Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide

Datasheet

Brachypodium sylvaticum
(slender false brome)

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Datasheet

Brachypodium sylvaticum (slender false brome)

Summary

  • Last modified
  • 20 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Preferred Scientific Name
  • Brachypodium sylvaticum
  • Preferred Common Name
  • slender false brome
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Plantae
  •     Phylum: Spermatophyta
  •       Subphylum: Angiospermae
  •         Class: Monocotyledonae
  • Summary of Invasiveness
  • B. sylvaticum is a bunchgrass naturally occurring in old world temperate forests and temperate zones of tropical Asian mountains. Its extensive native range includes most of Eurasia (e.g. Europe, Russia, China, J...

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Pictures

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PictureTitleCaptionCopyright
Brachypodium sylvaticum habit. Note that the grass is bright green, despite the fact that this photograph was taken mid-August (2009) and it last rained substantially in June 2009.  Also, the leaf blades are wide and lax, and the racemes are nodding.  The species is a bunchgrass, but in the invaded range there are so many seedlings and vegetative tillers that it can look like an unmown lawn, as in this picture.
TitleGrowth habit
CaptionBrachypodium sylvaticum habit. Note that the grass is bright green, despite the fact that this photograph was taken mid-August (2009) and it last rained substantially in June 2009. Also, the leaf blades are wide and lax, and the racemes are nodding. The species is a bunchgrass, but in the invaded range there are so many seedlings and vegetative tillers that it can look like an unmown lawn, as in this picture.
CopyrightBitty A. Roy
Brachypodium sylvaticum habit. Note that the grass is bright green, despite the fact that this photograph was taken mid-August (2009) and it last rained substantially in June 2009.  Also, the leaf blades are wide and lax, and the racemes are nodding.  The species is a bunchgrass, but in the invaded range there are so many seedlings and vegetative tillers that it can look like an unmown lawn, as in this picture.
Growth habitBrachypodium sylvaticum habit. Note that the grass is bright green, despite the fact that this photograph was taken mid-August (2009) and it last rained substantially in June 2009. Also, the leaf blades are wide and lax, and the racemes are nodding. The species is a bunchgrass, but in the invaded range there are so many seedlings and vegetative tillers that it can look like an unmown lawn, as in this picture.Bitty A. Roy
Brachypodium sylvaticum ligule and pubescence. Note that the leaf sheath is open, the ligule is membranous and ciliate and that leaf blades and internodes can be quite pubescent.
TitleLigule and pubescence
CaptionBrachypodium sylvaticum ligule and pubescence. Note that the leaf sheath is open, the ligule is membranous and ciliate and that leaf blades and internodes can be quite pubescent.
CopyrightBitty A. Roy
Brachypodium sylvaticum ligule and pubescence. Note that the leaf sheath is open, the ligule is membranous and ciliate and that leaf blades and internodes can be quite pubescent.
Ligule and pubescenceBrachypodium sylvaticum ligule and pubescence. Note that the leaf sheath is open, the ligule is membranous and ciliate and that leaf blades and internodes can be quite pubescent.Bitty A. Roy
Brachypodium sylvaticum spikelets. Note that the lemma awns are as long or longer than the lemmas (7-15mm long).
TitleSpikelets
CaptionBrachypodium sylvaticum spikelets. Note that the lemma awns are as long or longer than the lemmas (7-15mm long).
CopyrightBitty A. Roy
Brachypodium sylvaticum spikelets. Note that the lemma awns are as long or longer than the lemmas (7-15mm long).
SpikeletsBrachypodium sylvaticum spikelets. Note that the lemma awns are as long or longer than the lemmas (7-15mm long).Bitty A. Roy
Brachypodium sylvaticum infected by Epichloë sylvatica in Switzerland. The white patches on the stems of this false-brome grass are the symptoms of 'choke' disease caused by Epichloë sylvatica.  Infected stems will not flower, but instead form these patches where the fungus is reproducing.  Note that the leaves are broad, soft and bright green.
TitleNatural enemy
CaptionBrachypodium sylvaticum infected by Epichloë sylvatica in Switzerland. The white patches on the stems of this false-brome grass are the symptoms of 'choke' disease caused by Epichloë sylvatica. Infected stems will not flower, but instead form these patches where the fungus is reproducing. Note that the leaves are broad, soft and bright green.
CopyrightBitty A. Roy
Brachypodium sylvaticum infected by Epichloë sylvatica in Switzerland. The white patches on the stems of this false-brome grass are the symptoms of 'choke' disease caused by Epichloë sylvatica.  Infected stems will not flower, but instead form these patches where the fungus is reproducing.  Note that the leaves are broad, soft and bright green.
Natural enemyBrachypodium sylvaticum infected by Epichloë sylvatica in Switzerland. The white patches on the stems of this false-brome grass are the symptoms of 'choke' disease caused by Epichloë sylvatica. Infected stems will not flower, but instead form these patches where the fungus is reproducing. Note that the leaves are broad, soft and bright green.Bitty A. Roy
Brachypodium sylvaticum infected by a rust fungus in Switzerland. Leaf showing rust infection caused by Puccinia brachypodii var. brachypodii
TitleNatural enemy
CaptionBrachypodium sylvaticum infected by a rust fungus in Switzerland. Leaf showing rust infection caused by Puccinia brachypodii var. brachypodii
CopyrightBitty A. Roy
Brachypodium sylvaticum infected by a rust fungus in Switzerland. Leaf showing rust infection caused by Puccinia brachypodii var. brachypodii
Natural enemyBrachypodium sylvaticum infected by a rust fungus in Switzerland. Leaf showing rust infection caused by Puccinia brachypodii var. brachypodiiBitty A. Roy
Photomicrograph of Puccinia brachypodii var. brachypodii. This rust fungus is found on Brachypodium sylvaticum.  The round yellowish objects [arrowed] are urediniospores and the club-shaped objects are paraphyses. (Note scale bar)
TitleUrediniospores and paraphyses
CaptionPhotomicrograph of Puccinia brachypodii var. brachypodii. This rust fungus is found on Brachypodium sylvaticum. The round yellowish objects [arrowed] are urediniospores and the club-shaped objects are paraphyses. (Note scale bar)
CopyrightBitty A. Roy
Photomicrograph of Puccinia brachypodii var. brachypodii. This rust fungus is found on Brachypodium sylvaticum.  The round yellowish objects [arrowed] are urediniospores and the club-shaped objects are paraphyses. (Note scale bar)
Urediniospores and paraphysesPhotomicrograph of Puccinia brachypodii var. brachypodii. This rust fungus is found on Brachypodium sylvaticum. The round yellowish objects [arrowed] are urediniospores and the club-shaped objects are paraphyses. (Note scale bar)Bitty A. Roy
Photomicrograph of Puccinia brachypodii var. brachypodii. This rust fungus is found on Brachypodium sylvaticum.  The round yellowish objects [arrowed] are urediniospores and the club-shaped objects are paraphyses. (Note scale bar)
TitleUrediniospores and paraphyses
CaptionPhotomicrograph of Puccinia brachypodii var. brachypodii. This rust fungus is found on Brachypodium sylvaticum. The round yellowish objects [arrowed] are urediniospores and the club-shaped objects are paraphyses. (Note scale bar)
CopyrightBitty A. Roy
Photomicrograph of Puccinia brachypodii var. brachypodii. This rust fungus is found on Brachypodium sylvaticum.  The round yellowish objects [arrowed] are urediniospores and the club-shaped objects are paraphyses. (Note scale bar)
Urediniospores and paraphysesPhotomicrograph of Puccinia brachypodii var. brachypodii. This rust fungus is found on Brachypodium sylvaticum. The round yellowish objects [arrowed] are urediniospores and the club-shaped objects are paraphyses. (Note scale bar)Bitty A. Roy

Identity

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Preferred Scientific Name

  • Brachypodium sylvaticum (Huds.) P. Beauv. 1812

Preferred Common Name

  • slender false brome

Other Scientific Names

  • Agropyron sylvaticum (Huds.) Chevall.
  • Brevipodium sylvaticum (Huds.) Á. Löve & D. Löve
  • Bromus sylvaticus (Huds.) Lyons
  • Festuca sylvatica Huds. 1762
  • Triticum sylvaticum (Huds.) Moench

International Common Names

  • English: false wood brome (UK); perennial false brome; slender false broome
  • French: brachypode des bois; brachypode sylvestre

Local Common Names

  • Germany: Zwenke, Wald-
  • Italy: brachipodio selvatico
  • Japan: ezoyamakamojigusa
  • Korea, DPR: sup-gae-mil
  • Netherlands: boskortsteel
  • Sweden: lundlosta

EPPO code

  • BRCSI (Brachypodium sylvaticum)

Summary of Invasiveness

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B. sylvaticum is a bunchgrass naturally occurring in old world temperate forests and temperate zones of tropical Asian mountains. Its extensive native range includes most of Eurasia (e.g. Europe, Russia, China, Japan, India, Indonesia) as well as the Middle East (e.g. Lebanon, Syria, Iran) and North Africa (e.g. Algeria, Eritrea). It is invasive in North America (Piep, 2003), South America (Zuloaga et al., 1994), New Zealand (Edgar and Connor, 2000), and Australia (IBIS, 2009). It is shade tolerant (Murchie and Horton, 1998), spreads rapidly by seeds (Petersen and Philipp, 2001), has a persistent seed bank (Donelan and Thompson, 1980; Buckley et al., 1997) and is long-lived (Haeggström and Skytén, 1996). It forms monocultures and crowds out native plants and rare butterflies (Kaye and Blakeley-Smith, 2006; Severns and Warren, 2008). Furthermore, grasses significantly reduce recruitment of conifers (Powell et al., 1994; Lehmkuhl, 2002; Kruse et al., 2004). It is on noxious weed lists for three USA States: California, Oregon and Washington (CDFA, 2009; NWCB, 2009; ODA, 2009).

Taxonomic Tree

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

Notes on Taxonomy and Nomenclature

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Hudson originally described B. sylvaticum, commonly known as false brome or slender false brome, as Festuca sylvatica in 1762 (Flora Anglica 1: 38). It was later transferred to Bromus sylvaticus (Huds.) by Lyons in 1763 and then by Palisot de Beauvois (P. Beauv.) to Brachypodium in 1812 (Essai d'une Nouvelle Agrostographie 101, 155, pl. 3, f. 11). B. sylvaticum is closely related to Brachypodium pinnatum (Catalán and Olmstead 2000), with which it sometimes hybridizes (Khan and Stace, 1999).

Thus far, interspecific hybrids between B. sylvaticum and B. pinnatum have not been reported from the invaded range in the USA. However, there is extensive intraspecific hybridization in the USA among ecotypes of B. sylvaticum naturalized from different regions in Europe (Rosenthal et al., 2008).

Numerous varieties of B. sylvaticum have been described, and there is great need for a revision of the genus and of this species in particular (Khan and Stace, 1999; Piep, 2003; Shouliang and Phillips, 2006). There has never been a concerted study of specimens from across its extensive natural range and it is not known, for example, how similar the Asian and European plants actually are (Shouliang and Phillips, 2006). Furthermore, the named varieties are often based on pubescence traits, which can vary within single populations (Paszko, 2008). Pubescence can vary both as a result of the environment (Roy et al., 1999; Shouliang and Phillips, 2006) and genotype (Davies and Long, 1991; Paszko, 2008).

