Echium plantagineum (Paterson's curse)

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Plant in full flower

Echium plantagineum (Paterson's curse); plant in full flower. Portugal, April 2010.

©Joaquim Alves Gaspar, Lisboa, Portugal/CC BY-SA 3.0

Flowers

Echium plantagineum (Paterson's curse); close-up of flowers. Portugal. April, 2010.

©Joaquim Alves Gaspar, Lisboa, Portugal/CC BY-SA 3.0

Identity

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Preferred Scientific Name

Echium plantagineum L. (1771)

Preferred Common Name

Paterson's curse

Other Scientific Names

Echium latifolium Bubani (1897) nom. illeg.
Echium lycopsis L. (1754)
Echium maritimum Willd. (1798) sensu stricto
Echium murale Hill (1756) nom. illeg.
Echium plantaginoides Roem. & Schult. (1819)
Echium sennenii Pau (1924)
Echium violaceum L. (1767)

International Common Names

English: blue weed; dwarf blue bedder; Lady Campbell weed; purple bugloss; purple echium; purple viper's bugloss; riverina bluebell; salvation Jane
Spanish: borraja del campo; flor morada; flor morada; hierba azul; viborera morada
French: vipérine faux-plantain
Portuguese: soagem

Local Common Names

Austria: Blauer Natternkopf; Wegerrichblättriger Natternkopf
Brazil: borracha-chimarrona
Finland: ratamoneidonkieli
Germany: Blauer Natternkopf; Wegerrichblättriger Natternkopf
Italy: viperina piantaginea
South Africa: bloudissel; pers echium
Spain: llengua de bou; tapabraguetes

EPPO code

EHIPL (Echium plantagineum)

Summary of Invasiveness

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E. plantagineum is mainly known as a weed of pastures. It is found in parts of the world with a Mediterranean climate. Where introduced it has established and spread successfully in pastures where it can out-compete native species. It is listed as a noxious weed in all states of Australia, and described as invasive in Ethiopia, Kenya, South Africa and Tanzania, and countries of South America.

Taxonomic Tree

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Domain: Eukaryota
Kingdom: Plantae
Phylum: Spermatophyta
Subphylum: Angiospermae
Class: Dicotyledonae
Order: Boraginales
Family: Boraginaceae
Genus: Echium
Species: Echium plantagineum

Notes on Taxonomy and Nomenclature

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The nomenclature of E. plantagineum has been confused. At various times, the synonyms E. violaceum and E. lycopsis have been used. Lacaita (1919) and Ewart and Tovey (1920), for E. violaceum, and Gibbs (1971), for E. lycopsis, showed that this material was identical to E. plantagineum, the correct name for the species. Misidentifications of Echium material in the literature and herbaria have also been common. E. plantagineum has been determined incorrectly as E. creticum, E. italicum and E. vulgare (Piggin and Sheppard, 1995). Pérez Lara (1889) described var. megalanthus in Spain, but this distinction was not adopted.

Description

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E. plantagineum is a softly hairy, winter annual species which produces a bushy paniculate inflorescence of occasionally-branched unilateral cymes, often much enlarging in fruit. In sparse vegetation, E. plantagineum has a rosette of basal leaves and a more prostrate habit compared with the erect habit where plant densities are high. It has one to many erect flowering branches, 20-60 (sometimes to 200) cm tall; indumentum of appressed white setae (1-3 mm) and sometimes, especially on the stems and leaf margins, a sparse underlayer of softer, white hairs. Rosette and basal leaves 50-200+ mm long x 15-100 mm wide, broadly ovate, petiolate, with prominent lateral veins on upper and lower surfaces; cauline leaves oblong to lanceolate, the uppermost more or less cordate at the base. Calyx of five sepals, 7-10 mm at anthesis, up to 15 mm in fruit; corolla of five fused petals, 18-30 mm, broadly funnel-shaped, usually purple but also blue, pink or, very occasionally, white, glabrous except for sparse long hairs on the veins and margins; five stamens, attached to corolla, two long-exserted, two intermediate-enclosed, and one short-enclosed; filaments reddish-blue with occasional long hairs; pollen bluish-grey; style hairy, divided at the tip into two short branches each with terminal stigmas. Fruit (nutlet) 2 x 2.5 mm, triquarous, tuberculate, pale brownish-grey or black, usually in fours, three dropping readily from, and one remaining attached to, the receptacle. Seeds of E. plantagineum have a straight embryo, elliptic cotyledons, a short conical radicle, no endosperm and a hard seed coat. Individual seed weight varies from 360 to 390 mg per 100 seeds (Piggin and Sheppard, 1995). The root system usually consists of one taproot, or sometimes several taproots, branching freely into a network of finer roots. The floral formula is K(5) C(5) G(5) (Piggin and Sheppard, 1995).

Plant Type

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Annual
Broadleaved
Herbaceous
Seed propagated

Distribution

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Wapshere (1985) reported that the evolutionary centre of the genus occurs in southern Spain, Portugal and Morocco, where the genus and its natural enemies show highest diversity. The genus has a secondary evolutionary centre of mainly monocarpic and polycarpic perennial species on the Canary Islands and Madeira that developed following the arrival of ancestral Echium species (Lems and Holzapfel, 1968a,b; Carlquist, 1970). E. plantagineum originates from the western Mediterranean region, but local colonization has led to a broadly circum-Mediterranean distribution, including also England and the Channel Islands, Madeira and the Canary Islands (Gibbs, 1972).

E. plantagineum has been spread widely throughout the world and has become particularly well established in countries with Mediterranean-type climates including South Africa, southwest Russia, the western USA, Uruguay, Argentina and Brazil, and New Zealand (Holm et al., 1979). In addition to the listed distribution, there are unpublished records for Saudi Arabia and United Arab Emirates (AW Sheppard, CSIRO European Laboratory, Montferrier-sur-Lez, France, personal communication, 2005) and, in the USA, Oregon and Washington (Isaacson, personal communication, 2005).

The extent and abundance of E. plantagineum is often greater in the exotic range than in the Mediterranean regions where it originated for a number of reasons: a) extensive agriculture, particularly pasture-based, is now more commonly practised in other Mediterranean parts of the world than in Europe; b) native vegetation in the regions is usually poorly adapted to compete with E. plantagineum in the presence of livestock grazing; and c) there are relatively few native natural enemies damaging the plant in the exotic range (Piggin and Sheppard, 1995). Nonetheless, E. plantagineum can be a significant component of the flora in its native range, for example being described as a 'principal' weed in Tunisia (Holm et. al., 1979) and occasionally requiring some management in Spain (Zulueta, 1974).

E. plantagineum is now common, and often abundant, throughout southwest Western Australia, central and southern South Australia, southeast Queensland, central and southern New South Wales, Victoria, Tasmania and the Northern Territory. At the edge of its range it is mostly limited to disturbances on roadsides. In New Zealand, E. plantagineum is abundant north of the volcanic plateau of the North Island and scattered to rare elsewhere (Webb et al., 1988). It is regarded as a weed in many places in south Africa (Smith, 1964). There are few reports on the significance of E. plantagineum in pastures in other parts of the exotic range.

