Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide

Datasheet

Isatis tinctoria
(dyer's woad)

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Datasheet

Isatis tinctoria (dyer's woad)

Summary

  • Last modified
  • 20 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Host Plant
  • Preferred Scientific Name
  • Isatis tinctoria
  • Preferred Common Name
  • dyer's woad
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Spermatophyta
  •       Subphylum: Angiospermae
  •         Class: Dicotyledonae
  • Summary of Invasiveness
  • Isatis tinctoria is a herbaceous dye plant which has spread, often by human intervention, widely from its area of origin in central Asia or southeast Russia. Long valued for the dye it produces, it is of pest s...

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Pictures

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PictureTitleCaptionCopyright
Isatis tinctoria (dyer's woad); habit, showing leaves and flowers. This putative invasive species grows strictly in highly disturbance-prone settings. Box Elder County, Utah, USA. May, 2011.
TitleHabit
CaptionIsatis tinctoria (dyer's woad); habit, showing leaves and flowers. This putative invasive species grows strictly in highly disturbance-prone settings. Box Elder County, Utah, USA. May, 2011.
Copyright©Prof Matt Lavin/Bozeman, Montana, USA - CC BY-SA 2.0
Isatis tinctoria (dyer's woad); habit, showing leaves and flowers. This putative invasive species grows strictly in highly disturbance-prone settings. Box Elder County, Utah, USA. May, 2011.
HabitIsatis tinctoria (dyer's woad); habit, showing leaves and flowers. This putative invasive species grows strictly in highly disturbance-prone settings. Box Elder County, Utah, USA. May, 2011.©Prof Matt Lavin/Bozeman, Montana, USA - CC BY-SA 2.0
Isatis tinctoria (dyer's woad); habit, ca.160m a.s.l. Neue Donau, Vienna-Floridsdorf, Austria. -  April 2014.
TitleHabit
CaptionIsatis tinctoria (dyer's woad); habit, ca.160m a.s.l. Neue Donau, Vienna-Floridsdorf, Austria. - April 2014.
Copyright©Stefan.lefnaer-2014/via wikipedia - CC BY-SA 4.0
Isatis tinctoria (dyer's woad); habit, ca.160m a.s.l. Neue Donau, Vienna-Floridsdorf, Austria. -  April 2014.
HabitIsatis tinctoria (dyer's woad); habit, ca.160m a.s.l. Neue Donau, Vienna-Floridsdorf, Austria. - April 2014.©Stefan.lefnaer-2014/via wikipedia - CC BY-SA 4.0
Isatis tinctoria (dyer's woad); habit, showing leaves. This putative invasive species grows strictly in highly disturbance-prone settings. Box Elder County, Utah, USA.
TitleHabit
CaptionIsatis tinctoria (dyer's woad); habit, showing leaves. This putative invasive species grows strictly in highly disturbance-prone settings. Box Elder County, Utah, USA.
Copyright©Prof Matt Lavin/Bozeman, Montana, USA - CC BY-SA 2.0
Isatis tinctoria (dyer's woad); habit, showing leaves. This putative invasive species grows strictly in highly disturbance-prone settings. Box Elder County, Utah, USA.
HabitIsatis tinctoria (dyer's woad); habit, showing leaves. This putative invasive species grows strictly in highly disturbance-prone settings. Box Elder County, Utah, USA.©Prof Matt Lavin/Bozeman, Montana, USA - CC BY-SA 2.0
Isatis tinctoria (dyer's woad); habit, showing leaves. This putative invasive species grows strictly in highly disturbance-prone settings. Box Elder County, Utah, USA.
TitleHabit
CaptionIsatis tinctoria (dyer's woad); habit, showing leaves. This putative invasive species grows strictly in highly disturbance-prone settings. Box Elder County, Utah, USA.
Copyright©Prof Matt Lavin/Bozeman, Montana, USA - CC BY-SA 2.0
Isatis tinctoria (dyer's woad); habit, showing leaves. This putative invasive species grows strictly in highly disturbance-prone settings. Box Elder County, Utah, USA.
HabitIsatis tinctoria (dyer's woad); habit, showing leaves. This putative invasive species grows strictly in highly disturbance-prone settings. Box Elder County, Utah, USA.©Prof Matt Lavin/Bozeman, Montana, USA - CC BY-SA 2.0
Isatis tinctoria (dyer's woad); basal rosette. USA.
TitleRosette
CaptionIsatis tinctoria (dyer's woad); basal rosette. USA.
Copyright©Steve Dewey/Utah State University/Bugwood.org - CC BY 3.0 US
Isatis tinctoria (dyer's woad); basal rosette. USA.
RosetteIsatis tinctoria (dyer's woad); basal rosette. USA.©Steve Dewey/Utah State University/Bugwood.org - CC BY 3.0 US
Isatis tinctoria (dyer's woad); seedpods. USA.
TitleSeedpods
CaptionIsatis tinctoria (dyer's woad); seedpods. USA.
Copyright©Steve Dewey/Utah State University/Bugwood.org - CC BY 3.0 US
Isatis tinctoria (dyer's woad); seedpods. USA.
SeedpodsIsatis tinctoria (dyer's woad); seedpods. USA.©Steve Dewey/Utah State University/Bugwood.org - CC BY 3.0 US

Identity

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Preferred Scientific Name

  • Isatis tinctoria L.

Preferred Common Name

  • dyer's woad

Other Scientific Names

  • Isatis indigotica Fortune

International Common Names

  • English: asp of Jerusalem; dyers woad; dyers' woad; glastum
  • Spanish: glasto; gualda; hierba pastel
  • French: bleu de Picardie; guède; pastel des teinturiers; vouède; waide; wède
  • Portuguese: pastel-dos-tintureiros

Local Common Names

  • Germany: Faerber- Waid; Färberwaid
  • Netherlands: wede
  • Sweden: vejde

EPPO code

  • ISATI (Isatis tinctoria)

Summary of Invasiveness

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Isatis tinctoria is a herbaceous dye plant which has spread, often by human intervention, widely from its area of origin in central Asia or southeast Russia. Long valued for the dye it produces, it is of pest status in alkaline soils of some western states of the United States. It is the main biotic threat to the endangered Siskiyou Mariposa Lily (Calochortus persistens) found on the California-Oregon border and also threatens the endangered Yreka phlox, Phlox hirsuta. It responds well to high inputs of fertilizer and water, but also competes well in disturbed, dry and nutrient poor conditions. Its fruits have germination-inhibiting factors and the plant produces glucosinates which may alter soil microbiological properties. It causes considerable production losses of desired plants in rangelands. Control methods are effective, but with logistical challenges, especially in more remote areas. A native rust, Micropuccinia thlaspeos, limits, but does not prevent spread. Arthropod biological control agents are being researched in Europe, but to date none are specific enough.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Plantae
  •         Phylum: Spermatophyta
  •             Subphylum: Angiospermae
  •                 Class: Dicotyledonae
  •                     Order: Capparidales
  •                         Family: Brassicaceae
  •                             Genus: Isatis
  •                                 Species: Isatis tinctoria

Notes on Taxonomy and Nomenclature

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Isatis is a difficult genus requiring revision according to Ball and Akeroyd (2010) who state that I. leavigata, I. costata, I. praecox and I. arenaria are all very similar to, and some probably subspecies of, I. tinctoria, in addition to listing four recognized subspecies; athoa, tinctoria, tomentella and corymbosa. The plant can be very variable in the field (Callihan et al, 1984; Zouhar, 2009) creating further difficulties. One reported synonym seen quite frequently in literature is I. indigotica in China, although Gilbert et al (2002) separated out I. indigotica from I tinctoria on their genotypes. Many common names are given by Marhold (2011), but dyer’s woad, or dyers woad, is how it is best known in the USA, where it can be a pest species.

