Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide

Datasheet

Phleum pratense
(timothy grass)

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Datasheet

Phleum pratense (timothy grass)

Summary

  • Last modified
  • 19 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Host Plant
  • Preferred Scientific Name
  • Phleum pratense
  • Preferred Common Name
  • timothy grass
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Spermatophyta
  •       Subphylum: Angiospermae
  •         Class: Monocotyledonae
  • Summary of Invasiveness
  • Phleum pratense is a tufted or single-stemmed, short-lived, cool-season perennial grass that grows in stools or clumps, reaching up to 150 cm in height. It is an important forage grass that is grown in the cool...

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Pictures

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PictureTitleCaptionCopyright
Phleum pratense (timothy grass); habit. The stamens protruding from each flower render a distinctive 'fuzzy' appearance to the inflorescence spike. Bozeman, Montana, USA. July, 2009.
TitleHabit
CaptionPhleum pratense (timothy grass); habit. The stamens protruding from each flower render a distinctive 'fuzzy' appearance to the inflorescence spike. Bozeman, Montana, USA. July, 2009.
Copyright©Prof Matt Lavin-2009/Bozeman, Montana, USA - CC BY-SA 2.0
Phleum pratense (timothy grass); habit. The stamens protruding from each flower render a distinctive 'fuzzy' appearance to the inflorescence spike. Bozeman, Montana, USA. July, 2009.
HabitPhleum pratense (timothy grass); habit. The stamens protruding from each flower render a distinctive 'fuzzy' appearance to the inflorescence spike. Bozeman, Montana, USA. July, 2009.©Prof Matt Lavin-2009/Bozeman, Montana, USA - CC BY-SA 2.0
Phleum pratense (timothy grass); habit. This introduced species forms dense stands especially in open dry areas in the montane forest zone. 
Gallatin, Montana, USA. August, 2009.
TitleHabit
CaptionPhleum pratense (timothy grass); habit. This introduced species forms dense stands especially in open dry areas in the montane forest zone. Gallatin, Montana, USA. August, 2009.
Copyright©Prof Matt Lavin-2009/Bozeman, Montana, USA - CC BY-SA 2.0
Phleum pratense (timothy grass); habit. This introduced species forms dense stands especially in open dry areas in the montane forest zone. 
Gallatin, Montana, USA. August, 2009.
HabitPhleum pratense (timothy grass); habit. This introduced species forms dense stands especially in open dry areas in the montane forest zone. Gallatin, Montana, USA. August, 2009.©Prof Matt Lavin-2009/Bozeman, Montana, USA - CC BY-SA 2.0
Phleum pratense (timothy grass); habit. Timothy co-occurring with Tragopogon dubius, Lupinus argenteus, Helianthella uniflora, Artemisia ludoviciana, etc. Bozeman, Montana, USA. July, 2009.
TitleHabit
CaptionPhleum pratense (timothy grass); habit. Timothy co-occurring with Tragopogon dubius, Lupinus argenteus, Helianthella uniflora, Artemisia ludoviciana, etc. Bozeman, Montana, USA. July, 2009.
Copyright©Prof Matt Lavin-2009/Bozeman, Montana, USA - CC BY-SA 2.0
Phleum pratense (timothy grass); habit. Timothy co-occurring with Tragopogon dubius, Lupinus argenteus, Helianthella uniflora, Artemisia ludoviciana, etc. Bozeman, Montana, USA. July, 2009.
HabitPhleum pratense (timothy grass); habit. Timothy co-occurring with Tragopogon dubius, Lupinus argenteus, Helianthella uniflora, Artemisia ludoviciana, etc. Bozeman, Montana, USA. July, 2009.©Prof Matt Lavin-2009/Bozeman, Montana, USA - CC BY-SA 2.0
Phleum pratense (timothy grass); inflorescences. The tight cylindrical arrangement of spikelets, combined with the short stout awns arising from each of the glume tips is diagnostic. Gallatin, Montana, USA. July, 2009.
TitleInflorescences
CaptionPhleum pratense (timothy grass); inflorescences. The tight cylindrical arrangement of spikelets, combined with the short stout awns arising from each of the glume tips is diagnostic. Gallatin, Montana, USA. July, 2009.
Copyright©Prof Matt Lavin-2009/Bozeman, Montana, USA - CC BY-SA 2.0
Phleum pratense (timothy grass); inflorescences. The tight cylindrical arrangement of spikelets, combined with the short stout awns arising from each of the glume tips is diagnostic. Gallatin, Montana, USA. July, 2009.
InflorescencesPhleum pratense (timothy grass); inflorescences. The tight cylindrical arrangement of spikelets, combined with the short stout awns arising from each of the glume tips is diagnostic. Gallatin, Montana, USA. July, 2009.©Prof Matt Lavin-2009/Bozeman, Montana, USA - CC BY-SA 2.0
Phleum pratense (timothy grass); the stout awns from the glume tip renders a bristly flowering head.Bozeman, Montana, USA. September, 2009.
TitleAwns
CaptionPhleum pratense (timothy grass); the stout awns from the glume tip renders a bristly flowering head.Bozeman, Montana, USA. September, 2009.
Copyright©Prof Matt Lavin-2009/Bozeman, Montana, USA - CC BY-SA 2.0
Phleum pratense (timothy grass); the stout awns from the glume tip renders a bristly flowering head.Bozeman, Montana, USA. September, 2009.
AwnsPhleum pratense (timothy grass); the stout awns from the glume tip renders a bristly flowering head.Bozeman, Montana, USA. September, 2009.©Prof Matt Lavin-2009/Bozeman, Montana, USA - CC BY-SA 2.0
Phleum pratense (timothy grass); The well developed ligule is conspicuous from most angles. Gallatin, Montana, USA. June, 2005.
TitleLigule
CaptionPhleum pratense (timothy grass); The well developed ligule is conspicuous from most angles. Gallatin, Montana, USA. June, 2005.
Copyright©Prof Matt Lavin-2005/Bozeman, Montana, USA - CC BY-SA 2.0
Phleum pratense (timothy grass); The well developed ligule is conspicuous from most angles. Gallatin, Montana, USA. June, 2005.
LigulePhleum pratense (timothy grass); The well developed ligule is conspicuous from most angles. Gallatin, Montana, USA. June, 2005.©Prof Matt Lavin-2005/Bozeman, Montana, USA - CC BY-SA 2.0
Phleum pratense (timothy grass); the bulbous stem bases are covered with scaley leaves. Bozeman, Montana, USA. September, 2009.
TitleStem base
CaptionPhleum pratense (timothy grass); the bulbous stem bases are covered with scaley leaves. Bozeman, Montana, USA. September, 2009.
Copyright©Prof Matt Lavin-2009/Bozeman, Montana, USA - CC BY-SA 2.0
Phleum pratense (timothy grass); the bulbous stem bases are covered with scaley leaves. Bozeman, Montana, USA. September, 2009.
Stem basePhleum pratense (timothy grass); the bulbous stem bases are covered with scaley leaves. Bozeman, Montana, USA. September, 2009.©Prof Matt Lavin-2009/Bozeman, Montana, USA - CC BY-SA 2.0