Description

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B. sylvaticum is a cespitose perennial bunchgrass that is sometimes very weakly rhizomatous. It ranges in height from a few centimeters up to about 200 cm. Sheaths are open and the nodes are typically pubescent. The leaf blades are bright green and remain green throughout the summer, even in dry Mediterranean climates (BA Roy, University of Oregon, USA, personal observation, 2009). The blades are 4-15 mm wide, flat and lax, with variable pubescence. Ligules are variable in size (1-6 mm) and are generally pubescent and ciliate. Plant size and pubescence depend both on habitat (Shouliang and Phillips, 2006) and genotype (Davies and Long, 1991). Racemes are nodding with an average of 9 spikelets, each with 3-24 florets. Lemma awns are 7-15 mm. Excellent full descriptions are published in the Flora of North America (Piep, 2003) and Flora of China (Shouliang and Phillips, 2006), both of which can also be found on line at www.efloras.org/index.aspx.

Distribution

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B. sylvaticum is native to the Old World. To date it is known to be invading five states in the USA, New Zealand and small areas in Argentina and Australia. The sources of the invasions in the Pacific Northwest of the USA are European (Rosenthal et al., 2008).

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

AzerbaijanPresentNative Not invasive Euro+Med Plantbase, 2009
BhutanWidespreadNative Not invasive Shouliang and Phillips, 2006Mountain slopes, forest understory
ChinaWidespreadNative Not invasive Shouliang and Phillips, 2006Mountain slopes, forest understory
-AnhuiWidespreadNative Not invasive Shouliang and Phillips, 2006Mountain slopes, forest understory
-GansuWidespreadNative Not invasive Shouliang and Phillips, 2006Mountain slopes, forest understory
-GuizhouWidespreadNative Not invasive Shouliang and Phillips, 2006Mountain slopes, forest understory
-HunanWidespreadNative Not invasive Shouliang and Phillips, 2006Mountain slopes, forest understory
-JiangsuWidespreadNative Not invasive Shouliang and Phillips, 2006Mountain slopes, forest understory
-LiaoningWidespreadNative Not invasive Shouliang and Phillips, 2006Mountain slopes, forest understory
-QinghaiWidespreadNative Not invasive Shouliang and Phillips, 2006Mountain slopes, forest understory
-ShanxiWidespreadNative Not invasive Shouliang and Phillips, 2006Mountain slopes, forest understory
-SichuanWidespreadNative Not invasive Shouliang and Phillips, 2006Mountain slopes, forest understory
-TibetWidespreadNative Not invasive Shouliang and Phillips, 2006Mountain slopes, forest understory
-XinjiangWidespreadNative Not invasive Shouliang and Phillips, 2006Mountain slopes, forest understory
-YunnanWidespreadNative Not invasive Shouliang and Phillips, 2006Mountain slopes, forest understory
-ZhejiangWidespreadNative Not invasive Shouliang and Phillips, 2006Mountain slopes, forest understory
IndiaWidespreadNative Not invasive Shouliang and Phillips, 2006Northern India
-AssamPresentNative Not invasive Veldkamp and Vanscheindelen, 1989
-Indian PunjabPresentNative Not invasive Bor and Guest, 1968
-SikkimPresentNative Not invasive Bor and Guest, 1968
-Tamil NaduPresentNative Not invasive Veldkamp and Vanscheindelen, 1989
IndonesiaWidespreadNative Not invasive Shouliang and Phillips, 2006Mountain slopes, forest understory
-Irian JayaPresentNative Not invasive Veldkamp and Vanscheindelen, 19891500-3830 m
-JavaPresentNative Not invasive Hara, 1966Glabrous form called B. sylvaticum var. luzoniense
-MoluccasPresentNative Not invasive Veldkamp and Vanscheindelen, 19891500-3830 m
-Nusa TenggaraPresentNative Not invasive Veldkamp and Vanscheindelen, 19891500-3830 m
-SulawesiPresent, few occurrencesNative Not invasive Veldkamp and Vanscheindelen, 19891500-3830 m
-SumatraPresentNative Not invasive Hara, 1966Glabrous form called B. sylvaticum var. luzoniense
IranPresentNative Not invasive Bor, 1970Shady woods, streamside, 300-2300 m
IraqPresentNative Not invasive Bor, 1970Damp, shady places in the mountains, 700-1500 m
IsraelPresentNative Not invasive Danin, 2000Originally listed incorrectly as B. pinnatum
JapanPresentNative Not invasive Oi, 1965Woods, common
-HokkaidoPresentNative Not invasive Oi, 1965Woods, common
-HonshuPresentNative Not invasive Oi, 1965Woods, common
-KyushuPresentNative Not invasive Oi, 1965Woods, common
-ShikokuPresentNative Not invasive Oi, 1965Woods, common
JordanPresentAl-Eisawi and Al-Khader, 1998
KazakhstanPresentNative Not invasive Rosenthal et al., 2008
Korea, Republic ofPresentKPNI, 2009
KyrgyzstanPresentNative Not invasive Shouliang and Phillips, 2006Mountain slopes, forest understory
LebanonPresentNative Not invasive Euro+Med Plantbase, 2009
NepalPresentNative Not invasive Shouliang and Phillips, 2006Mountain slopes, forest understory
PakistanPresentNative Not invasive Bor, 1970Shady woods, streamside, 900-2100 m
PhilippinesPresentNative Not invasive Hara, 1966Glabrous form called B. sylvaticum var. luzoniense
Saudi ArabiaPresentNativeScholz and König, 1985
Sri LankaPresentSoreng et al., 2009Horton plains, 7000 feet
SyriaPresentNative Not invasive Euro+Med Plantbase, 2009
TaiwanPresentNative Not invasive Shouliang and Phillips, 2006Mountain slopes, forest understory
TajikistanPresentNative Not invasive Shouliang and Phillips, 2006Mountain slopes, forest understory
TurkeyWidespreadNative Not invasive Davis, 198810-2440 m forested slopes, meadows, river terraces
TurkmenistanPresentNative Not invasive Bor, 1970Shady woods, streamside
United Arab EmiratesPresentNative Not invasive Bor, 1970Bahrain
UzbekistanWidespreadNative Not invasive Shouliang and Phillips, 2006Mountain slopes, forest understory

Africa

AlgeriaPresentNative Not invasive Euro+Med Plantbase, 2009
EritreaPresentSoreng et al., 20092500 m
MoroccoPresentNative Not invasive Euro+Med Plantbase, 2009
Spain
-Canary IslandsPresentNative Not invasive Euro+Med Plantbase, 2009
TunisiaPresentNative Not invasive Euro+Med Plantbase, 2009

North America

CanadaPresentPresent based on regional distribution.
-AlbertaAbsent, intercepted onlyBarkworth et al., 2007
-British ColumbiaAbsent, intercepted onlyBarkworth et al., 2007
-Newfoundland and LabradorAbsent, intercepted onlyBarkworth et al., 2007
-OntarioPresentIntroducedMiller et al., 2011
USAPresent, few occurrencesEPPO, 2014
-CaliforniaLocalised2009Introduced2003 Invasive Amme, 2006Found in several locations in San Mateo County, including near Stanford and in the Santa Cruz Mountains
-MissouriLocalisedIntroducedSoreng et al., 2009There are two specimens at Missouri Botanic Gardens from Missouri, one from St Louis City (1977) and one without date or locality
-OregonWidespread2009Introduced< 1939Chambers, 1966; EPPO, 2014Thoroughly naturalized in Oregon's Willamette Valley by 1966, now common throughout Western Oregon from Portland to Rogue River Valley and east of Cascades on rivers (e.g. Metolius, John Day)
-UtahAbsent, formerly present Invasive Barkworth et al., 2007Planted in the 1970's, but not found since then
-VirginiaLocalised2009Introduced1992 Invasive Virginia Botanical Associates, 2009First collected by C. E. Stevens, Aug. 1992 at Preston Place, Charlottesville. Has persisted and slowly spread
-WashingtonLocalised2009Introduced2007 Invasive False Brome Working Group, 2009The first sighting in Washington State was in 2007 from Beacon Rock State Park, Skamania County, in the Columbia River Gorge

Central America and Caribbean

BarbadosAbsent, intercepted onlyGooding et al., 1965
Costa RicaAbsent, intercepted onlyHammel et al., 2003
NicaraguaAbsent, intercepted onlyStevens et al., 2001

South America

ArgentinaPresentIntroduced Invasive Zuloaga et al., 1994
EcuadorAbsent, intercepted onlyJørgensen and León-Yánez, 1999

Europe

AlbaniaPresentNative Not invasive Euro+Med Plantbase, 2009
AndorraLocalisedNative Not invasive Ninot et al., 2000Submontane-montane, shaded, mild clearings in the domain of Pinus sylvestris forests, irregularly mown or grazed
AustriaPresentNative Not invasive GBIF, 2009
BelarusPresentCBG NASB MSKH, 2009
BelgiumPresentNative Not invasive Euro+Med Plantbase, 2009
BulgariaPresentNative Not invasive GBIF, 2009
CroatiaPresentNative Not invasive GBIF, 2009
CyprusPresentNative Not invasive Euro+Med Plantbase, 2009
Czech RepublicPresentNative Not invasive GBIF, 2009
DenmarkPresentNative Not invasive Euro+Med Plantbase, 2009
EstoniaWidespreadNative Not invasive Palo et al., 2008
FinlandPresentNative Not invasive GBIF, 2009
FrancePresentNative Not invasive GBIF, 2009
-CorsicaPresentNative Not invasive Euro+Med Plantbase, 2009
GermanyPresentNative Not invasive GBIF, 2009
GreecePresentNative Not invasive GBIF, 2009
HungaryPresentNative Not invasive GBIF, 2009
IrelandPresentNative Not invasive GBIF, 2009
ItalyPresentNative Not invasive GBIF, 2009
LithuaniaLocalisedNative Not invasive Marozas, 2004
MaltaPresentNative Not invasive Euro+Med Plantbase, 2009
MoldovaPresentNative Not invasive Euro+Med Plantbase, 2009
NetherlandsPresentNative Not invasive GBIF, 2009
NorwayPresentNative Not invasive Euro+Med Plantbase, 2009
PolandPresentNative Not invasive GBIF, 2009
PortugalPresentNative Not invasive Euro+Med Plantbase, 2009
-AzoresPresentNative Not invasive Euro+Med Plantbase, 2009
-MadeiraPresentNative Not invasive Euro+Med Plantbase, 2009
RomaniaPresentNative Not invasive GBIF, 2009
Russian FederationPresentNative Not invasive Czerepanov, 1995Lists many varieties
-Central RussiaPresentNative Not invasive Czerepanov, 1995Lists many varieties
-Eastern SiberiaPresentNative Not invasive Czerepanov, 1995Lists many varieties
-Northern RussiaPresentNative Not invasive Czerepanov, 1995Lists many varieties
-Russian Far EastPresentNative Not invasive Czerepanov, 1995Lists many varieties
-Southern RussiaPresentNative Not invasive Czerepanov, 1995Lists many varieties
-Western SiberiaPresentNative Not invasive Czerepanov, 1995Lists many varieties
SerbiaPresentNative Not invasive Euro+Med Plantbase, 2009
SlovakiaPresentNative Not invasive Euro+Med Plantbase, 2009
SloveniaPresentNative Not invasive GBIF, 2009
SpainPresentNative Not invasive GBIF, 2009
-Balearic IslandsPresentNative Not invasive Euro+Med Plantbase, 2009
SwedenPresentNative Not invasive GBIF, 2009
SwitzerlandWidespreadNative Not invasive Lauber and Wagner, 1996Montane to subalpine
UKWidespreadNative Not invasive Stace, 1997Native; woods, scrub and shady wood-borders and hedgerows, in open grassland mainly in the north; common throughout the British Isles except much of northern Scotland
-Channel IslandsPresentNative Not invasive Euro+Med Plantbase, 2009
UkrainePresentNative Not invasive Czerepanov, 1995Lists many varieties
Yugoslavia (Serbia and Montenegro)PresentNative Not invasive Euro+Med Plantbase, 2009

Oceania

AustraliaPresentPresent based on regional distribution.
-Australian Northern TerritoryAbsent, never occurredIBIS, 2009
-VictoriaLocalised Invasive IBIS, 2009Sparingly naturalized
New ZealandWidespreadIntroduced Invasive Edgar and Connor, 2000Lowland to montane in shade of trees or shrubs
Papua New GuineaPresentNative Not invasive Hara, 1966Glabrous form called B. sylvaticum var. luzoniense

History of Introduction and Spread

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No information was found concerning the initial invasions in New Zealand or Australia. For Argentina, AA Beetle collected the oldest dated specimen in the Missouri Botanical Garden herbarium on February 21, 1952 in Buenos Aires (Soreng et al., 2009).

