Brachypodium sylvaticum (slender false brome)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Plant Type
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Habitat List
- Hosts/Species Affected
- Host Plants and Other Plants Affected
- Growth Stages
- Biology and Ecology
- Latitude/Altitude Ranges
- Soil Tolerances
- Natural enemies
- Notes on Natural Enemies
- Means of Movement and Dispersal
- Pathway Causes
- Pathway Vectors
- Impact Summary
- Economic Impact
- Environmental Impact
- Threatened Species
- Risk and Impact Factors
- Uses List
- Detection and Inspection
- Similarities to Other Species/Conditions
- Prevention and Control
- Gaps in Knowledge/Research Needs
- Links to Websites
- Distribution Maps
Don't need the entire report?
Generate a print friendly version containing only the sections you need.Generate report
PicturesTop of page
IdentityTop of page
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
- BRCSI (Brachypodium sylvaticum)
Summary of InvasivenessTop of page
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 TreeTop of page
- Domain: Eukaryota
- Kingdom: Plantae
- Phylum: Spermatophyta
- Subphylum: Angiospermae
- Class: Monocotyledonae
- Order: Cyperales
- Family: Poaceae
- Genus: Brachypodium
- Species: Brachypodium sylvaticum
Notes on Taxonomy and NomenclatureTop of page
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).
DescriptionTop of page
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.
Plant TypeTop of page
DistributionTop of page
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 TableTop of page
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.Last updated: 23 Apr 2020
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Algeria||Present||Native||Original citation: Euro+Med Plantbase (2009)|
|Morocco||Present||Native||Original citation: Euro+Med Plantbase (2009)|
|Tunisia||Present||Native||Original citation: Euro+Med Plantbase (2009)|
|Azerbaijan||Present||Native||Original citation: Euro+Med Plantbase (2009)|
|Bhutan||Present, Widespread||Native||Mountain slopes, forest understory|
|China||Present, Widespread||Native||Mountain slopes, forest understory|
|-Anhui||Present, Widespread||Native||Mountain slopes, forest understory|
|-Gansu||Present, Widespread||Native||Mountain slopes, forest understory|
|-Guizhou||Present, Widespread||Native||Mountain slopes, forest understory|
|-Hunan||Present, Widespread||Native||Mountain slopes, forest understory|
|-Jiangsu||Present, Widespread||Native||Mountain slopes, forest understory|
|-Liaoning||Present, Widespread||Native||Mountain slopes, forest understory|
|-Qinghai||Present, Widespread||Native||Mountain slopes, forest understory|
|-Shanxi||Present, Widespread||Native||Mountain slopes, forest understory|
|-Sichuan||Present, Widespread||Native||Mountain slopes, forest understory|
|-Tibet||Present, Widespread||Native||Mountain slopes, forest understory|
|-Xinjiang||Present, Widespread||Native||Mountain slopes, forest understory|
|-Yunnan||Present, Widespread||Native||Mountain slopes, forest understory|
|-Zhejiang||Present, Widespread||Native||Mountain slopes, forest understory|
|India||Present, Widespread||Native||Northern India|
|Indonesia||Present, Widespread||Native||Mountain slopes, forest understory|
|-Irian Jaya||Present||Native||1500-3830 m|
|-Java||Present||Native||Glabrous form called B. sylvaticum var. luzoniense|
|-Lesser Sunda Islands||Present||Native||1500-3830 m|
|-Maluku Islands||Present||Native||1500-3830 m|
|-Sulawesi||Present, Few occurrences||Native||1500-3830 m|
|-Sumatra||Present||Native||Glabrous form called B. sylvaticum var. luzoniense|
|Iran||Present||Native||Shady woods, streamside, 300-2300 m|
|Iraq||Present||Native||Damp, shady places in the mountains, 700-1500 m|
|Israel||Present||Native||Originally listed incorrectly as B. pinnatum|
|Kyrgyzstan||Present||Native||Mountain slopes, forest understory|
|Lebanon||Present||Native||Original citation: Euro+Med Plantbase (2009)|
|Nepal||Present||Native||Mountain slopes, forest understory|
|Pakistan||Present||Native||Shady woods, streamside, 900-2100 m|
|Philippines||Present||Native||Glabrous form called B. sylvaticum var. luzoniense|
|Sri Lanka||Present||Horton plains, 7000 feet|
|Syria||Present||Native||Original citation: Euro+Med Plantbase (2009)|