Identity

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Preferred Scientific Name

  • Phleum pratense L.

Preferred Common Name

  • timothy grass

Other Scientific Names

  • Phleum nodosum L.
  • Phleum pratense subsp. nodosum (L.) Dumort

International Common Names

  • English: common cat's tail; common cat's-tail; herd grass; herd’s grass; meadow cat's tail; meadow cat's-tail; timothy
  • Spanish: cola de gato; cola de rato; cola de topo; fleo de los prados; fleo pratense; timoti
  • French: fléole des prés; timothée
  • Chinese: ti mu cao

Local Common Names

  • Germany: Timotheegras; Wiesenlieschgras
  • Italy: coda di topo; codolina; fleolo; fleolo dei prati
  • Japan: o-awagaeri; ooawagaeri
  • Netherlands: timothee; timotheegras
  • Portugal: capim-timóteo; rabo-de-gato
  • Sweden: timotej

EPPO code

  • PHLPR (Phleum pratense)

Summary of Invasiveness

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Phleum pratense is a tufted or single-stemmed, short-lived, cool-season perennial grass that grows in stools or clumps, reaching up to 150 cm in height. It is an important forage grass that is grown in the cool temperate regions of the world. Native to Europe and Asia, it has been widely introduced and grown worldwide as a highly palatable pasture grass, being particularly suitable for hay making. However, it has become invasive and a noxious weed in parts of its introduced range, dominating other plants, altering native plant communities and often forming monocultures. Control of this grass in conservation areas is difficult.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Plantae
  •         Phylum: Spermatophyta
  •             Subphylum: Angiospermae
  •                 Class: Monocotyledonae
  •                     Order: Cyperales
  •                         Family: Poaceae
  •                             Genus: Phleum
  •                                 Species: Phleum pratense

Notes on Taxonomy and Nomenclature

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The genus Phleum contains over 15 species (The Plant List, 2013), with P. pratense the only species widely cultivated, although the diploid species P. bertololonii is cultivated in some countries, especially as an amenity grass (Jensen, 2005).

Phleum pratense L. is an accepted name originally published by Linnaeus in 1753 in volume I of his Species Plantarum. According to the Plant List (2013) the species has 46 synonyms and no infraspecific taxa.

P. pratense was reputedly given the common name timothy or timothy grass after Timothy Hanson, a US farmer and agriculturalist who is said to have introduced the species from New England to Maryland and other southern states of the USA from about 1720 (Lacefield et al., 2002).

Description

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According to Peeters (2004) and with additional material from Hubbard (1968)P. pratense is a tall, tufted or single-stemmed, short-lived, cool-season perennial grass. The plant is robust, hairless and caespitose. Stems are erect, 20-100(-150) cm tall, often bulging at the base and forming a small bulb. The blade is rolled when young, large (3-10 mm wide), flat, slightly rough on the margin, rather long (reaching 45 cm), pale green to greyish green. The ligule is strong, obtuse and white, and there are no auricles. The spike-like panicle is cylindrical, 6-20(-30) cm long. Spikelets are 1-flowered, breaking up at maturity above the glumes. Glumes persistent, narrowly oblong, truncate, keeled, 3-nerved, the keels fringed with soft spreading hairs and produced at the tip into a rigid awn 1-2 mm long. Lemma and palea one-third to three-quarters the length of the glumes. Anthers 2 mm long . Seeds are small, 2 mm long, with a 1000-seed weight of 0.3 to 0.7 g. P. pratense produces few tillers (4000 to 10,000 tillers/m²) compared to perennial ryegrass (Lolium perenne; 6000 to 15,000 tillers/m²). However, leaves are produced at a slightly faster rate than in perennial ryegrass, and the maximum number of leaves per tiller is up to 6-7 (rarely 8) against only 3 for the ryegrass. Leaf lifetime is also longer, so that timothy is able to accumulate a lot of standing biomass before senescence commences. 

Plant Type

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Herbaceous
Perennial
Seed propagated

Distribution

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According to the National Plant Germplasm System of the USDA-ARS (2015), timothy is native to Europe, northern Africa and temperate Asia, as well as parts of India and Pakistan. It is naturalized in Australia, New Zealand, Canada, Greenland, Mexico and the USA, including Hawaii and Alaska, as well as Mauritius, the West Indies, Argentina, Chile, Uruguay and Peru. It has also been introduced and is under cultivation in Japan. According to Kew (2015), the species is introduced in western, eastern and northern China, although the USDA-ARS (2015) considers it native to that country. Conversly, while Kew treats it as native to India, Bor (1960) has it as an introduction.

In North America, timothy is found in all 50 states of the USA and all Canadian provinces except Nunavut (USDA-NRCS, 2015). In the USA, timothy is most widely cultivated in the northeastern states, south to the Cotton Belt and west to the 100th meridian, in humid regions of Puget Sound and in mountainous regions (Esser, 1993). It is the most important forage grass in the northern part of the Nordic countries of Europe (Larsen and Marum, 2006).