Description

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The following paragraph is taken from Zouhar (2009, and references therein which relate largely to the USA), who notes that there exist substantial morphological variations in the species globally.

I. tinctoria is typically a biennial or short-lived usually monocarpic perennial, sometimes described as a winter annual, but this may be occasional or very localized. The plant begins as a rosette with long-petioled basal leaves usually 4-10 cm long and 0.8-4 cm wide, usually covered with simple leaves. Leaves are blue-green and slightly pubescent (Varga and Evans, 1978). Some sources suggest that around 20 stalks begin to develop from each rosette with seven or fewer maturing, while others consider that the plant usually has one main stem, simple below and branched above. Stems are erect, ranging from 35 to 120 cm tall, but mostly in the 50-90 cm range, with stem leaves, narrower than basal leaves, reducing as they rise up the stem. The plants are glabrous or hirsute. The flowers are about 6mm wide with petals about 3.5cm long. Flowers are borne in numerous, compound racemes forming a large, terminal panicle. The root system is dominated by a substantial taproot which can exceed 1.5 m in depth, with lateral roots on the upper 20-30 cm that spread laterally about 40 cm.

I. tinctoria fruits are samaroid, indehiscent silicles ranging from 8 to 18 mm long and 2.5 to 7 mm wide with a single, median seed. Fruits are dark to black at maturity and droop from a short, slender pedicel that is ascending to reflexed or recurved.

Kizil (2006) gives details of morphological and agronomic characteristics of I. tinctoria and other Isatis species. Plant height ranged between 73 and 112cm, averaging at 91cm, perhaps slightly taller than figures above.  

Plant Type

Top of page Annual
Biennial
Broadleaved
Herbaceous
Perennial
Seed propagated
Shrub
Vegetatively propagated

Distribution

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The centre of origin of I. tinctoria is thought to be Central Asia (Spataro et al., 2007). It is also reported as a native of southeastern Russia (Hurry, 1930; McConnell et al., 1999), to south-west Asia and perhaps to some parts of south-eastern Europe (Ball and Akeroyd, 2010) and to Afghanistan, western Tibet and China (Anon, 2015a). Hegi (1986) reports it to be native to the steppe area around the Caucasus, Central Asia to eastern Siberia and the Middle East. It is recorded as present in all European countries other than Ireland and Iceland and islands such as the Balearics and Crete (Ball and Akeroyd, 2010, Marhold, 2011), although Reynolds (2002) also reports it, as rare, in Ireland. It has been widely introduced globally.

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

AfghanistanPresentNative Not invasive Moazzeni and Zarre, 2006
ArmeniaPresentNative Not invasive USDA-ARS, 2015
AzerbaijanPresentNative Not invasive Marhold, 2011
ChinaPresentIntroducedUSDA-ARS, 2015Introduced thousands of years BP
-FujianPresentIntroduced Not invasive USDA-ARS, 2015
-GansuPresentIntroduced Not invasive USDA-ARS, 2015
-GuizhouPresentIntroduced Not invasive USDA-ARS, 2015
-HebeiPresentIntroduced Not invasive USDA-ARS, 2015
-HenanPresentIntroduced Not invasive USDA-ARS, 2015
-HubeiPresentIntroduced Not invasive USDA-ARS, 2015
-JiangxiPresentIntroduced Not invasive USDA-ARS, 2015
-LiaoningPresentIntroduced Not invasive USDA-ARS, 2015
-Nei MengguPresentIntroduced Not invasive USDA-ARS, 2015
-ShaanxiPresentIntroduced Not invasive USDA-ARS, 2015
-ShandongPresentIntroduced Not invasive USDA-ARS, 2015
-ShanxiPresentIntroduced Not invasive USDA-ARS, 2015
-SichuanPresentIntroduced Not invasive USDA-ARS, 2015
-TibetPresentNativeAnon, 2015a
-XinjiangPresentIntroduced Not invasive USDA-ARS, 2015
-YunnanPresentIntroduced Not invasive USDA-ARS, 2015
Georgia (Republic of)PresentNative Not invasive Marhold, 2011
IndiaPresentIntroduced Not invasive USDA-ARS, 2015Introduced thousands of years BP
-Himachal PradeshPresentIntroduced Not invasive USDA-ARS, 2015
-Jammu and KashmirPresentIntroduced Not invasive USDA-ARS, 2015
IranPresentIntroduced Not invasive Moazzeni and Zarre, 2006
JapanPresentIntroduced Not invasive USDA-ARS, 2015
-HokkaidoPresentIntroduced Not invasive USDA-ARS, 2015
-HonshuPresentIntroduced Not invasive USDA-ARS, 2015
KazakhstanPresentNative Not invasive USDA-ARS, 2015
Korea, Republic ofPresentIntroduced Not invasive USDA-ARS, 2015
LebanonPresentIntroduced Not invasive USDA-ARS, 2015
MongoliaPresentIntroduced Not invasive Encyclopedia of Life, 2015
PakistanPresentIntroduced Not invasive USDA-ARS, 2015
TajikistanPresentNative Not invasive Encyclopedia of Life, 2015
TurkeyPresentNative Not invasive USDA-ARS, 2015
UzbekistanPresentNative Not invasive USDA-ARS, 2015

Africa

AlgeriaPresentIntroduced Not invasive USDA-ARS, 2015
MoroccoPresentIntroduced Not invasive USDA-ARS, 2015
Spain
-Canary IslandsPresentIntroduced Not invasive Browne, 1834

North America

CanadaPresentIntroduced Not invasive USDA-ARS, 2015
-British ColumbiaPresentIntroduced Not invasive USDA-ARS, 2015
-Newfoundland and LabradorPresentIntroduced Not invasive USDA-ARS, 2015
-OntarioPresentIntroduced Not invasive USDA-ARS, 2015
-QuebecPresentIntroduced Not invasive USDA-ARS, 2015
USAWidespreadIntroduced16th-17th C Invasive USDA-ARS, 2015Invasive in some western states, not harmful in eastern states.
-ArizonaPresentIntroduced20th CUSDA-NRCS, 2015Declared prohibited noxious weed
-CaliforniaWidespreadIntroducedEarly 20th C Invasive USDA-ARS, 2015Declared B list noxious weed
-ColoradoPresentIntroduced20th CUSDA-NRCS, 2015Declared A list noxious weed
-IdahoWidespreadIntroduced20th C Invasive USDA-ARS, 2015Declared noxious weed
-IllinoisWidespreadIntroduced20th C Invasive USDA-ARS, 2015Not a significant pest species
-MissouriWidespreadIntroduced20th C Invasive USDA-ARS, 2015Not a significant pest species
-MontanaWidespreadIntroduced20th C Invasive USDA-ARS, 2015Declared category 2 noxious weed. Successful control programme undertaken
-NevadaWidespreadIntroducedEarly 20th C Invasive USDA-ARS, 2015Declared noxious weed
-New MexicoWidespreadIntroduced20th C Invasive USDA-ARS, 2015Declared Class A noxious weed
-New YorkPresentIntroducedlate 17th –early 18th C Not invasive Flora of North America Editorial Committee, 2015Not a significant pest species
-OregonWidespreadIntroducedEarly 20th C Invasive USDA-ARS, 2015Declared B designated weed
-UtahWidespreadIntroducedlate 19th C Invasive USDA-ARS, 2015Declared noxious weed
-VirginiaPresentIntroducedlate 17th –early 18th C Not invasive USDA-ARS, 2015Not a significant pest species
-WashingtonWidespreadIntroducedEarly 20th C Invasive USDA-ARS, 2015Declared Class A noxious weed
-West VirginiaPresentIntroducedlate 17th –early 18th C Not invasive USDA-ARS, 2015Not a significant pest species
-WyomingWidespreadIntroducedEarly 20th C Invasive USDA-ARS, 2015Declared noxious weed

