Primula veris L., Cowslip
Account Summary
Native and/or possibly accidentally introduced, rare. Eurosiberian temperate.
1881-2; Barrington, R.M.; Belleisle House estate, Upper Lough Erne.
April to August.
Growth form and preferred habitats
Cowslip is a rosette forming, early flowering, shortly rhizomatous, potentially long-lived, patch-forming perennial herb of damp but well-drained, mesic to calcareous, species-rich grassland soils in the pH range 5 to 8 (Grime et al. 1988).
Although often reputed to favour constantly moist, lightly-shaded sites on the margins of woods, along paths or other openings, or by open scrub, none of the Fermanagh P. veris records are associated with such habitats. In many areas of B & I and Europe the restriction of Cowslip to relatively unproductive, nutrient-poor, neutral or mildly acidic to lime-rich soil and to light- or half-shade, may simply be a mechanism of avoiding excessive competition (Tamm 1972). Apart from this list of relatively open inland habitats, P. veris also occurs in coastal sites on stabilised sand dunes and sea cliffs and shows a greater tolerance of drought than P. vulgaris (Whale 1984; Grime et al. 1988). Prof Grime and his Sheffield co-workers classified P. veris as a stress-tolerator with some competitive ability.
As is the case for the common Bluebell (Hyacinthoides non-scripta) and Bracken (Pteridium aquilinum), the grey cloudy skies of the NW of Ireland may provide sufficient shade for Cowslip, so that it can compete better and survive in fully-lit, unshaded sites.
Fermanagh occurrence
The very pretty Cowslip has certainly declined in well-drained, species-rich 'unimproved' limestone pastures in Fermanagh since 1970, when it was found in the Doagh Lough, Knockmore and above the Cladagh River Glen (ie the Marble Arch) areas. RHN and the current author (RSF) no longer know of it in this semi-natural habitat anywhere in the county. However, P. veris does also occur in the seasonally flooded, more or less organic soils in water meadows around sections of the shore of both Upper and Lower Lough Erne, which is an occasionally noted habitat for this species, provided it is not permanently water-logged. Of the three yellow Primula species that occur in Britain (P. vulgaris (Primrose), P. veris and P. elatior (Oxlip)), P. veris is the least shade tolerant and the most drought tolerant (Whale 1984).
There are records from a total of 23 Fermanagh tetrads (4.4%). As the distribution map shows, records are scattered across the VC, although the species is mainly found around the Lough Erne basin and on the limestones of the Western Plateau. Only 16 of the 23 tetrads have post-1975 records, indicating a very definite decline in Fermanagh populations. The map indicates that the most widespread loss occurred in the Clyhannagh and Coaghan Tds, above and to the west of the Marble Arch caves (ie in the Marlbank region), where Lynn Farrell recorded them in August 1970 and where the species has not been seen since.
Flowering reproduction
Unsurprisingly, Cowslips appear to flower more profusely amongst rank grasses and scrub than amongst large flocks of spring grazing sheep, but easing of grazing pressure tends to lead to masses of flowers reappearing on English downland, eg in Hertfordshire (VC 20) (Mabey 1996). P. veris bears up to 30 flowers in a 1-sided, drooping umbel on top of a tall (10-20 cm), erect, solitary scape (leafless flowering branch) in April, but chiefly in May (Richards 1993; Sell & Murrell 2014). The flowers are heterostylous as in Primrose (P. vulgaris), hang slightly downwards, are somewhat cup-shaped, about half the diameter of the Primrose and are clear golden yellow with an orange to reddish spot at the base of each petal. Like P. vulgaris, the flowers are mostly self-incompatible and are dependent on 'long-tongued' insect visitors (principally honey bees and bumblebees) for their cross-pollination (Richards 1989). However, it has been found that even relatively low levels of shading (41%-36% of full sunlight), can drastically reduce seed set in P. veris (Whale 1984).
In many areas of B & I, seed-set is good despite the requirement for cross-pollination. Plants with long-styled 'Pin' flowers can, however, self-fertilise (14%) in the absence of 'Thrum' pollen, while only 0.6% of thrums can do likewise (Lees 1971; Richards 1993). After fertilisation, the ovoid fruit capsule containing around 50 seeds (as large as those of P. vulgaris) is retained within the enlarged calyx, which must delay, restrict and limit seed dispersal distances achieved. Seed dispersal by gravity was found to be normally limited to around 50 cm from the parent plant (Richards & Ibrahim 1978). The seed is dormant when shed and primary seed dormancy is overcome by cold stratification, so that germination should occur in spring following dispersal, or later, if they survive soil burial (Milberg 1994).