|Taiwan||Present||Native||Mountain slopes, forest understory|
|Tajikistan||Present||Native||Mountain slopes, forest understory|
|Turkey||Present, Widespread||Native||10-2440 m forested slopes, meadows, river terraces|
|Turkmenistan||Present||Native||Shady woods, streamside|
|United Arab Emirates||Present||Native||Bahrain|
|Uzbekistan||Present, Widespread||Native||Mountain slopes, forest understory|
|Albania||Present||Native||Original citation: Euro+Med Plantbase (2009)|
|Andorra||Present, Localized||Native||Submontane-montane, shaded, mild clearings in the domain of Pinus sylvestris forests, irregularly mown or grazed|
|Belgium||Present||Native||Original citation: Euro+Med Plantbase (2009)|
|Cyprus||Present||Native||Original citation: Euro+Med Plantbase (2009)|
|Denmark||Present||Native||Original citation: Euro+Med Plantbase (2009)|
|-Corsica||Present||Native||Original citation: Euro+Med Plantbase (2009)|
|Malta||Present||Native||Original citation: Euro+Med Plantbase (2009)|
|Moldova||Present||Native||Original citation: Euro+Med Plantbase (2009)|
|Norway||Present||Native||Original citation: Euro+Med Plantbase (2009)|
|Portugal||Present||Native||Original citation: Euro+Med Plantbase (2009)|
|-Azores||Present||Native||Original citation: Euro+Med Plantbase (2009)|
|-Madeira||Present||Native||Original citation: Euro+Med Plantbase (2009)|
|Russia||Present||Native||Lists many varieties|
|-Central Russia||Present||Native||Lists many varieties|
|-Eastern Siberia||Present||Native||Lists many varieties|
|-Northern Russia||Present||Native||Lists many varieties|
|-Russian Far East||Present||Native||Lists many varieties|
|-Southern Russia||Present||Native||Lists many varieties|
|-Western Siberia||Present||Native||Lists many varieties|
|Serbia||Present||Native||Original citation: Euro+Med Plantbase (2009)|
|Serbia and Montenegro||Present||Native||Original citation: Euro+Med Plantbase (2009)|
|Slovakia||Present||Native||Original citation: Euro+Med Plantbase (2009)|
|-Balearic Islands||Present||Native||Original citation: Euro+Med Plantbase (2009)|
|-Canary Islands||Present||Native||Original citation: Euro+Med Plantbase (2009)|
|Switzerland||Present, Widespread||Native||Montane to subalpine|
|Ukraine||Present||Native||Lists many varieties|
|United Kingdom||Present, Widespread||Native||Native; 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 Islands||Present||Native||Original citation: Euro+Med Plantbase (2009)|
|Barbados||Absent, Intercepted only|
|Canada||Present||Present based on regional distribution.|
|-Alberta||Absent, Intercepted only|
|-British Columbia||Absent, Intercepted only|
|-Newfoundland and Labrador||Absent, Intercepted only|
|Costa Rica||Absent, Intercepted only|
|Nicaragua||Absent, Intercepted only|
|United States||Present, Few occurrences|
|-California||Present, Localized||2009||Introduced||2003||Invasive||Found in several locations in San Mateo County, including near Stanford and in the Santa Cruz Mountains|
|-Missouri||Present, Localized||Introduced||There are two specimens at Missouri Botanic Gardens from Missouri, one from St Louis City (1977) and one without date or locality|
|-Oregon||Present, Widespread||2009||Introduced||Naturalized||Thoroughly 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); First reported: < 1939|
|-Utah||Absent, Formerly present||Planted in the 1970's, but not found since then|
|-Virginia||Present, Localized||2009||Introduced||1992||Invasive||First collected by C. E. Stevens, Aug. 1992 at Preston Place, Charlottesville. Has persisted and slowly spread|
|-Washington||Present, Localized||2009||Introduced||2007||Invasive||The first sighting in Washington State was in 2007 from Beacon Rock State Park, Skamania County, in the Columbia River Gorge|
|Australia||Present||Present based on regional distribution.|
|-Northern Territory||Absent, Never occurred|
|-Victoria||Present, Localized||Invasive||Sparingly naturalized|
|New Zealand||Present, Widespread||Introduced||Invasive||Lowland to montane in shade of trees or shrubs|
|Papua New Guinea||Present||Native||Glabrous form called B. sylvaticum var. luzoniense|
|Ecuador||Absent, Intercepted only|
History of Introduction and SpreadTop of page
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).
