Plantago lanceolata L., Ribwort Plantain

Account Summary

Growth form and preferred habitats

P. lanceolata forms a rosette plant from a short, thick, erect rhizome. The root system is commonly represented by a few deep roots capable of descending over a metre and a mass of surface fibrous roots 5-10 cm in length suggesting physiological differentiation into water and nutrient-absorbing systems. The leaves are simple, 10-15(-45) cm long, 2-3.5 cm wide, linear-lanceolate to elliptic-spathulate, margins usually entire and with 3-5-longitudinal ribbed veins (raised on the leaf underside). It is the conspicuous ribs on the leaves that give the plant its English common name 'Ribwort Plantain'. When it has more linear leaves, the blade narrows gradually into the leaf base, the petiole length apparently depending on the height of the surrounding vegetation (Sagar & Harper 1964).

As with P. major (Greater Plantain), there is a great amount of phenotypic variation in many Ribwort Plantain characters and undoubtedly the species embraces a number of genetically different strains. Generally, the plant follows a perennial life history and reproductive strategy, but rarely (as in cultivation experiments), it can behave as either an annual or a biennial. Overwintering rosettes have wider leaves than spring and summer ones and those growing in exposed sites are more prostrate, so they can sometimes be confused with P. major.

P. lanceolata occurs in a very wide range of habitats and although it is largely absent from deeply shaded woodland, wetlands and the most acidic ground, it does commonly occur on their margins. The species is most commonly found in pastures, rough grassland and disturbed ground, including in gardens, lawns, roadsides and waste places. P. lanceolata is frost-tolerant and also very drought resistant. It competes best in drier environments and in hard-packed neutral to basic soils. It is also well able to cope with shade and can germinate and establish even when sited in closed turf vegetation among tall grasses (Blom 1978; Cavers et al. 1980).

Ribwort Plantain is fast growing and very competitive, being quickly able to exploit increased fertility either as a result of additional water or nutrients or both of these factors. It is also extremely successful in exploiting gaps in turf by both vegetative expansion and seedling recruitment from a persistent buried soil seed bank (Thompson et al. 1997). The deep taproot allows the plant to be highly drought-resistant and to survive on very dry soils, even in crevices in rocks and on walls. It is one of the world's most successful (or worst) weeds (Holm et al. 1977) and is ranked in the top dozen most successful colonising plant species in B & I. The established strategy is categorised as 'C-S-R', meaning it achieves a balance between or involving all three ecological strategies, Competitor, Stress-tolerator and Ruderal (Grime et al. 1988, 2007).

Variation

It is well recognised that P. lanceolata is extremely variable, much of the variation being with respect to the particular environment in which it grows (ie it is phenotypically plastic). However there is a clinal aspect to the variation also, reflecting genetic change across its geographical range. In the past, the variation within the species has frequently been split into a series of subspecies and varieties. As an example of this, Druce (1928) distinguished as many as twelve varieties, and Pilger (1937) set up just two varieties, var. communis Schlechtend and var. dubia Wahlenberg. However, the first of these he spilt further into five sub-varieties, and in the second variety he distinguished two more sub-varieties (Sagar & Harper 1964).

In the account in Flora Europaea 4, D.M. Moore came to the conclusion that formal recognition of any of the suggested variants as subspecies was impractical (Tutin et al. 1976). Having said this, in their critical Flora of Great Britain and Ireland 3, Sell & Murrell (2009) distinguish five varieties, based largely on habitat or geography, but recognised by differences in leaf shape, size and hairiness together with inflorescence shape. None of these forms have been looked for in Fermanagh.

Palatability and benefits to grazing animals

P. lanceolata is nutritionally valuable for stock animals, providing a particularly good source of essential minerals, including calcium, iron, sodium, chlorine, nitrogen, potassium, phosphate and cobalt. It also provides small amounts of magnesium but is poor in manganese (Thomas & Thompson 1948; Wilman & Derrick 1994). Ribwort Plantain has in the past been sown in herb strips along with ley grass and legume mixtures in order to provide forage variety to stock and to supplement their mineral nutrition. In Germany, P. lanceolata has been a recommended constituent of pasture mixtures because it is known to improve animal health and milk flavour (Heeger 1949). In 2021, Ribwort Plantain is again being introduced to cattle and sheep grazing experimental multi-species herb and legume swards in Co Down (NI) and elsewhere in B & I as part of an ongoing government sponsored European Innovation Partnership Project. This experiment is investigating both meat production and additional benefits to soil and animals from a reduced need for fertiliser nitrogen, extension of the growing season, drought resistance, reduced parasite burden and enhanced biodiversity above and below ground.

