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Lolium
Perennial ryegrass (Lolium perenne)
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Monocots
Clade: Commelinids
Order: Poales
Family: Poaceae
Subfamily: Pooideae
Supertribe: Poodae
Tribe: Poeae
Subtribe: Loliinae
Genus: Lolium
L.
Synonyms[1]
  • Arthrochortus Lowe
  • Bucetum Parn.
  • Craepalia Schrank
  • Crypturus Link
  • Drymonaetes Ehrh.
  • Gnomonia Lunell
  • Loliola Dubois
  • Micropyropsis Romero Zarco & Cabezudo
  • Schedonorus P.Beauv.

Lolium is a genus of tufted grasses in the bluegrass subfamily (Pooideae).[2][3] It is often called ryegrass, but this term is sometimes used to refer to grasses in other genera.

They are characterized by bunch-like growth habits. Lolium is native to Europe, Asia and northern Africa, as well as being cultivated and naturalized in Australia, the Americas, and various oceanic islands. Ryegrasses are naturally diploid, with 2n=14, and are closely related to the fescues (Festuca).[4][5][6][7]

Ryegrass should not be confused with rye, which is a grain crop.

Species

[edit]

As of February 2024 the species of Lolium listed by Plants of the World Online include:[1]

Formerly included

Several former Lolium species now regarded as part of other genera: Castellia, Enteropogon, × Festulolium, Hainardia, Lepturus, Melica, and Vulpia.[8]

Cultivation and uses

[edit]

Lolium contains some species which are important grasses for lawns, and as pasture and for grazing and hay for livestock, being a highly nutritious stock feed. Ryegrasses are also used in soil erosion control programs. It is the principal grazing grass in New Zealand where some 10 million kilograms of certified seed are produced every year. There is a large range of cultivars. The primary species found worldwide and used both for lawns and as a forage crop is perennial ryegrass (Lolium perenne). Like many cool-season grasses of the Poaceae, it harbors a symbiotic fungal endophyte, either Epichloë or its close relative Neotyphodium, both of which are members of the fungal family Clavicipitaceae.[10][11]

Some species, particularly L. temulentum, are weeds which can have a severe impact on the production of wheat and other crops. Annual ryegrass (L. rigidum) is one of the most serious and costly weeds of cropping systems in southern Australia, and herbicide resistance is a frequent problem.[12] Ryegrass pollen is also one of the major causes of hay fever. Tennis courts, including those at the All England Lawn Tennis and Croquet Club, the venue for Wimbledon, are planted with ryegrass.[13][14] Glyphosate-resistant Lolium has been reported from Mississippi in 2006.[15]

References

[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Lolium

View on Grokipedia
from Grokipedia
Lolium is a genus of approximately 8–10 species of cool-season, annual and perennial grasses in the family Poaceae, subfamily Pooideae, tribe Poeae, and subtribe Loliinae. Native primarily to Europe, temperate Asia, and North Africa, the genus has been widely introduced to other regions, including North and South America, Australia, and New Zealand, where it is commonly cultivated. Members of Lolium are typically cespitose (tufted) or sometimes rhizomatous, with slender to stout culms that grow 10–150 cm tall, erect to decumbent, and featuring open sheaths, membranous ligules up to 4 mm long, and flat, linear leaf blades. Their inflorescences are distinctive distichous spikes, with solitary, radially oriented, laterally compressed spikelets containing 2–22 florets, the distal ones often reduced, and lower glumes typically absent except in terminal positions. The five main species—L. perenne (perennial ryegrass), L. multiflorum (Italian ryegrass), L. rigidum (stiff ryegrass), L. temulentum (darnel), and L. persicum—are all diploids with 2n=14 chromosomes and exhibit morphological intergradation, leading to taxonomic debates about merging Lolium with related genera like Festuca or Schedonorus. Species in the genus are renowned for their high productivity, nutritional quality, and adaptability to diverse environments, making them economically vital as forage, pasture, and turfgrasses in temperate regions worldwide. Perennial ryegrass (L. perenne) and Italian ryegrass (L. multiflorum) are the most cultivated, providing rapid establishment, excellent palatability for livestock, and soil stabilization benefits, though they can become invasive in some introduced areas. Other species like L. temulentum have historical notoriety as weeds contaminating grain crops, with seeds sometimes containing toxic ergot alkaloids from fungal infection. Taxonomically, Lolium is closely allied to Festuca, sharing phylogenetic ties within the Loliinae subtribe, and molecular studies support potential inclusion in an expanded Festuca genus.