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

ArmeniaPresentNativeUSDA-ARS, 2015
AzerbaijanPresentNativeUSDA-ARS, 2015
BhutanPresent only in captivity/cultivationIntroducedUSDA-ARS, 2015
ChinaPresentIntroducedKew, 2015; USDA-ARS, 2015Introduced to north-central, south-central, southeast and Manchuria, but considered native by USDA-ARS
-XinjiangPresentIntroducedKew, 2015
Georgia (Republic of)PresentNativeUSDA-ARS, 2015
IndiaPresentNativeBor, 1960; Kew, 2015Western Himalayas
-Himachal PradeshPresentNativeShukla, 1996
-NagalandPresentNativeShukla, 1996
IranPresentNativeUSDA-ARS, 2015
IraqPresentNativeUSDA-ARS, 2015
JapanPresentIntroduced Invasive USDA-ARS, 2015
-HokkaidoPresentIntroduced Invasive Invasive Species of Japan, 2015
KazakhstanPresentNativeUSDA-ARS, 2015
Korea, DPRPresentIntroducedKew, 2015
Korea, Republic ofPresentIntroducedKew, 2015
KyrgyzstanPresentNativeUSDA-ARS, 2015
LebanonPresentNativeKew, 2015
MongoliaPresentNativeUSDA-ARS, 2015
PakistanPresentNativeUSDA-ARS, 2015
SyriaPresentNativeKew, 2015
TajikistanPresentNativeKew, 2015
TurkeyPresentNativeKew, 2015Including Turkey in Europe
UzbekistanPresentNativeKew, 2015

Africa

Crozet IslandsPresentIntroducedKew, 2015
EgyptPresentIntroducedKew, 2015
MauritiusPresentIntroduced Invasive PIER, 2016
MoroccoPresentNativeKew, 2015

North America

BermudaPresentIntroducedKew, 2015
CanadaPresentPresent based on regional distribution.
-AlbertaPresentIntroducedKew, 2015
-British ColumbiaPresentIntroducedKew, 2015
-ManitobaPresentIntroducedKew, 2015
-New BrunswickPresentIntroducedKew, 2015
-Newfoundland and LabradorPresentIntroducedKew, 2015
-Northwest TerritoriesPresentIntroducedKew, 2015
-Nova ScotiaPresentIntroducedKew, 2015
-OntarioPresentIntroducedKew, 2015
-Prince Edward IslandPresentIntroducedKew, 2015
-QuebecPresentIntroducedKew, 2015
-SaskatchewanPresentIntroducedKew, 2015
-Yukon TerritoryPresentIntroducedKew, 2015
GreenlandPresentIntroducedKew, 2015
MexicoPresentIntroducedKew, 2015Present in south-west
USAPresentPresent based on regional distribution.
-AlabamaPresentIntroducedKew, 2015
-AlaskaPresentIntroduced Invasive Kew, 2015Including the Aleutian Islands
-ArizonaPresentIntroducedKew, 2015
-ArkansasPresentIntroducedKew, 2015
-CaliforniaPresentIntroducedKew, 2015
-ColoradoPresentIntroducedKew, 2015
-ConnecticutPresentIntroducedKew, 2015
-DelawarePresentIntroducedKew, 2015
-District of ColumbiaPresentIntroducedKew, 2015
-FloridaPresentIntroducedKew, 2015
-GeorgiaPresentIntroducedKew, 2015
-HawaiiPresentIntroducedKew, 2015; PIER, 2016
-IdahoPresentIntroducedKew, 2015
-IllinoisPresentIntroducedKew, 2015
-IndianaPresentIntroducedKew, 2015
-IowaPresentIntroducedKew, 2015
-KansasPresentIntroducedKew, 2015
-KentuckyPresentIntroducedKew, 2015
-LouisianaPresentIntroducedKew, 2015
-MainePresentIntroducedKew, 2015
-MarylandPresentIntroducedKew, 2015
-MassachusettsPresentIntroducedKew, 2015
-MichiganPresentIntroducedKew, 2015
-MinnesotaPresentIntroducedKew, 2015
-MississippiPresentIntroducedKew, 2015
-MissouriPresentIntroducedKew, 2015
-MontanaPresentIntroduced Invasive Kew, 2015In Glacier National Park
-NebraskaPresentIntroducedKew, 2015
-NevadaPresentIntroducedKew, 2015
-New HampshirePresentIntroducedKew, 2015
-New JerseyPresentIntroducedKew, 2015Listed as a noxious weed
-New MexicoPresentIntroducedKew, 2015
-New YorkPresentIntroducedKew, 2015
-North CarolinaPresentIntroducedKew, 2015
-North DakotaPresentIntroducedKew, 2015
-OhioPresentIntroducedKew, 2015
-OklahomaPresentIntroducedKew, 2015
-OregonPresentIntroducedKew, 2015
-PennsylvaniaPresentIntroducedKew, 2015
-Rhode IslandPresentIntroducedKew, 2015
-South CarolinaPresentIntroducedKew, 2015
-South DakotaPresentIntroducedKew, 2015
-TennesseePresentIntroducedKew, 2015
-TexasPresentIntroducedKew, 2015
-UtahPresentIntroducedKew, 2015
-VermontPresentIntroducedKew, 2015
-VirginiaPresentIntroducedKew, 2015Listed as a noxious weed
-WashingtonPresentIntroducedKew, 2015
-West VirginiaPresentIntroducedKew, 2015
-WisconsinPresentIntroducedKew, 2015
-WyomingPresentIntroduced Invasive Kew, 2015In Yellowstone National Park

Central America and Caribbean

CaribbeanPresentIntroducedUSDA-ARS, 2015
JamaicaPresentIntroducedKew, 2015

South America

ArgentinaPresentIntroducedKew, 2015In north-east and south
BrazilPresentIntroducedKew, 2015In south
ChilePresentIntroducedKew, 2015In south and central regions
ColombiaPresentIntroducedKew, 2015
Falkland IslandsPresentIntroducedKew, 2015
PeruPresentIntroducedUSDA-ARS, 2015
South Georgia and the South Sandwich IslandsPresentIntroducedKew, 2015
UruguayPresentIntroducedUSDA-ARS, 2015

