Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide

Datasheet

Prunella vulgaris
(self-heal)

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Datasheet

Prunella vulgaris (self-heal)

Summary

  • Last modified
  • 16 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Preferred Scientific Name
  • Prunella vulgaris
  • Preferred Common Name
  • self-heal
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Spermatophyta
  •       Subphylum: Angiospermae
  •         Class: Dicotyledonae
  • Summary of Invasiveness
  • P. vulgaris, commonly known as self-heal, is a herbaceous perennial considered to be native to Europe, Eurasia and America but now with a large geographical range across all continents. It was introduced to man...

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Pictures

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PictureTitleCaptionCopyright
Prunella vulgaris (self-heal); flower spike. Keila, Northwestern Estonia. July, 2013.
TitleFlower spike
CaptionPrunella vulgaris (self-heal); flower spike. Keila, Northwestern Estonia. July, 2013.
Copyright©Ivar Leidus-2013/via wikipedia - CC BY-SA 3.0
Prunella vulgaris (self-heal); flower spike. Keila, Northwestern Estonia. July, 2013.
Flower spikePrunella vulgaris (self-heal); flower spike. Keila, Northwestern Estonia. July, 2013.©Ivar Leidus-2013/via wikipedia - CC BY-SA 3.0
Prunella vulgaris (self-heal); flower spikes, normal and white morph. Puu Nianiau, Maui, Hawaii, USA. December, 2004.
TitleFlower spikes
CaptionPrunella vulgaris (self-heal); flower spikes, normal and white morph. Puu Nianiau, Maui, Hawaii, USA. December, 2004.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Prunella vulgaris (self-heal); flower spikes, normal and white morph. Puu Nianiau, Maui, Hawaii, USA. December, 2004.
Flower spikesPrunella vulgaris (self-heal); flower spikes, normal and white morph. Puu Nianiau, Maui, Hawaii, USA. December, 2004.©Forest Starr & Kim Starr - CC BY 4.0
Prunella vulgaris (self-heal); flower spike, white morph. Puu Nianiau, Maui, Hawaii, USA. December, 2004.
TitleFlower spike
CaptionPrunella vulgaris (self-heal); flower spike, white morph. Puu Nianiau, Maui, Hawaii, USA. December, 2004.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Prunella vulgaris (self-heal); flower spike, white morph. Puu Nianiau, Maui, Hawaii, USA. December, 2004.
Flower spikePrunella vulgaris (self-heal); flower spike, white morph. Puu Nianiau, Maui, Hawaii, USA. December, 2004.©Forest Starr & Kim Starr - CC BY 4.0
Prunella vulgaris (self-heal); seed-bearing spikes. Polipoli, Maui, Hawaii, USA. May, 2009.
TitleSeed-bearing spikes
CaptionPrunella vulgaris (self-heal); seed-bearing spikes. Polipoli, Maui, Hawaii, USA. May, 2009.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Prunella vulgaris (self-heal); seed-bearing spikes. Polipoli, Maui, Hawaii, USA. May, 2009.
Seed-bearing spikesPrunella vulgaris (self-heal); seed-bearing spikes. Polipoli, Maui, Hawaii, USA. May, 2009.©Forest Starr & Kim Starr - CC BY 4.0
Prunella vulgaris (self-heal); flowering habit. Polipoli, Maui, Hawaii, USA. May, 2009.
TitleHabit
CaptionPrunella vulgaris (self-heal); flowering habit. Polipoli, Maui, Hawaii, USA. May, 2009.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Prunella vulgaris (self-heal); flowering habit. Polipoli, Maui, Hawaii, USA. May, 2009.
HabitPrunella vulgaris (self-heal); flowering habit. Polipoli, Maui, Hawaii, USA. May, 2009.©Forest Starr & Kim Starr - CC BY 4.0
Prunella vulgaris (self-heal); flowering habit. Redwood trail, Polipoli, Maui, Hawaii, USA. December, 2004.
TitleHabit
CaptionPrunella vulgaris (self-heal); flowering habit. Redwood trail, Polipoli, Maui, Hawaii, USA. December, 2004.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Prunella vulgaris (self-heal); flowering habit. Redwood trail, Polipoli, Maui, Hawaii, USA. December, 2004.
HabitPrunella vulgaris (self-heal); flowering habit. Redwood trail, Polipoli, Maui, Hawaii, USA. December, 2004.©Forest Starr & Kim Starr - CC BY 4.0
Prunella vulgaris (self-heal); flowering habit. Polipoli, Maui, Hawaii, USA. May, 2009.
TitleHabit
CaptionPrunella vulgaris (self-heal); flowering habit. Polipoli, Maui, Hawaii, USA. May, 2009.
Copyright©Forest Starr & Kim Starr - CC BY 4.0
Prunella vulgaris (self-heal); flowering habit. Polipoli, Maui, Hawaii, USA. May, 2009.
HabitPrunella vulgaris (self-heal); flowering habit. Polipoli, Maui, Hawaii, USA. May, 2009.©Forest Starr & Kim Starr - CC BY 4.0

Identity

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Preferred Scientific Name

  • Prunella vulgaris

Preferred Common Name

  • self-heal

Other Scientific Names

  • Prunella asiatica Nakai
  • Prunella japonica Makino
  • Prunella vulgaris f. albiflora (Koidz.) Kudô, nom. illeg.
  • Prunella vulgaris f. candida Yonek., nom. illeg.
  • Prunella vulgaris f. leucocephala K.Ohashi, H.Ohashi & Yonek.
  • Prunella vulgaris f. taiwanalpina T.Yamaz.
  • Prunella vulgaris subsp. parviflora Ehrh.
  • Prunella vulgaris var. albiflora Koidz., nom. illeg.
  • Prunella vulgaris var. aleutica Fernald
  • Prunella vulgaris var. nanhutashanense S.S.Ying
  • Prunella vulgaris var. taiwaniana T.Yamaz., nom. illeg.

