Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide

Datasheet

Plantago coronopus
(Buck's-horn plantain)

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Datasheet

Plantago coronopus (Buck's-horn plantain)

Summary

  • Last modified
  • 08 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Host Plant
  • Preferred Scientific Name
  • Plantago coronopus
  • Preferred Common Name
  • Buck's-horn plantain
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Spermatophyta
  •       Subphylum: Angiospermae
  •         Class: Dicotyledonae
  • Summary of Invasiveness
  • In its native range of Europe, northern Africa and Central and West Asia, P. coronopus is an inoffensive low-growing herb in coastal saline situations. It is rarely a weed of crops though

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    Compendia
    CAB International
    Wallingford
    Oxfordshire
    OX10 8DE
    UK
    compend@cabi.org
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Identity

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Preferred Scientific Name

  • Plantago coronopus L.

Preferred Common Name

  • Buck's-horn plantain

International Common Names

  • English: Buckhorn plantain; Buck's-horn plantain

Local Common Names

  • : koronopus
  • : wedaina
  • Sweden: strandkämpar
  • USA: minutina

Summary of Invasiveness

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In its native range of Europe, northern Africa and Central and West Asia, P. coronopus is an inoffensive low-growing herb in coastal saline situations. It is rarely a weed of crops though Holm et al. (1979) list it as a ‘common’ weed in Spain. In recent years, however, it has been recorded as invasive in Australia and in California, forming dense mats which displace native vegetation, including endangered species in California (Weber, 2003; United States Fish and Wildlife Service, 2008a, b). Furthermore, it is reported as a weed problem in non-tilled orchards, irrigated pastures, and alfalfa and clover fields in California.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Plantae
  •         Phylum: Spermatophyta
  •             Subphylum: Angiospermae
  •                 Class: Dicotyledonae
  •                     Order: Plantaginales
  •                         Family: Plantaginaceae
  •                             Genus: Plantago
  •                                 Species: Plantago coronopus

Notes on Taxonomy and Nomenclature

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Plantago coronopus was named by Linnaeus and this name has survived, with no alternatives being proposed. Hence there are no synonyms listed, but it is a highly variable species and there are several recognized subspecies and many varieties described, differing in leaf shape, etc. P. coronopus subsp. commutata differs in being tetraploid and has previously been treated as a separate species, P. commutata.

Description

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P. coronopus may behave as an annual, a biennial or a perennial. The plant can develop axillary offsets, hence reproducing vegetatively. A rosette of leaves develops, remaining flat or largely close to the soil. Leaves are up to 20 cm long by 2 cm wide, variously entire or shallowly or deeply toothed, somewhat pubescent. Under saline conditions the leaves may show distinct succulence. Flowering stems are numerous from each rosette, up to 20 cm high, carrying a dense spike of flowers 2-5 cm long. Each flower is subtended by a bract and consists of four sepals, the two on the posterior side conspicuously keeled and hairy. Alternating with the sepals are four whitish transparent petals. The flowers are normally hermaphrodite, having four stamens with long filaments, and large yellow versatile anthers and a syncarpous ovary surmounted by a long, hairy simple style. However, male sterility does occur. The capsule has two lower chambers with two seeds each, while there is usually an upper chamber containing a single smaller seed. The larger seeds are black, shiny, boat-shaped and 1.0-1.5 mm long (mean weight 0.20 mg) while the smaller are 0.7-0.9 mm long (mean weight 0.13 mg). The larger seeds are released when the capsule dehisces, while the smaller seed is usually retained (Rowling, 1933).

Pollen in P. coronopus is highly annulate but not highly operculate. In P. lanceolata it is both, while three other species in Pakistan, including P. major differ in not being highly annulate (Al-Quran, 2004).

Plant Type

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Biennial
Broadleaved
Herbaceous
Perennial
Seed propagated
Succulent
Vegetatively propagated

Distribution

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P. coronopus is native to Europe, northern Africa and Central and West Asia, but has been introduced to parts of North America, South America, South Africa, Australia and New Zealand.