South America

ChilePresentIntroduced Not invasive USDA-ARS, 2015
PeruPresentIntroduced Not invasive Peru Checklist, 2015

Europe

AlbaniaPresentNative Not invasive USDA-ARS, 2015
AndorraPresentIntroduced Not invasive Ball and Akeroyd, 2010
AustriaPresentIntroduced Not invasive Ball and Akeroyd, 2010
BelarusPresentNative Not invasive USDA-ARS, 2015
BelgiumPresentIntroducedBall and Akeroyd, 2010
Bosnia-HercegovinaPresentNative Not invasive USDA-ARS, 2015
BulgariaPresentNativeUSDA-ARS, 2015
CroatiaPresentNative Not invasive USDA-ARS, 2015
CyprusPresentIntroduced Not invasive Ball and Akeroyd, 2010
Czech RepublicPresentNative Not invasive USDA-ARS, 2015
Czechoslovakia (former)PresentNative Not invasive Ball and Akeroyd, 2010
DenmarkPresentIntroducedUSDA-ARS, 2015
EstoniaPresentIntroduced Not invasive USDA-ARS, 2015
FinlandPresentIntroduced Not invasive USDA-ARS, 2015
FrancePresentIntroduced Not invasive USDA-ARS, 2015
-CorsicaPresentIntroduced Not invasive Ball and Akeroyd, 2010
GermanyPresentIntroduced Not invasive USDA-ARS, 2015
GibraltarPresentIntroduced Not invasive Ball and Akeroyd, 2010Assumed present
GreecePresentNative Not invasive Ball and Akeroyd, 2010; USDA-ARS, 2015Absent from Crete, Ball and Akeroyd 2010
HungaryPresentIntroduced Not invasive USDA-ARS, 2015
IrelandPresentIntroduced Not invasive Reynolds, 2002Ball and Akeroyd 2010 regarded it as absent
ItalyPresentIntroduced Not invasive USDA-ARS, 2015
LatviaPresentIntroduced Not invasive USDA-ARS, 2015
LiechtensteinPresentIntroduced Not invasive Ball and Akeroyd, 2010
LithuaniaPresentIntroduced Not invasive USDA-ARS, 2015
LuxembourgPresentIntroduced Not invasive Ball and Akeroyd, 2010
MacedoniaPresentNative Not invasive USDA-ARS, 2015
MaltaPresentIntroduced Not invasive Marhold, 2011
MoldovaPresentNative Not invasive Ball and Akeroyd, 2010
MonacoPresentIntroduced Not invasive Ball and Akeroyd, 2010
MontenegroPresentNative Not invasive USDA-ARS, 2015
NetherlandsPresentIntroduced Not invasive USDA-ARS, 2015
NorwayPresentIntroduced Not invasive USDA-ARS, 2015
PolandPresentBall and Akeroyd, 2010
PortugalPresentBall and Akeroyd, 2010
-MadeiraPresentIntroduced Not invasive Marhold, 2011
RomaniaPresentNative Not invasive USDA-ARS, 2015
Russian FederationPresentNative Not invasive USDA-ARS, 2015
-Southern RussiaPresentNative Not invasive USDA-ARS, 2015
San MarinoPresentIntroduced Not invasive Ball and Akeroyd, 2010
SerbiaPresentNative Not invasive USDA-ARS, 2015
SlovakiaPresentNative Not invasive USDA-ARS, 2015
SloveniaPresentNative Not invasive USDA-ARS, 2015
SpainPresentIntroduced Not invasive USDA-ARS, 2015
SwedenPresentIntroduced Not invasive USDA-ARS, 2015
SwitzerlandPresentIntroduced Not invasive Ball and Akeroyd, 2010
UKPresentIntroduced Not invasive USDA-ARS, 2015
UkrainePresentNative Not invasive USDA-ARS, 2015
Yugoslavia (former)PresentIntroduced Not invasive Ball and Akeroyd, 2010
Yugoslavia (Serbia and Montenegro)PresentIntroduced Not invasive

Oceania

AustraliaPresentIntroduced Not invasive Callihan et al., 1984Grown and sold in Australia, for artisan and hobby dyeing

History of Introduction and Spread

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I. tinctoria has been long established in Europe, largely due to its use in the dyeing industry; Zech-Matterne and Leconte (2010) suggest that it was probably cultivated around 2400-2500 BP in the north of what is now France. Its cultivation in Britain at that time is likely, as the inhabitants were alleged to dye their bodies with woad, a practice disputed (Anon, 2015b). The blue impression may have been from tattooing or scarification rituals according to Carr (2005), who believes that it is unknown whether I. tinctoria was used in the Iron Age as a body dye, but that it can function as a body dye (G. Carr pers.comm). It was introduced into Italy in “remote times” with the Romans first using it for dyeing (Guarino et al., 2000). Major woad producers were found in England, France, Germany, Italy and the Netherlands (Hurry, 1930).

In northwest Iran, clothes from workers (and hence likely to be wearing locally produced clothes rather than expensive imported clothes) buried in a salt mine accident around 2000 BP contained dye probably from woad (Mouri et al., 2014). Its spread eastwards, including to China and Japan, are also likely to have occurred thousands of years ago (Zouhar, 2009); its medicinal uses are valued in these countries. It is grown in Australia, seeds and plants being sold by nurseries, but it is not a declared noxious weed (Australian Weeds Committee, 2015). It occurs in Algeria and Morocco, but apparently nowhere else on the African mainland (African Plant Database, 2015). Its long history of use confuses its definition as a native or introduced plant in many European and Asian countries. Webb (1985) defines a native plant as one that evolved or arrived in a region before the Neolithic period, or arrived since without any human activity being involved, a view supported by Richardson et al. (2000). All plants reaching an area through the intervention of neolithic or post-neolithic humans or their domestic animals are introduced plants. On that view, I tinctoria is introduced in most countries of north and south-west Europe and probably to most of central Europe as well. The records of Introduced or Native in the table reflect more the location of a country rather than confirmed knowledge of how I. tinctoria became established. Marhold (2011) records the plant as native to all of these countries, possibly because of their presence in these countries for at least 2000 years. Reynolds (2002) describes the plant as an alien in Ireland.

I. tinctoria was taken from Europe into North America by early colonists as a textile dye crop, possibly limited to Virginia in colonial times (Varga and Evans, 1978). It was then accidentally spread as a crop seed contaminant, from Ireland (despite the plant being rare in Ireland), in the early part of the twentieth century. According to McConnell et al. (1999) who also cite Roché (1992) and Callihan (1990), the introduction into the western USA resulted from contaminated alfafa (lucerne) seed, into California and Utah respectively. Finlay (2009) and Sagers (1991), however, state that it was deliberately introduced by Mormon pioneers into Utah in the nineteenth century, although this species is not noted by Wheeler (2011) who studied the plants used by Mormon settlers. Woad had largely been replaced by indigo from Indigofera tinctoria, by the eighteenth century, which became a major crop, especially in South Carolina (West, 2015; Martin, 2015), so there would appear to be no reason why settlers would deliberately carry woad with them.