In dry pastures, the plant is palatable to livestock (especially sheep) and provides, "fair quality fodder, although inferior to grasses" (Clark & Fletcher 1906). An animal grazing study by Milton (1933) found it, "definitely palatable and may even be sought after". Sheep are so keen on it that they, "will actually chisel the crowns out of the ground, using the lower incisors" (Sagar & Harper 1964). Grazing by cattle in short herbage is less important than by sheep as the plantain plants react to being browsed by developing a ± appressed rosette habit and the cow's tongue cannot then pull the leaves off (Sagar & Harper 1964). In grassland communities, P. lanceolata is reduced by grazing more than by mowing (Tamm 1956). Unsurprisingly, P. lanceolata is, therefore, less common in pastures than in ungrazed, mown meadows. It is also less tolerant of trampling than P. major, although like it, the small seed does colonise compacted dry soil very well (Cavers et al. 1980).

Cattle and sheep will remove flowering spikes of P. lanceolata and P. media (Hoary Plantain), but apparently not of P. major (Sagar & Harper 1964), obviously negatively affecting their seed production. A new crop of leaves is produced by plants in the autumn and the plant is wintergreen and frost resistant. Since these leaves have appreciable nutritive value, they serve as useful winter fodder (Salisbury 1964).

Flowering reproduction

From April to September, or until the first frost or competition prevents anthesis. The rosette plant produces one or more leafless flowering stems or peduncles (scapes) longer than the leaves, 10-60(-100) cm tall, strongly ribbed or furrowed and appressed-hairy. The terminal spike inflorescence is dense, ovoid to cylindrical in shape, 0.7-5.0 cm, lengthening up to 12 cm in fruit. The much reduced, densely packed flowers are sessile and are subtended by dark green-black, ovate-acuminate bracts with membranous points, the lower bracts larger than the upper ones. The calyx of four sepals is c 4 mm in diameter, brownish-white or yellow with a prominent green or brownish midrib reaching the tip on each sepal. The corolla has a tube 1.5-3.0 mm long and is black, with lobes broad, lanceolate and acute. The five stamens are yellowish-white, with slender filaments equalling or exceeding the corolla tube and the anthers are yellowish. The style is c 2 mm, the stigma filiform and feathery.

The flowers are protogynous, wind pollinated and are described as being 99% self-incompatible. They are visited by insects collecting pollen, including the Honey Bee (Apis mellifera). The insects do not pollinate the flowers as they do not touch the stigmas of flowers that have no anthers exerted. Male sterility is not infrequent.

The fruit capsule is a dehiscent pyxidium, ie it splits around the equator (circumscissile) and opens like a box with a lid. It is 3-6 mm long and contains one or two seeds only, each 1.2 × 0.5 mm, half-spherical (Butcher 1961; Sagar & Harper 1964; Grime et al. 1988, 2007; Sell & Murrell 2009).

Seed production in natural habitats in Europe for most plants is rather small. In pastures grazed by sheep and cattle, seed production will usually be seriously curtailed by animals removing the flowering stems as well as the leaves. Salisbury (1964) estimated the normal level of annual production per plant was probably around 2,500 seeds, but large plants developing more than 30 spikes could generate over 10,000 seeds.

As in P. major, the seeds of P. lanceolata are not actively dispersed after the capsule splits. They are coated with mucilage that swells when wet, making them adhesive and helping to disperse them on soil particles and by attachment to hair, feathers and skin, again as happens in P. major (Sagar & Harper 1964). Cattle and birds, including pigeons, sparrows, bullfinch and greenfinch, also feed on the seed and probably are important dispersal agents (Salisbury 1964; Sagar & Harper 1964). Animals brushing against a ripe spike may scatter seeds from the open capsule, or scapes may rot at the base and fall to the ground, shedding the seed. Some seed may be retained in open capsules for eight or more months.

Seed matures rapidly after fertilisation takes place, in some situations after two or three weeks, depending upon the weather (Sagar & Harper 1964). As mentioned above, seed is long-persistent in the soil seed bank, evidence suggesting up to 60 years (Holm et al. 1977). Germination normally ranges from 60 to 90% and seeds that have passed through birds have displayed 100% germination (Salisbury 1964). Germination occurs both in the autumn of production and in the following spring, contrasting with P. major and P. media, both of which germinate only in spring (Sagar & Harper 1964).

Vegetative reproduction

Individual plants can survive for twelve or more years and older plants tend to develop a thick rhizome which can sometimes produce side buds that develop new aerial shoots in the field leading to clumping of the species. In experimental studies, early autumn (September) planted P. lanceolata was able to regenerate buds from root fragments, but not from November or February plantings. It is not clear to what extent vegetative reproduction occurs under field conditions, but it is probably of limited and minor importance in comparison with colonisation by seed (Sagar & Harper 1964).