Europe

AlbaniaPresentNativeKew, 2015
AustriaPresentNativeKew, 2015
BelarusPresentNativeUSDA-ARS, 2015
BelgiumPresentNativeKew, 2015
BulgariaPresentNativeKew, 2015
CroatiaPresentNativeUSDA-ARS, 2015
Czech RepublicPresentNativeUSDA-ARS, 2015
DenmarkPresentNativeKew, 2015
EstoniaPresentNativeUSDA-ARS, 2015
Faroe IslandsPresentNativeKew, 2015
FinlandPresentNativeKew, 2015
FrancePresentNativeKew, 2015Including Corsica
GermanyPresentNativeKew, 2015
GreecePresentNativeKew, 2015
HungaryPresentNativeKew, 2015
IcelandPresentNativeKew, 2015
IrelandPresentNativeKew, 2015
ItalyPresentNativeKew, 2015Including Sardinia and Sicily
LatviaPresentNativeUSDA-ARS, 2015
LithuaniaPresentNativeUSDA-ARS, 2015
MoldovaPresentNativeUSDA-ARS, 2015
NetherlandsPresentNativeKew, 2015
NorwayPresentNativeKew, 2015
PolandPresentNativeKew, 2015
PortugalPresentNativeKew, 2015
-AzoresPresentNativeKew, 2015
RomaniaPresentNativeKew, 2015
Russian FederationPresentPresent based on regional distribution.
-Central RussiaPresentNativeKew, 2015
-Eastern SiberiaPresentNativeKew, 2015
-Northern RussiaPresentNativeKew, 2015
-Russian Far EastPresentIntroducedKew, 2015Including Sakhalin and the Kuril Islands
-Southern RussiaPresentNativeKew, 2015Caucasus region
-Western SiberiaPresentNativeKew, 2015
SerbiaPresentNativeUSDA-ARS, 2015
SlovakiaPresentNativeUSDA-ARS, 2015
SloveniaPresentNativeUSDA-ARS, 2015
SpainPresentNativeKew, 2015
SwedenPresentNativeKew, 2015
SwitzerlandPresentNativeKew, 2015
UKWidespreadNativeKew, 2015
UkrainePresentNativeUSDA-ARS, 2015
Yugoslavia (Serbia and Montenegro)PresentNativeKew, 2015Native to “Yugoslavia”

Oceania

AustraliaPresentIntroducedUSDA-ARS, 2015
-New South WalesPresentIntroduced Invasive Kew, 2015
-South AustraliaPresentIntroducedKew, 2015
-TasmaniaPresentIntroducedKew, 2015
-VictoriaPresentIntroduced Invasive Kew, 2015
-Western AustraliaPresentIntroduced Invasive Kew, 2015
New ZealandPresentIntroducedKew, 2015; PIER, 2016North, South and Chatham Islands

History of Introduction and Spread

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Although P. pratense is of Eurasian origin, it was first cultivated on any scale as a pasture grass in the USA. Known in Britain as common or meadow cat’s tail, it was probably introduced unintentionally to North America by early settlers. In North America it was termed herd’s grass after John Herd (or Hurd) who found it naturalized in New Hampshire in the early 1700s, while the more popular designation "timothy" may be derived from the name of Timothy Hansen, who apparently brought the grass from New England and popularized it in Maryland and other Atlantic states in the 1720s. By 1747, timothy had spread from New England north to Canada and westward, and by 1807 was the most important hay grass in the USA (Hoover et al., 1948; Encyclopedia of Life, 2015). Following its success in North America, it was promoted and introduced as a pasture grass to Britain in the 1760s and from there to other European countries (Larsen and Marum, 2006), as well as to Britain’s overseas colonies. P. pratense was introduced as a pasture grass to Japan (Hokkaido) in 1874 (Invasive Species of Japan, 2015).

Introductions

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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
Canada USA 1747 Forage (pathway cause) Yes Hoover et al. (1948)
Japan 1874 Forage (pathway cause) Yes Invasive Species of Japan (2015)
UK USA 1760s Forage (pathway cause) Yes Larsen and Marum (2006) Re-introduction of improved pasture types
USA Europe 1700-1720 Forage (pathway cause) Yes Hoover et al. (1948)

Risk of Introduction

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Timothy is relatively short-lived and spreads via seeds, which are easily dispersed by wind (NatureServe, 2015). According to Ogle et al. (2011), timothy is not considered a weedy or invasive species, but can spread into adjoining vegetative communities under ideal climatic and environmental conditions. However, it is listed as a noxious weed in several US states, including New Jersey and Virginia (Klein, 2011). It is considered as invasive in Wyoming’s Yellowstone National Park (Wallace and Macko, 1993), Alaska (Tyser, 1992) and Hawaii (PIER, 2016), as well as in Japan (Invasive Species of Japan, 2015), New Zealand and Australia (PIER, 2016). In Australia it is classed as an environmental weed (Queensland Government, 2015).

As P. pratense is widely grown as a pasture crop, especially in cool temperate regions, the risk of escape from cultivation is relatively high due to the species’ profuse production of light, wind-dispersed seeds (1000-seed weight = 0.3-0.7 g (Peeters, 2015)). It is known to have escaped cultivation and become established in grass and forb meadows in Alaska, and as its use for reclamation and erosion control increases, with planting for the rehabilitation of sites altered by recreational activities or disturbed by the construction of railroads, canals, trails and highways (Elliott et al. 1987), so the risk of introduction into the wild increases. The Alaska Natural Heritage Program has given P. pratense an invasiveness ranking of 54, which is calculated according to a species’ ecological impacts, biological attributes, distribution and response to control measures. The ranks are scaled from 0 to 100, with 0 representing a plant that poses no threat to native ecosystems and 100 representing a plant that poses a major threat (Klein, 2011).

Habitat

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In its native range, P. pratense was probably confined to the moist soils of water meadows and other low-lying grasslands. Since its widespread cultivation it is now common on a range of soils, as well as in field margins, roadsides and waste places (Hubbard, 1968). In North America it can be found growing in waterways, dry to wet meadows and other mesic environments. It is commonly found volunteering in canals and roadside borrow ditches (Ogle et al., 2011). It is encountered as an invasive species in abandoned fields (invasive.org, 2010), forests, meadows and alpine tundra (Esser, 1993), forest understorey (Houston, 1973), montane grasslands (Wallace and Macko, 1993) and early- to mid-successional grasslands and early-seral mixed forests (NatureServe, 2015). In Australia when grown as a forage crop it prefers heavy soils in moist habitats, but has spread into native vegetation, notably dry coastal vegetation, dry sclerophyll forests and woodlands (Queensland Government, 2015). In Japan it has invaded crop fields, farms, grasslands, streams, urban areas, vacant lots and waterfronts (Invasive Species of Japan, 2015).