Taxonomy and etymology

Etymology

The genus name Lolium derives from the Latin lolium, an ancient term for darnel (Lolium temulentum), a poisonous grass weed that contaminated grain fields in antiquity. This classical name, denoting a troublesome and inedible contaminant in crops, was formalized by in his seminal work (1753), where he established Lolium as the generic name for tufted grasses including ryegrasses. In Roman agriculture, lolium specifically referred to darnel as a harmful intruder in and fields, known for its toxic seeds that could induce intoxication, , or even blindness when accidentally milled with grain. The plant is mentioned in classical , such as Virgil's and , where it symbolizes agricultural misfortune and degeneration of crops, and in Pliny the Elder's , which describes its pestilent nature and occasional medicinal use despite its dangers. The term also carries biblical significance, with lolium (as darnel) identified as the "tares" (zizania in Greek) in the (:24–30), representing weeds sown by an enemy among wheat to illustrate and eschatological themes. No modern taxonomic revisions have altered the validity or application of the name Lolium, which continues to encompass 32 accepted species in the family.

Classification and synonyms

Lolium is classified within the kingdom Plantae, phylum Tracheophyta, class , subclass Commelinidae, order , family , subfamily , tribe Poeae, and L. (1753). Species in the Lolium are diploid with a base number of 2n=142n = 14. Historical synonyms for the include Arthrochortus Lowe, Bucetum Parn., Craepalia Schrank, and Crypturus Link. The is closely related to L., with molecular evidence indicating that Lolium is embedded within the Festuca lineage; some species have been historically transferred to other genera such as Castellia T.D.McMahon or Enteropogon Endl. Recent taxonomic treatments recognize 32 accepted species in the genus as of 2024, based on the World Checklist of Vascular Plants and regional updates.

Description

Morphology

Lolium species are annual or tufted (cespitose) grasses, typically 10–150 cm tall, with slender to stout, erect to decumbent culms and a that supports tillering growth. Perennials often form dense bunches through vegetative tillering, while annuals exhibit more ephemeral growth. Vegetatively, Lolium plants feature linear leaf blades that are flat or rolled, rolled in the bud stage, with sheaths that are open, rounded, and generally glabrous or slightly scabrous. Auricles are often present at the leaf sheath-blade , appearing as small, falcate, or clasping structures, though absent in some species; ligules are membranous, 0.5–4 mm long, and glabrous. The surfaces of stems and leaves are typically glabrous or scabrous, contributing to their smooth texture. The is a solitary, terminal, stiff, two-sided spike (distichous ) 5–30 cm long, with arranged edgewise in two opposite rows along the rachis, often sunken into its hollows. Each is laterally compressed, sessile, and contains 2–22 florets, with the lower typically absent except in the terminal ; lemmas are membranous to coriaceous, 5–9-nerved, and either awned (with straight, subterminal awns) or awnless depending on the . Paleas are membranous with ciliolate keels, and occurs above the and beneath successive florets. Distinguishing morphological traits of the genus include the two-ranked arrangement of spikelets on a compressed rachis, the radial orientation of spikelets to rachis concavities, and the overall lack of branches in the , setting Lolium apart from related genera like . are dorsally compressed, oblong to ovate, with a linear hilum in a longitudinal furrow and embryos comprising 1/5 to 1/3 of the length.