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.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

AfghanistanPresentNativeUSDA-ARS, 2012
AzerbaijanPresentNativeUSDA-ARS, 2012
IndiaPresentGBIF, 2013
IranPresentNativeUSDA-ARS, 2012
IraqPresentNativeUSDA-ARS, 2012
IsraelPresentNativeUSDA-ARS, 2012
JordanPresentNativeUSDA-ARS, 2012
KuwaitPresentNativeUSDA-ARS, 2012
LebanonPresentNativeUSDA-ARS, 2012
PakistanPresentNativeUSDA-ARS, 2012
Saudi ArabiaPresentGBIF, 2013
SyriaPresentNativeUSDA-ARS, 2012
TajikistanPresentNativeUSDA-ARS, 2012
TurkeyPresentNativeUSDA-ARS, 2012; GBIF, 2013
TurkmenistanPresentNativeUSDA-ARS, 2012; GBIF, 2013
UzbekistanPresentNativeUSDA-ARS, 2012

Africa

AlgeriaPresentNativeUSDA-ARS, 2012
EgyptPresentNativeUSDA-ARS, 2012; GBIF, 2013
LibyaPresentNativeUSDA-ARS, 2012
MoroccoPresentNativeUSDA-ARS, 2012
South AfricaPresentIntroducedMissouri Botanical Garden, 2012
Spain
-Canary IslandsPresentNativeUSDA-ARS, 2012
TunisiaPresentNativeSassi et al., 2008

North America

CanadaPresentPresent based on regional distribution.
-British ColumbiaPresentIntroduced
-ManitobaPresentIntroducedUSDA-NRCS, 2012
-New BrunswickPresentIntroducedUSDA-NRCS, 2012
GreenlandPresentUSDA-NRCS, 2012
USAPresentPresent based on regional distribution.
-CaliforniaPresentIntroducedUSDA-NRCS, 2012
-MainePresentIntroducedUSDA-NRCS, 2012
-New JerseyPresentUSDA-NRCS, 2012
-New YorkPresentIntroducedUSDA-NRCS, 2012
-OregonPresentIntroducedUSDA-NRCS, 2012
-PennsylvaniaPresentIntroducedUSDA-NRCS, 2012
-TexasPresentIntroducedUSDA-NRCS, 2012
-WashingtonPresentIntroducedUSDA-NRCS, 2012

South America

BrazilPresentIntroducedMissouri Botanical Garden, 2012; GBIF, 2013
-Rio Grande do SulPresentIntroducedMissouri Botanical Garden, 2012New record 2003
ChilePresentUSDA-NRCS, 2012
UruguayPresentGBIF, 2013; GBIF, 2013

Europe

AlbaniaPresentNativeUSDA-ARS, 2012
AustriaPresentNativeGerstberger, 2001
BelgiumPresentNativeUSDA-ARS, 2012
BulgariaPresentNativeUSDA-ARS, 2012
CroatiaPresentGBIF, 2013
CyprusPresentNativeUSDA-ARS, 2012
DenmarkPresentNativeUSDA-NRCS, 2012
Faroe IslandsPresentNativeUSDA-ARS, 2012
FinlandPresentGBIF, 2013; GBIF, 2013
FrancePresentNativeUSDA-ARS, 2012
GermanyPresentNativeUSDA-ARS, 2012
GreecePresentNativeUSDA-ARS, 2012Including Crete
IrelandPresentNativeUSDA-ARS, 2012
ItalyPresentNativeUSDA-ARS, 2012
MontenegroPresentGBIF, 2013
NetherlandsPresentNativeUSDA-ARS, 2012
NorwayPresentGBIF, 2013
PolandPresentNativeUSDA-ARS, 2012
PortugalPresentNativeUSDA-ARS, 2012
-AzoresPresentUSDA-ARS, 2012
-MadeiraPresentUSDA-ARS, 2012
RomaniaPresentNativeUSDA-ARS, 2012
Russian FederationPresentNativeUSDA-ARS, 2012
SpainPresentNativeUSDA-ARS, 2012
-Balearic IslandsPresentNativeUSDA-ARS, 2012
SwedenPresentNativeUSDA-ARS, 2012
SwitzerlandPresentGBIF, 2013
UKPresentNativeUSDA-ARS, 2012
UkrainePresentNativeUSDA-ARS, 2012
Yugoslavia (former)PresentNativeUSDA-ARS, 2012