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Last updated: 21 Jul 2022

Continent/Country/Region	Distribution	Origin	First Reported	Invasive	Reference	Notes
Africa
Algeria	Present, Widespread	Native			CABI (Undated)	Original citation: Greuter et al. (1984)
Egypt	Present, Widespread	Native			CABI (Undated)	Original citation: Greuter et al. (1984)
Ethiopia	Present	Introduced		Invasive	Witt and Luke (2017)
Kenya	Present	Introduced		Invasive	Witt and Luke (2017)
Libya	Present, Widespread	Native			CABI (Undated)	Original citation: Greuter et al. (1984)
Morocco	Present, Widespread	Native			CABI (Undated)	Original citation: Greuter et al. (1984)
South Africa	Present, Widespread	Introduced		Invasive	Adamson and Salter (1950) Seebens et al. (2017)
Tanzania	Present	Introduced		Invasive	Witt and Luke (2017)
Tunisia	Present, Widespread	Native			CABI (Undated)	Original citation: Greuter et al. (1984)
Zimbabwe	Present	Introduced	1918		Seebens et al. (2017)
Asia
Georgia	Present	Native			CABI (Undated)	Original citation: Greuter et al. (1984)
Israel	Present, Widespread	Native			CABI (Undated)	Original citation: Greuter et al. (1984)
Jordan	Present, Widespread	Native			CABI (Undated)	Original citation: Greuter et al. (1984)
Lebanon	Present, Widespread	Native			CABI (Undated)	Original citation: Greuter et al. (1984)
Syria	Present, Widespread	Native			CABI (Undated)	Original citation: Greuter et al. (1984)
Turkey	Present, Widespread	Native			Gibbs (1972) CABI (Undated)
Europe
Albania	Present, Widespread	Native			CABI (Undated)	Original citation: Greuter et al. (1984)
Austria	Present	Introduced	1960		Seebens et al. (2017)
Belgium	Present	Introduced	1890		Seebens et al. (2017)
Bosnia and Herzegovina	Present, Widespread	Native			CABI (Undated)	Original citation: Greuter et al. (1984)
Bulgaria	Present, Widespread	Native			Gibbs (1972) CABI (Undated)
Croatia	Present, Widespread	Native			CABI (Undated)	Original citation: Greuter et al. (1984)
Cyprus	Present, Widespread	Native			CABI (Undated)	Original citation: Greuter et al. (1984)
Czechia	Present	Introduced	1960		Seebens et al. (2017)
Federal Republic of Yugoslavia	Present	Native			CABI (Undated)	Original citation: Greuter et al. (1984)
France	Present, Localized	Native			Gibbs (1972) CABI (Undated)
-Corsica	Present, Widespread	Native			CABI (Undated)	Original citation: Greuter et al. (1984)
Gibraltar	Present, Localized	Native			Valdés et al. (1987)
Greece	Present, Widespread	Native			Gibbs (1972) CABI (Undated)
Ireland	Present	Introduced	2001		Seebens et al. (2017)
Italy	Present, Widespread	Native			Gibbs (1972) Pignatti (1982) CABI (Undated)
Malta	Present, Widespread	Native			Gibbs (1972)
North Macedonia	Present, Widespread	Native			Gibbs (1972)
Norway	Present	Introduced	1872		Seebens et al. (2017)
Poland	Present	Introduced	1866		Seebens et al. (2017)
Portugal	Present, Widespread	Native			Gibbs (1972) Couthinho (1995) CABI (Undated)
-Azores	Present, Widespread				Gibbs (1972) Seebens et al. (2017)
-Madeira	Present, Widespread				Couthinho (1995)
Russia	Present				CABI (Undated a)	Present based on regional distribution.
-Central Russia	Present				Gibbs (1972)
-Southern Russia	Present, Widespread	Native			Gibbs (1972)
Serbia	Present	Native			CABI (Undated)	Original citation: Greuter et al. (1984)
Slovenia	Present, Widespread	Native			CABI (Undated)	Original citation: Greuter et al. (1984)
Spain	Present, Widespread	Native			Gibbs (1972) CABI (Undated)
-Balearic Islands	Present, Widespread	Native			Gibbs (1972) CABI (Undated)
-Canary Islands	Present, Widespread	Native			Bramwell and Bramwell (1974)
Sweden	Present	Introduced	1921		Seebens et al. (2017)
Ukraine	Present				USDA-ARS (2005)
United Kingdom	Present, Localized	Native			Gibbs (1972) Seebens et al. (2017)
-Channel Islands	Present, Widespread	Native			Clapham et al. (1962)
North America
United States	Present				CABI (Undated a)	Present based on regional distribution.
-California	Present, Localized	Introduced		Invasive	Jepson (1979) Hickman (1993)
-Oregon	Present	Introduced			Oregon State University Herbarium (2005)
Oceania
Australia	Present				CABI (Undated a) Seebens et al. (2017) Michael et al. (2020)	Present based on regional distribution.
-New South Wales	Present, Widespread	Introduced		Invasive	Parsons and Cuthbertson (1992)
-Northern Territory	Present, Localized	Introduced			Parsons and Cuthbertson (1992)
-Queensland	Present, Localized	Introduced			Parsons and Cuthbertson (1992)
-South Australia	Present, Widespread	Introduced		Invasive	Parsons and Cuthbertson (1992)
-Tasmania	Present, Widespread	Introduced		Invasive	Parsons and Cuthbertson (1992)
-Victoria	Present, Widespread	Introduced		Invasive	Parsons and Cuthbertson (1992)
-Western Australia	Present, Widespread	Introduced		Invasive	Parsons and Cuthbertson (1992) Michael et al. (2020)
New Zealand	Present, Localized	Introduced			Webb et al. (1988)
South America
Argentina	Present, Widespread	Introduced		Invasive	Holm et al. (1979)
Brazil	Present, Widespread	Introduced		Invasive	Holm et al. (1979)
Chile	Present, Widespread	Introduced		Invasive	Holm et al. (1979) Seebens et al. (2017)
Uruguay	Present	Introduced		Invasive	Piggin (1976)

History of Introduction and Spread

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E. plantagineum was almost certainly first exported outside its native range intentionally as a garden species. For example, it was probably brought to Australia in the early 19th century as seed, shipped from England, Madeira, the Canary Islands or South Africa on the main shipping route prior to the Suez Canal opening in 1869. E. plantagineum is listed under the name E. violaceum or E. violaceum grandiflorum in the 19th century catalogues of plants growing in botanical gardens and nurseries across southern Australia (Piggin and Sheppard, 1995). Deliberate introduction into and spread from the gardens of early settlers around the early colonies in Australasia is the most likely path. A Mr Paterson in the Riverina district of New South Wales in about 1880, and Lady Campbell in Western Australia in about 1900, are reported to have been the first to cultivate the plant in gardens in what are now key infested areas in these states. The widely accepted common name 'Paterson's curse' originates from the first of these (Carne and Gardner, 1925). The earliest herbarium specimen was collected in New South Wales in 1875, with records of occurrence dating back to 1858 from Victoria (Kloot, 1982; Piggin and Sheppard, 1995). Forcella et al. (1986) considered that there were four or more isolated infestation points in Victoria/New South Wales, Tasmania, South and Western Australia by 1900, with later movement as migrating fronts leading to amalgamation. The plant escaped from settler gardens and quickly became established on stock routes, reserves and paddocks throughout southeast and southwest Australia (Maiden, 1905). It arrived latest in the Northern Territory in 1956. Accidental introductions also occurred through contamination in canary and cumin seed imported into New South Wales from Morocco and France (Piggin and Sheppard, 1995).