Reproduction and life cycle

Lolium species exhibit diverse life cycles, ranging from to forms, which influence their reproductive strategies. species, such as , complete their life cycle in a single through production and are predominantly self-pollinating via cleistogamous flowers, where occurs within unopened florets to ensure self-fertilization. In contrast, species like persist for multiple years, up to 30 or more, relying on both and vegetative propagation through tillering and short rhizomes, which allow clonal spread and persistence in established stands. Italian ryegrass (), often treated as an or short-lived , primarily reproduces by but can overwinter in favorable conditions. Reproduction in Lolium is predominantly sexual and anemophilous, with facilitating cross-pollination among species. Most species display gametophytic controlled by a two-locus , promoting and high , though rare self-fertilization can occur in hermaphroditic flowers under stress or in self-compatible variants. Flowering typically occurs from late spring to summer, triggered by and cool, moist conditions, with spikelets opening for during midday or early afternoon; optimal temperatures for flowering range from 15–25°C, and may become dormant in hot, dry summers. In cleistogamous annuals like L. temulentum, flowers remain closed, bypassing the need for dispersal and ensuring reproductive assurance in sparse populations. Seed production is prolific across the , supporting rapid establishment, with yields reaching up to 45,000 per square meter in dense stands of species like Lolium rigidum. exhibit primary in fresh harvests, varying from 0% to 70% , which is released through after-ripening under dry, warm conditions or cold stratification; this mechanism prevents premature and enhances survival in variable environments. is favored by cool, moist conditions above 5°C, with optimal rates at 20–26°C under alternating temperatures and light exposure, particularly for shallowly buried (1–5 cm depth), leading to seedling emergence within weeks in suitable habitats. Seed banks persist for 16–18 months typically, with low viability beyond four years. Hybridization is a key aspect of Lolium , particularly with closely related species, resulting in fertile intergeneric hybrids known as Festulolium. These crosses, such as between and Festuca pratensis, involve mixing and are facilitated by overlapping flowering periods and wind , producing viable progeny with combined traits like enhanced stress tolerance. Interspecific hybrids within Lolium, such as L. multiflorum × L. perenne, also occur readily and contribute to genetic variation in natural and agricultural settings.

Species

Accepted species

The genus Lolium comprises five accepted species, all of which are grasses in the family Poaceae, characterized by their temperate distributions and agricultural significance. Lolium perenne L., known as perennial ryegrass, is a perennial bunchgrass native to Europe, temperate Asia, and North Africa. It typically reaches heights of 50–100 cm and features spikelets with awned lemmas, making it a prominent species for forage and turf applications due to its persistence and nutritional value. Lolium multiflorum Lam., or Italian ryegrass, is an or biennial grass native to and . It shares morphological similarities with L. perenne, including a similar height range, but is shorter-lived and valued for its rapid growth in forage production. Lolium rigidum Gaudin, stiff ryegrass, is an species native to the Mediterranean region, encompassing , , and western Asia. Growing 20–60 cm tall, it exhibits variable awns on its spikelets and is recognized for its adaptability in disturbed habitats. Lolium temulentum L., commonly called darnel, is an annual grass native to and . Its spikelets bear long awns, and historically, its seeds have been noted for when contaminating crops. Lolium persicum Boiss. & Hohen., known as Persian darnel, is an annual grass native to southwest , from to and the . It grows 14–60 cm tall, with spikelets 9–27 mm long containing 4–9 florets and awned lemmas up to 18 mm; it is a in fields and waste places, introduced to where it ranks among major crop weeds in . As of 2025, none of the Lolium species are considered endangered; they are widely cultivated, naturalized, or established as common elements in agricultural and disturbed landscapes globally.

Formerly included species

Several species once classified within the genus Lolium have been reclassified to other genera based on molecular phylogenetic analyses and morphological evidence, refining the circumscription of Lolium within the subtribe Loliinae of the Pooideae subfamily. For example, Lolium arundinaceum (formerly known as tall fescue) is now placed in the genus Schedonorus as Schedonorus arundinaceus, reflecting its polyploid nature (2n=42–84) and distinct evolutionary position outside the core Lolium clade (diploids with 2n=14), as determined by ITS and other genetic data that highlight differences in genome size and inflorescence morphology. These reclassifications stem from broader analyses showing that such species exhibit polyphyletic relationships with true Lolium, often aligning more closely with broad-leaved fescues or other grass lineages based on combined nuclear and chloroplast data. The taxonomic revisions have significantly narrowed Lolium to its monophyletic core, now comprising only five accepted species (L. multiflorum, L. perenne, L. persicum, L. rigidum, and L. temulentum), as recognized in treatments like the Flora of North America, enhancing clarity in grass systematics and reducing historical misplacements driven by superficial similarities in spikelet structure.