Oceania

AustraliaPresentIntroduced Invasive USDA-ARS, 2012
-Australian Northern TerritoryPresentIntroducedCouncil of Heads of Australasian Herbaria, 2012
-New South WalesPresentIntroducedCouncil of Heads of Australasian Herbaria, 2012
-QueenslandPresentIntroducedCouncil of Heads of Australasian Herbaria, 2012
-South AustraliaPresentIntroducedCouncil of Heads of Australasian Herbaria, 2012
-TasmaniaPresentIntroducedCouncil of Heads of Australasian Herbaria, 2012
-VictoriaPresentIntroducedCouncil of Heads of Australasian Herbaria, 2012
-Western AustraliaPresentIntroducedCouncil of Heads of Australasian Herbaria, 2012
New ZealandPresentIntroducedUSDA-ARS, 2012

History of Introduction and Spread

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Although introduced to a number of countries, the history of this species introduction is not well documented.

Risk of Introduction

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There is some risk of introduction via trade in the seeds for cultivation as an ornamental but this is not extensive. The main risk would seem to be via the importation of grass seed from contaminated pastures.

Habitat

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In the UK, the species is found mainly in maritime habitats including the lower rocks immediately above high tide mark, rock crevices and ledges, the face of cliffs where detritus has collected, and grassy slopes leading up to, and above, the cliff face. It is also found on some fixed sand dunes and often in sandy soils when present inland (Dodds, 1953). Weber (2003) indicates occurrence in grassland, along rivers, in fresh and saline wetlands and on coastal dunes. Given its wide native range, however, the species is likely to inhabit a broader range of environments than reported here.  

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Brackish
Inland saline areas Principal habitat
Terrestrial
 
Terrestrial – ManagedCultivated / agricultural land Secondary/tolerated habitat
Managed grasslands (grazing systems) Secondary/tolerated habitat
Disturbed areas Secondary/tolerated habitat
Rail / roadsides Secondary/tolerated habitat
Terrestrial ‑ Natural / Semi-naturalNatural grasslands Secondary/tolerated habitat
Riverbanks Secondary/tolerated habitat
Wetlands Secondary/tolerated habitat
Littoral
Coastal areas Principal habitat
Coastal dunes Principal habitat
Mud flats Principal habitat
Salt marshes Principal habitat

Hosts/Species Affected

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In California, USA, P. coronopus is a weed problem in non-tilled orchards, irrigated pastures, and alfalfa (Medicago sativa) and clover (Trifolium spp.) fields. It is also reported as a threat to two endangered plant species in California; Trifolium amoenum (US Fish and Wildlife Service, 2008a) and Astragalus robbinsii var. jesupii (US Fish and Wildlife Service, 2008b).

Host Plants and Other Plants Affected

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Plant nameFamilyContext
Medicago sativa (lucerne)FabaceaeMain
Trifolium spp.FabaceaeMain

Growth Stages

Top of page Flowering stage, Vegetative growing stage

Biology and Ecology

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Genetics

The genetics of P. coronopus are complex. The plant is normally diploid (2n=10), but in a detailed study, Böcher et al. (1955) found one strain with 2n=11 and a single hexaploid plant with 2n=30. They treat the tetraploid form (2n=20) as a separate species P. commutata, presumably corresponding to P. coronopus subsp. commutata. A range of sources quoted by Missourri Botanical Garden (2012) confirm most forms being diploid (2n = 10 +/-1 or 2) but subsp. commutata tetraploid (2n = 20). Mohsenzadeh et al. (2008) conversely found the coronopus species to be tetraploid and the commutatus species diploid, but admit that this does not agree with previous publications.