International Common Names

  • English: carpenter weed; common heal all; common selfheal; heal all; healall; heal-all; self heal

Local Common Names

  • Australia: brunella
  • Azerbaijan: adi bogazotu
  • Brazil/Acre: busuioc de câmp
  • China: xia ku cao
  • Czech Republic: cernohlávek obecný
  • France: brunelle commune
  • Germany: gemeine braunelle; kleine braunelle
  • Iceland: blákolla
  • Italy: prunella commune
  • Netherlands: bijenkorfje; gewone brunel
  • Norway: blåkoll
  • Poland: Glowienka pospolita
  • Romania: busuioc de cîmp
  • Russian Federation: cernogolovka Obyknovennaja
  • Spain: consuelda menor
  • Sweden: brunört
  • Turkey: yara otu

Summary of Invasiveness

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P. vulgaris, commonly known as self-heal, is a herbaceous perennial considered to be native to Europe, Eurasia and America but now with a large geographical range across all continents. It was introduced to many countries (and islands) in the 1800s and since then has been recorded as invasive on many Pacific islands, as well as New Zealand (PIER, 2016). Along with other nonnative plants, it is thought to be threatening habitats and native plants on the Hawaiian Islands. In its native range it can be invasive in lawns and is often considered a weed, although in some places it has been deliberately grown as an ornamental.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Plantae
  •         Phylum: Spermatophyta
  •             Subphylum: Angiospermae
  •                 Class: Dicotyledonae
  •                     Order: Lamiales
  •                         Family: Lamiaceae
  •                             Genus: Prunella
  •                                 Species: Prunella vulgaris

Notes on Taxonomy and Nomenclature

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There appear to be at least two subspecies of Prunella vulgaris that are sometimes simply referred to as P. vulgaris (self-heal). These are P. vulgaris subsp. vulgaris, sometimes referred to as the Eurasian or European species, and P. vulgaris subsp. lanceolata, often considered to be a native North American species. See distribution section for more information and references.

Although recorded as an accepted name by the Plant List (2013), there is some confusion over the subspecies Prunella vulgaris subsp. aleutica (Aleutian selfheal). Some sources suggest that this subspecies is an incorrectly named variety, Prunella vulgaris var. aleutica, which is currently regarded as a synonym for Prunella vulgaris subsp. asiatica (WCSP, 2016).

P. vulgaris hybridises with P. grandiflora and P. laciniata in the UK and Northern Europe (Fitter et al., 1980). It also hybridizes with P. laciniata in New Zealand, with the hybrids (P. × hybrida Knaf) having pinnately lobed leaves (Webb et al., 1988).

Description

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The following description is adapted from Fitter et al. (1980), Rose (2006), Bojnanský and Fargašová (2007) and ZipcodeZoo (2014) and the Encyclopedia of Life (2016).

P. vulgaris is a low, creeping, non-aromatic perennial that is 5–30 cm tall with square stems. Leaves are in opposite pairs along the stem, and are approx. 2.5 cm long and 1.5 cm wide. Leaves are oval to lanceolate, untoothed or serrated. Petioles are generally short but can be up to 5 cm in lower leaves. The inflorescence is a dense, oblong or square, whirled cluster with a pair of stalkless leaves below. There are 3 flowers per bract, and the bracts and calyx are purplish. Flowers are violet (rarely white or pink) and are two-lipped and tubular. The top lip is a concave purple hood. The bottom lip may be white, and has three lobes, with a larger fringed middle lobe. The corolla is 10-14 cm long. Flowering time varies depending on the climate, but is throughout the summer (June-August in the USA; June-November in the UK). Seeds are smooth, shiny, brown nutlets that are obovate, oblong, with a convex dorsal side and a roof-like ventral side. Each flower can produce 4 seeds. P. vulgaris reproduces by seed, and vegetatively via creeping stems that root at the nodes.

Plant Type

Top of page Herbaceous
Perennial
Seed propagated
Vegetatively propagated

Distribution

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P. vulgaris is found in Europe, North Africa, Asia temperate and tropical, Australasia, North, Central and South America and in the sub-Antarctic islands (WCSP, 2016).

Some sources list P. vulgaris as a native of North America (e.g., Enclyclopaedia Britannica, 2016). Other sources suggest that the self-heal population in the USA consists of the native subspecies, P. vulgaris subsp. lanceolata and the introduced P. vulgaris subsp. vulgaris (Wilson, 2006). Some sources suggest that P. vulgaris is most likely originally from Europe (Godoy et al., 2011) or Eurasia (Bojnanský and Fargašová, 2007).

It has also been suggested that the variety of P. vulgaris that is observed in lawns in the USA is the Eurasian variety that roots at the nodes of leaves (Encyclopedia of Life, 2016).

P. vulgaris subsp. aleutica (Aleutian selfheal) is found in Alaska (USDA-NRCS, 2016) although other sources suggest that the variety P. vulgaris var. aleutica is a synonym for P. vulgaris subsp. asiatica (WCSP, 2016).

Although only fairly recently introduced to New Zealand, it is now a common species. P. vulgaris has been found in Australia for at least 200 years (before 1801) (Low, 1999) and some sources consider it to be native to Australia (Bean, 2007). Bean (2007) recommends that given it is the only species of Prunella in Australia, and given that Australia is so far from its considered natural range, it is likely to be non-native there (Low, 1999).

P. vulgaris hybridises with P. grandiflora and P. laciniata in the UK and Northern Europe (Fitter et al., 1980). It also hybridizes with P. laciniata in New Zealand, with the hybrids (P. × hybrida Knaf) having pinnately lobed leaves (Webb et al., 1988). 

