impact of introduced plants on foredune formation in south-eastern Australia. In ‘Are Australian ecosystems different? Proceedings of a Symposium, 1984.’.. eds J.R. Dodson and M. Westoby, pp. 23-41 (Ecological Society of Australia, Sydney).
Heyligers, P.C. (2006). Primary vegetation development on the sand spit of Shallow Inlet, Wilsons Promontory, southern Victoria. Cunninghamia 9, 571-96.
Heyligers, P.C. (2007). The role of currents in the dispersal of introduced seashore plants around Australia. Cunninghamia 10, 167-88.
Hilton, M., Harvey, N., Hart, A., James, K. and Arbuckle, C. (2006). The impact of exotic dune grass species on foredune development in Australia and New Zealand: a case study of Ammophila arenaria and Thinopyrum junceiforme. Australian Geographer 37, 313-34.
Hilton, M., Harvey, N. and James, K. (2007). The impact and management of exotic dune grasses near the mouth of the Murray River, South Australia. Australasian Journal of Environmental Management 14, 220-30.
Ievinsh, G. (2006). Biological basis of biological diversity: physiological adaptation of plants to heterogeneous habitats along a sea coast. Acta Universitatis Latviensis 710, 53-79.
James, K.F. (2012). Gaining new ground: Thinopyrum junceiforme, a model of success along the south eastern Australian coastline. PhD thesis, School of Social Sciences, University of Adelaide, Adelaide.
Jauhar, P.P. and Peterson, T.S. (2001). Hybrids between durum wheat and Thinopyrum junceiforme: Prospects for breeding for scab resistance. Euphytica 118, 127-36.
Koske, R.E., Gemma, J.N., Corkidi, L., Siguenza, C. and Rincon, E. (2004). Arbuscular mycorrhizas in coastal dunes. In ‘Ecological studies Volume 171, Coastal dunes, ecology and conservation’, eds M.L. Martinez and N.P. Psuty, pp. 173-87. (Springer-Verlag, Berlin).
Löve, Á. (ed.) (1980). Chromosome number reports: LXVII. Taxon 29, 347-67.
Löve, Á. (1984). Conspectus of Triticeae. Feddes Repertorium 95, 425-521.
Martinez, M.L., Vazquez, G. and Sanchez, C.S. (2001). Spatial and temporal variability during primary succession on tropical coastal sand dunes. Journal of Vegetation Science 12, 361-72.
Maun, M.A. (2009). ‘The biology of coastal sand dunes’. (Oxford University Press, New York). 265 pp.
Merker, A. and Lantai, K. (1997). Hybrids between wheats and perennial Leymus and Thinopyrum species. Acta Agriculturae Scandinavica Section B-Soil and Plant Science 47, 48-51.
Moustakas, M., Symeonidis, L. and Coucoli, H. (1986). Seed protein electrophoresis in Agropyron junceum (L.) P.B. complex. Annals of Botany 57, 35-40.
Niento-Lopez, R.M., Casanova, C. and Soler, C. (2000). Evaluation and characterisation of a collection of wild Spanish populations of the genera Elymus and Thinopyrum using morphological and agronomical traits. Agronomie 20, 111-22.
Niento-Lopez, R.M., Soler, C. and Garcia, P. (2003). Genetic diversity in wild Spanish populations of Thinopyrum junceum and Thinopyrum junceiforme using endosperm proteins and PCR-based markers. Hereditas 139, 18-27.
Olsen, A.M. and Shepherd, S.A. (2006). Historic drift bottle experiments show reversing surface water masses in western Bass Strait waters: implications for lobster larval dispersal. Transactions of the Royal Society of South Australia 130, 113-22.
Perumal, V.J. and Maun, M.A. (2006). Ecophysical response of dune species to experimental burial under field and controlled conditions. Plant Ecology 184, 89-104.
Ramos-Zapata, J.A., Zapata-Trujillo, R., Ortiz-Diaz, J.J. and Guadarrama, P. (2011). Arbuscular mycorrhizas in a tropical coastal dune system in Yucatan, Mexico. Fungal Ecology 4, 256-61.
Refoufi, A. and Esnault, M.-A. (2006). Genetic diversity and population structure of Elytrigia pycnantha (Godr.) (Triticeae) in Mont Saint-Michel Bay using microsatellite markers. Plant Biology 8, 234-42.
Rudman, T. (2003). Tasmanian beach weed strategy for marram grass, sea spurge, sea wheatgrass, pyp grass and beach daisy. (Department of Primary Industries, Water and Environment, Hobart, Tasmania). 17 pp.
South Australia Maritime Museum. (n.d.) ‘The last of the mosquito fleet’. http://maritime.historysa.com.au/research/projects/last-mosquito-fleet (accessed 17 December 2013).
Stearn, W.T. (2004). ‘Botanical Latin’. (Timber Press, Oregon). 560 pp.
Sykes, M.T. and Wilson, J.B. (1991). Vegetation of a coastal sand dune system in southern New Zealand. Journal of Vegetation Science 2, 531-8.
United States Department of Agriculture. (n.d.) Thinopyrum junceiforme (Á. Löve & D. Löve) Á. Löve. Russian wheatgrass. http://plants.usda.gov/core/profile?symbol=THJU3 (accessed 12 December 2013).
Valdes, B., Scholz, H., von Raab-Straube, E. and Parolly, G. (2009). Elytrigia juncea subsp. boreoatlantica http://ww2.bgbm.org/EuroPlusMed/PTaxonDetail.asp (accessed 6 January 2014).
Wang, R.R.-C. and Jensen, K.B. (1994). Absence of the J genome in Leymus species (Poaceae: Triticeae): evidence from DNA hybridization and meiotic pairing. Genome 37, 231-5.
Wang, R.R.-C., Li, X.-M., Hu, Z.-M., Zhang, J.-Y., Larson, S.R., Zhang, X.-Y., Grieve, C.M. and Shannon, M.C. (2003). Development of salinty-tolerant wheat recombinant lines from a wheat disomic addition line carrying a Thinopyrum junceiforme chromosome. International Journal of Plant Science 164, 25-33.
Woodell, S.R.J. (1985). Salinity and seed germination patterns in coastal plants. Vegetatio 61, 223-9.
Yamato, M., Yagame, T., Yoshimura, Y. and Iwase, K. (2012). Effect of environmental gradient in coastal vegetation on communities of arbuscular mycorrhizal fungi associated with Ixeris repens (Asteraceae). Mycorrhiza 22, 623-30.
Yuan, T., Maun, M.A. and Hopkins, W.G. (1993). Effects of sand accretion on photosynthesis, leaf-water potential and morphology of two dune grasses. Functional Ecology 7, 676-82.
Zhang, H.-B. and Dvorak, J. (1991). The genome origin of tetraploid species of Leymus (Poaceae: Triticeae) inferred from variation in repeated nucleotide sequences. American Journal of Botany 78, 871-84.
Zhang, J. (1996). Interactive effects of soil nutrients, moisture and sand burial on the development, physiology, biomass and fitness of Cakile edentula. Annals of Botany 78, 591-8.