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial
Terrestrial – ManagedCultivated / agricultural land Principal habitat Productive/non-natural
Managed forests, plantations and orchards Secondary/tolerated habitat Harmful (pest or invasive)
Managed forests, plantations and orchards Secondary/tolerated habitat Natural
Disturbed areas Principal habitat Harmful (pest or invasive)
Disturbed areas Principal habitat Natural
Rail / roadsides Secondary/tolerated habitat Natural
Urban / peri-urban areas Secondary/tolerated habitat Productive/non-natural
Terrestrial ‑ Natural / Semi-naturalNatural forests Secondary/tolerated habitat Harmful (pest or invasive)
Natural grasslands Principal habitat Harmful (pest or invasive)
Natural grasslands Principal habitat Natural
Riverbanks Principal habitat Natural
Wetlands Principal habitat Natural
Cold lands / tundra Secondary/tolerated habitat Natural
Freshwater
Irrigation channels Secondary/tolerated habitat Natural

Hosts/Species Affected

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Timothy is of great concern to wildland managers because it often dominates areas it invades. It has the potential to inhibit secondary successional processes and modify native communities (Rutledge and McLendon, 1996). The species can cause declines in and competitively exclude native grasses (NatureServe, 2015). In Yellowstone National Park, Wyoming, Houston (1973) observed that timothy had dominated and completely altered the composition of the understorey vegetation in remnant aspen (Populus tremuloides) stands in the northern range. Tyser (1992) observed that extensive tiller mats of timothy limited cryptogam colonization sites and reduced native graminoid colonization in invaded fescue (Festuca) grassland in Glacier National Park, Montana. 

Timothy can also be detrimental in young conifer plantations, where its seedlings compete strongly with conifer seedlings, especially those which are not fully established, by pre-emption of resources, allelopathy, attraction of insect pests and browsing animals, and increased fire potential. However, after conifer establishment (approximately 5 years), timothy may aid conifer growth by excluding shrub competition (Esser, 1993).

Host Plants and Other Plants Affected

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Plant nameFamilyContext
Festuca (fescues)PoaceaeMain

Biology and Ecology

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Genetics

P. pratense is a hexaploid species, with 2n = 6x = 42 (Joachimiak and Grabowska-Joachimiak, 2000). Using RAPD and UP-PCR molecular fingerprinting methods, Guo et al. (2003) found that the species’ genetic variation was largely dependent on geographic diversity; different geographic genotype groups were distinguished.

Reproductive Biology

Timothy mainly reproduces by seeds, which can remain viable for at least 4-5 years in dry, cool conditions. Baskin and Baskin (1998) include records of seed longevity to 20 and 39 years. Germination does not require alternating temperatures or light, but may be slow. Timothy can also reproduce vegetatively through tillering (Esser, 1993).

Environmental Requirements

According to Ogle et al. (2011), timothy is adapted to soils with a pH of 5.5 to 7.0. It is tolerant of partially shaded conditions and is very winter-hardy, exhibiting tolerance of both cold temperature and ice encasement. It does not tolerate drought or prolonged high temperatures, nor alkaline conditions. Thakur et al. (2014) demonstrated that P. pratense in the temperate-boreal forest ecotone of Minnesota significantly increased shoot biomass production in soils with a +3.4ºC warming history compared to those with +1.7 ºC or ambient warming histories, indicating that such +3.4ºC soils would favour invasive P. pratense establishment.

Climate

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ClimateStatusDescriptionRemark
Cf - Warm temperate climate, wet all year Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year
Cs - Warm temperate climate with dry summer Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers
Cw - Warm temperate climate with dry winter Tolerated 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)

Latitude/Altitude Ranges

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Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
65 45

Soil Tolerances

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Soil drainage

  • free
  • impeded

Soil reaction

  • acid
  • neutral

Soil texture

  • heavy
  • medium

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Alternaria arbusti Pathogen Leaves not specific
Alternaria viburni Pathogen Leaves not specific
Apiospora montagnei Pathogen Leaves not specific
Aureobasidium pullulans Pathogen Leaves not specific
Bacillus thuringiensis kurstaki Pathogen
Bacillus thuringiensis thuringiensis Pathogen
Cladosporium phlei Pathogen Leaves
Drechslera phlei Pathogen Leaves
Epicoccum nigrum Pathogen Leaves not specific
Fusarium sporotrichioides Pathogen Leaves not specific
Gibberella avenacea Pathogen Leaves not specific
Monographella Pathogen Leaves not specific
Paraphaeosphaeria michotii Pathogen Leaves not specific
Phaeosphaeria herpotrichoides Pathogen Leaves not specific
Puccinia graminis Pathogen Leaves not specific
Thymelicus lineola Herbivore Leaves not specific

Notes on Natural Enemies

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Stem rust (Puccinia graminis) is one of the most important diseases of timothy, causing loss of vigour and forage quality. Purple eyespot (Cladosporium phlei) and brown leaf blight (Drechslera phlei) are diseases commonly found across western Canada. Timothy is damaged by grasshoppers, and newly sown fields are also susceptible to wireworms and cutworms. Larvae of the European skipper butterfly (Thymelicus lineola) are a pest of timothy in eastern Canada (Ogle et al., 2011).

A preliminary study of endophytic fungi on P. pratense growing in Estonia found that half of the taxa isolated in this study (Aureobasidium pullulans, Epicoccum nigrum, Gibberella avenacea, Monographella sp. and Phaeosphaeria herpotrichoides) had been identified from timothy earlier. Five species (Alternaria arbusti, Lewia viburni [Alternaria viburni], Apiospora montagnei, Fusarium sporotrichioides and Paraphaeosphaeria michotii) were new reports for P. pratense. Two of these, A. arbusti and L. viburni, were also new species for Estonia (Varvas et al., 2013).

 

Means of Movement and Dispersal

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Natural Dispersal

P. pratense spreads via seeds, which are easily dispersed by wind (NatureServe, 2015).

Vector Transmission (Biotic)

Seed is reported to be transported by livestock (USDA Forest Service, 1937), although seeds lack any specific adaptations for animal dispersal (Klein, 2011).