The plant was introduced into Argentina in the late 1800s and increased the carrying capacity of pastures (Roseveare, 1948). In Uruguay, it was a browse species eaten mainly by sheep and was regarded as a weedy invader of pastures (Roseveare, 1948). Its importance might be on the increase, however, with more recent records of poisoning of cattle in Brazil following ingestion of the weed (Mendez et al. 1985). In South Africa, it was introduced as a stock feed and spread along roadsides (Retief and Wyk, 1998).

Risk of Introduction

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Risk of invasion and spread remains in those countries where the plant is still grown for its flowers, but the weed has not yet naturalized. Another risk of introduction comes from the movement of hay fodder (Thomas et al., 1984). Local spread risk comes from grazing stock that can ingest and pass seed of E. plantagineum. This can be reduced by a quarantine period (Piggin and Sheppard, 1995). E. plantagineum is a proclaimed noxious weed in all states of Australia.

Habitat

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In its native range, E. plantagineum is typical of species-rich semi-natural Mediterranean climate pasture communities on neutral to acid sandy soils (Noy-Meir et al., 1989; Fernández Alés et al., 1993). Such communities are widespread throughout the Mediterranean basin in areas with significant winter rainfall and have been maintained by long established grazing systems. Disrupted by only infrequent cultivation to remove shrub regeneration, these pastures are dominated by annual ruderal species which form a 'grazing disclimax' that may persist for many years (Noy-Meir et al., 1989). The soil remains parched throughout the Mediterranean summer, and most of the community germinates after the first heavy autumn rains (October-November). In such communities, E. plantagineum has been characterized as being relatively medium-sized, large-seeded, long-lived and late flowering (Fernández Alés et al., 1993), and occurring most abundantly under moderate grazing intensity (Noy-Meir et al., 1989), with abundance peaking several years after cultivation (Bernáldez et al., 1980). In one sampled community, E. plantagineum co-occurred with over 120 other species from 85 genera (Grigulis, 1999). E. plantagineum-rich communities in pasture (Fernández Alés et al. 1993), crops (Fernandez-Caro et al,. 1992), or orchards (Sa et al. ,1989) are generally species-rich compared to those in the exotic range.

In the exotic range, E. plantagineum occurs mainly in perennial and annual pastures on roadsides and in waste places, where it can often be dominant. It sometimes occurs in crops if sowing is early and/or land preparation is poor. In the annual pasture-cropping zone in southern Australia, E. plantagineum is a major component of an introduced pasture agro-ecosystem. Co-occurring species in such communities include Trifolium subterraneum, Sisymbrium officinale, Vulpia myuros, Bromus diandrus and Lolium rigidum plus 19 other species (Smyth et al. 1992). Other introduced species commonly found in association with E. plantagineum in Australia are Acetosella vulgaris [Rumex acetosella subsp. acetosella], Arctotheca calendula, Avena spp., Bromus molliformis [B. hordeaceus subsp. divaricatus], Emex australis, Erodium spp., Heliotropium europaeum, Hordeum spp., Oxalis pes-caprae, Raphanus raphanistrum, Romulea spp. and Tribulus terrestris. Annual pastures are usually rotated with cereal or fodder crops allowing E. plantagineum to carry over as a weed into these crops. In perennial pasture systems in southern Australia, E. plantagineum can be locally abundant and associated also with perennial species such as Phalaris aquatica, Dactylis glomerata, Lolium perenne, Medicago sativa and Trifolium repens (Friend, 1991, Dellow et al., 2002). At the northern end of its distribution in Australia (in Queensland), E. plantagineum grows mainly in winter crops such as Avena sativa and M. sativa (Piggin and Sheppard, 1995). In these situations, occurrence is related more to land use and history than to plant associations with the plant growing where it has been introduced as a garden species, as a contaminant of seed, by livestock around old homestead sites, poultry operations, saleyards and trucking yards, and in sown pastures and crops. E. plantagineum-dominated communities in other parts of the exotic range have not been documented.

Comparisons between associated plant communities in the native and exotic range show that many of the associated species are the same, being similarly adapted to common agricultural practice. In addition to exotic communities tending to be species poor assemblages compared to the native communities, these native communities always have a much higher diversity and community dominance from other broadleaf annual rosette-forming species that are in the same functional group as E. plantagineum (Grigulis, 1999).

Habitat List

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Category	Sub-Category	Habitat	Presence	Status
Terrestrial
Terrestrial	Managed	Cultivated / agricultural land	Present, no further details	Harmful (pest or invasive)
Terrestrial	Managed	Managed grasslands (grazing systems)	Present, no further details	Harmful (pest or invasive)
Terrestrial	Managed	Disturbed areas	Present, no further details	Harmful (pest or invasive)
Terrestrial	Managed	Rail / roadsides	Present, no further details	Harmful (pest or invasive)
Terrestrial	Managed	Urban / peri-urban areas	Present, no further details	Harmful (pest or invasive)
Terrestrial	Natural / Semi-natural	Natural grasslands	Present, no further details	Harmful (pest or invasive)
Terrestrial	Natural / Semi-natural	Riverbanks	Present, no further details	Harmful (pest or invasive)
Littoral		Coastal areas	Present, no further details	Harmful (pest or invasive)

Hosts/Species Affected

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In mixed pasture swards, E. plantagineum has been shown to compete with and reduce production from T. subterraneum. In an experiment on seedling removal of E. plantagineum in two pastures, 52-60% of the extra space was occupied by T. subterraneum and the rest by annual grasses (Smyth et al., 1997). Competition is most severe in under-grazed situations, if soil fertility is high, if E. plantagineum establishes before T. subterraneum, and as swards mature. Competition with T. subterraneum appears to be between shoots rather than roots (Piggin and Sheppard, 1995).