Baun (2009) states that I. tinctoria was believed to have been introduced into the Scott Valley, in Siskiyou County California, in the mid 1880s, through contaminated packing material protecting a piano shipped from the Eastern USA. The contaminated seed route seems certain, but whether that was the first introduction into the west cannot be confirmed. Extensive personal investigations carried out by Dr Ralph Whitesides with veteran scientists in Utah confirm that it was most likely introduced on a number of occasions, both intentionally and non-intentionally (RE Whitesides pers comm.). Rydberg (1919) recorded Isatis tinctoria in his Key to the Rocky Mountain Flora which perhaps indicates that it was present earlier than suggested by the contaminated seed route. The history is important as it is likely that plants conveyed to the USA by early colonists may have more variable, and certainly different, genotypes than those carried as seed contamination.

I. tinctoria is a declared noxious weed in 11 western US states: Arizona, California, Colorado, Idaho, Montana, Nevada, New Mexico, Oregon, Utah, Washington and Wyoming (USDA-NRCS, 2015), but also occurs in a number of the eastern states: Illinois, Missouri, New York, Virginia and West Virginia (Flora of North America Editorial Committee, 2015; Missouri Botanical Garden, 2015), as well as in a number of Canadian provinces (USDA-NRCS, 2015).

The species has also been introduced into Chile and Peru in the Americas.

Introductions

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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
Canada Europe 17th C Industrial purposes (pathway cause) Yes No USDA-ARS (2015) Carried with early colonists to North America for domestic and small scale community use
USA Europe 17th C Industrial purposes (pathway cause) Yes No USDA-ARS (2015) Carried with early colonists to North America for domestic and small scale community use

Risk of Introduction

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It is probable that I. tinctoria has already established itself in appropriate habitats through many centuries of trade, travel and colonization. Renewed interest in the use of woad may lead to wider distribution through trade routes, but at a low level compared with previous times. Where it is a serious weed (largely some states in the western USA) legislation should prevent this movement. 

Habitat

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It has been suggested that I. tinctoria is well adapted to the western USA, because the plant is adapted to alkaline conditions and an arid climate (McConnell et al., 1999). The species thrives on rocky soil with limited water-holding capacity (McConnell et al., 1999), but it can grow well under a wide variety of habitats. This includes special habitats where rare plants are found – the endangered Siskiyou Mariposa Lily (Calochortus persistens) was thought to exist in one site, in Siskiyou County, California, where it is found on open ridgetops and north-facing slopes, with bedrock of gray phylite and sandstones high in potassium and low in calcium, iron and magnesium (Diggles et al., 2003). An Oregon population also exists (Anon, 2012). Equally I. tinctoria can dominate plant communities in pastures, forests and rangelands when it occurs in pest status in various western states of the United States. Reeves (2010) who recorded its growth on dry, coarse textured rocky soils also noted it capable of invading undisturbed and disturbed sites such as roadsides, railroad right of ways, pastures, grain and alfalfa fields, forests and rangelands, especially big sagebrush (Artemisia tridentata) communities. In Utah, I. tinctoria prefers loose, alkaline soils (Varga and Evans, 1978). It also grows on mining spoil heaps (Durães et al, 2015), lava (Anon, 2015c) and under cultivation with the addition of high levels of fertilizer (Howard, 2010). Hurry (1930) noted that it responds well to intensive cultivation.

The wide geographic range of I. tinctoria in Europe and Asia may be an important feature of its success in the western USA. It has been suggested that widely distributed species have developed the ability to tolerate varied biotic and resource conditions and to resist many potential enemies (Pyšek et al., 2015). I. tinctoria does show a high degree of genetic diversity (Gilbert et al., 2002).

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial
Terrestrial – ManagedCultivated / agricultural land Secondary/tolerated habitat Harmful (pest or invasive)
Cultivated / agricultural land Secondary/tolerated habitat Productive/non-natural
Managed forests, plantations and orchards Present, no further details Harmful (pest or invasive)
Managed grasslands (grazing systems) Secondary/tolerated habitat Harmful (pest or invasive)
Disturbed areas Principal habitat Harmful (pest or invasive)
Rail / roadsides Principal habitat Harmful (pest or invasive)
Terrestrial ‑ Natural / Semi-naturalNatural forests Secondary/tolerated habitat Harmful (pest or invasive)
Natural grasslands Principal habitat Harmful (pest or invasive)
Arid regions Principal habitat Harmful (pest or invasive)

Hosts/Species Affected

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I. tinctoria competes with plants of pastures, forests and rangeland and with cereal crops and alfalfa. It is most troublesome on rangeland, in disturbed non-crop sites and in undisturbed natural areas in the intermountain west (DiTomaso et al., 2013). Callihan et al. (1984) list 23 plants of rangeland, 13 of pastures and cropland and 12 of disturbed areas that were observed, collected or associated with I. tinctoria. The rangeland habitats were dominated by Artemisia tridentata (big sagebrush). I. tinctoria is reported to quickly suppress annual grasses in rangeland, but perennial grasses appear to co-exist moderately well (Callihan et al., 1984). Perennial grasses were the main native grass species in western rangeland before being replaced by introduced annual grasses (DiTomaso, 2000) which were aided by overgrazing by introduced domestic animals. These livestock tend to avoid grazing I. tinctoria (DiTomaso, 2000).

Growth Stages

Top of page Pre-emergence, Seedling stage, Vegetative growing stage

Biology and Ecology

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Genetics

A tetraploid outbreeding plant (Spataro and Negri, 2008a) (2n = 4x =28). The limits of the genus Isatis are debatable and it is likely that the number of species will be greatly reduced (Al-Shehbaz et al., 2006). Spataro et al. (2007) carried out a study of fifteen Eurasian populations of I. tinctoria and demonstrated wide variation within and between populations. This was reinforced with studies showing wide adaptability to different mountain conditions and populations with high morphologic and genetic variability (Spataro and Negri, 2008b). Polyploid plants (with three or more complete chromosome sets) have been obtained and subsequent lines were characterized by high fertility and yields (Qiao et al., 1989).

Reproductive Biology

This paragraph is taken from Zouhar (2009 and references therein) unless stated.

I. tinctoria requires vernalization to achieve flowering, with crown rosettes (second year) and seedling rosettes responding differently to temperature regimes. Seed production is high, at 350-500 per plant, although there are reports of 10,000 from selected plants (McConnell et al., 1999). Kizil (2006) reported a much lower average of 88 seeds per plant from Turkey. They are still contained within fruits (silicles) when these are shed, usually at maturity although some remain on the plants until winter. Fruits generally drop close to the plant, mostly within 50cm, with longer distances being achieved through ant dispersal, human and animal activity, and water. Much dispersal has been through contaminated seed or baled alfalfa hay. The fruits contain water soluble germination inhibitors which may reduce inter- and intra-specific competition; seeds outside the fruits germinate easily (Young and Evans, 1971). Seed viability within the soil appears limited, with most seeds non-viable within 12 months, but this may be related to levels of soil moisture. Seedling establishment seems to be promoted by close proximity to sagebush plants, but in the field 97% of seeds fail to achieve successful establishment. Final numbers of established plants appear to be similar regardless of initial seed levels. Typically, germination occurs in the spring or autumn, the plant remains as a rosette of basal leaves the following summer and winter, and flowers in the second year with seeds ripening in June and July (McConnell et al., 1999). Plants die after producing seed according to Jacobs and Pokorny (2007), but Callihan (1990) states that the crown buds often produce new growth for additional years.