Distribution and habitat

Native distribution

The genus Lolium is native to , extending to and Korea, and from (including ) to the and the . These regions encompass temperate and Mediterranean climates, where the genus has evolved in diverse Eurasian and African landscapes. Among the accepted species, L. perenne is native to , , , , and the Himalaya. L. multiflorum occurs natively in , the , the Mediterranean Basin, , and the Himalaya. L. rigidum is found in , southern and central , and extends to the western Himalaya and parts of ( and provinces). L. temulentum has a native range covering , , and temperate . L. persicum is native to the , from through to northern and the western Himalaya. In their native habitats, Lolium species primarily occupy temperate grasslands, disturbed soils, and roadsides, thriving in open, fertile areas with moderate moisture. These grasses employ the cool-season C3 photosynthetic pathway, which supports efficient growth in cooler temperatures typical of their Eurasian and African origins.

Introduced ranges

Lolium species, particularly L. perenne and L. multiflorum, have been widely introduced to regions beyond their native Eurasian and ranges through human activities, establishing populations in , , , , , and various oceanic islands. These introductions began with and pastoralists who transported the grasses to support agriculture in new territories, including the , , and . The primary mechanisms of spread include intentional sowing for , turf, and cover cropping, as well as accidental dispersal via contaminated commercial , shipments, and . In , for example, ryegrass is deliberately sown across extensive areas, with annual production exceeding 29 million kilograms of in recent years to meet domestic and export demands. As of 2025, Lolium species are naturalized across temperate zones globally, forming a key component of managed pastures that span millions of hectares, including over 1.7 million effective hectares of farmland in alone where perennial ryegrass predominates. This widespread establishment reflects their adaptation to human-modified landscapes in cool, moist climates suitable for .

Ecology

Interactions with animals and humans

Lolium species, particularly L. perenne and L. multiflorum, serve as important forage for grazing animals such as cattle and sheep, owing to their high palatability, digestibility, and nutritional value including elevated protein and energy content. These grasses support livestock production in temperate regions, where they are grazed at heights of 6-8 inches to optimize regrowth and productivity. However, L. temulentum (darnel) presents risks due to loline alkaloids produced by endophytic fungi and ergot alkaloids from associated fungal infections such as Claviceps purpurea, which can cause fatal poisoning in livestock through symptoms such as neurological distress and digestive issues. In humans, Lolium pollen, especially from L. perenne, acts as a potent aeroallergen responsible for hay fever () and can exacerbate , with symptoms including , eye irritation, and respiratory distress during peak seasons from May to July. Historically, L. temulentum contaminated grain crops like , leading to darnel intoxication when ingested in or porridge, causing effects such as drowsiness, blurred vision, and skin ulcers as documented in ancient texts from and . This weed's of cereals made separation difficult, resulting in widespread in antiquity across regions like and . Within grassland ecosystems, Lolium species contribute to structure by providing cover and that support such as small mammals and , enhancing local food webs in temperate areas. However, as introduced plants in many regions, they engage in strong interactions with , often reducing native perennial grass productivity and limiting by dominating resources in open s like roadsides and abandoned fields. This competition favors exotic grasses over natives, altering composition in grasslands.