Opinions are divided on the weed status of E. plantagineum in crops. It is a weed that significantly reduces yield in lucerne (Medicago sativa) and wild oats where present (Piggin and Sheppard, 1995). In the Western Australian cereal belt, E. plantagineum can crowd out growing crops, reduce grain yield and make harvesting difficult (Carne and Gardner 1925). Pearce (1972), however considered that it was not a serious crop weed because it is controlled by normal cropping techniques and, if necessary, spraying with 2,4-D. Instances of herbicide resistance have since been reported. Piggin (1976) showed that the plant is rare in crops because most seed germinates in the summer and autumn when soil temperatures are high and seedlings are killed by cultivation before the crop is sown. The plant became a more important crop weed with the adoption of minimum-tillage crop establishment techniques.

Host Plants and Other Plants Affected

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Plant name	Family	Context
cereals		Other
Hordeum vulgare (barley)	Poaceae	Main
Triticum aestivum (wheat)	Poaceae	Main

Growth Stages

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Flowering stage, Fruiting stage, Post-harvest, Pre-emergence, Seedling stage, Vegetative growing stage

Biology and Ecology

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Genetics

The basic chromosome number of E. plantagineum is 8 (2n=16) (Britton, 1951). E. plantagineum exhibits some variation in vegetative and reproductive characters. The shape of rosette leaves varies from ovate to lanceolate and flower colour varies from pink to blue to purple to white. The unusual white-flowered and fasciated forms which occur occasionally in natural populations have been mentioned by Ewart (1916), Anon. (1919) and Piggin and Sheppard (1995). Isozyme studies have shown that E. plantagineum exhibits high levels of genetic variation (isozyme polymorphism) compared to other species with similar breeding systems present at a similar stage in succession. Australian populations possess 2.7 alleles per locus, a gene diversity of 34-38% and heterozygosity of 32-35%. These values are very similar to those measured from populations in the native range of the plant (Brown and Burdon, 1983; Burdon and Brown, 1986). This results from a high outcrossing rate which outweighs limitations placed on the diversity of this weed by long distance colonization. Flower colour most likely results from the complementary interaction of at least two loci. Moreover, genes controlling the colour of the corolla tube may act independently of genes controlling the colour of petal tips (Piggin and Sheppard, 1995). White is the recessive colour. Although rare white forms were consistently found to be poorer competitors compared to the wild type, variation in this result suggests that not all white individuals result from the inbreeding of existing white individuals (Burdon et al. 1983).

Physiology and Phenology

E. plantagineum has a winter annual life history strategy. Germination is epigeal. The radicle emerges through a split in the pointed end of the seed coat and forms the primary root, whilst the cotyledons, often carrying the split seed coat, are pushed to the soil surface. Seed germination occurs from 12° to 40°C, favoured by high constant (20° to 30°C) or alternating (15/40°C) temperatures in the presence of moisture; light is not a requirement (Ballard and Grant Lipp, 1970; Piggin and Sheppard, 1995). This is typical of winter annual species, where spring germination is inhibited by the preceding low winter temperatures. Seeds have an after-ripening period with germination occurring several months after maturity in spring and increasing to 30% by autumn. The remaining seeds germinate at a similar rate per year in subsequent years (Grigulis, 1999). The seed coat controls germination; its complete removal allows almost 100% germination (Ballard, 1970). Germination is depressed by a water-soluble inhibitor present in the aerial parts of the plant (Ballard and Grant Lipp, 1959). Emergence is abundant after rainfall in summer and autumn when temperatures are high, and it is not uncommon for over 50% of seedlings to emerge before the main autumn rains. Cultivation reduces emergence and this is associated with seed burial and reduced soil moisture. E. plantagineum emergence declines with burial depth and does not occur below 8 cm (Piggin and Sheppard, 1995). In the field, most seedlings of E. plantagineum emerge in late summer and early autumn.

The rate of leaf appearance is temperature related. The high relative growth rate, leaf-area ratio and leaf-weight ratio of E. plantagineum leads to large quantities of standing biomass in pastures, especially when nitrogen and phosphorus fertilizers are applied (Trumble and Fraser, 1932) and grazing pressure is low (Davies and Sim, 1931; Pearce, 1972). A glasshouse study showed that swards of E. plantagineum produce the equivalent of up to 6 t/ha of herbage and 6.5 t/ha of roots during the first 12 weeks of growth (Piggin and Sheppard, 1995). The above-ground biomass of E. plantagineum in the field measured 3.1 t/ha when E. plantagineum was 70% of the total spring pasture production with 121 plants/m² (Smyth et al., 1997).

Plants elongate and flower in late winter and early spring in response to increasing photoperiods enhanced by vernalization in winter (Ballard and Grant Lipp, 1959; Piggin and Sheppard, 1995). This flowering response limits the distribution of the species to warm-temperate and Mediterranean climate regions. Rosettes will flower slightly later if drought stressed (Wood and Degabriele, 1985). These plants normally die in summer, although those growing near drains, roadsides, rivers and dams, may continue flowering through summer and die in late summer/early autumn. Plants generally flower for 2-5 months. The flowering period is shortened by drought conditions particularly on shallow soil (Burdon et al., 1988) and lengthened if defoliated and if moisture is available. All these factors allow it to be possible to see the odd flowering plant during most of the year.

The seedbank decays at a peak rate of about 35% per annum such that it may last 10 years, but is highly affected by field conditions. Most seedbank losses are through germination (Sheppard and Smyth, 2002). Successful germination is higher in the exotic range due to lower competition from other species for microsites and more seeds make it into the seedbank (Grigulis et al., 2001)

Reproductive Biology

E. plantagineum reproduces only by seed. Each flowering cyme will open 0-3 flowers (mean 0.82-1.28) per 24-hour period (Núñez, 1977; Corbet and Delfosse, 1984). The flower is protandrous, the anthers dehiscing in the bud which favours cross-pollination. The conspicuous colour of the flower and the accessibility and abundance of nectar and pollen encourage insect visitors, mainly honey bees (93.5%; Davis 1992). Flower density peaks at ca. 450 flowers/m² and an average density of 250 flowers/m² represents a nectar sugar yield of 300 kg/ha/season. Nectar production is coupled to photosynthesis and its relationship with insect visitors has been described by Corbet and Delfosse (1984). Flowers are visited by a wide range of insects in addition to honeybees, including native ants, butterflies, moths, hover flies, bees and thrips (Kirk, 1984). The outcrossing rate is very high in E. plantagineum. The multilocus estimates range from 0.73 in low density <10 plants/m²) populations (Burdon and Brown, 1986) to between 0.81 and 1.05 in high density (>150 plants/m²) populations (Burdon et al. 1988). This high rate of outcrossing probably results more from the frequency of inter-plant movements by pollinating insects than the protandrous floral behaviour. At high plant densities the flowering shoot architecture encourages the intermingling of flower cymes on adjacent plants, which maximizes the chance that adjacent flowers visited by a pollinator are from different individuals (Burdon et al., 1988). At the start of the season, when fewer individuals are in flower, the outcrossing rate is lower (0.85; Burdon et al., 1988).