Paliwal and Hyde (1959) found that male sterility in P. coronopus was associated with a single extra chromosome which is largely heterochromatic, shorter, and not homologous with any of the other chromosomes. No male-fertile plants contained this B-chromosome. However, this finding was contradicted by Raghuvanshi and Kumar (1983) who found no relationship between B chromosomes and male sterility. Male sterility has been further studied by Koelewijn and Damme (1995b; 1996) and by Damme et al. (2004) who found that both nuclear and cytoplasmic genes are involved in the inheritance of the male sterile character. They describe the nuclear (restorer) genetics of two cytoplasmic types. In both types, at least five multiple interacting nuclear genes are demonstrated, three with dominant and two with recessive restorer allele action. Intermediate partially male sterile plants can also occur (Koelewijn and Damme, 1996).

Other studies on the genetics of P. coronopus include a genetic diversity analysis of Plantago species and their interspecific hybrids through RAPD markers (Samarth and Vala, 2012) and the interspecific hybridization and induction of variability in Plantago species (Samarth and Fougat, 2012).

Reproductive Biology

P. coronopus is self-compatible (Samarth et al., 2009) but protogynous, meaning that the ovaries are receptive before pollen shedding (Bos, 1992). This facilitates out-crossing.

The large seeds are shed in the autumn and germinate almost immediately. The small seeds are retained in the capsule and although they are capable of germinating immediately, their germination is generally delayed (Rowling, 1933; Braza and Garcia, 2011). The seeds differ also in the fact that the larger seeds develop mucilage on wetting while the small seeds do not. One consequence of this is that the larger seeds rapidly sink in water while the small ones float indefinitely (Rowling, 1933). There may be an initial dormant period, which is longer, up to two months, at higher temperatures. Germination occurs between 10 and 25°C. This may require light. Germination of P. coronopus subsp. crassipes was 96% after 72 h at 20°C in light, but only 2% in the dark (Shem-Tov et al., 1999). Rowling (1933) also showed that light was needed in May, but there was little difference between dark and light germination in September.

Male steriles produced more (41% to 148%) and heavier (12% to 22%) seeds than hermaphrodites (Koelewijn and Damme, 1996). In the field, the contributions of seed size variation (15%) and inbreeding (9%) combine with 48% higher seed production to give a total fitness advantage of 70% to male steriles relative to hermaphrodites. This is probably sufficient for maintenance of gynodioecy under the nuclear-cytoplasmic inheritance of male sterility (Koelewijn and Damme, 2005).

Germination is reduced under saline conditions and completely prevented in 0.2 M NaCl solutions (Luciani et al., 2001). The importance of mucilaginous seeds for the survival of the plant species under desert conditions is discussed by Gutterman and Shem-Tov (1996). The mucilage causes the seeds to stick to the crust of the desert soil and prevents the seeds being washed away by showers.

Physiology and Phenology

The relative growth rate (RGR) during the vegetative stage is 300 mg g-1d-1 and drops at the initiation of flowering buds to 60 mg g-1d-1, a fivefold reduction. The initiation of flowering is stage-dependent. After the formation of about 14 leaves, plants start to form flowering buds, independent of weight or leaf size (Koelewijn, 2004a). Seed size effects lasted until the end of the experiment and were reflected in all morphological measurements. The proportion of small verses large seeds is influenced to some degree by the environment (Braza et al., 2010). When grown intermixed, selfed offspring were always inferior to their out-crossed relatives (Koelwijn, 2004b).

In east Poland (Sotek, 2007) populations of P. coronopus are mainly composed of perennial plants with a significant proportion of biennial and a small contribution of annual plants. The shorter cycle was characteristic of the individuals growing in the conditions of lack of competition or strongly limited competition of the co-existing plants.

Under desert conditions P. coronopus showed plasticity according to the day length; plants under long days were larger and produced more seeds that germinated to higher percentages (Shem-Tov and Gutterman, 2003).

Longevity

Established plants may apparently survive for several years, while Dodds (1953) refers to high germination from 3-year-old seeds.

Associations

Dodds (1953) reported a number of associations in a range of habitats: in salt marsh - Armeria maritima, Glaux maritima, Plantago maritima, Cochlearia officnialis and Triglochin maritime; and in transitional salt marshes or dunes - Agrostis stolonifera, P. maritima and G. maritime; in less saline conditions a much wider range of species was recorded.