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

AfghanistanPresentNativeWCSP, 2016
ArmeniaPresentNativeWCSP, 2016
AzerbaijanPresentNativeWCSP, 2016; WCSP, 2016; WCSP, 2016
BhutanPresentNativeWCSP, 2016
ChinaPresentNativeWCSP, 2016
-AnhuiPresentNativeWCSP, 2016
-BeijingPresentNativeWCSP, 2016
-ChongqingPresentNativeWCSP, 2016
-FujianPresentNativeWCSP, 2016
-GansuPresentNativeWCSP, 2016
-GuangdongPresentNativeWCSP, 2016
-GuangxiPresentNativeWCSP, 2016
-GuizhouPresentNativeWCSP, 2016
-HainanPresentNativeWCSP, 2016
-HebeiPresentNativeWCSP, 2016
-HeilongjiangPresentNativeWCSP, 2016
-HenanPresentNativeWCSP, 2016
-Hong KongPresentNativeWCSP, 2016
-HubeiPresentNativeWCSP, 2016
-HunanPresentNativeWCSP, 2016
-JiangsuPresentNativeWCSP, 2016
-JiangxiPresentNativeWCSP, 2016
-JilinPresentNativeWCSP, 2016
-LiaoningPresentNativeWCSP, 2016
-MacauPresentNativeWCSP, 2016
-Nei MengguPresentNativeWCSP, 2016
-NingxiaPresentNativeWCSP, 2016
-QinghaiPresentNativeWCSP, 2016
-ShaanxiPresentNativeWCSP, 2016
-ShandongPresentNativeWCSP, 2016
-ShanghaiPresentNativeWCSP, 2016
-ShanxiPresentNativeWCSP, 2016
-SichuanPresentNativeWCSP, 2016
-TianjinPresentNativeWCSP, 2016
-TibetPresentNativeWCSP, 2016
-XinjiangPresentNativeWCSP, 2016
-YunnanPresentNativeWCSP, 2016
-ZhejiangPresentNativeWCSP, 2016
IndiaPresentNativeWCSP, 2016
-Andaman and Nicobar IslandsPresentNativeWCSP, 2016
-Andhra PradeshPresentNativeWCSP, 2016
-Arunachal PradeshPresentNativeWCSP, 2016
-AssamPresentNativeWCSP, 2016
-BiharPresentNativeWCSP, 2016
-ChandigarhPresentNativeWCSP, 2016
-ChhattisgarhPresentNativeWCSP, 2016
-Dadra and Nagar HaveliPresentNativeWCSP, 2016
-DamanPresentNativeWCSP, 2016
-DelhiPresentNativeWCSP, 2016
-DiuPresentNativeWCSP, 2016
-GoaPresentNativeWCSP, 2016
-GujaratPresentNativeWCSP, 2016
-HaryanaPresentNativeWCSP, 2016
-Himachal PradeshPresentNativeWCSP, 2016
-Indian PunjabPresentNativeWCSP, 2016
-Jammu and KashmirPresentNativeKour et al., 2014; WCSP, 2016
-JharkhandPresentNativeWCSP, 2016
-KeralaPresentNativeWCSP, 2016
-Madhya PradeshPresentNativeWCSP, 2016
-MaharashtraPresentNativeWCSP, 2016
-ManipurPresentNativeWCSP, 2016
-MeghalayaPresentNativeWCSP, 2016
-MizoramPresentNativeWCSP, 2016
-NagalandPresentNativeWCSP, 2016
-PuducherryPresentNativeWCSP, 2016
-RajasthanPresentNativeWCSP, 2016
-SikkimPresentNativeWCSP, 2016
-Tamil NaduPresentNativeWCSP, 2016
-TripuraPresentNativeWCSP, 2016
-Uttar PradeshPresentNativeWCSP, 2016
-UttarakhandPresentNativeWCSP, 2016
-West BengalPresentNativeWCSP, 2016
IranPresentNativeWCSP, 2016
IraqPresentNativeWCSP, 2016
IsraelPresentNativeWCSP, 2016
JapanPresentNativeWCSP, 2016
KazakhstanPresentNativeWCSP, 2016; WCSP, 2016
Korea, DPRPresentNativeWCSP, 2016
Korea, Republic ofPresentNativeWCSP, 2016
KyrgyzstanPresentNativeWCSP, 2016
LebanonPresentNativeWCSP, 2016
MongoliaPresentNativeWCSP, 2016
NepalPresentNativeWCSP, 2016
PakistanPresentNativeWCSP, 2016
SyriaPresentNativeWCSP, 2016
TaiwanPresentNativeWCSP, 2016
TajikistanPresentNativeWCSP, 2016
TurkeyPresentNativeWCSP, 2016
TurkmenistanPresentNativeWCSP, 2016
UzbekistanPresentNativeWCSP, 2016
VietnamPresentNativeWCSP, 2016

Africa

AlgeriaPresentNativeWCSP, 2016
Cape VerdePresentNativeWCSP, 2016
EgyptPresentNativeWCSP, 2016
MoroccoPresentNativeWCSP, 2016
Spain
-Canary IslandsPresentNativeWCSP, 2016
TunisiaPresentNativeWCSP, 2016

North America

BermudaPresentIntroducedWCSP, 2016
CanadaPresentNativeWCSP, 2016
-British ColumbiaPresentNativeWCSP, 2016
-ManitobaPresentNativeWCSP, 2016
-New BrunswickPresentNativeWCSP, 2016
-Newfoundland and LabradorPresentNativeWCSP, 2016
-Nova ScotiaPresentNativeWCSP, 2016
-OntarioPresentNativeWCSP, 2016
-Prince Edward IslandPresentNativeWCSP, 2016
-QuebecPresentNativeWCSP, 2016
-SaskatchewanPresentNativeWCSP, 2016
MexicoPresentNativeWCSP, 2016
USAPresentPresent based on regional distribution.
-AlabamaPresentNativeWCSP, 2016
-AlaskaPresentNativeWCSP, 2016
-ArizonaPresentNativeWCSP, 2016
-ArkansasPresentNativeWCSP, 2016
-CaliforniaPresentNativeWCSP, 2016
-ColoradoPresentNativeWCSP, 2016
-ConnecticutPresentNativeWCSP, 2016
-DelawarePresentNativeWCSP, 2016
-District of ColumbiaPresentNativeWCSP, 2016
-FloridaPresentNativeWCSP, 2016
-GeorgiaPresentNativeWCSP, 2016
-HawaiiPresentIntroducedFlora of the Hawaiian Islands, 2016; USDA-NRCS, 2016
-IdahoPresentNativeUSDA-NRCS, 2016
-IllinoisPresentNativeWCSP, 2016
-IndianaPresentNativeWCSP, 2016
-IowaPresentNativeWCSP, 2016
-KansasPresentNativeWCSP, 2016
-KentuckyPresentNativeWCSP, 2016
-LouisianaPresentNativeWCSP, 2016
-MainePresentNativeWCSP, 2016
-MarylandPresentNativeWCSP, 2016
-MassachusettsPresentNativeWCSP, 2016
-MichiganPresentNativeWCSP, 2016
-MinnesotaPresentNativeWCSP, 2016
-MississippiPresentNativeWCSP, 2016
-MissouriPresentNativeWCSP, 2016
-MontanaPresentNativeWCSP, 2016
-NebraskaPresentNativeWCSP, 2016
-NevadaPresentNativeWCSP, 2016
-New HampshirePresentNativeWCSP, 2016
-New JerseyPresentNativeWCSP, 2016
-New MexicoPresentNativeWCSP, 2016
-New YorkPresentNativeWCSP, 2016
-North CarolinaPresentNativeWCSP, 2016
-North DakotaPresentNativeWCSP, 2016
-OhioPresentNativeWCSP, 2016
-OklahomaPresentNativeWCSP, 2016
-OregonPresentNativeClark and Wilson, 2003; WCSP, 2016
-PennsylvaniaPresentNativeWCSP, 2016
-Rhode IslandPresentNativeWCSP, 2016
-South CarolinaPresentNativeWCSP, 2016
-South DakotaPresentNativeWCSP, 2016
-TennesseePresentNativeWCSP, 2016
-TexasPresentNativeWCSP, 2016
-UtahPresentNativeWCSP, 2016
-VermontPresentNativeWCSP, 2016
-VirginiaPresentNativeWCSP, 2016
-WashingtonPresentNativeWCSP, 2016
-West VirginiaPresentNativeWCSP, 2016
-WisconsinPresentNativeWCSP, 2016
-WyomingPresentNativeWCSP, 2016