Seed dispersal and its survival in soil
In 1992, Cowslip occurred as a 'weed' in a newly sown lawn at the Erne Hospital, Enniskillen and four or five of its previous Fermanagh sites were on established grassland on roadside verges, where its origin was something of a mystery. The seed is without a gelatinous nutritive elaiosome to attract animal vectors and thus dispersal must be entirely wind-, water- or soil-borne. Wind is likely to be an important means of seed dispersal, although the capsule is not elevated, which must limit distance travelled. Experiments indicate few seeds disperse more than 1 m (Richards 1989).
One possible explanation of the rather transient occurrences observed in Fermanagh is that viable buried seed might have persisted in the soil seed bank and have been induced to germinate and establish after road making or repairing disturbance brought them to the surface. However, the seed of P. veris is generally regarded as being only transient or short-term persistent in the soil, which rather knocks this hypothesis on the head (Thompson et al. 1997). However, experimental work in Sweden showed that P. veris seed can survive in soil for more than one year and might germinate in the second or later spring after release from the parent plant, ie following two winter cold stratification periods. P. veris could easily be overlooked in soil bank studies if a cold treatment is not included (Milberg 1994).
Another totally unsupported suggestion is that seed or rhizomatous material may have been imported with fresh topsoil, or on the tyres of vehicles involved in the road works at some stage.
Role of vegetative reproduction
Field observations in England in fully illuminated, short-sward meadows and pastures suggest seed set of Cowslips is normally good, but can be variable from year to year, sometimes poor or sporadic. In these latter circumstances, existing plants must rely on their limited potential for vegetative spread and rhizome growth for some of their increase and regeneration. In stable, favourable growing conditions, such as a meadow or pasture on deep soil that is capable of retaining moisture and with some marginal, light tree shade, P. veris can survive for many years without great change in number. In such situations, vegetative propagation by the rhizome branching and extending, compensates for the few individuals that die. Regular removal of vegetation by mowing or grazing keeps a sward low, reduces competition and helps P. veris survive (Tamm 1972).
In open, dry, limestone sites, Cowslip plants remain small and are normally short-lived, seedlings surviving an average of only two years. In closed grassland vegetation on a fertile, heavy soil the average lifespan of surviving seedlings is much longer, up to ten years and allows plants to develop large clumps. Naturally, seedling colonisation and establishment is much more difficult in closed vegetation than in open or disturbed sites (Richards 1989).
Herbivory, toxicity and vegetation disturbance
As is also the case with P. vulgaris, the leaves of P. veris, which are often appressed to the ground, tend to be ignored by browsing stock animals, very probably due to their saponin content (Grime et al. 1988; Cooper & Johnson 1998). Flowers and fruits are eaten by various animals and the latter must assist in seed dispersal. Regular or occasional disturbance, such as cattle trampling and dung, certainly affects pasture vegetation and may provide spots favourable for seedling development (Tamm 1972). Seedlings most frequently establish in small pockets of bare ground such as around molehills, anthills and hoofmarks, often clustered around the parent plant (Richards 1989).
Possible introduction?
In more recent years, Cowslip has become a popular constituent of so-called 'wild flower' seed mixtures, increasingly sown by contractors to re-vegetate disturbed verges after roadworks. However, while this has occurred in recent years, RHN and the current author (RSF) do not believe that the situation arose in Fermanagh at the earlier dates in question when records were made. The possibility of deliberate introduction is strongly supported by the fact that the New Atlas distribution map for P. veris quite accurately plots the major road system in NI at least and it may do likewise in other regions of B & I (Preston et al. 2002). The fact that a high proportion of the Fermanagh sites for this species are in or near large landed estates, eg Belleisle, Castle Coole, Crom and Florencecourt, might also be significant in this respect. There again, the occurrences might only prove what we already know, that the early planters (ie human settlers), recognised good, well-watered, fertile ground when they saw it!
In terms of conservation protection, the Beagh meadow site is an ASSI, and Cowslip is well established on the main drive at the Castle Coole estate, a National Trust property, while the meadows at Clonatty Bridge deserve protection. At the Roosky turlough, Cowslip clings on in a meadow which has already been 'improved' − at least in terms of NPK fertility, if not of biodiversity.
Variation
Three subspecies are recognised by Sell & Murrell (2014), distinguished by the near absence of glandular hairs (subsp. veris), or an abundance of glandular hairs, and then either a cordate leaf base (subsp. columnae (Ten.) Lüdi), or a gradually decurrent leaf base (subsp. canescens (Opiz) Hayck ex Lüdi). Subsp. veris is the form native to B & I and which has the widest range of the three, while subsp. columnae is a plant of the mountains of C Spain, C Italy, N Greece and NE Turkey, and subsp. canescens occurs in the Alps, Pyrenees and mountains of N Spain.