In the Swina valley, Poland, extensive grazing by horses and cattle on salted meadows contributes to the protection of 21 plant species growing there, including a rare stand of P. coronopus (Warda and Rogalski, 2004).

Environmental Requirements

P. coronopus favours situations with light sandy soils and high light intensity; withstanding high summer temperatures and moderately low rainfall. It is adapted to highly saline conditions (Apaydin et al., 2009). A study recorded the salinity threshold of P. coronopus being reached at 25% of sea water levels and growth was strongly depressed by higher salinities (Koyro, 2006).

Growth stimulation by carbon dioxide enrichment beyond close-to-current concentrations is only likely to be seen under nutrient-rich conditions in semi-arid and possibly other drought-stressed grasslands (Grünzweig and Körner, 2003).

Tánczos and Hasselt (1992) reported that P. coronopus was able to tolerate prolonged freezing at -4°C but given its presence in Norway and Greenland it may be able to tolerate even lower temperatures. The plant has also been shown to tolerate high levels of iron (Schmidt and Fühner, 1998) and compacted soils (Popay et al., 1995).

Climate

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ClimateStatusDescriptionRemark
Cf - Warm temperate climate, wet all year Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year
Cs - Warm temperate climate with dry summer Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers
Cw - Warm temperate climate with dry winter Preferred Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)
Ds - Continental climate with dry summer Tolerated Continental climate with dry summer (Warm average temp. > 10°C, coldest month < 0°C, dry summers)
Dw - Continental climate with dry winter Tolerated Continental climate with dry winter (Warm average temp. > 10°C, coldest month < 0°C, dry winters)

Latitude/Altitude Ranges

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Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
60 40

Rainfall Regime

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Summer
Uniform
Winter

Soil Tolerances

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Soil drainage

  • free
  • seasonally waterlogged

Soil reaction

  • alkaline
  • neutral

Soil texture

  • light

Special soil tolerances

  • infertile
  • saline
  • shallow

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Cadophora luteo-olivacea Pathogen Agustí-Brisach et al., 2011
Capra hircus Herbivore Whole plant not specific Dodds, 1953
Ceuthorhynchidius dawsoni Predator Seeds Dodds, 1953
Equus caballus Herbivore Whole plant not specific Dodds, 1953
Mecinus collari Predator Stems Dodds, 1953
Messor barbarus Predator Seeds Detrain and Pasteels, 2000
Messor rugosus Predator Seeds Gutterman and Shem-Tov, 1997
Metzneria littorella Predator Seeds Dodds, 1953
Oryctolagus cuniculus Herbivore Whole plant not specific Dodds, 1953
Ovis aries Herbivore Whole plant not specific Dodds, 1953
Phthorimaea plantaginella Predator Roots Dodds, 1953
Phytomyza plantaginis Predator Leaves Dodds, 1953

Notes on Natural Enemies

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Natural enemies of P. coronopus include the fungal pathogen Cadophora luteo-olivacea, which occurs in Spain (Agustí-Brisach, et al., 2011) and the ants Messor barbarus in France (Detrain and Pasteels, 2000) and M. rugosus in Israel (Gutterman and Shem-Tov, 1997). These authors noted that the larger mucilaginous ombrohydrochoric seeds were less likely to be taken by the ants. Lepidopterans Metzneria littorella and Phthorimaea plantaginella, the coleopterans Ceuthorhynchidius dawsoni and Mecinus collari and the dipteran Phytomyza plantaginis are all listed by Dodds (1953) as affecting P. coronopus in the UK.

Dodds (1953) also recorded that rabbits (Oryctolagus cuniculus) graze on the plants, especially the more succulent forms in the regions of salt marshes. Soay sheep (Ovis aries), goats (Capra hircus) and ponies (Equus caballus) have also been recorded as grazing on the succulent form.

Means of Movement and Dispersal

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Natural Dispersal (Non-Biotic)

While the larger, mucilaginous seeds are adapted to stick where they fall onto wet soil, the smaller seeds may be carried in the capsule by wind or water movement. 