B. sylvaticum was first found naturalized in North America near Eugene, Oregon in 1939 and was thoroughly naturalized in Oregon’s Willamette Valley by 1966 (Chambers, 1966). There are two different hypotheses for the source of the initial invasion in Oregon. Hitchcock (1950) said that it is sometimes cultivated as an ornamental, which led some later authors to say that it escaped from gardens (e.g. Severns and Warren, 2008). However, recent genetic work suggests that the original invasion in Oregon resulted from escape from the United States Department of Agriculture (USDA) seed trials near Corvallis and Eugene, Oregon (Rosenthal et al., 2008). The genetic data is consistent with records showing that B. sylvaticum was one of a suite of non-native grasses that were actively planted around the Western USA by USDA scientists, who were seeking to improve rangelands (Hull, 1974). The invasion is extensive in western Oregon (Kaye and Blakeley-Smith, 2006; False Brome Working Group, 2009), where it is listed as a noxious weed.

In California, USA, B. sylvaticum has thus far been found in small, localized populations (Amme, 2006). The first Jepson Herbarium records are from 2003. It has been found in several locations in San Mateo County. Including near Stanford and in the Santa Cruz Mountains (Jepson Herbarium, 2009). In California it seems to grow best around lake margins, and in clearings in redwood forest, although it has also been found in dense shaded forest (Hrusa, 2003). B. sylvaticum is listed as a noxious weed in California (CDFA, 2009).

The first sighting in Washington State, USA was in 2007 from Beacon Rock State Park, Skamania County, in the Columbia River Gorge (False Brome Working Group, 2009). As of 2009, it was listed as a noxious weed in Washington (NWCB, 2009).

Introductions

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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
Argentina < 1952 Yes No Soreng et al. (2009)
Australia   Yes No IBIS (2009)
New Zealand   Yes No Edgar and Connor (2000)
USA Europe <1939 Forage (pathway cause) ,
Horticulture (pathway cause)
Yes No Chambers (1966)

Risk of Introduction

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B. sylvaticum is a quarantined weed in three states in the USA: Washington (NWCB, 2009), Oregon (ODA, 2009), and California (CDFA, 2009). There is high risk of further spread because there are multiple pathways for it to do so (see the text section 'Means of Movement and Dispersal'). It can be carried by animals and humans, on machinery, by rivers and streams, it may also spread as a result of increasing popularity as a model organism, through intentional introductions and the horticultural trade.

Habitat

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Forests

- Temperate conifer forests (e.g. Scots pine, Pinussylvestris: Ninot et al., 2000; coast redwood, Sequoia sempervirens:Hrusa, 2003; and Douglas fir, Pseudotsugamenziesii:False Brome Working Group, 2009). Found in both disturbed and undisturbed forests, and under both open and closed canopies (Hrusa, 2003; Parks et al., 2005; BA Roy, University of Oregon, USA, personal communication, 2009).

- Temperate deciduous forests (including Quercus, Fagus, Fraxinus), in both forest and open areas (Szujkó-Lacza and Fekete, 1974; Petersen and Philipp, 2001; Safaian et al., 2005; Chaideftou et al., 2009; BA Roy, University of Oregon, USA, personal observation, 2009).

- Mediterranean forests (Safaian et al., 2005; Chaideftou et al., 2009).

- Boreal forests (Holten, 1980; Haeggström and Skytén, 1996; Aarrestad, 2000; Petersen and Philipp, 2001).

Meadows/Openings

- Stream edges and lakesides, including in arid regions (Hrusa, 2003; False Brome Working Group, 2009; BA Roy, University of Oregon, USA, personal communication, 2009: in Europe).

- High altitude meadows such as the Tibetan steppe and the tops of tropical mountains (Smith, 1975; Roder et al., 2007; Singh et al., 2008).

- Temperate meadows (Davies and Long, 1991; Stace, 1997; US Fish and Wildlife Service, 2006; Severns and Warren, 2008).

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Terrestrial
 
Terrestrial – ManagedManaged forests, plantations and orchards Principal habitat Harmful (pest or invasive)
Managed forests, plantations and orchards Principal habitat Natural
Managed grasslands (grazing systems) Principal habitat Harmful (pest or invasive)
Managed grasslands (grazing systems) Principal habitat Natural
Disturbed areas Secondary/tolerated habitat Harmful (pest or invasive)
Disturbed areas Secondary/tolerated habitat Natural
Rail / roadsides Principal habitat Harmful (pest or invasive)
Rail / roadsides Principal habitat Natural
Urban / peri-urban areas Secondary/tolerated habitat Harmful (pest or invasive)
Urban / peri-urban areas Secondary/tolerated habitat Natural
Terrestrial ‑ Natural / Semi-naturalNatural forests Principal habitat Harmful (pest or invasive)
Natural forests Principal habitat Natural
Natural grasslands Secondary/tolerated habitat Harmful (pest or invasive)
Natural grasslands Secondary/tolerated habitat Natural
Riverbanks Principal habitat Harmful (pest or invasive)
Riverbanks Principal habitat Natural
Wetlands Principal habitat Harmful (pest or invasive)
Wetlands Principal habitat Natural
Cold lands / tundra Present, no further details
Freshwater
Irrigation channels Principal habitat Harmful (pest or invasive)
Irrigation channels Principal habitat Natural
Lakes Principal habitat Harmful (pest or invasive)
Lakes Principal habitat Natural
Reservoirs Principal habitat Harmful (pest or invasive)
Reservoirs Principal habitat Natural
Rivers / streams Principal habitat Harmful (pest or invasive)
Rivers / streams Principal habitat Natural
Ponds Principal habitat Harmful (pest or invasive)
Ponds Principal habitat Natural

Hosts/Species Affected

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The Pacific Northwest is world renowned for its timber production from conifers. It has not been established through a controlled study that B. sylvaticum in particular competes with conifers; however, several lines of evidence suggest that this grass will reduce survival and growth of conifer seedlings. First, a number of studies have established that the germination, growth and survival of conifers are negatively affected by competition with grasses (e.g. Powell et al., 1994; Lehmkuhl, 2002; Kruse et al., 2004). Second, at least one timber company (Starker Forests Inc.) has noticed that dense patches of false-brome provide safe cover for voles, which girdle conifer seedlings (G Fitzpatrick, The Nature Conservancy, Oregon, USA, personal communication, 2009). Third, none of the native grasses affected form a solid carpet in the forest, whereas B. sylvaticum does. These dense carpets will compete with seedlings in both logged and unlogged forests, and the build up of thatch may increase fire risk (Anzinger and Radosevich, 2008; False Brome Working Group, 2009). On the other hand, because B. sylvaticum remains green throughout the summer it may decrease fire risk (Anzinger and Radosevich, 2008; False Brome Working Group, 2009). These questions about fire need to be addressed with further research because this grass has the potential to cause ecosystem change if it alters fire behaviour.

The native grasses and herbs that live in the habitats being invaded are likely to diminish in cover and may face local extinction as a result of competition. A recent study showed that under shady high nutrient conditions, B. sylvaticum is a superior competitor to a native prairie grass (Festuca roemeri), a native forest grass (Elymus glaucus) as well as to another aggressive invasive Schedonorus arundinaceus (previously known as Festuca arundinacea) (BA Roy, University of Oregon, USA, personal observation, 2009). Two other grasses that occur sporadically in the forest (Melica subulata and Bromus carinatus) are also quite likely to be negatively affected. In its native range, B. sylvaticum is known for its ability to out compete other species due to its rapid relative growth rate (RGR) and ability to form persistent leaf litter (Grime et al., 1988; Haeggström and Skytén, 1996; Alonso et al., 2001).

Growth Stages

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

Biology and Ecology

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Genetics

B. sylvaticum is a perennial, self-compatible, wind-pollinated grass that can also outcross (Khan and Stace, 1999; Rosenthal et al., 2008). It is a diploid, but variable in chromosome number, typically 2n=14, 16 or 18 (Veldkamp and Vanscheindelen, 1989; Khan and Stace, 1999; Piep, 2003). It is known to form interspecific hybrids with Brachypodium pinnatum (Khan and Stace, 1999), but has not done so in the invaded range (Rosenthal et al., 2008). However, it has formed intraspecific hybrids among individuals from different European origins (Rosenthal et al., 2008).

Plants in the genus Brachypodium are rapidly becoming model organisms because of their close relationship with wheat, and to a lesser extent rice, and because they have a much smaller genome size than these grains (Opanowicz et al., 2008). Most of the research activity is focusing on the annual Brachypodium distachyon (Garvin, 2007; Opanowicz et al., 2008; Bakker et al., 2009), but B. sylvaticum is also receiving significant attention (e.g. Foote et al., 2004; Wolny and Hasterok, 2007; Faris et al., 2008).

Reproductive Biology

B. sylvaticum is a perennial species that typically flowers late in the season relative to other temperate grasses. For example, in the Willamette Valley of Oregon, USA all the other grasses flower in May and June, whereas B. sylvaticum does not flower until early July. It is also late flowering in the native range of Europe (BA Roy, University of Oregon, USA, personal observation, 2009). B. sylvaticum produces seeds from both sexually produced tillers (that germinate from seeds) and vegetative tillers (asexual). A recent demographic study monitored population growth rates at four sites in the native range (Switzerland) and four sites in the invaded range (Oregon) (BA Roy, University of Oregon, USA, personal observation, 2009). In both ranges, tillers from seeds generally flowered 1 year post germination, whereas vegetative tillers could flower in the year produced, provided they were produced early in the year (BA Roy, University of Oregon, USA, personal observation, 2009). Population growth rates were significantly higher in the invaded range than the native range because asexual tillering, seed production from all tillers (sexual and asexual) and seed germination were all higher in the introduced range.