Physiology and Phenology

The combination of the tap root and lateral roots enables moisture to be obtained from an extensive soil profile, increasing its competitive fitness relative to annual and perennial plants in semi-arid areas (Jacobs and Pokorny, 2007). This two layered rooting capacity has been postulated to provide I. tinctoria with a strong competitive advantage in sagebrush (Artemisia spp.) in the Intermountain West (Sturges, 1977). Woad fruits contain water-soluble germination-inhibiting substances (Young and Evans, 1971); these have an effect on a restricted group of competing seeds, both of I. tinctoria and other species, but the reliance on rainfall to leach away these substances presumably enhances the chances of germinating with suitable soil moisture.

Brassica crops often appear to have an inhibitory effect on subsequent crops and this has been linked to levels of indole glucosinates or their breakdown products; I. tinctoria specifically has been implicated in reducing soil fertility (Iqbal, 2012). Related brassicas are known to be heavy users of nitrogen and phosphorus, as well as water (Varga and Evans, 1978). I. tinctoria seeds contain glucobrassicin and shoots contain this and also two other indole glucosinates (Elliott and Stowe, 1971b) which, the authors suggested, may explain the deleterious effect of the species on subsequent plantings (and possibly on existing competing plants). Some products formed by glucobrassicin decomposition have high goiter-inducing activity for grazing animals (Elliott and Stowe, 1971a) and the chemical composition may inhibit extensive grazing. Consequently, it may be postulated that these compounds suppress competing plants and reduce levels of herbivory. Isothiocyanates, derived from glucosinolates, inhibit nitrification processes by direct effects on the size of communities of nitrifying bacteria and by reducing their nitrifying activities (Bending and Lincoln, 2000), further modifying the original environment. The plant also shows competitive physiological adaptations in relation to nitrogen, light and water.

Monaco et al (2005) hypothesized that I. tinctoria would demonstrate low plasticity in growth and physiology with variable soil nitrogen and high morphological or physiological plasticity in response to variable light and water. This was confirmed; the plant increased leaf area when grown under shade compared with greater light and increased overall productivity with additional water. The results supported the theory that early succession of plants on poor soils is based on colonizing, not competitive, ability (Huston and Smith, 1987; Tilman and Wedin, 1991).

Longevity

Although described as a winter annual, biennial or short-lived perennial (McConnell, 1990) I. tinctoria can switch to vegetative reproduction through buds on the root crown (Farah et al., 1988) and from the taproot if the rosette leaves are removed (Roché, 1992). Vorobiov (1960) describes I. tinctoria as a biennial requiring one hibernation to bolt and flower.

Population Size and Structure

I. tinctoria may occur in small patches or as scattered plants or occupy large areas virtually as monocultures (Varga and Evans, 1978), spreading rapidly. The spread of the species was estimated at 14% per year in the Pacific Northwest (Jacobs and Pokorny, 2007). In 1990, I. tinctoria was estimated to occur in 10,000 ha in Idaho (Callihan, 1990). In Montana, I. tinctoria populations were anticipated to cover 39,000 ha, but Montana’s Dyer’s Woad Cooperation Project reduced the area to 2.6 ha (Pokorny and Kreuger-Mangold, 2007).

Nutrition

When grown commercially, the addition of mineral or organic nitrogen fertilizer resulted in increased biomass (Orsini et al., 2012); nitrogen fertilizer requirements may be around 220 kg/hectare (Anon, 2004). Monaco et al (2005, see Physiology above) demonstrated that the plant has low plasticity in relation to soil nitrogen with relatively small differences occurring between different nitrogen levels at any specific water status of the soil. It can thrive on poor rocky soils with limited water availability (McConnell, 1990) including those high in potassium and low in calcium, iron and magnesium (Diggles et al., 2003); equally it can grow well in rangeland, agricultural fields and forests covering a range of fertility types.

Associations

Associations with Artemisia tridentata have been noted (Callihan et al., 1984).

Environmental Requirements

I. tinctoria is associated with loose, alkaline soils in Utah and, more generally, in the western United States (Varga and Evans, 1978; McConnell 1999) and for domestic cultivation a sandy-loamy or gritty-loamy soil of pH 8-10 is recommended (Hortipedia, 2015). When cultivated it can utilize high levels of fertilizer and water, but is capable of competitive growth under arid and nutrient poor conditions. There is no information available for temperature requirements, but its range and habitats, through Eurasian steppes to Morocco and Arizona, shows that it can survive very cold winters and also very hot conditions. I. tinctoria is often found in association with Artemisia tridentata which is capable of surviving -18°C to 38°C extremes in the field (Loik and Redar, 2004) and short term tolerance to 46°C was shown by leaves (Loik and Harte, 1996). It may be conjectured that I. tinctoria has a similar capacity to survive these conditions. 

Climate

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ClimateStatusDescriptionRemark
B - Dry (arid and semi-arid) Tolerated < 860mm precipitation annually
BS - Steppe climate Tolerated > 430mm and < 860mm annual precipitation
Cf - Warm temperate climate, wet all year Tolerated 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 Tolerated Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)
Df - Continental climate, wet all year Tolerated Continental climate, wet all year (Warm average temp. > 10°C, coldest month < 0°C, wet all year)
Ds - Continental climate with dry summer Preferred 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)
64 19

Rainfall

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ParameterLower limitUpper limitDescription
Mean annual rainfall250mm; lower/upper limits

Rainfall Regime

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Soil Tolerances

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Soil drainage

  • free

Soil reaction

  • alkaline
  • neutral

Soil texture

  • light
  • medium

Special soil tolerances

  • infertile

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Aulacobaris fallax Herbivore
Ceutorhynchus peyerimhoffi Herbivore Seeds
Ceutorhynchus rusticus Herbivore Roots
Psylliodes isatidis Herbivore
Puccinia thlaspeos Pathogen All Stages not specific USA

Notes on Natural Enemies

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Many incidental natural enemies occur; Hurry (1930) lists a few including the turnip fly, Phyllotreta nemorum and the cabbage caterpillar Pieris brassicae and fungal pathogens such as Aecidium isatidis, Ophiobolus herbarum, Cicinnobolus cesati [Ampelomyces quisqualis], Monilia glasti and Oidium erysiphoides [Podosphaera fuliginea]. Hinz et al. (2007) list 56 insect species, two nematodes and four fungal pathogens associated with I. tinctoria found from Eurasian literature. In China, Ma (2007) obtained 172 fungal isolates from I. tinctoria (I. indigotica) exhibiting root rot, comprised of Fusarium solani [Haematonectria haematococca], F. oxysporum and Rhizoctonia cerealis [Ceratobasidium cereale]. Alternaria brassicicola causing leafspot on I. tinctoria was reported by Gao et al (2014) who also noted Alternaria napiformis and A. brassicae that have previously been reported as causal agents of leaf spot on the plant. The eriophyid mite Metaculus rapistri was collected from I. tinctoria in Turkey (Monfreda and Lillo, 2012), however this was a misidentification and a description of the Metaculus sp. that was collected from I. tinctoria is currently underway (P. Weyl, CABI Switerland, personal communication, 2017) and Cortat et al. (2008) record nine insect species that caused sufficient impact as to be considered for biological control studies.