Accidental Introduction

As a major hay and silage species, P. pratense seed can easily be shed from hay used in animal feeding.

Intentional Introduction

Timothy is one of the more important perennial grass cultivated in North America primarily for hay. In the western ranges it is also important as a forage plant (USDA Forest Service, 1937). As such a valuable grass crop, it is intentionally planted not only in North America but throughout its introduced range. 

Pathway Causes

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CauseNotesLong DistanceLocalReferences
Crop production Yes Yes
Disturbance Yes
Escape from confinement or garden escape Yes
Forage Yes Yes
Habitat restoration and improvement Yes Yes
Seed trade Yes

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Livestock Yes
Plants or parts of plantsSeed contaminant of seed lots; seed shed from hay Yes Yes
Wind Yes

Plant Trade

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

Impact Summary

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

Economic Impact

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Seed of P. pratense is considered a contaminant of grass and other seed lots in the eastern US states of Delaware, Maryland, New Hampshire, New Jersey, Pennsylvania, Virginia and West Virginia (Ogle et al., 2011), thus reducing seed lot quality and price. Timothy is also a host to diseases, such as ergot (Claviceps purpurea), that are serious pathogens of cereal crops (Munkvold et al., 1997; Davies and Ballingall, 2008).

Environmental Impact

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Impact on Habitats

Under ideal climatic and environmental conditions P. pratense can spread from cultivation into adjoining native plant communities where it can coexist or form monocultures (Ogle et al., 2011). Timothy is one of the exotics of greatest concern to wildland managers in North America because it establishes quickly, spreads vigorously and usually escapes early detection. Of 34 exotic species tested, timothy had the highest ability to invade closed vegetation areas, including forest, meadow and alpine tundra (Weaver et al., 1990). The species is capable of invading early- to mid-successional grasslands and early-seral mixed forests (NatureServe, 2015).

P. pratense has invaded and is seen as a major problem in several national and state parks in the USA. It is the most widely distributed exotic in Glacier National Park in Montana and is one of the most disruptive factors in native fescue grasslands there, having been intentionally seeded following underground utility construction in the 1980s. Timothy is of great concern because it often dominates the areas it invades. Extensive tiller mats of timothy limit cryptogam colonization sites and reduce native grass colonization (Tyser, 1992). In Yellowstone National Park, Wyoming, Houston (1973) observed that timothy had dominated and completely altered the composition of the understorey vegetation in remnant aspen stands in the northern range and as a result these stands would not respond to burning. Wallace and Macko (1993) noted it as an alien invasive occupying large areas of the montane grasslands of Yellowstone’s northern range.

In Australia, P. pratense is regarded as an environmental weed in Victoria and as a minor environmental weed or "sleeper weed" in other temperate regions of southern Australia. It has spread from cultivation to invade natural vegetation, posing a significant threat to dry coastal vegetation, dry sclerophyll forests and woodlands in Victoria. It appears on some local and regional environmental weed lists in this state, including those of the Goulburn Broken Catchment and at Falls Creek. In southern New South Wales, timothy has become naturalized in subalpine and montane areas of Kosciuszko National Park. It has also been recorded along the Warren River in south-western Western Australia (Queensland Government, 2015).

Impact on Biodiversity

Timothy can cause declines in and competitively exclude native grasses. It has the potential to inhibit secondary successional processes and modify native plant communities (Rutledge and McLendon, 1996). 

Social Impact

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Allergens are released from timothy pollen grains (Behrendt et al., 1999). In fact, pollen of the genus Phleum is second only to Dactylis in allergenicity, causing widespread pollinosis in early summer. P. pratense pollen contains at least 28 antigens, of which 15 have been shown to bind to IgE. A number of major allergens have also been detected, including Phl p 1, Phl p 4, Phl p 5, Phl p 6, Phl p 7, Phl p 12 and Phl p 13, with Phl p 5 being the dominant allergen (Steinman, 2012).  

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Has a broad native range
  • Abundant in its native range
  • Tolerates, or benefits from, cultivation, browsing pressure, mutilation, fire etc
  • Pioneering in disturbed areas
  • Tolerant of shade
  • Benefits from human association (i.e. it is a human commensal)
  • Fast growing
  • Has propagules that can remain viable for more than one year
Impact outcomes
  • Damaged ecosystem services
  • Modification of fire regime
  • Modification of successional patterns
  • Monoculture formation
  • Negatively impacts forestry
  • Negatively impacts human health
  • Reduced native biodiversity
  • Threat to/ loss of native species
Impact mechanisms
  • Allelopathic
  • Causes allergic responses
  • Competition - monopolizing resources
  • Pest and disease transmission
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally
  • Highly likely to be transported internationally deliberately
  • Difficult to identify/detect as a commodity contaminant
  • Difficult/costly to control

Uses

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Economic Value

From an agricultural perspective timothy is a very persistent and hardy grass, largely free of pests and diseases, and an extremely valuable crop for pasture and the production of hay and silage, as well as seed. Timothy is preferred by cattle and horses, and timothy hay is a premium feed for horses. Sheep utilize timothy during the summer in mountainous areas. Timothy seed is commonly used in wildlife seed mixtures. Under irrigated conditions, seed yields average 450-670 kg/ha (Ogle et al., 2011). Timothy is grown alone or in combination with lucerne, birdsfoot trefoil or clovers; with an application of 60 kg N/ha, timothy and clover mixtures have yielded 9.41 t DM/ha in Lithuania (Kadziulis, 1974). Sown alone, it yielded 4.86 t DM/ha in Italy (Cenci and Pagiotti, 1979). P. pratense is the most popular grass species in Konsen district in Japan, helping to maintain high milk production in cows (Saigusa et al., 1997).

Environmental Services

According to Esser (1993), timothy is often used, particularly in plant mixtures, to stabilize soil against erosion and to provide cover for wildlife in clearcut areas that have been burned. It is widely used for rehabilitation of cutover, burned-over and overgrazed mountain rangelands. Timothy provides a highly palatable and nutritious forage, not just for domestic livestock, but also for big game animals. According to Ogle et al. (2011), timothy can be used with legumes and/or other grasses in seed mixtures for cover, filter strips, herbaceous buffers, waterways and other critical area applications.