IntroductionsTop of page
|Introduced to||Introduced from||Year||Reason||Introduced by||Established in wild through||References||Notes|
|Natural reproduction||Continuous restocking|
|Argentina||< 1952||Yes||No||Soreng et al. (2009)|
|New Zealand||Yes||No||Edgar and Connor (2000)|
|USA||Europe||<1939||Forage (pathway cause)
Horticulture (pathway cause)
Risk of IntroductionTop of page
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.
HabitatTop of page
- 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).
Habitat ListTop of page
|Terrestrial||Managed||Managed forests, plantations and orchards||Principal habitat||Harmful (pest or invasive)|
|Terrestrial||Managed||Managed forests, plantations and orchards||Principal habitat||Natural|
|Terrestrial||Managed||Managed grasslands (grazing systems)||Principal habitat||Harmful (pest or invasive)|
|Terrestrial||Managed||Managed grasslands (grazing systems)||Principal habitat||Natural|
|Terrestrial||Managed||Disturbed areas||Secondary/tolerated habitat||Harmful (pest or invasive)|
|Terrestrial||Managed||Disturbed areas||Secondary/tolerated habitat||Natural|
|Terrestrial||Managed||Rail / roadsides||Principal habitat||Harmful (pest or invasive)|
|Terrestrial||Managed||Rail / roadsides||Principal habitat||Natural|
|Terrestrial||Managed||Urban / peri-urban areas||Secondary/tolerated habitat||Harmful (pest or invasive)|
|Terrestrial||Managed||Urban / peri-urban areas||Secondary/tolerated habitat||Natural|
|Terrestrial||Natural / Semi-natural||Natural forests||Principal habitat||Harmful (pest or invasive)|
|Terrestrial||Natural / Semi-natural||Natural forests||Principal habitat||Natural|
|Terrestrial||Natural / Semi-natural||Natural grasslands||Secondary/tolerated habitat||Harmful (pest or invasive)|
|Terrestrial||Natural / Semi-natural||Natural grasslands||Secondary/tolerated habitat||Natural|
|Terrestrial||Natural / Semi-natural||Riverbanks||Principal habitat||Harmful (pest or invasive)|
|Terrestrial||Natural / Semi-natural||Riverbanks||Principal habitat||Natural|
|Terrestrial||Natural / Semi-natural||Wetlands||Principal habitat||Harmful (pest or invasive)|
|Terrestrial||Natural / Semi-natural||Wetlands||Principal habitat||Natural|
|Terrestrial||Natural / Semi-natural||Cold lands / tundra||Present, no further details|
|Freshwater||Irrigation channels||Principal habitat||Harmful (pest or invasive)|
|Freshwater||Irrigation channels||Principal habitat||Natural|
|Freshwater||Lakes||Principal habitat||Harmful (pest or invasive)|
|Freshwater||Reservoirs||Principal habitat||Harmful (pest or invasive)|
|Freshwater||Rivers / streams||Principal habitat||Harmful (pest or invasive)|
|Freshwater||Rivers / streams||Principal habitat||Natural|
|Freshwater||Ponds||Principal habitat||Harmful (pest or invasive)|
Hosts/Species AffectedTop of page
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).
