Echium plantagineum (Paterson's curse)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Plant Type
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Habitat List
- Hosts/Species Affected
- Host Plants and Other Plants Affected
- Growth Stages
- Biology and Ecology
- Air Temperature
- Rainfall Regime
- Soil Tolerances
- Natural enemies
- Notes on Natural Enemies
- Means of Movement and Dispersal
- Pathway Vectors
- Plant Trade
- Impact Summary
- Environmental Impact
- Impact: Biodiversity
- Social Impact
- Risk and Impact Factors
- Uses List
- Similarities to Other Species/Conditions
- Prevention and Control
- Distribution Maps
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PicturesTop of page
IdentityTop of page
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
- EHIPL (Echium plantagineum)
Summary of InvasivenessTop of page
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 TreeTop of page
- Domain: Eukaryota
- Kingdom: Plantae
- Phylum: Spermatophyta
- Subphylum: Angiospermae
- Class: Dicotyledonae
- Order: Boraginales
- Family: Boraginaceae
- Genus: Echium
- Species: Echium plantagineum
Notes on Taxonomy and NomenclatureTop of page 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.
DescriptionTop of page 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 TypeTop of page Annual
DistributionTop of page 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 TableTop of page
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.
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Georgia (Republic of)||Present||Native||Not invasive||Greuter et al., 1984|
|Israel||Widespread||Native||Not invasive||Greuter et al., 1984|
|Jordan||Widespread||Native||Not invasive||Greuter et al., 1984|
|Lebanon||Widespread||Native||Not invasive||Greuter et al., 1984|
|Syria||Widespread||Native||Not invasive||Greuter et al., 1984|
|Turkey||Widespread||Native||Not invasive||Gibbs, 1972; Greuter et al., 1984|
|Algeria||Widespread||Native||Not invasive||Greuter et al., 1984|
|Egypt||Widespread||Native||Not invasive||Greuter et al., 1984|
|Ethiopia||Present||Introduced||Invasive||Witt and Luke, 2017|
|Kenya||Present||Introduced||Invasive||Witt and Luke, 2017|
|Libya||Widespread||Native||Not invasive||Greuter et al., 1984|
|Morocco||Widespread||Native||Not invasive||Greuter et al., 1984|
|South Africa||Widespread||Introduced||Invasive||Adamson and Salter, 1950|
|-Canary Islands||Widespread||Native||Not invasive||Bramwell and Bramwell, 1974|
|Tanzania||Present||Introduced||Invasive||Witt and Luke, 2017|
|Tunisia||Widespread||Native||Not invasive||Greuter et al., 1984|
|USA||Present||Present based on regional distribution.|
|-California||Restricted distribution||Introduced||Invasive||Jepson, 1979; Hickman, 1993|
|-Oregon||Present||Introduced||Not invasive||Oregon State University Herbarium, 2005|
|Argentina||Widespread||Introduced||Invasive||Holm et al., 1979|
|Brazil||Widespread||Introduced||Invasive||Holm et al., 1979|
|Chile||Widespread||Introduced||Invasive||Holm et al., 1979|
|Albania||Widespread||Native||Not invasive||Greuter et al., 1984|
|Bosnia-Hercegovina||Widespread||Native||Not invasive||Greuter et al., 1984|
|Bulgaria||Widespread||Native||Not invasive||Gibbs, 1972; Greuter et al., 1984|
|Croatia||Widespread||Native||Not invasive||Greuter et al., 1984|
|Cyprus||Widespread||Native||Not invasive||Greuter et al., 1984|
|France||Restricted distribution||Native||Not invasive||Gibbs, 1972; Greuter et al., 1984|
|-Corsica||Widespread||Native||Not invasive||Greuter et al., 1984|
|Gibraltar||Restricted distribution||Native||Not invasive||Valdés et al., 1987|
|Greece||Widespread||Native||Not invasive||Gibbs, 1972; Greuter et al., 1984|
|Italy||Widespread||Native||Not invasive||Gibbs, 1972; Pignatti, 1982; Greuter et al., 1984|
|Macedonia||Widespread||Native||Not invasive||Gibbs, 1972|
|Malta||Widespread||Native||Not invasive||Gibbs, 1972|
|Portugal||Widespread||Native||Not invasive||Gibbs, 1972; Greuter et al., 1984; Couthinho, 1995|
|-Azores||Widespread||Not invasive||Gibbs, 1972|
|-Madeira||Widespread||Not invasive||Couthinho, 1995|
|Russian Federation||Present||Present based on regional distribution.|
|-Central Russia||Present||Not invasive||Gibbs, 1972|
|-Southern Russia||Widespread||Native||Not invasive||Gibbs, 1972|
|Serbia||Present||Native||Not invasive||Greuter et al., 1984|
|Slovenia||Widespread||Native||Not invasive||Greuter et al., 1984|
|Spain||Widespread||Native||Not invasive||Greuter et al., 1987; Greuter et al., Valdés et al.; Gibbs, 1972; Greuter et al., 1984|
|-Balearic Islands||Widespread||Native||Not invasive||Gibbs, 1972; Greuter et al., 1984|
|UK||Restricted distribution||Native||Not invasive||Gibbs, 1972|
|-Channel Islands||Widespread||Native||Not invasive||Clapham et al., 1962|
|Yugoslavia (former)||Present||Native||Not invasive||Greuter et al., 1984|
|Australia||Present||Present based on regional distribution.|