Endophytes and symbioses

Lolium species, particularly L. perenne (perennial ryegrass), commonly host symbiotic fungal from the genus Epichloë (previously classified under Neotyphodium), forming mutualistic, systemic, and often obligate relationships that colonize the intercellular spaces of host tissues. These endophytes are primarily transmitted vertically through seeds, achieving near 100% transmission efficiency in natural associations, though rates can be lower in certain European populations due to environmental factors. In L. perenne, the specific endophyte Epichloë festucae var. lolii is prevalent, enhancing host fitness without causing visible symptoms in asymptomatic infections. The symbiosis provides significant benefits to the host grass, including improved tolerance to biotic and abiotic stresses. Endophyte-infected exhibit enhanced resistance to insect pests, such as and Argentine stem , through the production of alkaloids like peramine and lolines, which deter herbivory without broad toxicity to non-target organisms. Additionally, the association boosts drought tolerance by promoting greater root biomass, better stomatal regulation, and accumulation of osmolytes like and , allowing infected L. perenne to maintain productivity under water stress. These defensive compounds, including alkaloids and indole-diterpenes, contribute to overall resilience in temperate grasslands. However, the alkaloids can pose risks to grazing , leading to conditions such as ryegrass staggers from lolitrem B and ergovaline-induced (analogous to fescue toxicosis), which affect health and productivity. Infection rates in Lolium cultivars vary widely, typically ranging from 20% to 90% in temperate regions, with commercial seed batches often requiring over 80% infection for endophyte-enhanced varieties. Since the , breeding programs have developed endophyte-free cultivars of L. perenne to mitigate risks, alongside novel endophytes engineered to produce beneficial alkaloids while minimizing animal harm, such as the AR1 strain released in 2001.

Cultivation and uses

Forage and pasture

Lolium perenne (perennial ryegrass) and Lolium multiflorum (Italian or annual ryegrass) are the primary used in and systems due to their rapid growth, palatability, and high nutritional quality. These grasses provide digestible exceeding 70% and crude protein levels typically ranging from 15% to 25% in vegetative stages, making them suitable for and production. They are often sown in mixtures with legumes such as white clover (Trifolium repens) or red clover (Trifolium pratense) to enhance , improve overall productivity, and balance animal while reducing needs. The cultivation of Lolium species for forage has a long history in European agriculture, with records of deliberate sowing for pastures dating back to the 17th century, when L. perenne was recognized as one of the first grasses intentionally planted for livestock feed. Over time, these grasses spread globally through agricultural expansion, becoming staples in temperate regions for their adaptability to cool, moist climates and ability to support intensive grazing. Modern breeding programs have focused on improving yield, disease resistance, and persistence, with selections for higher dry matter production—up to 3.2% per decade in late-heading cultivars—and enhanced tolerance to pests and environmental stresses. Endophyte-enhanced varieties, incorporating symbiotic fungi like Epichloë festucae var. lolii, have been developed to boost resistance and without compromising animal health, further advancing their agricultural utility. Globally, Lolium species occupy a significant portion of temperate , estimated at around 14 million hectares for L. perenne alone, comparable to other major forage grasses. Annual seed production for ryegrass exceeds 200,000 tons worldwide, supporting widespread establishment in regions like , , and .

Turf and lawns

Lolium perenne, commonly known as perennial ryegrass, is widely utilized in turf applications for its rapid establishment and high wear tolerance, making it a preferred choice for lawns, sports fields, and golf courses. This cool-season grass germinates quickly, often within 5 to 10 days under optimal conditions, allowing for swift coverage and stabilization of bare areas. Its bunch-type growth habit and fine leaf texture contribute to a uniform, dense sward that withstands foot traffic and mechanical stress effectively. In high-profile settings, such as the grass courts at Wimbledon, perennial ryegrass is sown exclusively to enhance durability and presentation. Since 2001, the All England Lawn and Club has used 100% L. perenne at an 8 mm cut height, following a previous mix that included 70% ryegrass and 30% creeping red fescue; this shift improves sward strength against the rigors of professional play. The grass's dark green color and fine texture provide an aesthetically pleasing surface, while its compatibility with fine fescues allows for blended turfs that offer superior resilience in mixed stands, combining ryegrass's quick recovery with fescues' shade and drought tolerance. Beyond ornamental and recreational uses, L. perenne plays a key role in , particularly in temperate climates where its optimal growth occurs at 18-20°C. It is commonly hydroseeded on eroded slopes, mine spoils, and roadside embankments to stabilize soil rapidly through its extensive . In revegetation projects, such as streambank stabilization and post-fire recovery in regions like the , perennial ryegrass establishes cover that reduces sediment loss and supports long-term site restoration when seeded at rates around 29 pounds per acre.