Timothy stands provide habitat and nesting cover for game birds, small mammals and waterfowl. The seeds are consumed by birds (Esser, 1993).

Uses List

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Animal feed, fodder, forage

  • Fodder/animal feed
  • Forage

Environmental

  • Amenity
  • Erosion control or dune stabilization
  • Revegetation
  • Wildlife habitat

Similarities to Other Species/Conditions

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Timothy is often confused with meadow foxtail (Alopecurus pratensis) and creeping foxtail (A. arundinaceus), but the foxtails have smoother, silky, flowering heads.

In timothy, the spikelets are strongly flattened, with a characteristic U-shaped outline and translucent bristles extending from the edges. The glumes extend into short stout awns that resemble horns. The tight cylindrical arrangement of spikelets, combined with the awns arising from each of the glume tips, is diagnostic of timothy. Compared to timothy, in the flowering heads of Alopecurus the spikelets are not strongly flattened and are more or less ovate in outline. The glumes do have long hairs but they lack awns at the tip and the lemmas have a delicate awn extending from the back. The glumes of timothy in contrast have strong awns and the lemmas lack awns entirely (Iowa State University, 2016).

Cured seed heads of creeping foxtail are dark or blackish, while those of timothy are tan to buff coloured. Timothy also resembles alpine timothy (P. alpinum), but plants of the latter are shorter and produce much shorter inflorescences. Additionally, alpine timothy is mainly found in montane meadows at higher altitudes than P. pratense and usually flowers in late July to early August, while timothy flowers from June until September (Ogle et al., 2011).

Timothy differs from most other grasses in that one or occasionally two of the basal internodes of the stem swell into a bulb-like growth called a haplacorm. This characteristic is often used for identification of the plant during its early stages of growth (Ogle et al, 2011).

Prevention and Control

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Prevention and Control

Various measures for controlling invasive P. pratense have met with moderate success, although control in areas of conservation concern is difficult as most control methods negatively affect native vegetation (NatureServe, 2015). According to Tyser (1992) working in the Glacier National Park in Montana, reduction of timothy is not a realistic option there or other natural areas; the most reasonable recommendation for resource managers is not to use the species for revegetating disturbed sites.

Eradication

According to Weaver et al. (1990) working in the Northern Rocky Mountains of the USA, timothy control should include both elimination and simultaneous introduction of a desirable competitor to minimize reinvasion.

Cultural Control and Sanitary Measures

Timothy stands become weak under close and continuous grazing (Ogle, 2011).

Physical/Mechanical Control

Hand pulling can be effective for controlling timothy infestations (Klein, 2011), while frequent cutting or mowing can weaken overall plant health and vigour (Rutledge and McLendon, 1996).

Chemical Control

Glyphosate is used to control timothy and other perennial grass weeds in cereal crops with preharvest and stubble/fallow applications (Davies and Ballingall, 2008). Because of the herbicide’s effects on other plant species, however, this means of control would only be effective for timothy monocultures.

References

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Baskin CC; Baskin JM, 1998. Ecology of seed dormancy and germination in grasses. In: Population biology of grasses [ed. by Cheplick, G. P.]. Cambridge, UK: Cambridge University Press, 30-83.

Behrendt H; Tomczok J; Sliwa-Tomczok W; Kasche A; Ebner Eschenbach Cvon; Becker WM; Ring J, 1999. Timothy grass (Phleum pratense L.) pollen as allergen carriers and initiators of an allergic response. International Archives of Allergy and Immunology, 118(2-4):414-418.

Bor NL, 1960. The grasses of Burma, Ceylon, India and Pakistan (excluding Bambuseae). Oxford, UK: Pergamon Press, xviii + 767 pp.

Cenci CA; Pagiotti R, 1979. Evaluation by grazing of Phleum pratense L. and Hordeum bulbosum L. ecotypes. (Valutazione per il pascolo di ecotipi de Phleum pratense L. ed Hordeum bulbosum L.) Annali della Facolta di Agraria, Universita degli Studi di Perugia, 33:707-726.

Davies K; Ballingall M, 2008. Management of grass weeds in winter cereals. SAC Technical Note TN609, 12 pp.

Elliott CL; McKendrick JD; Helm D, 1987. Plant biomass, cover, and survival of species used for stripmine reclamation in south-central Alaska, U.S.A. Arctic and Alpine Research, 19(4):572-577.

Encyclopedia of Life, 2015. Phleum pratense, herd's grass. http://eol.org/pages/1114570/overview

Esser LL, 1993. Phleum pratense. Fire Effects Information System. Fort Collins, CO, USA: USDA Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory. http://www.feis-crs.org/beta/

Guo YangDong; Yli-Mattila T; Pulli S, 2003. Assessment of genetic variation in timothy (Phleum pratense L.) using RAPD and UP-PCR. Hereditas (Lund), 138(2):101-113.

Hoover MM; Hein MA; Dayton WA; Erlanson CO, 1948. The main grasses for farm and home. In: Grass, the Yearbook of Agriculture 1948 [ed. by Stefferud, A.]. Washington, DC, USA: US Department of Agriculture, 639-700. http://naldc.nal.usda.gov/naldc/download.xhtml?id=IND43895002&content=PDF

Houston DB, 1973. Wild fires in northern Yellowstone National Park. Ecology, 54(5):1112-1117.

Hubbard CE, 1968. Grasses. A guide to their structure, identification, uses, and distribution in the British Isles. Harmondsworth, UK: Penguin Books, 463 pp.

Invasive Species of Japan, 2015. Phleum pratense. Tsukuba, Japan: National Institute for Environmental Studies. https://www.nies.go.jp/biodiversity/invasive/DB/detail/80730e.html

invasive.org, 2010. Timothy, Phleum pratense L. http://www.invasive.org/browse/subinfo.cfm?sub=6179

Iowa State University, 2016. Timothy, Phleum pratense L. Grasses of Iowa, a project of Iowa State University. Ames, IA, USA: Iowa State University Department of Ecology, Evolution, and Organismal Biology. http://www.eeob.iastate.edu/research/IowaGrasses/speciespages/PhleuPrate/PhleuPrate.html

Jensen JW, 2005. Varieties of amenity grasses 2005. (Sorter af plænegræsser 2005.) Grøn Viden, Havebrug, No.165:15 pp.