Full-sized, black seeds (or nutlets) take 26 days after pollination to develop at 17°C and 34 to 68 days at 10°C (Piggin and Sheppard, 1995). While four mature seeds could be produced in all flowers, this is never observed. Flowers produce, on average, 37% (max 51%) of potential seeds with a fairly constant percentage between plants at a given site. This percentage declines through the flowering period (Burdon et al., 1988). When damage or cold weather prevents seed production from a series of flowers on a cyme, later flowers immediately compensate producing the full complement of four seeds. Flower number (r²=0.88), total cyme length (r²=0.82), and tap root diameter (r²=0.72) per plant are good predictors of seed production across sites suggesting seed production was not limited by pollination (Piggin and Sheppard, 1995; Smyth et al., 1997). The weight and number of seeds produced per plant is also directly proportional to the plant straw weight (plant dry weight - seed weight) and therefore reproductive effort (seed weight produced per unit straw weight) is constant for plants of all sizes and there is no threshold size for seed production (Piggin and Sheppard, 1995). Individual plants can produce up to 10,000 seeds. However, in grazed paddocks, plants produce an average of 15-60 (max. 260) seeds (Smyth et al., 1997). The common abundance of E. plantagineum on roadsides can be related to prolific seeding where grazing pressure is light. Resulting seed rain (seeds produced in a season/m²) varies from 100 (at 0.8 plants/m²) to 12,000 (at 700 plants/m²) in grazed paddocks. Soil seed banks of up to 13,000-18,000 seeds/m² with grazing and of up to 30,000 seeds/m² without grazing have been recorded in Australia (Piggin and Sheppard, 1995), although most populations have between 2000 and 10,000 seeds/m². Most seeds fall from the plant at maturity, although the last seed produced per calyx may remain attached. The seeds are relatively small and heavy (Piggin and Sheppard, 1995).

Environmental Requirements

E. plantagineum has been recorded from 0 to 1300 m a.s.l. in its native and introduced range (Pignatti, 1982; Piggin and Sheppard, 1995). Moore (1967) suggested that dense stands of E. plantagineum in Australia occur under the following conditions: annual rainfall is less than 1270 mm; in the warmest month, rainfall is greater than 64 mm, mean temperature is 21.1-26.7°C, and daily temperature range is 13.9-19.4°C; and in the coldest month, rainfall is greater than 25 mm, mean temperature is 4.4-10.0°C, and daily range is 8.3-13.9°C. However, high rainfall during the warmest month does not characterize the climate of areas supporting dense stands of E. plantagineum and a more realistic limit would be less than 64 mm. Otherwise, such conditions do occur in the Mediterranean-type climatic areas where E. plantagineum is abundant. The plant also grows in areas where rainfall and daylengths are more even and temperatures are higher during the year, but it appears to be at the limit of its climatic range in this region. Short photoperiods and high temperatures retard flowering (Ballard and Grant Lipp, 1970; Piggin and Sheppard, 1995). Although E. plantagineum is a calcifuge and occurs most commonly on sandy soils, this does not prevent it from growing well on a wide range of other soil types (from light to heavy and red to grey in Australia).

Associations

E. plantagineum has vesicular-arbuscular mycorrhizas. In South Australia, studies during 1977 showed that 15-95% of the root length was colonized by hyphae, vesicles or arbuscules of mycorrhiza. The significance of these mycorrhizas is uncertain but they may assist in the uptake of phosphorus (Piggin and Sheppard, 1995).

Air Temperature

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Parameter	Lower limit	Upper limit
Absolute minimum temperature (ºC)	-15
Mean annual temperature (ºC)	10	20
Mean maximum temperature of hottest month (ºC)	20	40
Mean minimum temperature of coldest month (ºC)	2	10

Rainfall

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Parameter	Lower limit	Upper limit	Description
Dry season duration	0	12	number of consecutive months with <40 mm rainfall
Mean annual rainfall	300	1300	mm; lower/upper limits

Rainfall Regime

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Bimodal
Summer
Uniform
Winter

Soil Tolerances

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Soil drainage

free
impeded

Soil reaction

acid
neutral
very acid

Soil texture

light
medium

Special soil tolerances

infertile
shallow
sodic

Natural enemies

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Natural enemy	Type	Life stages	Biological control in
Aceria echii	Herbivore	Plants\|Inflorescence
Alfalfa mosaic virus	Pathogen
Austroicetes vulgaris	Herbivore
Cercospora echii	Pathogen	Plants\|Leaves
Dialectica scalariella	Herbivore	Plants\|Leaves	New South Wales; Western Australia
Erysiphe horridula f. echii-myosotidis	Pathogen	Plants\|Leaves
Ethmia bipunctella	Herbivore	Plants\|Inflorescence
Longitarsus aeneus	Herbivore	Plants\|Roots
Longitarsus echii	Herbivore	Plants\|Roots
Meligethes planiusculus	Herbivore	Plants\|Inflorescence; Plants\|Seeds
Mogulones geographicus	Herbivore	Plants\|Leaves; Plants\|Roots
Mogulones larvatus	Herbivore	Plants\|Growing point; Plants\|Leaves; Plants\|Roots
Phaulacridium vittatum	Herbivore
Phytoecia coerulescens	Herbivore	Plants\|Roots; Plants\|Stems
Tinagma balteolella	Herbivore	Plants\|Inflorescence
Tobacco mosaic virus	Pathogen
Tobacco necrosis virus	Pathogen
Tobacco ringspot virus	Pathogen

Notes on Natural Enemies

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In the native range, surveys listed 80 arthropods and six fungi as natural enemies of the genus Echium (Wapshere, 1985). Of these, eight Lepidoptera, eight Coleoptera, three Diptera, twelve Hemiptera and one gall mite are known to breed on E. plantagineum (Piggin and Sheppard, 1995). Not all of these species, however, use E. plantagineum as the primary/regular host. In addition to the main potential biological control agents listed, the moths Tinagma balteolella and Ethmia bipunctella, the fly Contarinia echii and the mite Aceria echii have regular resident populations on E. plantagineum in southern Europe. The plant pathogens Erysiphe horridula f. echii-myosotidis and Cercospora echii are commonly found on E. plantagineum in the native range, the former causing considerable loss of seed production.

Ten natural enemies from the native range of E. plantagineum have been researched as biological control agents for Australia. The moth Dialectica scalariella was the first agent (Wapshere and Kirk, 1977). It was successfully established in 1988 and has spread to all but the coldest parts of the distribution of the weed in Australia, but will attack native Boraginaceae. Its leaf mines are commoner where introduced than in the native range. Only under drought conditions has it killed plants. It appears to have blown across the Tasman sea to New Zealand.

All other biological control agents on E. plantagineum have only one generation per year. The two root weevils Mogulones larvatus and M. geographicus lay eggs in autumn in the leaf stalks of the larger young rosettes. The larvae (Vayssieres, 1986) burrow down the leaves and cause most damage in either the root crown (M. larvatus) or the root cortex (M. geographicus) (Vayssieres and Wapshere, 1983; Forrester, 1993). M. larvatus has now been released throughout Australia where widespread plant death has been observed (Sheppard et al., 1999; Shea et al., 2000; Sheppard et al., 2001; Buckley et al., 2005). Twenty percent of plants in one field in southern France had all root cortex destroyed by M. geographicus. Following release in Australia, M. geographicus is increasing in abundance and distribution.