Mammals have been reported as grazing on the plant; sheep have been deliberately used in grazing trials. Almost certainly a far wider range of pathogens and herbivores than noted at present, incidentally attack I. tinctoria. Further details on promising biological control agents are given in the Prevention and Control section of the datasheet.

Means of Movement and Dispersal

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I. tinctoria was dispersed by man throughout prehistory through Eurasia, and into the Americas with European colonization. Within the USA, spread into the western states was originally through contaminated alfalfa seed from Ireland (Roché, 1992; Callihan, 1990).

Natural Dispersal

Wind dispersal is very limited except in winter when fruits may be carried across snow for considerable distances. Although the plant grows frequently in arid conditions, water transport is important and the flattened shape of I. tinctoria may facilitate this (Farah et al., 1988).

Vector Transmission (Biotic)

The fruit has adaptations – a hook-like pedicel – which can stick to animals, both grazing stock and overwintering game animals such as deer, grouse, partridge and pheasant (Farah et al., 1988). Mud carrying seeds may also stick to animals and also to humans, their equipment and vehicles (Farah et al., 1988).

Accidental Introduction

As stated above, introduction into the western states of the USA has resulted via crop seed contaminant (McConnell et al., 1999) with Roché (1992) and Callihan (1990) also stating that introduction into the West resulted from contaminated alfalfa (lucerne) seed into California and Utah respectively, early in the 20th century. Although these introductions did occur, the plant may have already been deliberately introduced during the late 19th century.

Intentional Introduction

Throughout most of its present range I. tinctoria has probably been introduced deliberately for industrial purposes. There has been renewed interest recently with cultivar movement in Europe. In contrast to some other authors, Finlay (2009) and Sagers (1991), state that it was deliberately introduced by Mormon pioneers into Utah in the nineteenth century, although this is not substantiated by Wheeler (2011).  

Pathway Causes

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CauseNotesLong DistanceLocalReferences
Crop productionHistorically extensive deliberate introductions Yes Yes Hurry, 1930
Escape from confinement or garden escape Yes
Industrial purposesDyeing from pre-history though to industrial scale. Yes Yes

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Clothing, footwear and possessionsFruits, probably uncommon form of dissemination Yes Zouhar, 2009
Host and vector organismsSeeds carried by ants Yes Zouhar, 2009
Land vehiclesEspecially agricultural vehicles Yes Zouhar, 2009
LivestockFruits Yes Yes Farah et al., 1988
Machinery and equipmentSeeds carried on agricultural machinery and vehicles Yes Yes Zouhar, 2009
WaterSuggested fruit adapted for water transport Yes Yes Farah et al., 1988
WindSuggested as important under snow conditions Yes Yes Farah et al., 1988

Impact Summary

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

Economic Impact

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Where of pest status, I. tinctoria causes major economic losses. Infestation in rangelands of the Pacific Northwest, USA, reduced cattle grazing capacity by 38% (Jacobs and Pokorny, 2007).  Evans and Chase (1981) estimated that the plant reduced crop and rangeland production in Utah by US$2 million per year and McConnell et al (1990) noted a doubling of infestation during the 1980s and estimated losses of several million dollars annually in Utah alone. Pokorny and Krueger-Mangold (2007) estimated that I. tinctoria infestations in Montana would cost $1.9 million annually to manage if Montana’s Dyer’s Woad Cooperative Eradication Project, costing US$142,000 over seven years, had not been so effective.

I. tinctoria is of minor pest status in many parts of Europe and further east. It exists as a minor (usually) weed in some areas (Vorobiov, 1960) and may also be considered for enrichment of crop flora as new species for fodder (Varlamova, 1984).

Environmental Impact

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Impact on Habitats

I. tinctoria is highly invasive in a range of habitats and its long term occurrence may lead to harmful effects on subsequent or concurrent plants leading to marked changes in habitat. This may be associated with changes in microbial populations in the soil. I. tinctoria cultivation has a long history of association with declining soil fertility (Iqbal, 2012). In rangeland and in wild habitats, changes in grazing could also lead to changes in succession; grazing animals avoid I. tinctoria, preferentially eating other plants, reducing competition for the weed. The weed threatens protected habitats and, for example, is found at the southern edges of Yellowstone National Park (RE Whitesides, personal communication, 2015).

Impact on Biodiversity

The clearest specific threat from I. tinctoria is to the endemic Siskiyou Mariposa Lily (Calochortus persistens) which may only exist in one site, in Siskiyou County, California, where I. tinctoria affects 90% of the known habitat (Diggles et al., 2003 and references therein). A conservation agreement (Anon, 2012) has been required to protect the species. I. tinctoria is thought to compete with C. persistens for water and light as woad is the much taller plant.  

I. tinctoria is also seen as a significant threat to Phlox hirsuta, an endemic shrub in the Polemoniaceae family, found in Siskiyou County, California (Anon., 2015e; Baun 2015). This has been listed as endangered under the Endangered Species Act since 2000 and was listed as endangered by the State of California in 1986.

Leffler et al. (2014) suggested that I. tinctoria may be more effective at invading grass mixtures and monocultures than forb or shrub communities. It suppresses annual grasses more readily than perennial grasses.

Threatened Species

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Threatened SpeciesConservation StatusWhere ThreatenedMechanismReferencesNotes
Calochortus persistens (Siskiyou mariposa lily)NatureServe NatureServe; USA ESA candidate species USA ESA candidate speciesCalifornia; OregonCompetitionAnon, 2012; Diggles et al., 2003
Phlox hirsuta (Yreka phlox)USA ESA listing as endangered species USA ESA listing as endangered speciesCaliforniaBaun, 2009

Social Impact

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I. tinctoria is of limited negative social impact in most countries where it exists. In the western states of the USA, landowners are obliged to observe regulations of Noxious Weeds legislation and community groups may need to be involved in control programmes. Loss of the natural habitat may have some impact on amenity activities and the visual impact of large areas of plants bearing black fruits, blackening local environments, is likely to be upsetting to residents (Young and Evans, 1971). 

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Has a broad native range
  • Highly adaptable to different environments
  • Is a habitat generalist
  • Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
  • Pioneering in disturbed areas
  • Has propagules that can remain viable for more than one year
  • Reproduces asexually
  • Has high genetic variability
Impact outcomes
  • Damaged ecosystem services
  • Ecosystem change/ habitat alteration
  • Increases vulnerability to invasions
  • Modification of nutrient regime
  • Modification of successional patterns
  • Monoculture formation
  • Negatively impacts agriculture
  • Negatively impacts animal health
  • Negatively impacts livelihoods
  • Reduced amenity values
  • Reduced native biodiversity
  • Threat to/ loss of endangered species
  • Threat to/ loss of native species
Impact mechanisms
  • Allelopathic
  • Competition - monopolizing resources
  • Competition - shading
  • Poisoning
Likelihood of entry/control
  • Highly likely to be transported internationally deliberately

Uses

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In the many parts of the world where I. tinctoria is not of major pest status it has many uses such as the production of dyes. Dyer’s woad is a possible alternative crop to match the crop diversification policy of the EU (Spataro and Negri, 2008b). There is interest in the plant as a source of anti-inflammatory (Hamburger, 2002) and anti-tumoral compounds. The plant is also regarded of some value in animal feeds, fodder crops, ground cover, organic fertilizers, plant breeding (Xiang et al., 2013) and erosion control.