Host Plants and Other Plants AffectedTop of page
|Bromus carinatus (California brome grass)||Poaceae||Wild host|
|Danthonia californica||Poaceae||Wild host|
|Elymus glaucus||Poaceae||Wild host|
|Festuca roemeri||Poaceae||Wild host|
|Lupinus sulphureus||Wild host|
|Melica subulata||Poaceae||Wild host|
|Pseudotsuga menziesii (Douglas-fir)||Pinaceae||Wild host|
|Sequoia sempervirens (coast redwood)||Taxodiaceae||Wild host|
Growth StagesTop of page
Biology and EcologyTop of page
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).
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.
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.
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.
ClimateTop of page
|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 RangesTop of page
|Latitude North (°N)||Latitude South (°S)||Altitude Lower (m)||Altitude Upper (m)|
RainfallTop of page
|Parameter||Lower limit||Upper limit||Description|
|Dry season duration||1||4||number of consecutive months with <40 mm rainfall|
Soil TolerancesTop of page
Natural enemiesTop of page
|Natural enemy||Type||Life stages||Specificity||References||Biological control in||Biological control on|
|Agromyza albipennis||Herbivore||Leaves||not specific|
|Carterocephalus palaemon||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 EnemiesTop of page
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 DispersalTop of page
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.
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.
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.
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.
Pathway CausesTop of page
Pathway VectorsTop of page
|Aircraft||Likely infrequent. Seeds were likely imported via airmail for USDA work||Yes||Hull AC Jr, 1974; Rosenthal et al., 2008|
|Clothing, footwear and possessions||Possible anytime||Yes||Yes||Heinken and Raudnitschka, 2002|
|Floating vegetation and debris||Frequency is not documented, but common along rivers and streams||Yes||Yes|
|Germplasm||Seeds were imported for USDA plantings, frequency unknown||Yes||Hull AC Jr, 1974; Rosenthal et al., 2008|
|Hides, trophies and feathers||Possible - seeds have been found on animals shot by hunters||Yes||Yes||Heinken and Raudnitschka, 2002|
|Livestock||Seeds, likely||Yes||Yes||Heinken and Raudnitschka, 2002|
|Machinery and equipment||Seeds, likely||Yes||Yes||False Brome Working Group, 2009; Fletcher, 2009|
|Mulch, straw, baskets and sod||seeds and stems possible||Yes|
|Pets and aquarium species||Seeds, likely. Not documented but given that ungulates carry seeds, dogs could too||Yes|
|Water||Likely. Commonly grows streamside||Yes||Yes|
Impact SummaryTop of page
Economic ImpactTop of page
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 ), 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 ImpactTop of page
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 SpeciesTop of page
|Threatened Species||Conservation Status||Where Threatened||Mechanism||References||Notes|
|Euphydryas editha taylori||National list(s)||Oregon||Competition - monopolizing resources; Competition - smothering||Severns and Warren, 2008|
|Icaricia icarioides fenderi||USA ESA listing as endangered species||Oregon||Competition - monopolizing resources; Competition - smothering||US Fish and Wildlife Service, 2006|
|Lupinus sulphureus||No Details||Oregon||Competition - monopolizing resources; Competition - smothering||US Fish and Wildlife Service, 2006|
|Eremophila alpestris strigata (streaked horned lark)||USA ESA listing as threatened species||Oregon; Washington||Ecosystem change / habitat alteration||US Fish and Wildlife Service, 2013|
|Lupinus oreganus var. kincaidii (Kincaid's lupine)||NatureServe; USA ESA listing as endangered species||Oregon; Washington||Competition - strangling||US Fish and Wildlife Service, 2006|
|Solidago houghtonii (Houghton's goldenrod)||NT (IUCN red list: Near threatened); USA ESA listing as threatened species||Ontario; Michigan||Competition - monopolizing resources; Ecosystem change / habitat alteration||US Fish and Wildlife Service, 2011|
Risk and Impact FactorsTop of page
- 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
- 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
- Competition - monopolizing resources
- Competition - smothering
- Competition - strangling
- Rapid growth
- 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
UsesTop of page
B. sylvaticum has economic value for three uses:
- 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)
Uses ListTop of page
Animal feed, fodder, forage
- Invertebrate food
- Botanical garden/zoo
- Laboratory use
- Research model
- Seed trade
Detection and InspectionTop of page
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/ConditionsTop of page
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 ControlTop of page
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.
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).