|-Australian Northern Territory||Restricted distribution||Introduced||Not invasive||Parsons and Cuthbertson, 1992|
|-New South Wales||Widespread||Introduced||Invasive||Parsons and Cuthbertson, 1992|
|-Queensland||Restricted distribution||Introduced||Not invasive||Parsons and Cuthbertson, 1992|
|-South Australia||Widespread||Introduced||Invasive||Parsons and Cuthbertson, 1992|
|-Tasmania||Widespread||Introduced||Invasive||Parsons and Cuthbertson, 1992|
|-Victoria||Widespread||Introduced||Invasive||Parsons and Cuthbertson, 1992|
|-Western Australia||Widespread||Introduced||Invasive||Parsons and Cuthbertson, 1992|
|New Zealand||Restricted distribution||Introduced||Not invasive||Webb et al., 1988|
History of Introduction and SpreadTop of page 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 IntroductionTop of page 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.
HabitatTop of page 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 ListTop of page
|Terrestrial – Managed||Cultivated / agricultural land||Present, no further details||Harmful (pest or invasive)|
|Managed grasslands (grazing systems)||Present, no further details||Harmful (pest or invasive)|
|Disturbed areas||Present, no further details||Harmful (pest or invasive)|
|Rail / roadsides||Present, no further details||Harmful (pest or invasive)|
|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)|
|Riverbanks||Present, no further details||Harmful (pest or invasive)|
|Coastal areas||Present, no further details||Harmful (pest or invasive)|
Hosts/Species AffectedTop of page 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 AffectedTop of page
Growth StagesTop of page Flowering stage, Fruiting stage, Post-harvest, Pre-emergence, Seedling stage, Vegetative growing stage
Biology and EcologyTop of page 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)
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).
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).
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 TemperatureTop of page
|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|
RainfallTop of page
|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 RegimeTop of page Bimodal
Soil TolerancesTop of page
- very acid
Special soil tolerances
Natural enemiesTop of page
|Natural enemy||Type||Life stages||Specificity||References||Biological control in||Biological control on|
|Alfalfa mosaic virus||Pathogen|
|Dialectica scalariella||Herbivore||Leaves||New South Wales; Western Australia|
|Erysiphe horridula f. echii-myosotidis||Pathogen||Leaves|
|Mogulones larvatus||Herbivore||Growing point/Leaves/Roots|
|Tobacco mosaic virus||Pathogen|
|Tobacco necrosis virus||Pathogen|
|Tobacco ringspot virus||Pathogen|
Notes on Natural EnemiesTop of page 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 DispersalTop of page 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.
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.
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).
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).
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 VectorsTop of page
Plant TradeTop of page
|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||flowers||Yes||Pest or symptoms usually visible to the naked eye|
|True seeds (inc. grain)||seeds||Yes||Pest or symptoms usually visible to the naked eye|
|Plant parts not known to carry the pest in trade/transport|
|Fruits (inc. pods)|
|Growing medium accompanying plants|
|Stems (above ground)/Shoots/Trunks/Branches|
Impact SummaryTop of page
|Fisheries / aquaculture||None|
ImpactTop of page 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 ImpactTop of page 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: BiodiversityTop of page There are no impacts on biodiversity of any consequence, beyond the general effects of extensive livestock agriculture.
Social ImpactTop of page 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 FactorsTop of page 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
- Damaged ecosystem services
- Ecosystem change/ habitat alteration
- Negatively impacts agriculture
- Negatively impacts human health
- Negatively impacts animal health
- Negatively impacts tourism
- Reduced amenity values
- Competition - monopolizing resources
- Produces spines, thorns or burrs
- 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
UsesTop of page 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 ListTop of page
Human food and beverage
- Honey/honey flora
- Poisonous to mammals
Similarities to Other Species/ConditionsTop of page 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 ControlTop of page
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.
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 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).
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.
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).
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