Physiology and Phenology

There is considerable variation among localities in numerous traits in B. sylvaticum, from morphology to phenology, physiology and pathogen resistance. Some of this variation is the result of phenotypic plasticity related to climate or other abiotic conditions, and some is related to genetic differences among populations or ranges (Paszko, 2008).

In colder climates or at higher elevations, B. sylvaticum dies back to the ground and only begins to grow after snowmelt, which can vary depending on year (BA Roy, University of Oregon, USA, personal observation, 2009). Under more favourable conditions, such as in oak forests in Hungary (Szujkó-Lacza and Fekete, 1974) and Oregon (BA Roy, University of Oregon, USA, personal observation, 2009), some shoots are able to continue photosynthesizing even in the middle of winter. B. sylvaticum is capable of germinating and growing in soils drier than many species can tolerate, perhaps because they have relatively large seeds (Evans and Etherington, 1991). Seed production, on the other hand, is strongly dependent on climate, with spring droughts in temperate forests limiting seed set (BA Roy, University of Oregon, USA, personal observation, 2009).

Under common garden conditions, Blaser (2008) found consistent evidence for higher water use efficiency (WUE, estimated with d13C) in plants originating from North American seeds compared to those from Switzerland. Plants from Oregon populations had higher WUE, allocated more biomass to their roots, invested less biomass in leaf area and had thicker leaves with a lower specific leaf area (SLA) than Swiss plants. No differences between origins or among populations were found for leaf mass area (LMA), photosynthesis or transpiration.

A common garden in the invaded range was used to examine genetic variation in resistance to pests (BA Roy, University of Oregon, USA, personal observation, 2009). The garden included eight populations from the native European range (ranging from Spain to England) and eight from the invaded North American range. Populations differed in both the frequency of infection and the amount of damage by the pathogens Drechslera erythrospila and Alternaria sp. and by herbivores, indicating genetic variation in resistance. On average, North American plants had significantly more damage (were less resistant) than European plants. However, plants from invaded North America nonetheless had greater reproduction.

Associations

Besides pests, which are treated in another section, there are two important associations for B. sylvaticum: endophytic infections by Epichloësylvatica and vesicular-arbuscular mycorrhizae (VAM). Infection by E.sylvatica (asexual state=Neotyphodium) ranges in effects on the host plant from mutualistic to parasitic. Infection reduces herbivory (Brem and Leuchtmann, 2001), but some strains cause choke disease, and strongly reduce fitness as they keep the plant from flowering (Bucheli and Leuchtmann, 1996; Meijer and Leuchtmann, 2000; Brem and Leuchtmann, 2003). Epichloë/Neotyphodium infection rates are high; in Europe they are 100% infected (Bucheli and Leuchtmann, 1996; Brem and Leuchtmann, 2001; A Leuchtmann, Swiss Federal Institute of Technology, Zurich, Switzerland, personal communication, 2009), and preliminary data from the USA also suggests high rates of infection in the invaded range (BA Roy, University of Oregon, USA, personal observation, 2009).

Grass endophytes are well-known for producing toxins that are poisonous to some insects as well as to mammals such as sheep and cattle. Mammals that ingest these toxins have high abortion rates and can succumb to gruesome neurological disorders (Clay, 1996; Brem and Leuchtmann, 2001). B. sylvaticum is known to be toxic to army worm (Spodoptera) larvae in ways that are consistent with the presence of alkaloids (Brem and Leuchtmann, 2001). However, which alkaloids are present is not known (Leuchtmann et al., 2000).

As is true for many grasses, B. sylvaticum commonly forms associations with AM fungi (Abeyakoon and Pigott, 1975; BA Roy, University of Oregon, USA, personal observation, 2009), which likely aid in mineral nutrition (Smith and Read, 1997). Little is known about the particular associations with B. sylvaticum, except that they occur in both the native (Abeyakoon and Pigott, 1975) and invaded ranges (BA Roy, University of Oregon, USA, personal observation, 2009), and there is variation in infection rates among populations in the invaded range.

Environmental Requirements

The environmental requirements for this plant are broad. It ranges from arid regions such as the Middle East (Bor, 1970) to the boreal zone (Holten, 1980; Aarrestad, 2000) to high elevation tropical mountains (Hara, 1966), and from sea level to more than 4000 m (Sheehy et al., 2006; Roder et al., 2007). Based on the information from references cited in the distribution table, it is most common in areas receiving 40 cm or more of rain per year, but if rainfall levels are less that this, it will persist along river and lake margins, as for example in California, USA (Hrusa, 2003) and Eastern Oregon (False Brome Working Group, 2009). In the North temperate zone, it is most often found in shady forests, or in meadows adjacent to forests (Davies and Long, 1991; Stace, 1997; BA Roy, University of Oregon, USA, personal communication, 2009). However, it grows in open grasslands in the Northern part of the UK (Stace, 1997), in extensive grasslands in Tibet (Sheehy et al., 2006) and in tropical montane and alpine meadows (Hara, 1966; Roder et al., 2007; Singh et al., 2008).

In Europe, B. sylvaticum is often an indicator of prior disturbance such as logging or coppicing (Abeyakoon and Pigott, 1975; Rodwell, 1998; Ninot et al., 2000; Corney et al., 2008; Palo et al., 2008) and in invaded North America it is also often associated with logging (False Brome Working Group, 2009; Fletcher, 2009) and transportation corridors (BA Roy, University of Oregon, USA, personal communication, 2009). It is shade tolerant (Murchie and Horton, 1998), but tends to grow in areas where the canopy is more open and conditions are better lit (Corney et al., 2008). However, it can also be found in deep shade in undisturbed forest (Hrusa, 2003; Parks et al., 2005; Palo et al., 2008; BA Roy, University of Oregon, USA, personal observation, 2009). It has a broad tolerance of soil pH, and is even able to grow in limestone-rich, rocky areas (quarries and scree) and soils (Abeyakoon and Pigott, 1975), where it can be the first colonizer (Abeyakoon and Pigott, 1975).

There are mixed reports on how well B. sylvaticum tolerates fire. Two different studies in the native range (Canary Islands and Iran) compared the vegetation of recently (within 5 years) burned to unburned areas and found that there was no B. sylvaticum in the burned areas (Arévalo et al., 2001; Safaian et al., 2005). This information conflicts with anecdotal data on control in the invaded range, which suggested that fire was ineffective because the grass re-sprouted (False Brome Working Group, 2009). It is possible that a more intense fire was needed to kill the tussocks.

Climate

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ClimateStatusDescriptionRemark
BS - Steppe climate Tolerated > 430mm and < 860mm annual precipitation
C - Temperate/Mesothermal climate Preferred Average temp. of coldest month > 0°C and < 18°C, mean warmest month > 10°C
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)
D - Continental/Microthermal climate Preferred Continental/Microthermal climate (Average temp. of coldest month < 0°C, mean warmest month > 10°C)
Df - Continental climate, wet all year Preferred Continental climate, wet all year (Warm average temp. > 10°C, coldest month < 0°C, wet all year)
Ds - Continental climate with dry summer Tolerated Continental climate with dry summer (Warm average temp. > 10°C, coldest month < 0°C, dry summers)
Dw - Continental climate with dry winter Tolerated Continental climate with dry winter (Warm average temp. > 10°C, coldest month < 0°C, dry winters)
ET - Tundra climate Preferred Tundra climate (Average temp. of warmest month < 10°C and > 0°C)

Latitude/Altitude Ranges

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Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
59 34

Rainfall

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ParameterLower limitUpper limitDescription
Dry season duration14number of consecutive months with <40 mm rainfall

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Agromyza albipennis Herbivore Leaves not specific
Carterocephalus palaemon Herbivore Leaves not specific
Elachista Herbivore Leaves not specific
Helcystogramma rufescens Herbivore Leaves not specific
Maniola jurtina Herbivore Leaves not specific
Pararge aegeria Herbivore Leaves not specific
Pararge xiphia Herbivore Leaves not specific
Thymelicus acteon Herbivore Leaves not specific
Thymelicus sylvestris Herbivore Leaves not specific

Notes on Natural Enemies

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B. sylvaticum is attacked by a diversity of pathogens and herbivores in both the native and invaded ranges (BPI, 2009; Halbritter, 2009). A recent study compared attack rates in 10 populations in the native range (Switzerland) to 10 populations in the invaded range (Oregon), and found that although there was greater pathogen damage in the native range, the pathogens were mostly grass generalists and thus not useful as potential biocontrol agents (Halbritter, 2009). Common pathogen genera included: Ascochyta, Alternaria, Claviceps, Colletotrichum, Didymella, Drechslera, Phaeosphaeria, and Puccinia. It is not known whether these pathogens are native or invasive in either range. Given the long history of grass introductions to the USA, it is possible that the large degree of pathogen overlap between the native and invasive ranges is the result of accidental introductions; phylogenetic studies of molecular data and population genetic analyses are necessary to disentangle the origins of the pathogens.

Interestingly, although pathogen damage was higher in the native range, herbivore damage (percent area removed) was higher in the invaded range in the USA (Halbritter, 2009). Chewing damage (holes and bitten edges) is caused by non-specific grasshoppers and by caterpillars, which tend to be grass specialists, but are otherwise non-specific, for example: the chequered or arctic skipper, Carterocephaluspalaemon, which preferred B. sylvaticum to other grasses in England (Asher et al., 2001), Maniola jurtina (meadow brown), which doesn’t prefer B. sylvaticum, but will eat it (Asher et al., 2001), the skipperlings, Thymelicus acteon and Thymelicus sylvestris (Dennis, 1992) and the browns, Pararge aegeria and Pararge xiphia (Dennis, 1992). Leaf and stem mining has been reported by numerous species of Elachista moths, the moth Helcystogramma rufescens and by three agromyzids: Agromyza albipennis, Cerodonthapygmaea and Chromatomyianigra (Pitkin et al., 2009).

Means of Movement and Dispersal

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Natural Dispersal

Natural, normal dispersal distances have been measured in Denmark where the distance moved by B. sylvaticum seeds over a 10-year period was 10.2 m (Petersen and Philipp, 2001). This measurement puts the unassisted rate in terrestrial habitats at about 1 m/year. However, longer distances are also possible because B. sylvaticum commonly grows along rivers and streams (Bor, 1970; Davis, 1988; False Brome Working Group, 2009), and it is thus likely to be carried by currents to new locations downstream.

Vector Transmission

The grass has long awns that catch in animal fur. A study of animals shot by hunters found seeds on both types of animals examined (roe deer and boars) (Heinken and Raudnitschka, 2002). The seeds also readily catch in socks and shoes (BA Roy, University of Oregon, USA, personal communication, 2009) and invasive populations are strongly associated with trails and roads (BA Roy, University of Oregon, USA, personal communication, 2009). It does not survive digestion by sheep (Scholz, 2007), but it is not known whether it passes through other ungulates unharmed.

Accidental Introduction

It is moved about by logging equipment and other machinery used in infested areas (False Brome Working Group, 2009; Fletcher, 2009; A Smith, Sweet Home Ranger District (USFS), USA, personal communication, 2009). Based on genetic similarity, it has been hypothesized that seed lodged on logging equipment from McDonald-Dunn Forest near Corvallis was the source of the invasion near Sweet Home, Oregon (M Cruzan, Portland State University, Oregon, USA, personal communication, 2009). There is also some risk that it will be accidentally moved around and escape from scientists due to its increasing popularity as a model organism.