As a weed

Invasive species

Species of the genus Lolium, particularly L. rigidum (rigid ryegrass) and L. multiflorum (Italian ryegrass), are recognized as invasive in several non-native regions, including , , and , where they displace native vegetation in grasslands and other open habitats. In , L. rigidum invades natural ecosystems and is classified as an environmental in states such as Victoria and . Similarly, L. multiflorum is rated as a moderate invasive by the California Invasive Plant Council, occurring widely except in the and deserts, and outcompeting natives in fertile or clay soils. In , L. multiflorum naturalizes aggressively, reducing and diversity in grasslands. These invasions often stem from initial introductions as or cover crops, leading to unintended spread into conservation areas. The invasive potential of Lolium species is driven by prolific seed production and persistence, enabling rapid of disturbed sites. A single can produce up to 45,000 , with dense stands yielding 1000 or more seeds per plant, facilitating widespread dispersal via , , machinery, and animal vectors. exhibit and remain viable in the for 2-3 years, with annual decline rates around 70%, allowing staggered that enhances establishment in variable conditions. These grasses thrive in disturbed habitats like roadsides, overgrazed pastures, and post-fire landscapes, where they form dense monocultures that suppress native recruitment. Management of invasive Lolium emphasizes prevention and mechanical methods in sensitive conservation areas to minimize . Strategies include early detection, habitat restoration, and avoiding introductions through clean equipment and seed sources. Mechanical controls such as mowing, , or hand-pulling before seed set can reduce populations, though repeated applications are needed due to the plant's tolerance and resprouting ability. These invasions alter regimes by increasing fine fuel loads and continuity, promoting more frequent and intense that favor annual grasses over woody or natives, further degrading structure and function.

Herbicide resistance

Glyphosate-resistant biotypes of Lolium multiflorum (Italian ryegrass) were first identified in 2005 in , , marking an early case of resistance in North American populations of the genus. This resistance has since spread, with multiple herbicide-resistant populations now prevalent in the region. In contrast, L. rigidum (annual ryegrass) in exhibited the earliest global glyphosate resistance in 1996, and by the 2020s, widespread multiple resistance to acetyl-CoA carboxylase (ACCase) inhibitors and acetolactate synthase () inhibitors had evolved in Australian populations due to intensive use in cropping systems. These cases highlight Lolium ' propensity for rapid resistance under selective pressure from repeated applications. Resistance mechanisms in Lolium populations primarily involve target-site and enhanced metabolic . Target-site resistance includes point mutations in genes encoding EPSPS (for ), ACCase (for herbicides), and ALS (for group 2 herbicides), which alter the and reduce efficacy. Enhanced metabolism, often mediated by monooxygenases, allows plants to degrade herbicides before they reach lethal concentrations, contributing to non-target-site resistance. By 2025, over 125 unique cases of resistance across various modes of action have been documented globally in Lolium species, with multiple mechanisms co-occurring in many populations. In early 2025, the first case of resistance was confirmed in L. multiflorum populations in the . The evolution of resistance imposes significant economic burdens on farmers, with annual revenue losses exceeding AUD $93 million in Australian grain production alone due to increased control costs and yield reductions from uncontrolled L. rigidum. In response, integrated weed management strategies—combining diverse rotations, cultural practices like and , and mechanical control—are recommended to delay further resistance development and maintain effective suppression. Invasive spread of resistant biotypes can exacerbate these challenges by facilitating across agricultural landscapes.

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  58. https://www.fs.usda.gov/database/feis/plants/graminoid/lolperp/all.html
  59. https://www.cottoninc.com/wp-content/uploads/2017/03/Programs-For-Managing-Glyphosate.pdf
  60. https://www.cambridge.org/core/services/aop-cambridge-core/content/view/28A99023A321046E76CEA9FBCF289B4C/S0043174500018919a.pdf/decade_of_glyphosateresistant_lolium_around_the_world_mechanisms_genes_fitness_and_agronomic_management.pdf
  61. https://pmc.ncbi.nlm.nih.gov/articles/PMC7769757/
  62. https://www.frontiersin.org/journals/plant-science/articles/10.3389/fpls.2020.609209/full
  63. https://www.seedworld.com/europe/2025/01/16/first-case-of-glyphosate-resistance-in-a-uk-weed-species-confirmed/
  64. https://pmc.ncbi.nlm.nih.gov/articles/PMC8400064/
  65. https://www.publish.csiro.au/cp/pdf/CP22370
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