Central America and Caribbean

Costa RicaPresentNativeWCSP, 2016
Dominican RepublicPresentIntroducedWCSP, 2016
GuatemalaPresentNativeWCSP, 2016
HaitiPresentIntroducedWCSP, 2016
JamaicaPresentIntroducedIremonger, 2002; WCSP, 2016

South America

ArgentinaPresentIntroducedWCSP, 2016
BrazilPresentPresent based on regional distribution.
-Espirito SantoPresentIntroducedWCSP, 2016
-Minas GeraisPresentIntroducedWCSP, 2016
-Rio de JaneiroPresentIntroducedWCSP, 2016
-Rio Grande do SulPresentIntroducedWCSP, 2016
-Santa CatarinaPresentIntroducedWCSP, 2016
-Sao PauloPresentIntroducedWCSP, 2016
ChileWidespreadIntroduced1870 Invasive Godoy et al., 2011; WCSP, 2016Introduced from Europe
Falkland IslandsPresentIntroducedWCSP, 2016
PeruPresentIntroducedWCSP, 2016
VenezuelaPresentIntroducedWCSP, 2016

Europe

AlbaniaPresentNativeWCSP, 2016
AustriaPresentNativeWCSP, 2016
BelarusPresentNativeWCSP, 2016
BelgiumPresentNativeWCSP, 2016
BulgariaPresentNativeWCSP, 2016
CyprusPresentNativeWCSP, 2016
Czech RepublicPresentNativeWCSP, 2016
DenmarkPresentNativeWCSP, 2016
EstoniaPresentNativeWCSP, 2016
Faroe IslandsPresentNativeWCSP, 2016
FinlandPresentNativeWCSP, 2016
FranceWidespreadNativePolunin and Smythies, 1989; WCSP, 2016SW
-CorsicaPresentNativeWCSP, 2016
GermanyPresentNativeWCSP, 2016
GreecePresentNativeWCSP, 2016
HungaryPresentNativeWCSP, 2016
IrelandPresentNativeWCSP, 2016
ItalyPresentNativeWCSP, 2016
LatviaPresentNativeWCSP, 2016
LithuaniaPresentNativeWCSP, 2016
MoldovaPresentDatamining 2011 - Invasive Species Databases
NetherlandsPresentNativeWCSP, 2016
NorwayPresentNativeWCSP, 2016
PolandPresentNativeWCSP, 2016
PortugalPresentNativePolunin and Smythies, 1989; WCSP, 2016
-AzoresPresentNativeWCSP, 2016
-MadeiraPresentNativeWCSP, 2016
RomaniaPresentNativeWCSP, 2016
Russian FederationPresentNativeWCSP, 2016
SpainPresentNativePolunin and Smythies, 1989; WCSP, 2016
-Balearic IslandsPresentNativeWCSP, 2016
SwedenPresentNativeWCSP, 2016
SwitzerlandPresentNativeWCSP, 2016
UKWidespreadNative Invasive Fitter et al., 1980; Rose, 2006; WCSP, 2016Widespread, visible on lawns that have not been treated with weedkiller

Oceania

AustraliaPresentIntroducedLow, 1999; ZipcodeZoo, 2014A weed throughout Auatralia, probably originating in Europe
New ZealandPresentIntroducedWebb et al., 1988; Beckmann et al., 2013; WCSP, 2016North Isaland, South Island and Chatham Island
Norfolk IslandPresentIntroducedWCSP, 2016

History of Introduction and Spread

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P. vulgaris was introduced to Chile from Europe around 1870 for use in gardening, and is now found from 31°30′S to 42°53′S, from sea level to 1500 m (Godoy et al., 2011). It was also deliberately introduced to New Zealand in 1867 (Webb et al., 1988), although it was already present in Australia before 1801 (Low, 1999).

It is also recorded as introduced in some parts of the Caribbean and South America (WCSP, 2016).  

Introductions

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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
Chile Central Europe 1870 Horticulture (pathway cause) Yes Godoy et al. (2011)
New Zealand 1867 Yes Webb et al. (1988) Common throughout. A few specimens with some pinnately lobed leaves are probably referable to P. × hybrida Knaf (P. laciniata × P. vulgaris)

Risk of Introduction

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P. vulgaris subsp. lanceolata is available commercially as seed and container plants are occasionally sold at nurseries in the USA (Young-Mathews, 2012). Therefore it is possible that where seed/plants are sold there is the risk of accidental introduction into the wild. P. vulgaris subsp. lanceolata has been grown to produce lawns in the Pacific Northwest USA, providing a potential route for spreading to the wild (although it is considered a native subspecies by several sources) (Young-Mathews, 2012). There is a serious risk of it spreading to the Pacific and Indian Ocean islands where it is not yet present (PIER, 2016).  

Habitat

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P. vulgaris is found in grasslands, woods, bare ground and wasteland in the UK and Northern Europe (Fitter et al., 1980; Rose, 2006). It can also grow in wetlands in parts of North America (Alaska, Atlantic and Gulf Coastal Plain, Great Plains, Midwest, Northcentral and Northeast) and is occasionally found in the wetlands of Hawaii (USDA-NRCS, 2016). In New Zealand it tends to be found in damp, wet environments, including riverbeds, damp pastures and lawns and forest margins particularly when modified and open (PIER, 2016).
 