Intentional Introduction

B. sylvaticum was intentionally introduced in seed trials for “range improvement” throughout the Western USA (Hull, 1974), and it is listed as a fodder species at mid-elevations (1650 m) in the Himalayas (Singh et al., 2008). Hopefully the fact that this plant is a known invader will reduce the risk of future intentional forage introductions. Furthermore, its use as a forage species should be restricted until more is known about the toxins produced by Epichloë sylvatica, an endophytic fungus that almost always infects them (Bucheli and Leuchtmann, 1996; Brem and Leuchtmann, 2001; BA Roy, University of Oregon, USA, personal observation, 2009).

The grass is attractive, and is unfortunately sometimes recommended for ornamental cultivation, particularly for shady gardens (see for example King and Oudolf, 1998). Seeds are available online.

Seedborne Aspects of Disease

2052 seeds of B. sylvaticum were observed from both the native and invaded ranges and 12 fungal species were identified (BA Roy, University of Oregon, USA, personal communication, 2009). No specialized fungi were found on/in the seeds, and only five of the 12 were likely to be pathogens: a smut (likely to be Tilletia olida, Bipolaris sorokiniana, Ascochyta pinodella, Fusarium avenaceum and Monochaetia kansensis. Plants in the native range were more often infected by pathogenic fungi than those from the invaded range. In terms of fitness, in the native range B. sylvaticum produced 10% more aborted seeds and had a 30-50% lower germination rate.

Impact Summary

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CategoryImpact
Economic/livelihood Negative
Environment (generally) Negative

Economic Impact

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There are three kinds of economic impacts associated with the invasion of B. sylvaticum in the Northwestern USA:

- Cost of control for public lands (e.g. National Forest, BLM, and Parks (City, County, State, National)). These costs are substantial. For example, the Friends of Buford Park, a support organization for a large (956 hectares) County Park in Lane County, Oregon, USA, spends at least US $50,000 per year for manual and chemical control with glyphosate (V Rogers and J Blazar, Friends of Buford Park, Oregon, USA, personal communication, 2009). Costs include grant writing, reporting, and monitoring, outreach and education as well as control. After the initial eradication effort is completed (at least 5-7 years from now [2009]), the annual amount to maintain control should be much less. The good news is that they are having some success.

- Cost of control for private lands (for example, private logging companies, The Nature Conservancy (TNC), or landowners). According to the Oregon Department of Agriculture (ODA, 2009), private logging companies typically use herbicides on their lands to remove vegetation already, so the presence of false brome will not be adding to those costs unless they have to do additional spraying. However, other private landowners could have substantial additional costs. For example, Greg Fitzpatrick (formerly) of TNC reports that they have about 12 hectares infested with false-brome at Philomath Prairie, Philomath, Oregon and they spend about US $2667.00 per year to treat this relatively small area with herbicide. The treatments are working, and after 6 years, he says it is reduced to 60-70% of the original (G Fitzpatrick, The Nature Conservancy, Oregon, USA, personal communication, 2009).

- Potential for changed fire regime and associated costs: unknown as of yet, but could be huge.

Environmental Impact

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Impact on Habitats

There is considerable potential for this grass to cause ecosystem change by affecting fire regimes (Anzinger and Radosevich, 2008; False Brome Working Group, 2009). It is the only species in the Pacific Northwest that grows thickly in dense forest and it leaves behind copious litter. However, it is not yet known whether it will affect fire regimes.

The grass has a tendency to invade parks and nature preserves (for example, in the USA: Beacon Rock State Park in Washington State; Bald Hill Park, Corvallis, Oregon; Howard Buford Recreation Area (Mt. Pisgah), Eugene, Oregon; Philomath Prairie (TNC), Philomath, Oregon; the Wild and Scenic Rogue River, Oregon, and it was recently sighted in Redwood National Park in North California (M Cruzan, Portland State University, Oregon, USA, personal communication, 2009). The commonness in parks may be a function of propagule pressure as it is readily carried by people (BA Roy, University of Oregon, USA, personal communication, 2009), vehicles, and by animals (Heinken and Raudnitschka, 2002). Commonness in parks may also be related to disturbances, which can keep canopies a bit more open (Corney et al., 2008), or otherwise change the environment in ways that allow some invasions to occur (Hansen and Clevenger, 2005; Kalwij et al., 2008).

Impact on Biodiversity

If left unchecked in the invaded range, B. sylvaticum eliminates other native species and forms a virtual monoculture (Kaye and Blakeley-Smith, 2006; False Brome Working Group, 2009). It can also be aggressive under some conditions in the native range (Corney et al., 2008). It excludes co-occurring taxa through competition for resources, by over-topping them, and by smothering them with litter (Corney et al., 2008; BA Roy, University of Oregon, USA, personal communication, 2009). Furthermore, where native plant quantity and diversity is reduced, a “trophic cascade” can occur with a corresponding reduction of invertebrates and larger wildlife species dependent on them (Zuefle et al., 2008; Burghhardt et al., 2009).

B. sylvaticum negatively affects (via competition) rare plants and associated animals in its invaded range in the USA. The federally listed endangered Fendler’s blue butterfly (Icariciaicarioidesfenderi), which uses the federally listed threatened Lupinussulphureus ssp. kincaidii (Kincaid's lupine) as its primary host plant, is endangered primarily as a result of prairie loss. Several important remaining prairies where this butterfly and host plants live are in one of the two epicenters of the B. sylvaticum invasion and some of the prairies are being encroached upon by B. sylvaticum (US Fish and Wildlife Service, 2006). A second prairie inhabiting butterfly, Euphydryas editha taylori (Taylor’s checkerspot), a federal candidate endangered species, is also threatened by this invading grass (Severns and Warren, 2008).

Threatened Species

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Threatened SpeciesConservation StatusWhere ThreatenedMechanismReferencesNotes
Euphydryas editha tayloriNational list(s) National list(s)OregonCompetition - monopolizing resources; Competition - smotheringSeverns and Warren, 2008
Icaricia icarioides fenderiUSA ESA listing as endangered species USA ESA listing as endangered speciesOregonCompetition - monopolizing resources; Competition - smotheringUS Fish and Wildlife Service, 2006
Lupinus sulphureusNo DetailsOregonCompetition - monopolizing resources; Competition - smotheringUS Fish and Wildlife Service, 2006
Eremophila alpestris strigata (streaked horned lark)USA ESA listing as threatened species USA ESA listing as threatened speciesOregon; WashingtonEcosystem change / habitat alterationUS Fish and Wildlife Service, 2013
Lupinus oreganus var. kincaidii (Kincaid's lupine)NatureServe NatureServe; USA ESA listing as endangered species USA ESA listing as endangered speciesOregon; WashingtonCompetition - stranglingUS Fish and Wildlife Service, 2006
Solidago houghtonii (Houghton's goldenrod)NT (IUCN red list: Near threatened) NT (IUCN red list: Near threatened); USA ESA listing as threatened species USA ESA listing as threatened speciesOntario; MichiganCompetition - monopolizing resources; Ecosystem change / habitat alterationUS Fish and Wildlife Service, 2011

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
  • Pioneering in disturbed areas
  • Tolerant of shade
  • Benefits from human association (i.e. it is a human commensal)
  • Long lived
  • Fast growing
  • Has high reproductive potential
  • Has propagules that can remain viable for more than one year
  • Reproduces asexually
  • Has high genetic variability
Impact outcomes
  • Ecosystem change/ habitat alteration
  • Modification of successional patterns
  • Monoculture formation
  • Negatively impacts forestry
  • Reduced native biodiversity
  • Threat to/ loss of endangered species
  • Threat to/ loss of native species
  • Negatively impacts animal/plant collections
Impact mechanisms
  • Competition - monopolizing resources
  • Competition - smothering
  • Competition - strangling
  • Hybridization
  • Rapid growth
  • Rooting
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

B. sylvaticum has economic value for three uses:

- It is grown for fodder throughout its high elevation Asian distribution (Veldkamp and Vanscheindelen, 1989; Sheehy et al., 2006; Singh et al., 2008).

- It is a beautiful grass used in ornamental plantings (King and Oudolf, 1998).

- B. sylvaticum and the related Brachypodium distachyon are gaining popularity as a research model due to their similarity to the wheat and rice genomes, but they are easier to work with than these grains due to having smaller genomes (Foote et al., 2004; Garvin, 2007; Opanowicz et al., 2008)

Detection and Inspection

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For a grass, B. sylvaticum is relatively easy for botanists to identify because it grows in the shade of the forest (and often along trails, streams and rivers) and is quite pretty: it stays a bright vibrant green all season long, and has broad lax leaves and nodding inflorescences. Nonetheless, for identification it is best to involve a trained botanist as there are a couple of native forest grasses in North America with which it could be confused (for example, Bromus vulgaris, and possibly Melica subulata, Elymus glaucus and Bromus carinatus). Excellent identification information can be found at Piep (2003) and Shouliang and Phillips (2006).

Similarities to Other Species/Conditions

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B. sylvaticum is closely related to Brachypodium pinnatum (Catalán and Olmstead, 2000). However, the taxa are easily differentiated because B. sylvaticum has longer awns (7-15 mm vs. 0-7 mm in B. pinnatum), has nodding racemes instead of erect ones, and it does not have spreading rhizomes (Piep, 2003). Paszko (2008) adds that B. sylvaticum always has pubescent abaxial palea surfaces whereas B. pinnatum does not.

Prevention and Control

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Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

Prevention

SPS measures

The best prevention is to prohibit it, restrict its movement (including reducing sales over the internet), to never plant it, and to teach people how to identify and remove it. It has been designated as a quarantined “A & Q” weed in California (CDFA, 2009), it is a class ‘A’ quarantined weed in Washington State, USA (eradication is required), and a class ‘B’ quarantined weed (intensive control where needed) in Oregon (ODA, 2009).

Eradication

This grass has two weaknesses despite it’s overall strength. Firstly, it is susceptible to glyphosate (False Brome Working Group, 2009), and because it stays green late into the year this allows application of the herbicide with reduced consequences for the associated native herb layer, which typically dies back during the summer. Second, it is not rhizomatous, thus once the clumps have been removed, they stay removed.

Containment/Zoning

Containment is difficult to impossible with this species due to its ability to bank seeds (Donelan and Thompson, 1980; Buckley et al., 1997), and the fact that seeds are readily spread by wildlife (Heinken and Raudnitschka, 2002).

Control

Physical/Mechanical Control

If the invaded area is small, clumps can be removed by hand. However, it does have a persistent seed bank (Donelan and Thompson, 1980; Buckley et al., 1997), thus control measures need to be maintained for several years. Mowing can be used to reduce seed set, if timed properly. However, there are problems with mowing: mowing does not decrease competition with the native species and it is not very effective for seed control because timing is critical and many plants will be missed. If mowing happens too early, the plants will re-sprout to flower anyway, and if it is too late, the mowers will simply broadcast the seed. Hot foam ( (a system that uses a hot surfactant foam to deliver and trap superheated steam onto foliage) is effective, but is only practical on roadsides and is more expensive than herbicides (False Brome Working Group, 2009).