Economic Value

Once of immense value to European societies, providing great levels of employment and being the first crop that received systematic land cultivation, leading the way to other agricultural improvements (Hurry, 1930). Woad and woad merchants are found represented in sculpture and in glass in religious buildings in Europe, indicating the value of the crop, which has left some European town and cities with notable buildings from the time of cultivation. The modern value is very limited, but a quick internet search will show interest in traditional dyeing, the sale of woad and woad-dyed cloths, woad based cosmetics, woad themed tourism and the use of I. tinctoria preparations in wood preservation. Potential value may result from pharmacological research (Hamburger, 2002) or exploiting insecticidal or fungicidal properties (Seifert and Unger, 1994).

Social Benefit

I. tinctoria is an important medicinal plant and is part of the cultural appreciation of herbal medicines common, or previously common, in many if not all societies where it is present. Herbal medicines may play a useful role to link with ancestral cultures amongst immigrant groups (Jiang and Quave, 2013). Chinese immigrants in Atlanta, Georgia, use I. tinctoria root, whereas Taiwanese immigrants do not (Jiang and Quave, 2013). Other positive social aspects include renewed interests in botanical dyes, resurrecting traditional customs (Guarrera, 2006) and woad’s use for medicinal purposes.

Environmental Services

I. tinctoria is valuable for bees. It may also give protection against soil erosion on exceptionally steep hillsides (Varga and Evans, 1978). On the other hand, as erosion is associated with a loss of species diversity (Berendse et al., 2015) and I. tinctoria probably reduces diversity, this is a dubious environmental service claim.

Uses List

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Animal feed, fodder, forage

  • Fodder/animal feed
  • Forage

Environmental

  • Erosion control or dune stabilization

General

  • Ritual uses
  • Sociocultural value

Genetic importance

  • Gene source

Human food and beverage

  • Vegetable

Materials

  • Dye/tanning
  • Dyestuffs
  • Green manure
  • Pesticide

Medicinal, pharmaceutical

  • Source of medicine/pharmaceutical
  • Traditional/folklore

Detection and Inspection

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Largely achieved after growth of the plant, rather than through inspection. Detection in seed, forage or similar is achievable.

Similarities to Other Species/Conditions

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Ball and Akeroyd (2010) state that I. tinctoria is very variable, especially in the size, shape and pubescence of the silicula. The genus is difficult and requires revision and a number of the present species are probably subspecies of I. tinctoria (Ball and Akeroyd, 2010); Al-Shehbaz et al (2006) substantiate this view. Callihan et al. (1984) list species that may be confused with I. tinctoria including: Lomatium dissectum (biscuit-root), Descurainia sophia (flixweed), Brassica spp., Sisymbrium altissimum (Jim Hill mustard) and other mustards, Crepis acuminatus (tapertip hawksbeard) Senecio spp, Balsamorhiza sagittata (arrowleaf balsamroot), Wyethia amplexicaulis (mule’s ears), Camelina microcarpa (falsflax), Melitotus officinalis (yellow sweet-clover), Euphorbia esula (leafy spurge), Arnica spp and Eriogonum spp (wild buckwheat).

Prevention and Control

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SPS Measures

The affected western states of the USA maintain SPS measures with, for example, movement prohibitions, awareness raising and recommendations to prevent spread. The Utah Noxious Weed Act (Utah Department of Administrative Services, 2015) is an example of the procedures in place in the relevant states. The components of early warning, rapid response and public awareness are given by Pokorny and Kreuger-Mangold (2007), who described a control programme in Montana that effectively eliminated the plant as a pest species. Eradication is not feasible as the plant is too well established and widespread, but the plant can be managed.

Containment/Zoning

This is not applicable other than to improve the efficacy of other control measures. Mechanical pulling may be considered containment in the initial years, whereas if done repeatedly it becomes a successful control technique.

Control

A great many control measures are very effective, but logistical issues impact which are used, depending upon the nature of infestations, such as their occurrence in crops, on rangeland or in environmentally sensitive or difficult to reach areas. Consequently, an effective classical biological control would be valuable. As I. tinctoria is often found on or near marginal land, control measures may not be economic or there may be environmental considerations that will favour different control measures. Pokorny and Krueger-Mangold (2007) described a successful control project comprising of four key components –1) early detection, 2) treatment technologies including hand-pulling, digging and spot-spraying with metsulfuron plus nonionic surfactants and the removal of flowering or seeding plants in double-lined plastic bags, 3) repeated site visits with monitoring, and 4) education.

Sanitary Measures

I. tinctoria is a specified noxious weed in 11 western states of the USA and these states have sanitary methods to prevent dissemination within and between states; the requirements from Utah (Utah Department of Administrative Services, 2015) are an example. This requires persons, companies or corporations to clean transport and farm machinery free from all noxious weeds. Screenings for livestock feed, livestock feed itself, hay straw or similar, soil, sod and nursery stock and livestock are also listed with procedures to be followed.

Physical/Mechanical and Cultural Control

DiTomaso et al. (2013) gave recent advice on mechanical control. Hand pulling can be very effective if the crown is removed, which is easier after bolting but needs to be done before seed set. Repeat visits are necessary each season and for several years. Mowing, while it can reduce plant reserves and seed production if repeated, is not effective as there is resprouting from the crown. Clipping can be more effective at preventing resprouting, if close enough, but is ineffective if done too early in the season (West and Farah, 1989). Soil cultivation for field crops can be effective but, in alfalfa may require both spring and late fall/autumn cultivation (Jacobs and Pokorny, 2007). Sheep grazing is not very effective as the plant is not preferred and the required stocking densities would damage the rangeland (West and Farah, 1989). However, goat grazing has been recommended for noxious weeds (Lamming, 2001) and goats have been used effectively on I. tinctoria (L Malmberg, personal communication, 2015).

Movement Control

Within seriously affected states, movement of I. tinctoria is prevented. For example, in Utah it is unlawful to disseminate I. tinctoria and the articles of such dissemination are listed (Utah Department of Administrative Services, 2015,) and other states have similar legislation.

Biological Control

A plant pathogen, described as Puccinia thlaspeos (preferred name now Micropuccinia thlaspeos) was identified from I. tinctoria in Idaho in 1978. Phylogenetic studies of the rust showed that it is most probably native to North America (Kropp et al., 1997) and so was considered as a potential biological control agent. The systemic colonization may allow survival of the rust in root tissues over winter and after plant exposure to range fires (Kropp et al., 1996). Infection greatly reduces vegetative growth, reducing its competitive ability and also seed production; in most cases no seed is produced on infected plants, although those that are produced germinate as well as those from non-infected plants (Kropp et al., 1996).

A product, “Woad Warrior” based on I. tinctoria leaves infected with the rust was submitted for registration to the US Environmental Protection Agency (EPA) (Anon, 2002) who recommended registration. The EPA reviewed the evidence finding that there were no unreasonable adverse effect to humans or the environment and also noted that occasionally wild deer fed selectively on infected I. tinctoria plants, as did mice (Kough, 2002). Woad Warrier was available, in a limited fashion, for a few years but is no longer on the market (RE Whitesides, personal communication, 2015). The product was never commercially available due to a lack of a commercial backer, but after registration the fungus was spread by researchers (Bellgard, 2008). Woad Warrior was specified for use in Utah’s waterfowl management areas in 2009 (Berger, 2009), but this appears to be outdated advice. Although the rust has significant adverse effects on the plant and seed production it does not seem to be able to stop the spread of I. tinctoria (RE Whitesides, personal communication, 2015), although presumably it does slow the spread. Seedling mortality is normally extremely high (Farah et al., 1988) so reduced seed production may result in reduced seedling competition with a less than commensurate reduction in established plants.