It is found in USDA zones 04a–09b and at elevations from sea level to 1720 m (Wilson, 2006).

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial
 
Terrestrial – ManagedCultivated / agricultural land Principal habitat Natural
Managed grasslands (grazing systems) Principal habitat Natural
Disturbed areas Principal habitat Natural
Rail / roadsides Principal habitat Natural
Urban / peri-urban areas Principal habitat Natural
Terrestrial ‑ Natural / Semi-naturalNatural grasslands Principal habitat Natural
Wetlands Secondary/tolerated habitat Natural
Scrub / shrublands Secondary/tolerated habitat Natural

Biology and Ecology

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Genetics
 
The chromosome number of P. vulgaris is 2n=28 (Bojnanský and Fargašová, 2007).
 
ISSR and SRAP markers have been used to investigate genetic diversity between P. vulgaris populations in China (Li et al., 2012). It was found that the 26 populations tested had high levels of genetic diversity, and they were clustered into three groups by cluster analysis. 
 
Reproductive Biology
 
P. vulgaris can reproduce asexually via stolons and sexually via seed production. It is thought to spread most aggressively via stolons, producing roots wherever they are in contact with soil (Wilson, 2006). However, some studies have found that in some habitats P. vulgaris reproduces almost exclusively via seed (Winn, 1985). 
 
Seeds are smooth, shiny, brown nutlets that are obovate, oblong, with a convex dorsal side and a roof-like ventral side (Encyclopedia of Life, 2016). Seeds do not appear to form a consistent seed bank (Clark and Wilson, 2003). In a two-year field investigation, seeds were found to have mortality rates of 54.4 and 68.5% in years 1 and 2, respectively (Clark and Wilson, 2003).
 
Physiology and Phenology
 
Flowers are protandrous and self-compatible (Kuriya et al., 2015) and are pollinated by insects e.g., Apidae, Halictidae, Scollidae, Bombyliidae, Lycaenidae and Hesperiidae (Encyclopedia of Life, 2016). Flowering time varies depending on the climate but is throughout the summer (Fitter et al., 1980). Each flower produces four seeds (Encyclopedia of Life, 2016). The corolla length has been shown to correlate with the proboscis length of the pollinator type present (Kuriya et al., 2015).
 
The plant has a variable growth rate of 3–45 cm in a growing season and can remain evergreen in mild climates (Wilson, 2006). 
 
P. vulgaris has been shown to exhibit phenotypic plasticity, local adaptation and the potential for rapid evolution of shade tolerance (Godoy et al., 2011). 
 
Longevity
 
Plants can live from 3–10 years (Wilson, 2006).
 
Population Size and Structure
 
Typical forest edge density near Mount Rainier/White Pass (Washington State, USA) has been observed as 3-10 plants per m2 (at 800 meters elevation) (Wilson, 2006). Density is primarily dependent upon water availability, with higher plant densities found in areas with more soil moisture consistently available throughout the growing season (Wilson, 2006). 
 
Associations
 
Bacillus wuyishanensis sp. nov., has been found in the rhizosphere of P. vulgaris in China (Liu et al., 2015). 
 
Many insects are associated with visiting P. vulgaris flowers for nectar or pollen, e.g., Apidae (bees), Halictidae (bees), Scollidae (wasps), Bombyliidae (flies), Lycaenidae (butterflies) and Hesperiidae (skippers) (Encyclopedia of Life, 2016). 
 
P. vulgaris is a weed or colonizer and can be seral at many successional stages, or form part of a climax community, e.g., at south Puget Sound prairie, USA (Wilson, 2006).
 
For full details of species associated with P. vulgaris see Wilson (2006).
 
Environmental Requirements
 
P. vulgaris has been shown to exhibit phenotypic plasticity and local adaptation when colonizing new areas, and has the potential for rapid evolution of shade tolerance (Godoy et al., 2011).
 
It prefers full sun to partial shade, on basic and neutral soils (soil pH 6.1–7.8) (ZipcodeZoo, 2014).

Climate

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ClimateStatusDescriptionRemark
Aw - Tropical wet and dry savanna climate Tolerated < 60mm precipitation driest month (in winter) and < (100 - [total annual precipitation{mm}/25])
BS - Steppe climate Tolerated > 430mm and < 860mm annual precipitation
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 Preferred Continental climate with dry summer (Warm average temp. > 10°C, coldest month < 0°C, dry summers)

Soil Tolerances

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Soil drainage

  • free
  • impeded
  • seasonally waterlogged

Soil reaction

  • alkaline
  • neutral

Soil texture

  • heavy
  • light
  • medium

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Agriopodes teratophora Herbivore Leaves not specific Encyclopedia of Life, 2016
Colias philodice Herbivore Leaves not specific NPIN, 2016
Meligethes umbrosus Herbivore Whole plant not specific Encyclopedia of Life, 2016
Trifurcula headleyella Herbivore Leaves not specific Kuchlein and Kuchlein-Nijsten, 2001

Notes on Natural Enemies

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P. vulgaris is a food plant for the larvae of the nepticulid moth (Trifurcula headleyella) in the Netherlands (Kuchlein and Kuchlein-Nijsten, 2001) and for larvae of the clouded sulphur butterfly (Colias philodice) in North America (NPIN, 2016). The larvae of the beetle Meligethes umbrosus feed on P. vulgaris, and the moth Agriopodes teratophora occasionally feeds on the foliage (Encyclopedia of Life, 2016).

In the former Yugoslavia, the eriphoid mite Aculops clinopodii has been isolated, from P. vulgaris (Petanovic, 1985) and Erysiphe biocellata has been found on the plants (Rankovic and Comic, 1995).

Septoria coelomycetous (anamorph of Septoria prunellae) causes spots on the leaves of P. vulgaris (Encyclopedia of Life, 2016).

Means of Movement and Dispersal

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Natural Dispersal

P. vulgaris produces stolons that produce roots wherever they are in contact with soil (Wilson, 2006). This species also produces viable seed but investigations have found that the species did not produce a persistent seed bank (Clark and Wilson, 2003; Wilson, 2006).

Vector Transmission

Seeds may be dispersed by invertebrates and vertebrates, particularly as Clark and Wilson (2003) showed that the seeds are sticky when wet.