Movement Control

B. sylvaticum is known to be carried by animals in their fur (Heinken and Raudnitschka, 2002), and by humans in their socks and shoes. The strong correlation with roadsides (BA Roy, University of Oregon, USA, personal communication, 2009) and with logging (Corney et al., 2008; Palo et al., 2008; Fletcher, 2009) also strongly suggests that it is carried by vehicles. Implementing boot/shoe cleaning stations in public parks that are known to be infested is a good idea (False Brome Working Group, 2009), and all vehicles that have been in infested forest should be thoroughly cleaned before moving to new areas (False Brome Working Group, 2009; Fletcher, 2009).

Biological Control

Biological control is not likely to be possible given the relationship with species of agricultural importance, such as wheat [Triticum aestivum] (Catalán and Olmstead, 2000; Opanowicz et al., 2008; Rathore and Shekhawat, 2009), and the fact that most grass pathogens and herbivores are generalists that attack numerous species. Even if a specific agent were to be found, given the commonness of this family worldwide, the risk of host jumping at some point would be high.

There is one specialist that is absent from North America, and that is the sexual strain of the “mutualistic” endophyte Neotyphodium/Epichloë sylvatica. When this fungus reproduces asexually, it should be called Neotyphodium and it is likely to be of benefit to the host (mutualistic), but when it reproduces sexually it should be called Epichloë sylvatica and is a parasite. Plants of B. sylvaticum in Europe are always infected by the Neotyphodium/Epichloë fungus, and usually by the asexual, or Neotyphodium, strain. However, some European populations of false-brome endophyte, have infection by the sexual strain, which should be called Epichloë sylvatica in this case. The sexual strain causes ‘choke’ in false-brome. ‘Choke’ reduces and/or prevents seed production because the fungus reproduces instead of the host. Leuchtmann and colleagues have shown that the strain that sexually reproduces is genetically different from the one that asexually reproduces (Bucheli and Leuchtmann, 1996; Meijer and Leuchtmann, 2001). Furthermore, the sexual strain is quite rare in Europe (Bucheli and Leuchtmann, 1996; Meijer and Leuchtmann, 1999). North American populations are only infected by the asexual, or Neotyphodium strain, and are thus never choked by the fungus (Halbritter, 2009; BA Roy, University of Oregon, USA, personal observation, 2009). The sexual genotype, E. sylvaticum, causes choke and thus reduces host fitness, but we don’t advise introducing it to North America for biocontrol because the fungus might jump hosts, as it has done in the past (Brem and Leuchtmann, 2003). Furthermore, the fact that the sexual strain is rare in Europe suggests that it is not favoured in natural populations and thus may not sustain itself after release, nor be effective. Substantial screening of fungal genotypes and vulnerable native species, research, and planning are required before release would be appropriate or permitted.

Chemical Control

Chemical control is generally effective as the grass is susceptible to glyphosate. More information can be found at the False Brome Working Group website (False Brome Working Group, 2009).

Control by Utilization

Until more is known about the type and quantity of toxins produced by the endophyte that almost always infects this grass (Brem and Leuchtmann, 2001), grazing is not recommended. Grass endophytes are well known for producing toxins that are poisonous to some insects as well as to mammals such as sheep and cattle (Clay, 1990; Brem and Leuchtman, 2001). Mammals that ingest these toxins have high abortion rates and can succumb to gruesome neurological disorders (Clay, 1996; Brem and Leuchtmann, 2001). Probably due to its inedibility, animals, including sheep, generally avoid this grass. A study was done to determine whether sheep could be trained to eat B. sylvaticum; they found that when sheep were exposed early in life, they did eat more of it later (Scholz, 2007). Even if it is eventually found that grazing this grass is safe, grazing will not eliminate it, although it could reduce seed production if the timing of grazing was carefully controlled.

Monitoring and Surveillance (incl. remote sensing)

Monitoring in the invaded USA range is on going, including weed control personnel at the Federal State (e.g. US Forest Service and Bureau of Land Management) and local levels. Friends of Buford Park (non-profit stewards of a 956 hectare Lane County, Oregon park) have creatively engaged volunteers to adopt areas for patrolling and controlling this grass. Surveillance with remote sensing has not yet been developed.

Ecosystem Restoration

Restoration has been a priority with The Nature Conservancy (TNC) and with local parks in the Eugene, Oregon area (e.g. Elijah-Bristow State Park and the Howard Buford Recreation Area). For large areas, glyphosate is used (see False Brome Working Group, 2009, for recommended procedures) in the late-summer or autumn after most of the native vegetation has died back. Herbicide applications continue for at least 2 years, and the area is re-seeded with natives.

Gaps in Knowledge/Research Needs

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- The effects of B sylvaticum on fire behaviour in Pacific Northwest forests need to be evaluated because if this grass changes forest fire susceptibility, it has the potential to be an ecosystem changer (Anzinger and Radosevich, 2008). Furthermore, the potential for fire to control the grass in the invasive range needs to be re-evaluated in light of data from the native range showing a negative association with burned areas (Arévalo et al., 2001; Safaian et al., 2005).

- The effects of B. sylvaticum on germination, growth and survival of trees in the invaded range (e.g. Pseudotsuga menziesii, Quercus garryana, Sequoia sempervirens) needs to be evaluated. It is known that the presence of grasses in general decreases conifer fitness, but it is not known whether this grass has the same effect (it could even be worse given its shade tolerance).

- For detection in meadow areas, it would be extremely useful to determine whether B. sylvaticum has a unique signal that could be detected using remote sensing. The grass has a unique colour in the visible spectrum, but it is not yet known whether it can be screened for with remote sensing. This would facilitate tracking it in open portions of roadless areas and on private lands.

- The population dynamics within the invasive range need to be better understood to facilitate control. Current practice for controlling this grass is to focus on small satellite populations, which seem possible to control, rather than the large populations that seem like a lost cause. However, there may be reasons why the small populations are less worrisome (for example, if inbreeding depression depresses seed set in small populations). Work on population dynamics could address a number of useful questions, such as: Are there source and sink populations? Are there ways to identify source populations? Does endophyte infection affect herbivory and control of population growth rates by enemies? Does inbreeding depression depress seed set in small populations? Is disturbance necessary for invasion? Does invasion occur under all light levels?

- More work needs to be done on understanding the relationships between habitat and dispersal. In Europe, the grass is associated with logging and other disturbances and it has been assumed that this relationship is the result of opening the forest up to more light (Rodwell, 1998; Corney et al., 2008). There is evidence that light may be a limiting factor given the germination and growth patterns of the grass (Grime et al., 1988; Buckley et al., 1997). However, another hypothesis for which there is some evidence is that the logging trucks and equipment move seeds around. Two lines of evidence support the logging = moving around hypothesis. First, in the invaded range there is a strong association with B. sylvaticum and the proximity of roads and paths (BA Roy, University of Oregon, USA, personal communication, 2009). Second, genetic evidence suggests that the invasion near Sweet Home, Oregon is related to the one at McDonald-Dunn forest near Corvallis, and the most parsimonious idea for spread to Sweet Home was via seeds on logging equipment (M Cruzan, Portland State University, Oregon, USA, personal communication, 2009). A third hypothesis that deserves exploration is that disturbance of the litter is in someway necessary for the grass to get a foothold (M Cruzan, Portland State University, Oregon, USA, personal communication, 2009).

- The taxonomy of B. sylvaticum needs to be re-evaluated. Numerous varieties have been described, mostly on the basis of pubescence differences (Veldkamp and Vanscheindelen, 1989; Piep, 2003; Shouliang and Phillips, 2006). However, given that hairiness can vary even within populations (Paszko, 2008), these taxa may not be real. On the other hand, it seems possible that the high elevation grassland types found in tropical alpine Asia, may well be different from the B. sylvaticum found in low elevation North European forests. The best way to analyze the variation across the entire range of B. sylvaticum would be to combine data from a common garden (to hold the environment constant so that all the variation was genetic) with molecular data such as microsatellites.

References

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Aarrestad PA, 2000. Plant communities in broad-leaved deciduous forests in Hordaland county, Western Norway. Nordic Journal of Botany, 20(4):449-466.

Abeyakoon KF, Pigott CD, 1975. The inability of Brachypodium sylvaticum and other species to utilize apatite or organically bound phosphate in calcareous soils. New Phytologist, 74(1):147-154.

Al-Eisawi DM, Al-Khader IA, 1998. Checklist of plants of the Hashemite Kingdom of Jordan. Jordan country study on biological diversity. Checklist of plants of the Hashemite Kingdom of Jordan. Jordan country study on biological diversity. Amman, Jordan: The General Corporation for the Environment Protection, unpaginated.

Alonso I, Hartley SE, Thurlow M, 2001. Competition between heather and grasses on Scottish moorlands: interacting effects of nutrient enrichment and grazing regime. Journal of Vegetation Science, 12(2):249-260.

Amme D, 2006. Slender false brome (Brachypodium sylvaticum) spreading into California. Grasslands, 16:4.

Anzinger, Radosevich SR, 2008. Fire and nonnative invasive plants in the Northwest coastal bioregion. USDA Forest Service Gen. Tech. Rep. RMRS-GTR-42, 6. Washington, D. C., USA: USDA Forest Service, 197-223.

Arévalo JR, Fernández-Palacios JM, Jiménez MJ, Gil P, 2001. The effect of fire intensity on the understorey species composition of two Pinus canariensis reforested stands in Tenerife (Canary Islands). Forest Ecology and Management, 148(1/3):21-29.

Asher J, Warren M, Fox R, Harding P, Jeffcoate G, Jeffcoate S, 2001. The millennium atlas of butterflies in Britain and Ireland [ed. by Asher, J.\Warren, M.\Fox, R.\Harding, P.\Jeffcoate, G.\Jeffcoate, S.]. Oxford, UK: Oxford University Press, xx + 433 pp.

Bakker EG, Montgomery B, Nguyen T, Eide K, Chang J, Mockler TC, Liston A, Seabloom EW, Borer ET, 2009. Strong population structure characterizes weediness gene evolution in the invasive grass species Brachypodium distachyon. Molecular Ecology, 18(12):2588-2601. http://www.blackwell-synergy.com/loi/mec

Barkworth ME, Capels KM, Long S, Anderton LK, Piep MB, 2007. Flora of North America North of Mexico. New York, New York, USA: Oxford University Press.

Blaser W, 2008. Common gardens as a tool to address questions related to plant invasions. Zürich, Switzerland: ETH (Swiss Federal Institute of Technology).

Bor NL, 1970. Volume 70: Graminae. Flora Iranica: Flora des Iranischen Hochlandes und der umrahmenden Gebirge; Persien, Afganistan, Teile von West-Pakistan, Nord-Iraq, Azerbaidjan, Turkmenistan [ed. by Reichinger KH]. Graz, Austria: Akademische Druck und Verlagsanstalt, 573 pp.

Bor NL, Guest E, 1968. Flora of Iraq. Baghdad, Iraq: Ministry of Agriculture of the Republic of Iraq.

BPI (US National Fungus Collections), 2009. Fungal Databases - Specimens. Beltsville, USA: Systematic Mycology and Microbiology Laboratory, Agricultural Research Service, USDA. www.nt.ars-grin.gov/fungaldatabases/specimens/specimens.cfm

Brem D, Leuchtmann A, 2001. Epichloë grass endophytes increase herbivore resistance in the woodland grass Brachypodium sylvaticum. Oecologia, 126(4):522-530.

Brem D, Leuchtmann A, 2003. Molecular evidence for host-adapted races of the fungal endophyte Epichloë bromicola after presumed host shifts. Evolution, 57(1):37-51.