Richards (1993) and Flora Europaea 3 (Tutin et al. 1972) add subsp. macrocalyx (Bunge) Lüdi in Hegi, loc. cit. to these three, it being a plant of SE Russia, the Caucasus and SC Asia that extends to E Siberia (Richards 1993). This form is possibly regarded as a related, separate species by some taxonomists, which might account for its neglect or discard by Sell & Murrell (2014).
Although there are four recognised subspecies, the authors of the Flora Europaea account (D.H. Valentine and A. Kress) warn that there are many transitional plants (Tutin et al. 1972).
Irish occurrence
P. veris is very much more frequent and widespread in the RoI (especially on the Central Plain), than is the case in NI. Apart from Fermanagh, the few other occurrences in NI are all considered probable introductions or garden escapes (FNEI 3). Having said that, on account of the few perhaps indigenous Fermanagh stations, plus one site in Co Armagh (H37), P. veris retains Protected Species status in NI under Schedule 8 of the Wildlife (NI) Order, 1985. As BSBI VC Recorders, RHN and the current author (RSF) are sceptical as to the scientific basis for this protection.
British occurrence
The New Atlas hexad map shows that P. veris subsp. veris remains frequent and widely scattered throughout much of lowland Britain, especially in chalky, light-soiled areas, becoming much rarer and more coastal and eastern in distribution in Scotland (Preston et al. 2002). Many populations have survived the widespread destruction of calcareous meadows and pastures that resulted from the intensification of farming that took place over a half century in the 1930-1980 period, presumably persisting on adjacent marginal land. Pressure on Cowslip populations, especially near urban conurbations, from flower picking to satisfy deep-seated folk cultural traditions around Easter and other celebrations, also took its toll until 1975 when conservation legislation made clear the need to refrain from such collecting. There has been a considerable degree of population recovery from the 1990s onwards as a result of public awareness and conservation efforts.
Some of the currently mapped Cowslip populations may be derived from gardens and from deliberate plantation of 'wild seed mixtures' in more recent years. Plants of the other two subspecies from Europe are occasionally grown by specialist gardeners and some of these may escape into the wild, although they are likely to be rare and probably not long persisting (Preston et al. 2002; Sell & Murrell 2014).
European and world occurrence
P. veris is a polymorphic species with three or four subspecies depending upon authority consulted. It is mainly restricted to Europe, but extends eastwards into Asia. Hultén & Fries (1986, Map 1467) choose to map just two subspecies. Subsp. veris is shown to be widespread in much of Europe, becoming more coastal further north in Scandinavia and the Baltic area. Subsp. macrocalyx occurs in more eastern parts of C Europe and adjacent Asia, but occurs in a discontinuous manner. The distributions of the other two subspecies is given above under 'Variation'.
Uses and folklore
The flowers have a very distinctive, fresh fragrance similar to Anise or to some volatile oil like Mannite. The flowers contain narcotic juices, including saponin, which encouraged their use in wine-making. Cowslip wine was given to children in small doses as a medicine for various ailments and it was also prescribed for both jaundice and measles (Vickery 1995). Young cowslip leaves were used in spring salads and mixed with other herbs to stuff meat (Grieve 1931). The main medicinal uses were as a sedative and anti-spasmodic, their special value being for strengthening the nerves and the brain, relieving restlessness, giddiness, nervous excitement and insomnia. It was also greatly valued for relieving pains in the head and was formerly administered for palsy and all forms of paralysis, and names like 'Palsy Wort' or 'Herba paralysis' were given. The root was also called in old herbals 'Radix arthritica' from its use as a cure for muscular rheumatism (Grieve 1931).
Cowslip balls were made of flower stalks tied together and thrown up for love divination, the names of possible suitors recited until the ball landed (Vickery 1995). Cowslips were scattered on the altar in churches on May Day (the first Sunday in May) (Vickery 1995). This use in church may somehow connect with plant names that refer to the keys of St Peter (an allusion to the pendulous umbel of flowers), or a Welsh name that translates as 'Mary's tears' (Grigson 1987). Previously Cowslip must have been extremely common and abundant in at least some parts of England, as folklore has it that flowers were worn as necklaces on May Day and were scattered in front of newlyweds emerging from church, like confetti, practices that could not happen nowadays even without conservation legislation (Mabey 1996).
Names
The English common name 'Cowslip' is a polite euphemism for 'cow-slop' or 'cow-pat' which hints at the kind of company the plants often keep in pastures (Mabey 1996). In old herbals, the plant is referred to as 'Herb Peter' or 'Key Flower', the pendulous flowers being likened to a bunch of keys and hence with St Peter and his keys to the Kingdom of Heaven. They are also called 'Our Lady's Keys' and 'Keyflower' for similar reasons (Grieve 1931).
Threats
Further improvement of meadow fertility, causing decline of species diversity.