Accidental Introduction

P. vulgaris subsp. lanceolata is available commercially as seed and container plants are occasionally sold at nurseries in the USA (Young-Mathews, 2012). Therefore it is possible that where seed or plants are sold there is the risk of accidental introduction into the wild or introduction through garden waste and contaminated soils.  

Intentional Introduction

There is a risk of intentional introduction to new areas as P. vulgaris subsp. lanceolata is available commercially as seed and container plants and so it may be intentionally introduced to gardens.  

Pathway Causes

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CauseNotesLong DistanceLocalReferences
Hitchhiker Yes
Medicinal useUsed by herbalists for medicines in Europe and China. Also used as an anti-cancer drug in China Yes Yes WCSP, 2016
Ornamental purposes Yes Yes Godoy et al., 2011

Impact Summary

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CategoryImpact
Cultural/amenity Positive
Environment (generally) Positive and negative

Environmental Impact

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Impact on Habitats

In Hawaii, P. vulgaris is reported as an invasive species (US Fish and Wildlife Service, 2015). Along with other nonnative plants it is thought to be capable of displacing all of the riparian habitat elements including native plants such as Deparia kaalaana on the Hawaiian Islands (US Fish and Wildlife Service, 2015).

Reports suggest that P. vulgaris is modifying and destroying habitat on the Hawaiian Islands; competing with native species for water, nutrients, light and space, and possibly via allelopathy (US Fish and Wildlife Service, 2015).

P. vulgaris subsp. lanceolata may be used for erosion control along roadsides, streambanks and pond edges due to its fibrous roots and spreading growth habit (Young-Mathews, 2012).

Impact on Biodiversity

P. vulgaris exhibits phenotypic plasticity, local adaptation and the potential for rapid evolution of shade tolerance and therefore may pose a threat to understory flora where it is an introduced species (Godoy et al., 2011). In its native range it can be invasive in lawns and is often considered a weed, although in some places it has been deliberately grown as an ornamental. It presents a serious risk to islands in the Pacific and Indian Ocean as it has already been recorded as invasive on a number of Hawaiian islands, Reunion and the Juan Fernández Islands (US Fish and Wildlife Service, 2012; US Fish and Wildlife Service, 2015PIER, 2016). It has also been recorded as invasive in New Zealand (PIER, 2016). 

In the Dry Cliff Ecosystem in Hawaii, P. vulgaris and other nonnative species are thought to be threatening the native plants Nothocestrum latifolium, Ochrosia haleakalae and Sicyos lanceoloideus (US Fish and Wildlife Service, 2015).

P. vulgaris and other nonnative plants are thought to be threatening Deparia kaalaana and Sanicula sandwicensis on the Hawaiian Islands (US Fish and Wildlife Service, 2015). It is also reported to be impacting Bidens campylotheca subsp. pentamera, Phyllostegia haliakalae and Pleomele fernaldii on the Hawaiian islands of Lanai and Maui (US Fish and Wildlife, 2012).

Both P. vulgaris and P. vulgaris subsp. lanceolata provide nectar and pollen to many insects. P. vulgaris subsp. lanceolata flowers provide nectar and pollen to butterflies such as whites (Pierinae), sulfurs (Coliadinae) and skippers (Hesperiidae), as well as native bees such as bumblebees (Bombus spp.), digger bees (Anthophora spp.), small carpenter bees (Ceratina spp.), long-horned bees (Eucera spp.) and green sweat bees (Agapostemon spp.) (Young-Mathews, 2012).

P. vulgaris subsp. lanceolata also has the potential to be used as a cover crop to provide habitat for pollinators and other beneficial species in vineyards, orchards, cane fruit or blueberry plantations (Young-Mathews, 2012).

Threatened Species

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Threatened SpeciesConservation StatusWhere ThreatenedMechanismReferencesNotes
Bidens campylotheca subsp. pentamera (ko`oko`olau)CR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered); USA ESA listing as threatened species USA ESA listing as threatened speciesHawaiiCompetition - monopolizing resources; Competition - shadingUS Fish and Wildlife Service, 2012
Chrysodracon fernaldii (hala pepe)EN (IUCN red list: Endangered) EN (IUCN red list: Endangered); USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiCompetition - monopolizing resources; Competition - shadingUS Fish and Wildlife Service, 2012
Deparia kaalaanaUSA ESA species proposed for listing USA ESA species proposed for listing; USA ESA species proposed for listing USA ESA species proposed for listingHawaiiUS Fish and Wildlife Service, 2015
Nothocestrum latifolium (broadleaf aiea)EN (IUCN red list: Endangered) EN (IUCN red list: Endangered); USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiCompetitionUS Fish and Wildlife Service, 2015
Ochrosia haleakalae (island yellowwood)EN (IUCN red list: Endangered) EN (IUCN red list: Endangered); NatureServe NatureServe; USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiCompetitionUS Fish and Wildlife Service, 2015
Phyllostegia haliakalae (Lanai phyllostegia)CR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered); USA ESA listing as endangered species USA ESA listing as endangered speciesHawaiiCompetition - monopolizing resources; Competition - shadingUS Fish and Wildlife Service, 2012
Sanicula sandwicensisUSA ESA species proposed for listing USA ESA species proposed for listingAllelopathic; Competition - monopolizing resources; Competition - shadingUS Fish and Wildlife Service, 2015
Sicyos lanceoloideusUSA ESA species proposed for listing USA ESA species proposed for listingHawaiiCompetitionUS Fish and Wildlife Service, 2015

Risk and Impact Factors

Top of page Invasiveness
  • Invasive in its native range
  • Proved invasive outside its native range
  • Has a broad native range
  • Abundant in its 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
  • Tolerant of shade
  • Reproduces asexually
  • Has high genetic variability
Impact outcomes
  • Ecosystem change/ habitat alteration
Impact mechanisms
  • Allelopathic
  • Competition - monopolizing resources
  • Competition - shading
  • Competition

Uses

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Economic Value

Many studies on P. vulgaris have focused on plant cultivation and domestication, effective chemical compounds analysis and the pharmacology of P. vulgaris (Li et al., 2012). 

P. vulgaris subsp. lanceolata may be sown as ornamental lawns in the Pacific Northwest USA (Young-Mathews, 2012).

Social Benefit

P. vulgaris has been used as a traditional medicinal herb in Europe and China for thousand of years (Li et al., 2012). The inflorescence spikes have been used to treat sore throat, fever and wounds and there is an increasing commercial demand for this plant due to its potential anti-inflammatory, antiviral and antibacterial activity (Li et al., 2012).