A project to look for classical biological control agents in 2004 identified four species for detailed research, and further candidates were identified from extensive literature and field surveys (Cortat et al., 2008, Hinz et al., 2007). Of the original four species, the weevil Aulacobaris fallax and the flea beetle Psylliodes isatidis were found to be insufficiently specific (Cortat et al., 2008; Hinz et al., 2015) for consideration. As of 2018, the seed-feeding weevil Ceutorhynchus peyerimhoffi and the root crown-mining weevil Ceutorhynchus rusticus have been prioritized as promising agents.

The requirements for a successful biological control agent for I. tinctoria must take into account its biology. Farah et al. (1988) considered that an agent effective at the young rosette stage would be most effective as this stage is very vulnerable, whereas mature rosettes generally survive and reproduce. This could well be an argument to actively target the mature rosettes. The majority of seeds fail to establish, and markedly different seed production often results in similar levels of establishment, one reason why Puccinia thlaspeos has not been as successful as anticipated. The level of successful attack of the stems by P. thlaspeos appears low (H Hinz, CABI Switzerland, personal communication, 2015) but plants can be asymptomatic for many months, so infestation levels may be underestimated (Kropp et al., 1996).

Successful macrobial biological control agents are likely to share attributes promoting rapid population build up and phenotypic plasticity that are demonstrated by successful herbivorous invasive species (Nahrung and Swain, 2015). This may be of critical importance with I. tinctoria which also demonstrates marked plasticity and the ability to succeed under stress tolerant conditions.

Chemical Control

Metsulfuron or chlorsulfuron combined with 2,4-D and a nonionic surfactant are recommended as is the combination of imazic and methylated seed oil on rosettes or bolting plants (Jacobs and Pokorny, 2007). Berger (2009) described chemical treatments for Utah’s waterfowl management areas and lists dicamba plus 2,4-D, 2,4-D ester and amine and glyphosate. Other chemicals that can be appropriate for use in natural areas are given by Tu et al. (2001). The scale of I. tinctoria infestations in rangeland and difficulties of application in some areas combined with the cost:benefit aspect means that herbicide use is often impractical and/or uneconomic.

IPM

Control techniques can be used in various combinations for sensitive areas (Tu et al., 2001); Utah’s Waterfowl Management Areas’ plans for eradication involves herbicide treatments or biological control using Puccinia thlaspeos or a combined herbicide, mowing and sheep grazing strategy (Berger, 2009). Metsulfuron and chlorsulfuron did not adversely affect P. thlaspeos teliospore germination, whereas 2,4-D and some surfactants did (Jacobs and Pokorny, 2007).

Pokorny and Krueger-Mangold (2007) described a successful control project comprising of four key components –1) early detection, 2) treatment technologies including hand-pulling, digging and spot-spraying with metsulfuron plus nonionic surfactants and the removal of flowering or seeding plants in double-lined plastic bags, 3) repeated site visits with monitoring and 4) education.

Communication is an important component of IPM. A general strategic plan for managing noxious and invasive weeds in Utah is one example of IPM, covering education and research, mapping and monitoring, prevention, early detection and rapid response, control through integrated weed management and regulation and enforcement (Whitesides, 2004). A Noxious Weed Field Guide for Utah (Belliston et al., 2009) deals exclusively with Utah’s state noxious weeds, listing 27 weeds as noxious, and notes that individual counties may add county-listed weeds to the list. I. tinctoria is a Class B weed: “Control – Declared noxious weeds not native to the state that pose a threat to the state and should be considered a high priority for control”.  The Utah Noxious Weed Act (Utah Department of Administrative Services, 2015) designates the state noxious weeds (as above), groups ten articles as capable of disseminating noxious weeds (e.g. machinery, seed, manure and livestock) and gives prescribed treatments to prevent dissemination by these articles. In addition each county is required to submit an “Annual Progress Report of County Noxious Weed Control Program” to the Commissioner of Agriculture and Food and general and individual notices “pertaining to the control and prevention of noxious weeds” are specified. These are a) a General Notice to be posted in at least three public places within a county, stating the responsibility of every property owner to control and prevent the spread of noxious weeds on their land, b) an Individual Notice requiring a landowner to take (specified) actions to control weeds on the land and c) a Notification of Noxious Weed Lien Assessment whereby the Board of County Commissioners may carry out the control on land and require payment by the owner of that land.

Monitoring and Surveillance

Early detection is important; using Landsat-5 Thematic Mapper spectral data and field survey information Dewey et al (1991) determined specific land cover types that were associated with I. tinctoria infestation sites (highly suited types). Estimating the non-infested areas associated with the highly suited cover types gave an estimate of the potential for plant invasion. In a practical setting, the use of a Global Positioning System, with data subsequently recorded in the GPS data dictionary, assisted the implementation of a successful control programme that was largely achieved by manual removal and spot applications of herbicides (Pokorny and Kreuger-Mangold, 2007). Despite improved technologies for remote sensing (Huang and Asner, 2009; Naupari et al., 2013) this has not been reported as a major tool for I. tinctoria management.

Ecosystem Restoration

Competition from desired plants reduces I. tinctoria spread; reseeding after appropriate seed bed preparation is effective. On rough ground, fire (see Zouhar, 2009 for guidelines) or grazing may be used, followed by establishing competitive plants and maintaining grazing (Jacobs and Pokorny, 2007).

Gaps in Knowledge/Research Needs

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Determining which life stages to target for biological control is important. The merits of agents reducing seed production may be debated if this just reduces seedling competition (Farah et al., 1988), but seed persistence in the soil is limited, so constant seed attrition should have a positive control effect over time. Seed reduction may need to be accompanied by rosette control. Modelling of a related plant, garlic mustard (Allaria petiolata) suggests that reduced rosette survival and reduced seed output would control populations (Davis et al., 2006). 

References

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African Plant Database, 2015. African Plants Database (version 3.4.0). Geneva and Pretoria, Switzerland and South Africa: Conservatoire et Jardin botaniques de la Ville de Genève and South African National Biodiversity Institute. http://www.ville-ge.ch/musinfo/bd/cjb/africa/index.php?langue=an

Al-Shehbaz IA, Beilstein MA, Kellogg EA, 2006. Systematics and phylogeny of the Brassicaceae (Cruciferae): an overview. Plant Systematics and Evolution [Evolution and phylogeny of the Brassicaceae. Papers presented at the symposium "Systematics and Evolution in Brassicaceae", XVII International Botanical Congress, Vienna, Austria, 2005.], 259(2/4):89-120. http://www.springerlink.com/link.asp?id=104878

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Links to Websites

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WebsiteURLComment
GISD/IASPMR: Invasive Alien Species Pathway Management Resource and DAISIE European Invasive Alien Species Gatewayhttps://doi.org/10.5061/dryad.m93f6Data source for updated system data added to species habitat list.
Global register of Introduced and Invasive species (GRIIS)http://griis.org/Data source for updated system data added to species habitat list.

Contributors

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22/01/18 Updated by:

Philip Weyl, CABI, Delémont, Switzerland 

17/06/2015 Original text by:

Dr. Dave Moore, CABI, Egham, UK

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