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 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).
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).
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 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 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.
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 NeedsTop of page
- 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.
ReferencesTop of page
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.
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
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.
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
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.
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
Clay K, 1990. Fungal endophytes of grasses. Annual Review of Ecology and Systematics, 21:275-295.
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
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/
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
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.
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
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.
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
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.
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.
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.
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
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.
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/
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
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
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
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
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
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
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.
Al-Eisawi D M, Al-Khader I A, 1998. Checklist of plants of the Hashemite Kingdom of Jordan. In: Checklist of plants of the Hashemite Kingdom of Jordan. Amman, Jordan: The General Corporation for the Environment Protection. unpaginated.
Anon, 1994. Catálogo de la familia Poaceae en la República Argentina. [ed. by Zuloaga F O, Nicora E G, Rúgolo de Agrasar Z E, Morrone O, Pensiero J F, Cialdella A M]. St. Louis, Missouri, USA: Missouri Botanical Garden. xi + 178 pp.
Anon, 2003. Manual de plantas de Costa Rica, Vol. 3. Monocotyledoneas (Orchidaceae-Zingiberaceae). [ed. by Hammel B E, Grayum M H, Herrara C, Zamora N]. St. Louis, Missouri, USA: Missouri Botanical Garden.
Anon, 2007. Flora of North America North of Mexico, Vol. 24. Magnoliophyta: Commelinidae (in part): Poaceae, part 1. [ed. by Barkworth M E, Capels K M, Long S, Anderton L K, Piep M B]. New York, New York, USA: Oxford University Press. unpaginated.
Australia, IBIS, 2009. Brachypodium sylvaticum. In: Australian Plant Name Index - Integrated Botanical Information System. Australia: Australian National Botanic Gardens, Australian National Herbarium. unpaginated. http://www.anbg.gov.au
Belarus, CBG, Belarus, NASB, Belarus, MSKH, 2009. Brachypodium sylvaticum. In: Plants of Belarus. Belarus: Herbarium of CBG NASB MSKH supported by the Ministry of Natural Resources and Environmental Protection of the Republic of Belarus. http://hbc.bas-net.by/plantae/eng/allplantras.php
Bor N L, 1970. Volume 70: Graminae. (Volume 70: Graminae.). In: Flora Iranica: Flora des Iranischen Hochlandes und der umrahmenden Gebirge; Persien, Afganistan, Teile von West-Pakistan, Nord-Iraq, Azerbaidjan, Turkmenistan, [ed. by Reichinger K H]. Graz, Austria: Akademische Druck und Verlagsanstalt.
CABI, Undated. Compendium record. Wallingford, UK: CABI
CABI, Undated a. CABI Compendium: Status inferred from regional distribution. Wallingford, UK: CABI
Miller B M, Aitken R J, Oldham M J, Reznicek A A, 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
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 DOI:10.1016/j.jnc.2008.08.001
Rosenthal D M, Ramakrishnan A P, Cruzan M B, 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 DOI:10.1111/j.1365-294X.2008.03844.x
Soreng R J, Davidse G, Peterson P M, Zuloaga F O, Judziewicz F J, Filgueiras T S, Morrone O, 2009. Brachypodium sylvaticum. In: Catalogue of new world grasses (Poaceae), http://mobot.mobot.org/W3T/Search/nwgc.html
USA, False Brome Working Group, 2009. Brachypodium sylvaticum. In: Brachypodium sylvaticum, USA: USDA Forest Service, USDI Bureau of Land Management, Oregon Department of Agriculture, US Army Corps of Engineers, OSU College of Forestry, Institute. unpaginated. http://www.appliedeco.org/invasive-species-resources/FBWG
ContributorsTop of page
18/08/09 Original text by:
Bitty Roy, University of Oregon, Biology Department, University of Oregon, Eugene, OR 97403-1210, USA
Distribution MapsTop of page
Select a dataset
CABI Summary Records
Unsupported Web Browser:
One or more of the features that are needed to show you the maps functionality are not available in the web browser that you are using.
Please consider upgrading your browser to the latest version or installing a new browser.
More information about modern web browsers can be found at http://browsehappy.com/