Fossil history

Pollen grains are identifiable to species level and prove that P. lanceolata has been present in Britain since every zone of the Late Weichselian glacial onwards to the present day (Godwin 1975). In late glacial times, it may have been restricted to arctic-alpine grasslands, but as an indicator species of open ground, it has provided science with a dramatic picture of the extent and speed of deforestation following the arrival of Neolithic man clearing the landscape for farming. P. lanceolata spread with the grass, hay and the herds at that time in European history, and it did the same when the New World and other colonies were invaded and farming practices, fodder and seed-stocks exported (Holm et al. 1977).

Fermanagh occurrence

This is one of the most common and widespread species in the county; Ribwort Plantain occurs in 496 Fermanagh tetrads, 94% of the VC total and it might even then be ignored and overlooked elsewhere through it being considered by some recorders as an excessively common, weedy, perennial. Despite this reservation, the species ranks as 5th equal along with Crataegus monogyna (Hawthorn) in terms of tetrad frequency in the VC.

British and Irish occurrence

P. lanceolata is common and ± ubiquitous throughout B & I, being absent only from the most acidic soils, deep shade and wetlands. It achieves its greatest altitude (845 m) at Great Dun Fell in Westmorland (G.M. Kay, in: Preston et al. 2002).

European and world occurrence

While this very common species is now ubiquitous across the whole of Europe except the extreme north and is also considered native in N Africa and N & C Asia, its centre of origin remains unknown, although it may probably have been somewhere in C E & S Europe. P. lanceolata has been introduced to temperate areas across the whole world and may sensibly be referred to as 'cosmopolitan'.

Holm et al. (1977) stated in their book entitled The world's worst weeds, "On a world basis, P. lanceolata is said to be one of the 12 most successful non-cultivated colonising species." P. lanceolata and P. major are such successful worldwide weeds, since in disturbed agricultural areas where they enter a microsite, become established, spread out and form a rosette to occupy valuable space, they can then manage to hold it against almost all other species. In this fashion, they have invaded many of the world's agricultural fields and have been reported as weeds from more than 50 countries in 26 crops that are as different as alfalfa, coffee, rice, cereals, onions, cotton, small fruits as well as temperate orchards and pastures. The most serious competition takes place in cereals and in sugar beet fields and plantain seeds frequently contaminate small-seeded legume seed-stocks. These two plantains are thus a constant source of difficulty and the farmer must spend a great deal of time, energy and resources attempting to hold them back. However, Holm et al. (1977) concluded, "On account of their individual limited size, they are seldom reported as a principal weed for any particular crop."

Ribwort Plantain belongs to the southern temperate phytogeographical element, but is now circumpolar southern temperate in its distribution, although there is a wide gap between C Siberia and Alaska (Hultén & Fries 1986, Map 1724; Sell & Murrell 2009).

Medicinal and other uses

As mentioned under P. major (see this website), plantains, including P. lanceolata have a reputation as vulnerary herbs in quickly staunching bleeding wounds. Usually it is found sufficient for a fresh leaf to be crushed and applied (or chewed) for enough of the healing chemicals to be released, but for severe wounds the whole plant has been boiled. Alternatively, an ointment has been manufactured and applied. The herb has also been used to treat skin rashes, friction damage, burns, stings, varicose veins and piles, and for drawing out pus from infected wounds. In Ireland, plantain juice was used for coughs and the plant was used for very many other medicinal treatments including for gout, liver trouble, jaundice and even for treating sore eyes (Allen & Hatfield 2004).

Another quite different farm use made of Ribwort Plantain was as a means of testing the dryness of hay. The leaves of P. lanceolata dry more slowly than those of grass, so when making hay it was important to make sure that the plantain leaves were dry, or else the resulting hay stack might overheat and perhaps even catch fire. This gave rise to English common names such as 'Fire grass' and 'Fireweed' (Vickery 2019).

Weed control

As with P. major, cutting off the leaf rosette of P. lanceolata at ground level merely stimulates the formation of a ring of lateral buds that re-establish the plant. Both species can be best eliminated from areas where they are unwanted by spraying with an herbicide, such as 2-4-D (Salisbury 1964).

Names

No fewer than 46 English common names are listed by Grigson (1955, 1987). Vickery (2019) manages to beat this total with his 52 names. Twelve of these names include or infer 'black' as a word element, a probable reference to the black colour of the immature spike inflorescence or the more mature spike with a dusting of white from the stamens. Children, from time immemorial, have matched one 'Black Man', one 'Soldier', one 'Hardhead', one 'Fighting Cock', swinging its long peduncle against another until the weaker one loses its head. This game, which is widely played across Europe, has given rise to at least 15 of the English common names listed by Grigson and Vickery, including 'Conkers', 'Fighting Cocks', 'Fox and geese', 'Hard Heads', 'Knights' and 'Kings'.

Threats

None.

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