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Plantago maritima L., Sea Plantain

Account Summary

Native, very rare. Eurosiberian wide-boreal.

1947; MCM & D; on a clay bank by NE shore of Ross Lough.

April to July.

Growth form and preferred habitats

P. maritima is a rosette-forming perennial with a branched, woody rootstock and long, narrow and rather thick, fleshy leaves, rounded on the back and channelled on their upper surfaces. The greenish flowers are borne on a slender cylindrical stem in a narrow spike rather similar to that of P. major (Greater Plantain). Each plant rosette usually forms numerous flowering stems. When it flowers, from June to September, the yellow anthers on long filaments are very conspicuous and the plant is wind pollinated (Hepburn 1952; Clapham et al. 1987).

Like other Plantago species, Sea Plantain has no powers of increase or spreading other than by seed. Again, in common with all maritime species, Sea Plantain is a salt-tolerant (ie halophyte) plant of middle and upper reaches of salt-marshes, in the middle and upper zones of salt-marshes and in open, low-growing coastal turf, often alongside Armeria maritima (Sea Pink or Thrift). The leaves of P. maritima are relished by sheep and, like Thrift, it stands up well to natural grazing in a salt-marsh setting (Hepburn 1952). P. maritima can also grow in shingle at the top of beaches, in rock crevices and on short turf on sand near the sea. It is also frequent in sea cliff grassland, where it sometimes becomes dominant or co-dominant with other plantains or Thrift, especially if sheep grazing is taking place or other stress features including sea spray and wind exposure help reduce interspecific competition from grasses and other somewhat taller-growing or more aggressive plant species (Chapman 1976). Sometimes this form of plantain sward vegetation is extremely dwarf, being just a few cms in height (Hepburn 1952). Like Sea Aster (Aster tripolium), Sea Plantain can also grow in brackish situations in salinities of, "a quarter of sea water or less" according to Proctor (2013, p. 428).

P. maritima is a phenotypically very plastic species that can tolerate an extremely wide variety of soil physical and chemical conditions, ranging from clay-rich muds to sandy gravel marshes and pastures. However, it cannot tolerate much plant competition (Ranwell 1972). Nevertheless, Sea Plantain does survive considerable pressure from other forms of stress, enabling it to colonise, thrive and sometimes even dominate salt-laden, weather-exposed and sheep or rabbit grazed habitats, always in situations that minimise direct species competition (Hepburn 1952).

In addition to its common and very widespread coastal occurrences, inland populations of P. maritima occur beside streams or around wet rocks and in stony flushes in the higher mountains in the English Pennines, N Wales, NW Scotland and the N of Ireland, and on limestone in western parts of Ireland, sometimes at low altitude (Botanist in Ireland, paragraph 64; Godwin 1975; Garrard & Streeter 1983; Sell & Murrell 2009). It tends to frequent both acidic and limestone soils where there is some degree of flushing water moving through silty soil on cliff rock ledges and screes (Tansley 1939; Graham 1988; Halliday 1997; G.M. Kay, in: Preston et al. 2002).

In recent decades, and especially since the advent of motorways in B & I, P. maritima has been spreading inland along salted roads in many places and doing so particularly commonly in Scotland (G.M. Kay, in: Preston et al. 2002).

Variation

Various attempts have been made to subdivide this rather variable species, but the taxa proposed have proved difficult to define morphologically.

Plants from Arctic Europe have been named subsp. juncoides (Lam.) Hultén, which is believed to differ from the widespread form subsp. maritima in having wide bracts, a more ovid to globose capsule shape, the scapes not exceeding the leaves and more numerous seeds per capsule. However, these criteria for discrimination have not proved very satisfactory.

Another variant named subsp. serpentina (All.) Arcangeli is described as having long-acuminate bracts and the posterior sepal with the keel often narrowly winged. It occurs in S Europe, especially on mountains above 2,000 m. However, it is probably only an ecological variant and therefore not worthy of subspecific rank (D.M. Moore, in: Tutin et al. 1976).

A more complicated set of suggestions for subdivision of what is considered a polymorphic species to subspecific or varietal rank on a worldwide basis has been made by Hultén (1974) and applied to a map in Hultén & Fries (1986, Map 1726), without being any more convincing to the current author (RSF). Apart from the questionable taxonomy, the Hultén (1974) nomenclature is also confusing.

The more recently published British Floras do not recognise any taxonomic subdivisions in P. maritima (Sell & Murrell 2009; Stace 2019).