Bucheli E, Leuchtmann A, 1996. Evidence for genetic differentiation between choke-inducing and asymptomatic strains of the Epichloë grass endophyte from Brachypodium sylvaticum. Evolution, 50(5):1879-1887.

Buckley GP, Howell R, Anderson MA, 1997. Vegetation succession following ride edge management in lowland plantations and woods. 2. The seed bank resource. Biological Conservation, 82(3):305-316.

Burghardt KT, Tallamy DW, Shriver WG, 2009. Impact of native plants on bird and butterfly biodiversity in suburban landscapes. Conservation Biology, 23(1):219-224. http://www.blackwell-synergy.com/loi/cbi

Catalán P, Olmstead RG, 2000. Phylogenetic reconstruction of the genus Brachypodium P. Beauv. (Poaceae) from combined sequences of chloroplast ndhF gene and nuclear ITS. Plant Systematics and Evolution, 220(1/2):1-19.

CBG NASB MSKH, 2009. Herbarium of CBG NASB MSKH supported by the Ministry of Natural Resources and Environmental Protection of the Republic of Belarus. Plants of Belarus. http://hbc.bas-net.by/plantae/eng/allplantras.php

CDFA, 2009. California department of food and agriculture plant quarantine manual. California department of food and agriculture plant quarantine manual. unpaginated. http://pi.cdfa.ca.gov/pqm/manual/pdf/107.pdf

Chaideftou E, Thanos CA, Bergmeier E, Kallimanis A, Dimopoulos P, 2009. Seed bank composition and above-ground vegetation in response to grazing in sub-Mediterranean oak forests (NW Greece). Plant Ecology, 201(1):255-265. http://springerlink.metapress.com/link.asp?id=100328

Chambers KL, 1966. Notes on some grasses of the pacific coast. Madroño, 18:250-251.

Clay K, 1990. Fungal endophytes of grasses. Annual Review of Ecology and Systematics, 21:275-295.

Clay K, 1996. Interactions among fungal endophytes, grasses and herbivores. Researches on Population Ecology, 38(2):191-201.

Corney PM, Kirby KJ, Duc MGle, Smart SM, McAllister HA, Marrs RH, 2008. Changes in the field-layer of Wytham Woods - assessment of the impacts of a range of environmental factors controlling change. Journal of Vegetation Science, 19(3):287-298. http://www.opuluspress.se/index.php?option=com_phpshop&page=shop/flypage_journal&product_id=8&Itemid=56&cat=1&nr=1

Czerepanov SK, 1995. Vascular plants of Russia and adjacent states (the former USSR). Cambridge, UK: Cambridge University Press, x+516pp.

Danin A, 2000. The nomenclature news of flora Palaestina. Flora Mediterranea:109-172.

Davies MS, Long GL, 1991. Performance of two contrasting morphs of Brachypodium sylvaticum transplanted into shaded and unshaded sites. Journal of Ecology (Oxford), 79(2):505-517.

Davis PH, 1988. Flora of Turkey and the East Aegean Islands, vol. 9. Edinburgh, United Kingdom: University Press.

Dennis RLH, 1992. The ecology of butterflies in Britain. Oxford, United Kingdom: Oxford University Press.

Donelan M, Thompson K, 1980. Distribution of buried viable seeds along a successional series. Biological Conservation, 17(4):297-311.

Edgar E, Connor HE, 2000. Flora of New Zealand, volume V: grasses. Lincoln, New Zealand: Manaaki Wehnua Press.

eFloras, 2009. Efloras: Flora of Madagascar. Efloras: Flora of Madagascar. ST. Louis, MO: Missouri Botanical Garden, unpaginated. http://www.efloras.org

EPPO, 2014. PQR database. Paris, France: European and Mediterranean Plant Protection Organization. http://www.eppo.int/DATABASES/pqr/pqr.htm

Euro+Med Plantbase, 2013. The information resource for Euro-Mediterranean plant diversity. The information resource for Euro-Mediterranean plant diversity. unpaginated. http://ww2.bgbm.org/EuroPlusMed/

Evans CE, Etherington JR, 1991. The effect of soil-water potential on seedling growth of some British plants. New Phytologist, 118:571-579.

False Brome Working Group, 2009. USDA forest service, USDI bureau of land management, Oregon department of agriculture, US army corps of engineers, OSU college of forestry, institute for applied ecology, Starker forests, Inc., the nature conservancy, native plant society of Oregon. USDA forest service, USDI bureau of land management, Oregon department of agriculture, US army corps of engineers, OSU college of forestry, institute for applied ecology, Starker forests, Inc., the nature conservancy, native plant society of Oregon. unpaginated. http://www.appliedeco.org/invasive-species-resources/FBWG

Faris JD, Zhang ZC, Fellers JP, Gill BS, 2008. Micro-colinearity between rice, Brachypodium, and Triticum monococcum at the wheat domestication locus Q. Functional & Integrative Genomics, 8(2):149-164. http://www.springerlink.com/content/m42424r565087214/?p=451e016a58804ed48b10c72a7f7712a1&pi=5

Fletcher R, 2009. McDonald-Dunn forest plan revision: invasive plant management. McDonald-Dunn forest plan revision: invasive plant management. unpaginated. http://www.cof.orst.edu/cf/forests/mcdonald/plan/

Foote TN, Griffiths S, Allouis S, Moore G, 2004. Construction and analysis of a BAC library in the grass Brachypodium sylvaticum: its use as a tool to bridge the gap between rice and wheat in elucidating gene content. Functional & Integrative Genomics, 4(1):26-33. http://springerlink.metapress.com/app/home/contribution.asp?wasp=agxy6mvrqg3tul0vtrw6&referrer=parent&backto=issue,3,7;journal,3,16;linkingpublicationresults,1:105695,1

FRIM, 2009. Flora of Peninsular Malaysia online. Flora of Peninsular Malaysia online. unpaginated. http://www.tfbc.frim.gov.my/default.asp

Garvin DF, 2007. Brachypodium: a new monocot model plant system emerges. Journal of the Science of Food and Agriculture, 87(7):1177-1179. http://www.interscience.wiley.com/jsfa

GBIF, 2009. GBIF Data Portal. Copenhagen, Denmark: Global Biodiversity Information Facility. http://data.gbif.org

Gooding EGB, Loveless AR, Proctor GR, 1965. Flora of Barbados. London : HMSO, 502 pp.

Grime JP, Hodgson JG, Hunt R, 1988. Comparative plant ecology. A functional approach to common British species. London, UK: Unwin Hyman Ltd., 679 pp.

Halbritter AH, 2009. Testing assumptions of the enemy release hypothesis: Herbivore and pathogen damage in the native and invaded range of the grass Brachypodium sylvaticum. Masters. Zürich, Switzerland: Swiss Federal Institute of Technology (ETH).

Halbritter AH, Roy BA, Güsewell S, Carroll G, undated. Testing assumptions of the enemy release hypothesis: Herbivore and pathogen damage of Brachypodium sylvaticum ecology. Testing assumptions of the enemy release hypothesis: Herbivore and pathogen damage of Brachypodium sylvaticum ecology. unpaginated.

Hammel BE, Grayum MH, Herrera C, Zamora N, 2003. Manual de plantas de Costa Rica v. 3. Monocotyledoneas (Orchidaceae-Zingiberaceae). St. Louis, Missouri, USA: Missouri Botanical Garden.

Hansen MJ, Clevenger AP, 2005. The influence of disturbance and habitat on the presence of non-native plant species along transport corridors. Biological Conservation, 125(2):249-259. http://www.sciencedirect.com/science/journal/00063207

Hara H, 1966. The flora of Eastern Himalaya. Tokyo, Japan: University of Tokyo.

Heinken T, Raudnitschka D, 2002. Do wild ungulates contribute to the dispersal of vascular plants in Central European forests by epizoochory? A case study in north-east Germany. (Trägt Schalenwild durch Epizoochorie zur Ausbreitung von Gefäßpflanzen in mitteleuropäischen Wäldern bei? Eine Fallstudie aus Nordostdeutschland.). Forstwissenschaftliches Centralblatt, 121(4):179-194.

Hitchcock S, 1950. Manual of the Grasses of the United States. New York, USA: Dover Publications Inc.

Holten JI, 1980. Distribution and ecology of Brachypodium sylvaticum, Bromus benekeni and Festuca altissima in central Norway. (Utbredese og oekologi for Brachypodium sylvaticum, Bromus benekeni og Festuca altissima i midr-Norge.) Blyttia, 38(3):137-144.

Hrusa GF, 2003. Two potentially serious new weeds in California. Plant pest diagnostics laboratory annual report [ed. by Kodira UC]. http://www.cdfa.ca.gov/phpps/ppd/PDF/PPDC2003.pdf

Hull AC Jr, 1974. Species for seeding mountain rangelands in southeastern Idaho, northeastern Utah, and western Wyoming. Journal of Range Management, 27(2):150-153.

Hæggström CA, Skytén R, 1996. Flowering and individual survival of a population of the grass Brachypodium sylvaticum in Natö, Aland Islands, SW Finland. Annales Botanici Fennici, 33(1):1-10.

IBIS, 2009. Integrated botanical information system (Australian national botanic gardens, Australian national herbarium). Australian Plant Name Index. unpaginated. http://www.anbg.gov.au

Jepson Herbarium, 2009. University of California and Jepson Herbarium. University of California and Jepson Herbarium. unpaginated. http://ucjeps.berkeley.edu/cgi-bin/get_consort.pl?taxon_name=Brachypodium%20sylvaticum

Jørgensen PM, León-Yánez S, 1999. Catalogue of the vascular plants of Ecuador. Monographs in Systematic Botany from the Missouri Botanical Garden. 75:1-1182.

Kalwij JM, Milton SJ, McGeoch MA, 2008. Road verges as invasion corridors? A spatial hierarchical test in an arid ecosystem. Landscape Ecology, 23(4):439-451. http://springerlink.metapress.com/link.asp?id=103025

Kaye TN, Blakeley-Smith M, 2006. False-brome (Brachypodium sylvaticum). In: Invasive species in the Pacific Northwest [ed. by Boersma, P. D.\Reichard, S. E.\Buren, A. N. van]. Seattle: University of Washington Press, 80-81.

Khan MA, Stace CA, 1999. Breeding relationships in the genus Brachypodium (Poaceae: Pooideae). Nordic Journal of Botany, 19(3):257-269.

King M, Oudolf P, 1998. Gardening with grasses., Hong Kong: Francis Lincoln, Ltd.

KPNI, 2009. KPNI (Korean plant names index). KPNI (Korean plant names index). unpaginated. http://www.nature.go.kr

Kruse R, Bend E, Bierzychudek P, 2004. Native plant regeneration and introduction of non-natives following post-fire rehabilitation with straw mulch and barley seeding. Forest Ecology and Management, 196(2/3):299-310.

Lauber K, Wagner G, 1996. Flora Helvetica. Bern, Switzerland: Paul Haupt Berne.

Lehmkuhl JF, 2002. The effects of spring burning and grass seeding in forest clearcuts on native plants and conifer seedlings in coastal Washington. Northwest Science, 76(1):46-60.

Leuchtmann A, Schmidt D, Bush LP, 2000. Different levels of protective alkaloids in grasses with stroma-forming and seed-transmitted Epichloë/Neotyphodium endophytes. Journal of Chemical Ecology, 26(4):1025-1036.