Longitarsus aeneus and L. echii, halticine flea beetles, have larvae that feed in the roots and adults which feed on the leaves. The life cycle is similar to that of the root weevils except that the next generation adults do not emerge in spring but remain in the soil until autumn (Wapshere, 1982). L. aeneus, a root hair feeder was released in 1993, but failed to establish. L. echii is causing significant damage following release, especially in dry areas (Smyth and Sheppard, 2002; Smyth et al., 2004).

Phytoecia coerulescens has spread since release too although this species is likely to develop in other exotic and some native Boraginaceae (Kirk and Wapshere, 1979). The most damaging flower-feeding insect is the sap beetle Meligethes planiusculus (Swirepik et al., 1996). Three other species were studied but not released as biological control agents. These are the two shoot-sucking bugs Dictyla echii and D. nassata (Vayssieres, 1983) and the flower cyme-feeding moth Ethmia terminella. The pathogen Cercospora echii also occurs in Australia through accidental introduction (Floyd et al., 1996).

Native species in the exotic range in Australia that use E. plantagineum include the economically important Thrips imaginis and T. tabaci which breed in significant numbers in the flowers (Kirk, 1984; Milne and Walter, 1998). Larvae and adults of the native moth Utethesia pulchelloides are common on E. plantagineum. A native Longitarsus species spread from native Cynoglossum spp. to attack E. plantagineum and can kill plants in some circumstances (Piggin and Sheppard, 1995).

Means of Movement and Dispersal

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Natural Dispersal (Non-Biotic)

Because seeds are relatively heavy they are not spread over any great distance by wind. Seeds are spread by water where plants grow close to rivers and streams or in steep country prone to erosion and run-off.

Vector Transmission

Several species of ant (e.g., Messor sp. in the native range; Pheidole megacephala, Iridomyrmex discors, Prolasius sp. in Australia) collect seeds and store them in granaries above and below the ground, where many later germinate (Piggin and Sheppard, 1995), but ants probably do not transport seed over great distances. The seed can form a significant part of the diet of ground-feeding seed-eating birds (e.g., crested pigeons Ocyphaps lophotes), especially in drought years, and they may be responsible for dispersal over greater distances (Piggin and Sheppard, 1995). The seeds can also pass through the digestive tracts of grazing animals.

Agricultural Practices

Movement of soil, fodder, vehicles and livestock has probably caused the greatest spread within and between farms and districts. In addition to ingestion, seeds are often carried on the coats of livestock because the calyx and seed coat of E. plantagineum are rough and adhesive. Wool waste from affected areas can also contain seed of E. plantagineum (Piggin and Sheppard, 1995).

Movement in Trade

E. plantagineum seed is sometimes reported as a contaminant of harvested grain (Sahadeva Singh, 2001) because the plant can be a weed in cereal and pasture seed crops (Piggin and Sheppard, 1995).

Accidental Introduction

Dispersal may occur by seed movement in soil for road building or construction or in mud attached to all-terrain vehicles (Parsons and Cuthbertson, 1992).

Intentional Introduction

E. plantagineum was almost certainly first exported outside its native range intentionally as a garden species and remains a species of horticultural interest in parts of the world (e.g., USA).

Pathway Vectors

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Vector	Notes	Long Distance
Clothing, footwear and possessions		Yes
Land vehicles		Yes
Mail	Seed catalogues, internet	Yes
Plants or parts of plants	Grain and hay movement.	Yes
Soil, sand and gravel	River sand	Yes

Plant Trade

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Plant parts liable to carry the pest in trade/transport	Pest stages	Borne internally	Borne externally	Visibility of pest or symptoms
Flowers/Inflorescences/Cones/Calyx	weeds/flowers		Yes	Pest or symptoms usually visible to the naked eye
True seeds (inc. grain)	weeds/seeds		Yes	Pest or symptoms usually visible to the naked eye

Plant parts not known to carry the pest in trade/transport
Bark
Bulbs/Tubers/Corms/Rhizomes
Fruits (inc. pods)
Growing medium accompanying plants
Leaves
Roots
Seedlings/Micropropagated plants
Stems (above ground)/Shoots/Trunks/Branches
Wood

Impact Summary

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Category	Impact
Animal/plant collections	None
Animal/plant products	Negative
Biodiversity (generally)	None
Crop production	Negative
Environment (generally)	Negative
Fisheries / aquaculture	None
Forestry production	None
Human health	Negative
Livestock production	Negative
Native fauna	None
Native flora	Negative
Rare/protected species	None
Tourism	None
Trade/international relations	Negative
Transport/travel	None

Impact

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E. plantagineum competes strongly with more valuable pasture species, especially under light grazing which favours the development of large rosettes, but the high productivity of E. plantagineum can make it a useful forage species. Depending on the rainfall it either dries off in spring leaving little feed for the critical summer/early autumn period or continues to grow vigorously and provide some green feed over summer, especially when heavily grazed, at a time when other species have dried off (Piggin and Sheppard, 1995). Young growth is readily eaten and provides considerable sustenance to stock, whilst older growth is rough and hairy and generally avoided by stock. The plant appears to be eaten readily by sheep, less-readily by cattle and reluctantly by horses, but little data support this so it is possible that differences observed are due, at least partly, to differences in grazing pressure and pasture height. Rosettes contain 6-14% dry matter, 2.0 to 4.8% d.m. nitrogen and are 50 -75% d.m. digestible. Flowering plants contain 20-70% dry matter, 0.5-4.7% d.m. nitrogen and are 37-69% d.m. digestible (Piggin, 1976; 1977). Comparisons suggest that E. plantagineum is as nutritious as most recognized pasture species with low dry matter content (20-30%) and relatively high nitrogen content (4.1-4.3%) and digestibility (61-64%) and so should provide useful fodder. The hairiness can cause slavering, dermatitis, inflammation and itching to animals and man. The key problem associated with E. plantagineum, however is that it contains eight hepatotoxic pyrrolizidine alkaloids, particularly echiumine and echimidine (Culvenor, 1956), which are at higher levels in rosettes and also vary with soil type, soil fertility and climate. These alkaloids can cause cumulative chronic liver damage and animal mortality, especially if substantial amounts of herbage are eaten over prolonged periods. Monogastric grazing animals such as horses are more susceptible than ruminants because the alkaloids are largely broken down in the rumen. E. plantagineum was considered the primary cause of pyrrolizidine alkaloid poisoning in horses in Australia (Seaman, 1978; Dellow and Seaman, 1985) and toxicity in cattle in Brazil (Schild et al., 2004). E. plantagineum is also the major cause of sheep deaths from primary pyrrolizidine alkaloid and associated hepatogenous chronic copper poisoning in New South Wales (Seaman, 1987), particularly in crossbred ewes (Culvenor et al., 1984). Cross-breeds grazing pasture that was up to 61% E. plantagineum had reduced weight and wool growth, but no mortality after 19 months, while the same animals fed solely on fresh E. plantagineum showed some 40% mortality after 16 weeks due to progressive liver damage (Seaman et al., 1989; Seaman and Dixon, 1989). The plant has also been implicated in several reported cases of mortality in horses and occasionally young pigs (Bull et al., 1968) and even caged canaries (Hurst, 1942). Such effects can be minimized if good pasture and livestock management maintains the E. plantagineum percent pasture consumption below 50% (Piggin and Sheppard, 1995).