Accidental Introduction

Presumably P. coronopus could be accidently transported as a grass seed contaminant and also in garden waste. New infestations of P. coronopus subsp. commutata in south Germany and Austria have been associated with the use of salt on roads (Gerstberger, 2001).

Intentional Introduction

P. coronopus is available from some commercial suppliers as an ornamental or salad plant, but to what extent this has been responsible for its spread is not well documented.

Pathway Causes

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CauseNotesLong DistanceLocalReferences
Escape from confinement or garden escape Yes
Garden waste disposal Yes
Nursery trade Yes Yes
Seed trade Yes Yes

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Aircraft Yes
Floating vegetation and debris Yes
Mail Yes Yes
Plants or parts of plants Yes Yes

Plant Trade

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Plant parts liable to carry the pest in trade/transportPest stagesBorne internallyBorne externallyVisibility of pest or symptoms
True seeds (inc. grain)

Impact Summary

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CategoryImpact
Economic/livelihood Positive and negative
Environment (generally) Negative

Economic Impact

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In California, USA, P. coronopus is a weed problem in non-tilled orchards, irrigated pastures, and alfalfa and clover fields where mowing is used for weed control. Although slow to establish, P. coronopus is difficult to control when mature because of its extensive crown system.

It is a possible alternate host for the grapevive pathogen Cadophora luteo-olivacea (Agusti-Brisach et al., 2011).

Environmental Impact

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Impact on Biodiversity

It has been reported as forming dense mats that compete with native species and impact upon two threatened species in California, the showy Indian clover (Trifolium amoenum) (US Fish and Wildlife Service, 2008a) and Jesup's milkvetch (Astragalus robbinsii var. jesupii) (US Fish and Wildlife Service, 2008b).

P. coronopus is regarded as an environmental weed in Victoria and Western Australia (Queensland Government, 2012). Weber (2003) indicates it is invasive in Australia, but it has proved difficult to confirm the basis for this classification.

In Spain, Holm et al. (1979) list it as a ‘common’ weed.

Threatened Species

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Threatened SpeciesConservation StatusWhere ThreatenedMechanismReferencesNotes
Astragalus robbinsii var. jesupii (Jesup's milk-vetch)USA ESA listing as endangered species USA ESA listing as endangered speciesCaliforniaCompetition - smotheringUS Fish and Wildlife Service, 2008
Trifolium dichotomum (showy Indian clover)EN (IUCN red list: Endangered) EN (IUCN red list: Endangered); National list(s) National list(s); USA ESA listing as endangered species USA ESA listing as endangered speciesCaliforniaCompetition - smotheringUS Fish and Wildlife Service, 2008

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Has a broad native range
  • Highly adaptable to different environments
  • Highly mobile locally
  • Long lived
  • Has high reproductive potential
  • Has propagules that can remain viable for more than one year
  • Reproduces asexually
  • Has high genetic variability
Impact outcomes
  • Reduced native biodiversity
  • Threat to/ loss of endangered species
  • Threat to/ loss of native species
Impact mechanisms
  • Competition - smothering
Likelihood of entry/control
  • Difficult to identify/detect as a commodity contaminant

Uses List

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Animal feed, fodder, forage

  • Forage

Human food and beverage

  • Leaves (for beverage)
  • salad
  • Spices and culinary herbs
  • Vegetable

Medicinal, pharmaceutical

  • Source of medicine/pharmaceutical
  • Traditional/folklore

Similarities to Other Species/Conditions

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P. ovata (important as a medicinal plant in India) and P. lanceolata differ in having somewhat more erect, entire leaves and shorter inflorescences. The widely common P. major and P. media have much broader leaves.

Prevention and Control

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Physical/Mechanical Control

Control measures suggested for ornamental and perennial crops include cultivation, cover crops and the use of mulch (UC-IPM, 2012).