Marozas V, 2004. Diversity of lime forest in the different regions of Lithuania. Acta Biologica Universitatis Daugavpiliensis, 4(1):47-52.

Meijer G, Leuchtmann A, 1999. Multistrain infections of the grass Brachypodium sylvaticum by its fungal endophyte Epichloë sylvatica. New Phytologist, 141(2):355-368.

Meijer G, Leuchtmann A, 2000. The effects of genetic and environmental factors on disease expression (stroma formation) and plant growth in Brachypodium sylvaticum infected by Epichloë sylvatica. Oikos, 91(3):446-458.

Meijer G, Leuchtmann A, 2001. The effects of genetic and environmental factors on disease expression (stroma formation) and plant growth in Brachypodium sylvaticum infected by Epichloe sylvatica. Oikos, 91(3):446-458.

Miller BM, Aitken RJ, Oldham MJ, Reznicek AA, 2011. Slender false brome (Brachypodium sylvaticum, Poaceae), an invasive grass new to Ontario, Canada. Canadian Field-Naturalist, 125(3):235-240. http://www.canadianfieldnaturalist.ca/index.php/cfn/article/view/1226/1214

Missouri Botanical Garden, 2011. Tropicos database. St Louis, USA. http://www.tropicos.org

Murchie EH, Horton P, 1998. Contrasting patterns of photosynthetic acclimation to the light environment are dependent on the differential expression of the responses to altered irradiance and spectral quality. Plant, Cell and Environment, 21(2):139-148.

Ninot JM, Carreras J, Carrillo E, Vigo J, 2000. Syntaxonomic conspectus of the vegetation of Catalonia and Andorra. I.: Hygrophilous herbaceous communities. Acta Botanica Barcelona, 46:191-237.

NWCB, 2009. Washington state noxious weed control board. Washington state noxious weed control board. unpaginated. http://www.nwcb.wa.gov/

ODA, 2009. Oregon department of agriculture plant division, noxious weed control, Oregon, USA. Oregon department of agriculture plant division, noxious weed control, Oregon, USA. unpaginated. http://www.oregon.gov/ODA/PLANT/WEEDS/

Oi J, 1965. Flora of Japan. Washington, D. C., USA: Smithsonian Institution.

Opanowicz M, Vain P, Draper J, Parker D, Doonan JH, 2008. Brachypodium distachyon: making hay with a wild grass. Trends in Plant Science, 13(4):172-177. http://www.sciencedirect.com/science/journal13601385

Palo A, Linder M, Truu J, Mander Ü, 2008. The influence of biophysical factors and former land use on forest floristic variability on Saaremaa and Muhu islands, Estonia. Journal for Nature Conservation, 16(3):123-134. http://www.elsevier.de/jnc

Parks CG, Radosevich SR, Endress BA, Naylor BJ, Anzinger D, Rew LJ, Maxwell BD, Dwire KA, 2005. Natural and land-use history of the Northwest mountain ecoregions (USA) in relation to patterns of plant invasions. Perspectives in Plant Ecology, Evolution and Systematics, 7(3):137-158. http://www.sciencedirect.com/science/journal/14338319

Paszko B, 2008. The variability of natural populations of Brachypodium pinnatum and B. sylvaticum based on morphological features. Acta Societatis Botanicorum Poloniae, 77(3):255-262.

Petersen PM, Philipp M, 2001. Implantation of forest plants in a wood on former arable land: a ten year experiment. Flora (Jena), 196(4):286-291.

Piep MB, 2003. 11.01 Brachypodium. In: Flora of North America: North of Mexico, volume 24 [ed. by Barkworth ME, Capels KM, Anderton L, Long S] New York, USA: Oxford University Press, 187-192.

Pitkin B, Ellis W, Plant C, Edmunds R, 2009. Leaf and stem mines of British flies and other insects. Leaf and stem mines of British flies and other insects. unpaginated. http://www.nhm.ac.uk/research-curation/research/projects/british-insect-mines/index.html

Powell GW, Pitt MD, Wikeem BM, 1994. Effect of forage seeding on early growth and survival of lodgepole pine. Journal of Range Management, 47(5):379-384.

Rathore MS, Shekhawat NS, 2009. Smaller model genome for monocots - Brachypodium distachyon. Current Science, 96:876-876.

Roder W, Dorji K, Wangdi K, 2007. Implications of white clover introduction in East Himalayan grasslands. Mountain Research and Development, 27(3):268-273. http://www.bioone.org/doi/full/10.1659/mrd.0751

Rodwell J, 1998. Woodlands and scrub. Cambridge, United Kingdom: Cambridge University Press.

Rosenthal DM, Ramakrishnan AP, Cruzan MB, 2008. Evidence for multiple sources of invasion and intraspecific hybridization in Brachypodium sylvaticum (Hudson) Beauv. in North America. Molecular Ecology, 17(21):4657-4669. http://www3.interscience.wiley.com/cgi-bin/fulltext/120696752/HTMLSTART

Roy BA, Stanton ML, Eppley SM, 1999. Effects of environmental stress on leaf hair density and consequences for selection. Journal of Evolutionary Biology, 12(6):1089-1103.

Safaian N, Shokri M, Ahmadi MZ, Strakchali A, Tavili A, 2005. Fire influence on the grassland vegetation in Golestan National Park (Alborz Mts. Iran). Polish Journal of Ecology, 53:435-443.

Scholz H, König P, 1985. [English title not available]. (Ergänzungen zur Flora Saudi-Arabiens: Gramineae.) Willdenowia, 14:373-377.

Scholz R, 2007. Use of sheep in the control of an invasive grass. Corvallis, Oregon, USA: Oregon State University.

Severns PM, Warren AD, 2008. Selectively eliminating and conserving exotic plants to save an endangered butterfly from local extinction. Animal Conservation, 11(6):476-483. http://www.blackwell-synergy.com/loi/acv

Sheehy DP, Miller D, Johnson DA, 2006. Transformation of traditional pastoral livestock systems on the Tibetan steppe. Sécheresse, 17(1/2):142-151. http://www.secheresse.info

Shouliang C, Phillips SM, 2006. 13. Tribe Brachypodieae. In: Flora of China. Vol. 22. Poaceae [ed. by Wu, Z. Y.\Raven, P. H.\Hong, D. Y.]. Beijing, China: Science Press (Beijing), 368-369.

Singh V, Guaur RD, Bohra B, 2008. A survey of fodder plants in mid-altitude Himalayan rangelands of Uttarakhand, India. Journal of Mountain Science, 5:265-278.

Smith JMB, 1975. Notes on the distributions of herbaceous angiosperm species in the mountains of New Guinea. Journal of Biogeography, 2(2):87-101.

Smith SE, Read DJ, 1997. Mycorrhizal symbiosis, Ed. 2. San Diego, US: Academic Press, ix + 605 pp.

Soreng RJ, Davidse G, Peterson PM, Zuloaga FO, Judziewicz FJ, Filgueiras TS, Morrone O, 2009. Catalogue of new world grasses (Poaceae). http://mobot.mobot.org/W3T/Search/nwgc.html

Stace C, 1997. New flora of the British Isles. Second edition. Cambridge, UK: Cambridge University Press.

Stevens WP, Ulloa-Ulloa C, Pool A, Monhel OM, 2001. Flora de Nicaragua Angiospermes (Pandanaceae-Zygophyllaceae). St. Louis, Missouri, USA: Missouri Botanical Garden.

Szujkó-Lacza J, Fekete G, 1974. Examination of development and growth of Brachypodium silvaticum and Euphorbia cyparissias in oakwoods. Acta Botanica Academiae Scientiarum Hungaricae, 20(1/2):147-158.

US Fish and Wildlife Service, 2006. In: Endangered and Threatened Wildlife and Plants; Designation of Critical Habitat for the Fender's blue butterfly (Icaricia icarioides fenderi), Lupinus sulphureus ssp. kincaidii (Kincaid's lupine), and Erigeron decumbens var. decumbens (Willamette daisy). 71(210) US Fish and Wildlife Service, 63862-63977. http://www.gpo.gov/fdsys/pkg/FR-2006-10-31/pdf/06-8809.pdf

US Fish and Wildlife Service, 2011. In: Houghton's Goldenrod (Solidago houghtonii). 5-Year Review: Summary and Evaluation. US Fish and Wildlife Service, 23 pp. http://ecos.fws.gov/docs/five_year_review/doc3880.pdf

US Fish and Wildlife Service, 2013. In: Endangered and Threatened Wildlife and Plants; Determination of Endangered Status for the Taylor's Checkerspot Butterfly and Threatened Status for the Streaked Horned Lark; Final Rule. 78(192) US Fish and Wildlife Service, 61452-61503. https://www.gpo.gov/fdsys/pkg/FR-2013-10-03/pdf/2013-23567.pdf

Veldkamp JF, Vanscheindelen HJ, 1989. Australopyrum, Brachypodium, and Elymus (Gramineae) in Malesia. Blumea, 34:61-76.

Virginia Botanical Associates, 2009. Digital atlas of the Virginia flora. Digital atlas of the Virginia flora. unpaginated. http://www.biol.vt.edu/digital_atlas

Wolny E, Hasterok R, 2007. Comparative cytogenetic analysis of Brachypodium species. Chromosome Research, 15:30-30.

Zuefle ME, Brown WP, Tallamy DW, 2008. Effects of non-native palnts on the native insect community of Delaware. Biological Invasions, 10:1159-1169.

Zuloaga FO, Nicora EG, Rúgolo de Agrasae ZE, Piensiero J, Cialdella AM, 1994. [English title not available]. (Catálogo de la familia Poaceae en la república Argentina.) Catálogo de la familia Poaceae en la república Argentina. St. Louis, Missouri, USA: Missouri Botanical Garden, unpaginated.

Links to Websites

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WebsiteURLComment
California Department of Agriculture Quarantine Manualhttp://pi.cdfa.ca.gov/pqm/manual/htm/pqm_index.htm
Catalogue of New World Grasses (Poaceae)http://www.tropicos.org/Project/CNWG
Euro+Med Plantbase - the information resource for Euro-Mediterranean plant diversityhttp://ww2.bgbm.org/EuroPlusMed/
False-Brome Working Grouphttp://appliedeco.org/invasive-species-resources/FBWG
Friends of Buford Parkhttp://www.bufordpark.org/
GBIF Networkwww.gbif.net
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.
Integrated Botanical Information System (IBIS)http://www.anbg.gov.au/anbg/index-ibis.html
McDonald-Dunn forest plan revision: Invasion plant managementhttp://www.cof.orst.edu/cf/forests/mcdonald/plan/files/McDonald%20Forest%20Invasive%20Plant%20Management%20Plan.pdf
Oregon Department of Agriculture Plant Division, Noxious Weed Control (ODA)http://www.oregon.gov/ODA/PLANT/WEEDS/
University of California and Jepson Herbariumhttp://ucjeps.berkeley.edu/
USDA Germplasm Resources Information Networkhttp://www.ars-grin.gov/
Washington State Noxious Weed Control Board (NWCB)http://www.nwcb.wa.gov/

Contributors

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18/08/09 Original text by:

Bitty Roy, University of Oregon, Biology Department, University of Oregon, Eugene, OR 97403-1210, USA

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