Its medicinal properties are due to the presence of constituents like triterpenoids, flavonoids, anionic polysaccharides-prullin, betulinic acid, d-camphor, oleanolic acid, uroslic acid and tannins (Kour et al., 2014).

For example, its triterpenoids show anti-allergic and anti-inflammatory effects, polysaccharide extracts show anti-microbial activity and reduce the expression of HSV-1 and HSV-2 antigens in hosts (Kour et al., 2014). Other investigations on triterpenoid extracts have been shown to inhibit glycogen phosphorylase and improve insulin sensitivity in adipocytes (Yu et al, 2014). Extracts from P. vulgaris exhibit anti-oestrogenic activity and inhibit the proliferation of endometrial cancer cell line-1 (Kour et al., 2014). Other medicinal properties include: anti-pyretic, anti-spasmodic, carminative, diuretic, vermifuge, antioxidant and dermatological photoprotectant (Kour et al., 2014).

Environmental Services

Many insects are associated with visiting P. vulgaris flowers for nectar or pollen, e.g., Apidae (bees), Halictidae (bees), Scollidae (wasps), Bombyliidae (flies), Lycaenidae (butterflies) and Hesperiidae (skippers) (Encyclopedia of Life , 2016).

P. vulgaris subsp. lanceolata may be used for erosion control along roadsides, streambanks and pond edges due to its fibrous roots and spreading growth habit (Young-Mathews, 2012).

P. vulgaris subsp. lanceolata has the potential to be used as a cover crop to provide habitat for pollinators and other beneficial species in vineyards, orchards, cane fruit or blueberry plantations (Young-Mathews, 2012).

P. vulgaris subsp. lanceolata flowers provide nectar and pollen to butterflies such as whites (Pierinae), sulfurs (Coliadinae), and skippers (Hesperiidae), as well as native bees such as bumblebees (Bombus spp.), digger bees (Anthophora spp.), small carpenter bees (Ceratina spp.), long-horned bees (Eucera spp.), and green sweat bees (Agapostemon spp.) (Young-Mathews, 2012). Caterpillars of the gray marvel moth (Agriopodes teratophora) sometimes feed on the foliage (Young-Mathews, 2012).

Uses List

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Human food and beverage

  • Leaves (for beverage)

Materials

  • Chemicals

Medicinal, pharmaceutical

  • Source of medicine/pharmaceutical
  • Traditional/folklore

Ornamental

  • Potted plant

Detection and Inspection

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P. vulgaris is easy to identify in the field, particularly when the distinctive inflorescence spikes are produced.

Similarities to Other Species/Conditions

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P. vulgaris is similar to a number of related species, including P. laciniata, P. grandiflora and P. hyssopifolia but can be distinguished by various characteristics. Cut-leaved self-heal (P. laciniata), usually has cream flowers with the upper leaves being pinnately lobed (Fitter et al., 1980; Rose, 2006). Large self-heal (P. grandiflora) is larger with flowers of 20-25mm and flowerheads with no leaves at the base and P. hyssopifolia, is distinguished by its narrow, entire, stalkless leaves and absence of stolons, and its distribution, which is limited to areas in France and Spain (Polunin and Smythies, 1989; Schmidt-Lebuhn, 2013). 
 
P. vulgaris is also similar to bugle (Ajuga reptans) although bugle has oval, hairless and untoothed leaves, and the inflorescence is a more open spike with bluer flowers (Fitter et al., 1980; Rose, 2006). 
 
P. vulgaris hybridises with P. grandiflora and P. laciniata in the UK and Northern Europe (Fitter et al., 1980). It also hybridizes with P. laciniata in New Zealand, with the hybrids (P. × hybrida Knaf) having pinnately lobed leaves (Webb et al., 1988).

Prevention and Control

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Eradication

It appears that seeds of P. vulgaris do not persist in the soil for much longer than 1-2 years (Clark and Wilson, 2003). Therefore eradication is likely to involve physical removal followed by repeated chemical treatments of an area. Where chemical treatments are not possible, repeated physical removal of the plants for two seasons or more would be required for eradication.

Physical/Mechanical Control

P. vulgaris is a small herbaceous perennial that could be removed by hand from small areas such as lawns although this would be laborious if the area is large and all root material should be removed to prevent vegetative regeneration. If lawns are regularly mowed and flowering is prevented then this will prevent spread via seed, although it will not prevent vegetative spread via stolons. 

Chemical Control

In field studies, P. vulgaris showed some tolerance to glyphosate or aminopyralid treatment, although the response varied significantly between locations and years (Pfleeger et al., 2012). In greenhouse investigations, P. vulgaris was more sensitive to glyphosate (Pfleeger et al., 2012). P. vulgaris also showed sensitivity to tribenuron (Olszyk, 2013).

If P. vulgaris is invasive in lawns, broad leaved herbicides can be used to remove it and other weeds from the turf. It is likely that repeated applications would be required to completely eradicate the plants.

Gaps in Knowledge/Research Needs

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Further taxonomic work or phylogenetic studies are required to determine the status of the species and subspecies of P. vulgaris. This could also help elucidate the origins of species that occur where they are considered to be non-native.

The persistence of seeds in the soil could be investigated further, particularly in Hawaii where the species is considered to be invasive and a threat to some native species. Physical control methods could also be investigated in Hawaiian field trials to determine how best to eradicate the species where herbicides cannot be used.

There is little information in the literature that describes natural pests and disease of P. vulgaris and so biological control does not seem to be part of any potential control methods at present. 

References

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Bean T, 2007. A new system for determining which plant species are indigenous in Australia. Australian Systematic Botany, 20(1):1-43.

Beckmann M; Bruelheide H; Erfmeier A, 2013. Local performance of six clonal alien species differs between native and invasive regions in Germany and New Zealand. Austral Ecology, 39(4):378-387.

Bojnansky V; Fargašova A, 2007. Atlas of Seeds and Fruits of Central and East-European Flora: The Carpathian Mountains Region. Dordrech, Netherlands: Springer Science and Business Media, 1046 pp.

Clark DL; Wilson MV, 2003. Post-dispersal seed fates of four prairie species. American Journal of Botany, 90(5):730-735.