Fossil history

Fossil evidence proves that P. maritima has been present in B & I in every glacial and interglacial sub-stage or zone from the very ancient late Anglian to the present day (Godwin 1975). Thus the inland mountain and salt-marsh sites in B & I represent very long-established populations of this fleshy perennial. On the basis of fossil pollen and macrofossil remains (including in the case of P. maritima, seeds), this species, and a limited number of others like it, are recognised as relict survivors from Late-glacial vegetation populations in these isles that were established under totally different climate and soil conditions from those pertaining today (Godwin 1975). P. maritima and its previously widespread Full- and Late-glacial plant associates were forced to retreat to a range of tree-free coastal and high mountain habitats in order to avoid the shade, closed cover and severe competition of taller-growing, better adapted, immigrant colonising plant species that arrived when temperatures rose and glaciers melted 10,000 years BP. In later post-glacial stages and climates, these Late-glacial flora relicts also had to survive the development and expansion of acid peat deposits (Raven 1956).

The fact that some of these Late-glacial species populations survive as relicts today is quite amazing, and it is not surprising that after persistence of thousands of years, a number of them represent the rarest and most thinly scattered species in B & I. While inland mountain and salt-marsh populations of P. maritima belong to this group of rarities, the species is very common, widespread and indeed often super-abundant in coastal maritime sites.

Fermanagh occurrence

In land-locked Fermanagh, Sea Plantain is locally very rare, but it has been recorded in four tetrads in the VC over a 50 year period. The Fermanagh sites where this species has been recorded are unexpected and varied, including both relatively upland calcareous lakeshores around or close to Monawilkin Lough and Ross Lough (the VC has no high mountains) and in lowland grassland in Knockmore, Tonlisderrit and Tirmacspird townlands. At Ross Lough, it appears confined to the more or less steep slope of the old, superseded shoreline, particularly where cattle trample and poach the ground, keeping it open so that bare soil is always present. There is a danger that nearby gorse bushes may spread into this site.

The additional local record details are: around Monawilkin Lough, 10 July 1985, S.J. Leach & S.A. Wolfe-Murphy; W end of Monawilkin Lough (to N of track under cliffs), 1994 & 1996, I. Rippey; Tonlisderritt, 1 km SE of Cashel, 28 April 1996, RHN & HJN; lowland acid grassland, Tirmacspird Td, NW of Lack, 7 June 1997, RHN; and also at Meikle and co-workers' old site, the clay bank by NE shore of Ross Lough, 25 July 2000, RHN & RSF; also here 17 September 2006 & 19 September 2009.

Other normally maritime species which 'misbehave' in this ecological manner by occurring in land-locked Fermanagh include Carex distans (Distant Sedge), Asplenium marinum (Sea Spleenwort) and Viola tricolor subsp. curtisii (a seashore form of Wild Pansy, Seaside Pansy).

British and Irish occurrence

P. maritima is a very common and widespread species around almost the entire maritime coastline of B & I. While of a very much rarer occurrence, it is also found in upland species-rich pastures and in rocky mountain ground in the Pennines and scattered across the N & W of B & I. It is rarest in inland salt-marshes, which are themselves rare habitats, but it has more recently appeared along some stretches of salt-treated motorways, chiefly in Scotland (G.M. Kay, in: Preston et al. 2002; Sell & Murrell 2009).

European and world occurrence

On the European mainland, P. maritima extends in coastal habitats from the North Cape to Spain and it also occurs rarely in inland situations (Godwin 1975). It is rarer in the south of Europe and extends into N Africa. It is also present in Greenland (Böcher et al. 1966), Iceland (Löve 1983), N America and southern S America (Sell & Murrell 2009).

Uses

Apart from the fact that it was appreciated to be nutritious for sheep and pigs to graze, there does not appear to be much mention of folklore uses for the plant (Grieve 1931). It is mentioned in passing by Allen & Hatfield (2004) as having a reputation in the Cork region of Ireland as a general stauncher of wounds, in common with other plantains.

Names

The genus name 'Plantago' is from the Latin 'planta', 'the sole of the foot', referring to the broad, flat leaves that are often pressed to the ground and from the feminine termination 'āgo' (Gilbert-Carter 1964). The Latin specific epithet 'maritima' means 'belonging to the sea', or 'growing on the sea coast' ('mare') (Gilbert-Carter 1964).

English common names include 'Buck's-horn', Buck's-horn Plantain', 'Sea Plantain' and 'Gibbals Kemps': the first two of these is nowadays transferred to Plantago coronopus. "'Gibbals' is or was a name applied to P. maritima around Rommy marshes, about 2 miles [3.2 km] from Cardiff". A manuscript account of a botanical excursion in 1775 by Lightfoot recounts how hogs were encouraged to graze on the roots that they were reputed to be fond of, that were grubbed up for them, and on which they grew fat (Britten & Holland 1886). 'Kemps' ('kemp' is to fight in northern dialect and was more often applied to P. lanceolata (Grigson 1955, 1987). It is not known how it came to be applied to P. maritima.