E. plantagineum has been noted as a weed in pastures grown for the production of Trifolium subterraneum seed. Seeds of the two species are produced at the same time and are difficult to separate because they are of similar size and colour (Gardner, 1933).

Environmental Impact

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The environmental impacts of E. plantagineum are minimal as it is only a weed of ecosystems managed for agriculture and is a symptom of poor management of species-poor pasture systems in low to medium rainfall Mediterranean environments. In can spread as a casual into disturbed native grasslands or rough grassing grassland, but is not maintained there without at least the sporadic presence of livestock.

Impact: Biodiversity

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There are no impacts on biodiversity of any consequence, beyond the general effects of extensive livestock agriculture.

Social Impact

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E. plantagineum has been associated with human allergy problems. Use or contamination in foodstuffs can lead to human health risks from the pyrrolizidine alkaloids and threatens bans on contaminated cereal exports from affected areas. Its conspicuous flowers and high density also affect the aesthetics of regions within which large infestations occur.

Risk and Impact Factors

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Invasiveness

Proved invasive outside its native range
Highly adaptable to different environments
Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
Highly mobile locally
Has high reproductive potential
Has propagules that can remain viable for more than one year

Impact outcomes

Damaged ecosystem services
Ecosystem change/ habitat alteration
Negatively impacts agriculture
Negatively impacts human health
Negatively impacts animal health
Negatively impacts tourism
Reduced amenity values

Impact mechanisms

Competition - monopolizing resources
Produces spines, thorns or burrs

Likelihood of entry/control

Highly likely to be transported internationally accidentally
Highly likely to be transported internationally deliberately
Difficult to identify/detect as a commodity contaminant
Difficult/costly to control

Uses

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From early times E. plantagineum has been cited as a valuable honey plant both in Australia (Rayment 1925, Goodacre 1938), South Africa (Allsopp, 1993) and in the native region (i.e. Spain; Ortiz et al., 1990). The flowers produce good early pollen and nectar to stimulate brood-rearing in the hive. Light amber honey with a typical flavour is produced. More recently concerns have increased based on the levels of pyrrolizidine alkaloids that carry over into the honey and that such levels can be high relative to recommended international levels for human consumption. This has led to the mixing of honey produced from Echium species with honey from other sources to minimize such risks.

The seed oil of E. plantagineum is unusual in that it contains a unique ratio of omega-3 and omega-6 fatty acids which are valuable as health supplements. E. plantagineum is currently the best agricultural source of certain of these. The seed oil is also valued for its moisturizing and anti-inflammatory action (McEwen et al., 2003). E. plantagineum is also grown and marketed in some parts of the world a as garden species because of its attractive flowers. 'Blue bedder' and 'White Hills' varieties are available in USA (Britton, 1951). The plant has been well regarded as an attractive wildflower and considered a tourist attraction in parts of South Australia (Burt, 1978). It was also harvested and sold as a cut flower in the Sydney markets for many years.

Uses List

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General

Ornamental

Human food and beverage

Honey/honey flora

Materials

Poisonous to mammals

Similarities to Other Species/Conditions

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Of the 40 species in the genus Echium (all from Europe and Macaronesia), two other herbaceous species, both monocarpic perennials, have naturalized and spread outside their native range; Echium italicum and E. vulgare. E. italicum is a larger species with smaller pale flowers and occurs extremely locally in Australia and North America, while E. vulgare is most similar to E. plantagineum (except for smaller flowers and stamens more exserted) and quite widespread in temperate regions of Australasia, North and South America and South Africa. E. vulgare is more a cool temperate species, where it can be a common weed in some perennial pasture, roadsides and neglected areas (Piggin, 1977; Forcella et. al., 1986; Parsons and Cuthbertson, 1992). Piggin (1977) presents a key to distinguish them from E. plantagineum. E. plantagineum has more rapid germination over a wider temperature range, more rapid vegetative growth, and greater capacity to flower under a wide range of temperature and photoperiod, than either E. italicum or E. vulgare (Forcella et. al., 1986). Given that members of Echium have strong horticultural interest based on their flowers, that several of these have become garden escapes (e.g., E. simplex, E. candicans and E. pininana), and that other species are weeds in their native range, many of the species in the genus are considered a threat on a global scale (Randall, 2002).

Prevention and Control

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Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

Cultural Control

Heavy grazing has long been recognized as an effective means of controlling E. plantagineum. It should be concentrated when plants are young, continued at regular intervals during the year, and again concentrated at the end of the year when plants are seeding. Like mowing, it is useful in conjunction with other control measures such as cultivation, handpulling and spraying (Maiden, 1905; Carne and Gardner, 1925; Pearce, 1972). Some authors suggest that cattle are less effective than sheep because they do not graze sufficiently close to the ground. Pearce (1972) and Piggin (1979) considered that the level of control comes from the stocking rate not the livestock type and added that spray-grazing can be effective using sheep or cattle. Piggin (1979) reported that more or less continuous grazing by ca. nine sheep/ha reduced the cover of E. plantagineum from 36% to 3% for 6 months over winter. Pearce (1972) reported that the proportion and plant density of E. plantagineum were reduced as the stocking rate of sheep was increased. Response to grazing, however, can be complicated. In areas where E. plantagineum occurs with perennial grasses such as Phalaris aquatica and Dactylis glomeratus, grazing can increase the abundance of E. plantagineum through preferential grazing of the grasses (Piggin and Sheppard, 1995). Grazing with rabbits reduces available herbage but not percentage composition (Piggin and Sheppard, 1995).

Establishment of vigorous pasture with species such as Trifolium subterraneum and Phalaris aquatica, sown in autumn after fallowing over summer, has been effective in controlling E. plantagineum (Michael, 1970). Pasture establishment is usually recommended as a follow-up to other control measures. Top-dressing with fertilizers can be combined with pasture establishment, or used alone, to encourage T. subterraneum to control E. plantagineum, although nitrogen- and phosphate-rich fertilizers encourage, rather than control, E. plantagineum (Piggin and Sheppard, 1995). The destruction of plants by burning with a flame thrower has also been suggested as a useful method of control, especially along fences, roads and channels.