Enclyclopaedia Britannica, 2016. Enclyclopaedia Britannica. Chicago, USA: Encyclopaedia Britannica. https://www.britannica.com/

Encyclopedia of Life, 2016. Encyclopedia of Life. http://www.eol.org

Fitter R; Fitter A; Blamey M, 1980. The Wildflowers of Britain and Northern Europe. London, UK: William Collins Sons & Co Ltd, 336 pp.

Flora of the Hawaiian Islands, 2016. Flora of the Hawaiian Islands. USA: National Museum of National History, Smithsonian Institution. http://botany.si.edu/pacificislandbiodiversity/hawaiianflora/query2.cfm

Godoy O; Saldaña A; Fuentes N; Valladares F; Gianoli E, 2011. Forests are not immune to plant invasions: phenotypic plasticity and local adaptation allow Prunella vulgaris to colonize a temperate evergreen rainforest. Biological Invasions, 13(7):1615-1625. http://www.springerlink.com/content/r6j75u8013842h25/

Hwang YJ; Lee EJ; Kim HR; Hwang KA, 2013. In vitro antioxidant and anticancer effects of solvent fractions from Prunella vulgaris var. lilacina. BMC Complementary and Alternative Medicine, 13:310-319.

Iremonger S, 2002. A Guide to Plants in the Blue Mountains of Jamaica. Kingston, Jamaica: University of the West Indies Press, 280 pp.

Kour B; Azhar M; Kaul S; Dhar MK, 2014. In vitro regeneration and mass multiplication of Prunella vulgaris L. National Academy Science Letters, 37(1):81-86. http://rd.springer.com/article/10.1007/s40009-013-0200-9

Kuchlein JH; Kuchlein-Nijsten CM, 2001. Trifurcula headleyella, a new nepticulid moth for the Dutch fauna (Lepidoptera: Nepticulidae). Entomologische Berichten, 61(9):126-129.

Kuriya S; Hattori M; Nagano Y; Itino T, 2015. Altitudinal flower size variation correlates with local pollinator size in a bumblebee-pollinated herb, Prunella vulgaris L. (Lamiaceae). Journal of Evolutionary Biology, 28(10):1761-1769. http://onlinelibrary.wiley.com/journal/10.1111/(ISSN)1420-9101

Li L; Guo QS; Wang ZY; Li L; Zhu ZB, 2012. Genetic diversity analysis of Prunella vulgaris in China using ISSR and SRAP markers. Biochemical Systematics and Ecology, 45:209-217.

Liu Bo; Liu GuoHong; Sengonca C; Schumann P; Che JianMei; Zhu YuJing; Wang JiePing, 2015. Bacillus wuyishanensis sp. nov., isolated from rhizosphere soil of a medical plant, Prunella vulgaris. International Journal of Systematic and Evolutionary Microbiology, 65(7):2030-2035. http://ijs.sgmjournals.org

Low T, 1999. Feral Future: The Untold Story of Australia's Exotic Invaders. Chicago, USA: University of Chicago Press, 394 pp.

Missouri Botanical Garden, 2016. Tropicos database. St. Louis, Missouri, USA: Missouri Botanical Garden. http://www.tropicos.org/

NPIN, 2016. Native Plant Database, Lady Bird Johnson Wildflower Center. Texas, USA: The University of Texas at Austin. http://www.wildflower.org/plants/

Olszyk D; Blakeley-Smith M; Pfleeger T; Lee EH; Plocher M, 2013. Effects of low levels of herbicides on prairie species of the Willamette Valley, Oregon. Environmental Toxicology and Chemistry, 32(11):2542-2551. http://onlinelibrary.wiley.com/journal/10.1002/(ISSN)1552-8618

Petanovic R, 1985. Studies on eriophyid mites (Acarida: Eriophyoidea) of Yugoslavia, I. Acta Entomologica Jugoslavica, 21(1-2):43-48.

Pfleeger T; Blakeley-Smith M; King G; Lee EH; Plocher M; Olszyk D, 2012. The effects of glyphosate and aminopyralid on a multi-species plant field trial. Ecotoxicology, 21(7):1771-1787. http://www.springerlink.com/content/e7217663n1662159/

PIER, 2016. Pacific Island Ecosystems at Risk. Honolulu, USA: HEAR, University of Hawaii. http://www.hear.org/pier/index.html

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Rankovic B; Comic L, 1995. Erysiphe biocellata, Erysiphe buhrii and Erysiphe lythrii, new parasites of some plant species in Yugoslavia. Zastita Bilja, 46(2):135-143

Rose F, 2006. The Wild Flower Key. London, UK: Frederick Warne, The Penguin Group, 576 pp.

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US Fish and Wildlife Service, 2015. Federal register, 80(189/ Wednesday September 30) Proposed Rules: 58820. The Daily Journal of the United States Government, 80(189). Washington DC, USA: National Archives and Records Administration. https://www.gpo.gov/fdsys/pkg/FR-2015-09-30/pdf/2015-24305.pdf

USDA-ARS, 2016. Germplasm Resources Information Network (GRIN). National Plant Germplasm System. Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysearch.aspx

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Wilson WS, 2006. Prunella vulgaris. Washington, USA: College of Forest Resources, University of Washington. http://depts.washington.edu/propplnt/Plants/prunella_vulgaris.htm

Winn AA, 1985. Effects of seed size and microsite on seedling emergence of Prunella vulgaris in four habitats. Journal of Ecology, 73(3):831-840.

Young-Mathews A, 2012. Plant fact sheet for lance selfheal (Prunella vulgaris ssp. lanceolata). Oregon, USA: USDA Natural Resources Conservation Service, Corvallis Plant Materials Center. http://plants.usda.gov/factsheet/pdf/fs_prvul2.pdf

Yu Q; Qi J; Wang L; Liu SJ; Yu BY, 2014. Pentacyclic Triterpenoids from Spikes of Prunella vulgaris L. Inhibit Glycogen Phosphorylase and Improve Insulin Sensitivity in 3T3-L1 Adipocytes. Phytotherapy Research, 29(1):73-79.

ZipcodeZoo, 2014. ZipcodeZoo. ZipcodeZoo. http://www.zipcodezoo.com/

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.
Massey University - New Zealand weeds databasehttp://weeds.massey.ac.nz/database.asp

Contributors

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26/01/16 Original text by:

Vicki Cottrell, Consultant, UK

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