Joachimiak A; Grabowska-Joachimiak A, 2000. Stomatal cell length and ploidy level in four taxa belonging to the Phleum sect. Phleum. Acta Biologica Cracoviensia. Series Botanica, 42(1):103-107.

Kadziulis L, 1974. Three aspects of comparative evaluation of legumes and grasses. Vaxtodling, 29:126-132.

Kew, 2015. An online resource for the world's plants. Kew, UK: Royal Botanic Gardens. http://wfo.kew.org/;jsessionid=4F750221A431BB0EC3431CAA07E7A925.kppapp02

Klein H, 2011. Timothy, Phleum pratense L. Alaska Natural Heritage Program. Anchorage, AK, USA: University of Alaska, 3 pp. http://aknhp.uaa.alaska.edu/wp-content/uploads/2013/01/Phleum_pratense_BIO_PHPR3.pdf

Lacefield GD; Henning JC; Phillips TD; Rasnake M, 2002. Timothy. University of Kentucky Cooperative Extension Service, AGR-84. Lexington, KY, USA: University of Kentucky College of Agriculture, 2 pp. http://www2.edu/agc/pubs/agr/agr84/AGR84.pdf

Larsen A; Marum P, 2006. Breeding goals and possibilities in future timothy breeding. In: NJF Seminar 384: Timothy productivity and forage quality - possibilities and limitations, 10-12 August 2006, Akureyri, Iceland [ed. by Sveinsson, T.]. Akureyri, Iceland: Agricultural University of Iceland, 31-39. http://www1.foragebeef.ca/$foragebeef/frgebeef.nsf/all/frg109/$FILE/Species%20Iceland%20Timothy%20Conference%20Proceedings.pdf

Munkvold GP; Carson T; Thoreson D, 1997. Outbreak of ergot (Claviceps purpurea) in Iowa barley, 1996. Plant Disease, 81(7):830.

NatureServe, 2015. Phleum pratense - L. NatureServe Explorer: an online encyclopedia of life. Version 7. Arlington, Virginia, USA: NatureServe. http://explorer.natureserve.org

Ogle DG, St. John L, Tilley DJ, 2011. Timothy, Phleum pratense L. USDA-NRCS Plant Guide. Washington, DC, USA: USDA-NRCS, 4 pp. http://plants.usda.gov/plantguide/pdf/pg_phpr3.pdf

Peeters A, 2004. Wild and sown grasses. Profiles of a temperate species selection: ecology, biodiversity and use [ed. by Peeters, A.]. Rome, Italy: Food and Agriculture Organization of the United Nations (FAO), ix + 311 pp.

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

Queensland Government, 2015. Timothy grass, Phleum pratense. Weeds of Australia, Biosecurity Queensland edition. http://keyserver.lucidcentral.org/weeds/data/03030800-0b07-490a-8d04-0605030c0f01/media/Html/Phleum_pratense.htm

Rutledge CR; McLendon T, 1996. An assessment of exotic plant species of Rocky Mountain National Park: an evaluation of the potential impacts of known exotic plants and summary of management options for species of concern. Fort Collins, CO, USA: Colorado State University, 97 pp.

Saigusa T; Hojito M; Noshiro M, 1997. Effects of zinc and copper application on timothy (Phleum pratense L.) growth on andisol grasslands in Konsen district. In: Ando T, Fujita K, Mae T, Matsumoto H, Mori S, Sekiya J, eds. Plant nutrition for sustainable food production and environment. Proceedings of the XIII International Plant Nutrition Colloquium, 13-19 September 1997, Tokyo, Japan. Dordrecht, Netherlands: Kluwer Academic Publishers, 951-952.

Shukla U, 1996. The grasses of north-eastern India. Jodhpur, India: Scientific Publishers, 404 pp.

Steinman H, 2012. Timothy grass. Uppsala, Sweden: Thermo Fisher Scientific Inc.

Thakur MP; Reich PB; Eddy WC; Stefanski A; Rich R; Hobbie SE; Eisenhauer N, 2014. Some plants like it warmer: increased growth of three selected invasive plant species in soils with a history of experimental warming. Pedobiologia, 57(1):57-60. http://www.sciencedirect.com/science/journal/00314056

The Plant List, 2013. The Plant List: a working list of all plant species. Version 1.1. London, UK: Royal Botanic Gardens, Kew. http://www.theplantlist.org

Tyser RW, 1992. Vegetation associated with two alien plant species in a fescue grassland in Glacier National Park, Montana. Great Basin Naturalist, 52(2):189-193.

USDA Forest Service, 1937. Range plant handbook. Washington, DC, USA: USDA Forest Service.

USDA-ARS, 2015. Germplasm Resources Information Network (GRIN). Online Database. Beltsville, Maryland, USA: National Germplasm Resources Laboratory. https://npgsweb.ars-grin.gov/gringlobal/taxon/taxonomysearch.aspx

USDA-NRCS, 2015. The PLANTS Database. Baton Rouge, USA: National Plant Data Center. http://plants.usda.gov/

Varvas T; Kasekamp K; Kullman B, 2013. Preliminary study of endophytic fungi in timothy (Phleum pratense) in Estonia. Acta Mycologica, 48(1):41-49. https://pbsociety.org.pl/journals/index.php/am/article/view/am.2013.006/1603

Wallace LL; Macko SA, 1993. Nutrient acquisition by clipped plants as a measure of competitive success: the effects of compensation. Functional Ecology, 7(3):326-331.

Weaver T; Lichthart J; Gustafson D, 1990. Exotic invasion of timberline vegetation, Northern Rocky Mountains, USA. In: Proceedings - symposium on whitebark pine ecosystems: ecology and management of a high-mountain resource, 29-31 March 1989, Bozeman, MT, USA [ed. by Schmidt, W. C. \McDonald, K. J.]. Ogden, UT, USA: USDA Forest Service, Intermountain Research Station, 208-213. [USDA Forest Service General Technical Report INT-270.]

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.

Contributors

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08/09/2015    Original text by:

Daiva Kasulyte and Andrew Praciak, CABI, UK 

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