Chemical Control

Isoxaben, a relatively new broadleaf pre-emergent herbicide, has been effective in limiting germination of P. coronopus in turfgrass and some ornamental crops. Post-emergent 2,4-D, triclopyr, MCPA, and mecoprop can control seedlings, but control of established plants is much more difficult. For established plants, 2,4-D works best while triclopyr, MCPA, and mecoprop will only reduce vigour. Repeated applications to perennial plants with products containing 2,4-D or triclopyr can be helpful (UC-IPM, 2012). In lucerne or alfalfa, pre-emergence treatment with hexazinone has been successful. Glyphosate can be effective in plantation and orchard crops.

References

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Agustí-Brisach C; Gramaje D; León M; García-Jiménez J; Armengol J, 2011. Evaluation of vineyard weeds as potential hosts of black-foot and Petri disease pathogens. Plant Disease, 95(7):803-810. http://apsjournals.apsnet.org/loi/pdis

Al-Quran SA, 2004. Pollen morphology of Plantaginaceae in Jordan. Pakistan Journal of Biological Sciences, 7(9):1594-1602. http://www.ansinet.org/pjbs

Apaydin Z; Kutbay HG; Özbucak T; Yalçin E; Bilgin A, 2009. Relationship between vegetation zonation and edaphic factors in a salt-marsh community (Black Sea Coast). Polish Journal of Ecology, 57(1):99-112. http://psjc.icm.edu.pl/psjc/cgi-bin/pis_f.cgi?status=all

Bos M, 1992. Gene flow characters and population structure in Plantago lanceolata. Plantago: a multidisciplinary study [ed. by Kuiper, P. J. C.\Bos, M.]. Berlin, Germany: Springer-Verlag, 222-231.

Braza R; Arroyo J; García MB, 2010. Natural variation of fecundity components in a widespread plant with dimorphic seeds. Acta Oecologica, 36(5):471-476. http://www.sciencedirect.com/science/journal/1146609X

Braza R; García MB, 2011. Spreading recruitment over time to cope with environmental variability. Plant Ecology, 212(2):283-292. http://springerlink.metapress.com/link.asp?id=100328

Böcher TW; Larsen K; Rahn K, 1955. Experimental and cytological studies on plant species: III. Plantago coronopus and allied species. Hereditas, 41(3-4):423-453.

Council of Heads of Australasian Herbaria, 2012. Australia's Virtual Herbarium. http://avh.ala.org.au/

Damme JMMvan; Hundscheid MPJ; Ivanovic S; Koelewijn HP, 2004. Multiple CMS-restorer gene polymorphism in gynodioecious Plantago coronopus. Heredity, 93(2):175-181.

Detrain C; Pasteels JM, 2000. Seed preferences of the harvester ant Messor barbarus in a Mediterranean mosaic grassland (Hymenoptera: Formicidae). Sociobiology, 35(1):35-48; 23 ref.

Dodds JG, 1953. Plantago coronopus L. Journal of Ecology, 41(2):467-478.

Flowers of India, 2012. Flowers of India. http://www.flowersofindia.net/

GBIF, 2013. Global Biodiversity Information Facility. Global Biodiversity Information Facility (GBIF). http://data.gbif.org/species/

Gerstberger P, 2001. Plantago coronopus subsp. commutata introduced as a roadside halophyte in central Europe. (Plantago coronopus subsp. commutata als Strassenrandhalophyt eingebürgert in Mitteleuropa.) Tuexenia, No.21:249-256.

Grünzweig JM; Körner C, 2003. Differential phosphorus and nitrogen effects drive species and community responses to elevated CO2 in semi-arid grassland. Functional Ecology, 17(6):766-777.

Gutterman Y; Shem-Tov S, 1996. Structure and function of the mucilaginous seed coats of Plantago coronopus inhabiting the Negev desert of Israel. Israel Journal of Plant Sciences, 44(2/3):125-133.

Gutterman Y; Shem-Tov S, 1997. The efficiency of the strategy of mucilaginous seeds of some common annuals of the Negev adhering to the soil crust to delay collection by ants. Israel Journal of Plant Sciences, 45(4):317-327.

Holm LG; Pancho JV; Herberger JP; Plucknett DL, 1979. A geographical atlas of world weeds. New York, USA: John Wiley and Sons, 391 pp.

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03/12/2012 Original text by:

Chris Parker, Bristol, UK

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