Mechanical Control

Handpulling and cutting have been suggested for the control of single plants and small patches, especially where flowering plants have survived following the use of other control methods such as spraying, cultivation and grazing. After removal, plants should be piled up and burnt because seeds may continue to mature (Maiden, 1905; Carne and Gardner, 1925). E. plantagineum can be effectively controlled in arable areas by cultivation, especially if followed by cropping with root or cereal crops, establishment of a vigorous, smothering pasture, grazing or handpulling (Ewart and Tovey, 1909; Carne and Gardner, 1925; Piggin and Sheppard, 1995).

Mowing has been recommended, especially in combination with other control measures, as a means of controlling E. plantagineum in pastures and reducing seed production. It is especially useful where stocking pressure cannot be maintained (Piggin and Sheppard, 1995). Early hay cutting and silage making can also help control. Davies and Sim (1931) reported that in a pasture near Adelaide, South Australia, the dry matter yield and average composition of E. plantagineum were reduced from 435 kg/ha and 25% with a single cut in spring to 62 kg/ha and 7% with fortnightly cuts during 1928. Piggin (1978) showed that mowing reduced the herbage production of swards containing E. plantagineum and reduced shoot weight per plant, root weight and root:shoot ratios of E. plantagineum.

Biological Control

Biological control of E. plantagineum has only been attempted in Australia, where six agents have been released (Julien and Griffiths, 1998). The biological control programme became a landmark case in the history of weed biological control; conflicts of interest raised by graziers and beekeepers led to a full public enquiry and benefit:cost analysis that came out in favour of control by 10 to 1 and also led to the first biological control legislation (Delfosse and Cullen, 1981, 1985). Since then the impacts of two of the agents (Mogulones larvatus and Longitarsus echii) have been significant (Swirepik and Smyth, 2002) and sufficient for detailed economic evaluation of the project which is expected to save Australian agriculture nearly A1$ billion by 2050 (Nordblom et al., 2001).

Chemical Control

Herbicides, in the form of brine, petroleum and arsenical preparations, were first recommended for the control of E. plantagineum in Australia by Davey (1922). Since then, many have been found to be effective in controlling the plant: asulam; atrazine; atrazine-amitrole mixtures; amitrole; arsenic pentoxide; bromoxynil; chlorsulfuron; chlorthal-dimethyl; chlorthal-dimethyl-linuron mixtures; cotoran [fluometuron]; dicamba; diuron; 4-CPA; glyphosate mixtures; ioxynil; linuron; MCPA; metribuzin-methabenzthiazuron mixtures; metsulfuron methyl; paraquat; picloram; prometryne; propionic acid-MCPA mixtures; sodium chlorate; terbutryn; 2,3,6-TBA; 2,4-D ester and amine; 2,4-DB; 2,4-D-atrazine mixtures; and 2,4-D-picloram mixtures (Parsons and Cuthbertson, 1992; Piggin and Sheppard, 1995). Up until the 1970s, the plant was usually controlled by spraying with 2,4-D at the flowering stage in spring when the plant was most conspicuous. There are disadvantages, however, with spraying in spring: firstly, the high doses of herbicide necessary to kill flowering plants are expensive and severe on associated desirable species such as Trifolium subterraneum; secondly, by spring, E. plantagineum has competed in the pasture for most of the year; and, thirdly, recolonization by desirable species is poor after spraying. Herbicides are now considered to be most effective when applied at the young rosette stage. Wetting agents may improve control and spraying should be accompanied by adjustments in pasture and livestock management to prevent the return of the weed. A selective application technique, such as a carpet wiper, can target non-selective herbicides such as glyphosate on the weed.

IPM Programmes

To overcome disadvantages of herbicide-only strategies, a technique known as spray-grazing was developed which combined grazing with the application of low doses of 2,4-D in the autumn to control E. plantagineum with a minimum of damage to associated pasture (Pearce, 1972). Doses as low as 0.14 kg a.i./ha sprayed in May/June followed by grazing a week later with five to seven times the normal stocking rate have resulted in 100% control. The capacity of Trifolium subterraneum to continue to germinate through winter after E. plantagineum germination drops off, can allow this to lead to a balanced pasture of T. subterraneum and grasses in spring (Piggin et al., 1973; Piggin, 1979). Huwer et al. (2005) compare more complex integrated strategies where spray-grazing to remove E. plantagineum are combined with fertilizing and over-sowing infested pasture with desirable species to make the pasture more competitive and sustainable without the need for complete re-seeding. Such strategies were also considered in relation to how they interact with biological control agents (Smyth and Sheppard, 1996).

References

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Allsopp M, 1993. Biocontrol of bloublommetjies. South African Bee Journal, 65(2):32-36; [Bj]

Anon, 1919. Notes and exhibits: Ordinary monthly meeting, 24 September 1919. Proceedings of the Linnaean Society of N.S.W., 44:525-528

Ballard LAT, 1970. Germination behaviour of Echium lycopsis. Proceedings of the Australian Weeds Conference, Hobart, 1970, 5-9 & 5-11

Ballard LAT, Grant Lipp AE, 1959. Differential specificity exhibited by two germination inhibitors present in Echium plantagineum L. Australian Journal of Biological Science, 12:343-347

Ballard LAT, Grant Lipp AE, 1970. Flowering responses of Echium lycopsis to photoperiod and vernalization Proceedings of Australian Weeds Conference, Hobart, 1970, 5.11-5.13

Bernáldez FG, Haeger JF, Levassor C, Merino JA, Sancjo F, 1980. La prospection intégrée de paturages extensifs dans la Sierra Morena (Espagne). Espaniola Geographica, 3:241-252

Bramwell D, Bramwell ZI, 1974. Wild Flowers of the Canary Islands. Cabildo Insular de Tenerife, Canary Islands: Excmo

Britton DM, 1951. Cytogenetic studies on the Boraginaceae. Brittonia, 7:233-266

Brown AHD, Burdon JJ, 1983. Multilocus diversity in an outbreeding weed, Echium plantagineum L. Australian Journal of Biological Science, 36:503-509

Buckley YM, Rees M, Sheppard AW, Smyth M, 2005. Stable coexistence of an invasive plant and biocontrol agent: a coupled plant-herbivore model. Journal of Applied Ecology, 42:in press

Bull LB, Culvenor CCJ, Dick AT, 1968. The pyrrolizidine alkaloids. Amsterdam, Netherlands: North Holland Publishing Co

Burdon JJ, Brown AHD, 1986. Population genetics of Echium plantagineum L. target weed for biological control. Australian Journal of Biological Sciences, 39(4):369-378

Burdon JJ, Jarosz AM, Brown AHD, 1988. Temporal patterns of reproduction and outcrossing in weedy populations of Echium plantagineum. Biological Journal of the Linnean Society, 34(1):81-92

Burdon JJ, Marshall DR, Brown AHB, 1983. Demographic and genetic changes in populations of Echium plantagineum. Journal of Ecology, 71:667-679

Burt J, 1978. Magnificent Australia in Colour. Melbourne, Australia: Budget Books

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Echium plantagineum (Paterson's curse)

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