Fagus sylvatica L., Beech

Account Summary

Growth form and preferred habitats

Beech are large trees up to 30 m or more in height and they can grow on a very wide range of soil pH, from 3.5 at the extreme acid end of the spectrum, to very much rarer occurrences at pH 7.5 in alkaline rendzina calcareous soils, or even pH 7.9 on softer, chalk and oolite substrates (Tansley 1949, pp. 361-6 & 421-6; Rackham 1980, Chapter 19). Beech seedlings or saplings are occasionally found on acid bogland, and we have records of the tree in Fermanagh on or very close to raised-, blanket- and cut-over bogs. The tree also grows in calcareous areas, for instance in County Fermanagh (H33) at Hanging Rock NNR, at Marble Arch and on the Florencecourt estate. However, beech has not been planted, and it has not colonised in other limestone areas, including around Knockmore mountain, nor on Knockninny hill. In general, beech trees do not thrive on calcareous soils, and they certainly never approach anything like the dimensions they achieve on deeper, more fertile, free-draining moderately acid, brown earth soils.

Beech litter and gradual soil deterioration

Beech trees normally cast a heavy leaf, twig and branch litter each year, each tree typically shedding up to 10% of its terminal branch material annually (Thomas 2000). The slow decay and incorporation of beech litter gradually leads to the development of a strongly acidic, infertile, sour, mor type of humus in soils beneath beech trees. This degrades the soil through the leaching out of minerals, and particularly in our moist oceanic Irish climate, it creates a less fertile podsol-type of soil profile. Podsolization involves the formation of a rust coloured 'iron-pan' of re-precipitated iron at depth in the soil profile, and this impedes the previous usual free drainage of the substrate (Ingrouille 1995). As the soil gradually becomes podsolized, it turns wetter and becomes subject to waterlogging, so that ultimately, beech itself ends up at a disadvantage, since by this process it creates soil surface conditions that are unfavourable to the germination and establishment of its own seedlings (Tansley 1949, p. 361).

Environmental factors limiting beech longevity

The twin enemies of the mature beech are late frost, which can defoliate the tree and kill both buds and seedlings, and prolonged summer drought, which may do the same. In Fermanagh and in most of Ireland, long droughts are extremely rare. Beech is not a long-lived tree, and being shallow-rooted, it is susceptible to wind-throw during severe storms. Thus specimens over 250 years old really are exceptional. Wind-throw of large, old trees is especially likely if they are growing in open parkland or in an avenue rather than in the shelter of other woodland. Old trees generally become hollow, which makes them more resistant to wind since the tubular trunk is actually more stable and stronger than when it is solid (Milner 1992, p. 16). A moving account by Thomas Packenham in the introduction to his beautiful book, Meetings with remarkable trees, describes the effect on him of the toppling of several giant beeches at his Tullynally estate in Co Westmeath (H23). Accounts like his should make us realise the need for active replanting of beech for landscape reasons in many parts of these islands (Packenham 1996).

Native and alien (introduced) status in Britain and Ireland: Beech was one of the last woodland trees to expand in Britain after the last glacial period. Although present as a rare species from around 4450 BP in the Sub-boreal pollen zone VIIb, F. silvatica does not appear abundant in the fossil pollen record until around 3000 BP. Thus having increased and spread at a time when competing vegetation was well established, and agriculture had already been developing for around 2000 years, the pace of beech colonisation was fairly slow. Indeed, in many areas its natural dispersal may have absolutely required the destruction of pre-existing forest by man, a process that was already occurring at this time in Britain (Godwin 1975; Ingrouille 1995, p. 195) and also in other parts of Europe, eg S Sweden (Bjorkman 2001).

F. sylvatica plantation has occurred so much in Britain within and beyond the likely indigenous range in SE England and SE Wales, that native beechwoods and trees have become very local and erratically distributed. Planting has happened to the extent that, except in very general terms, it is impossible to accurately delimit beech's native occurrence (Rackham 1980; T.D. Dines, in: Preston et al. 2002). It is therefore puzzling as to why the editors of the prestigious BSBI New Atlas decided to publish a map of F. sylvatica that totally ignores the question of status, and misleadingly and knowingly and erroneously plots and displays beech as a native species throughout Britain and Ireland. In doing so, they include mainland Scotland, the western and northern isles and Ireland, all perfectly distinct geographical areas where nobody imagines the tree is indigenous. If for some production reason a single status map was obligatory, then surely an all introduced map would have been a more accurate picture than the one published in the New Atlas.

The same apparently nonsensical, misleading approach has been taken with Carpinus betulus (Hornbeam) and Tilia cordata (Small-leaved Lime), but NOT with Tilia platyphyllos (Large-leaved Lime) or Acer campestre (Field Maple), both of which have perfectly sensible maps that discriminate native from introduced areas. None of these trees are indigenous in Ireland, and obviously it is important for botanists in both Britain and Ireland to know this fact, as knowledge of it colours their impression of what types of woodland there will be on the smaller, more ancient island (Rackham 1986; McCracken 1971).

Woodland 'Climatic Climax' vegetation and dominant tree species

Despite its questionable status, beech is a tall, potentially dominant tree, and on the more fertile soils at least, it is capable of outgrowing and replacing old, well-established oak, Quercus robur (Pedunculate Oak) and Q. petraea (Sessile Oak). Oaks have been long regarded as the dominant native species of the 'Climatic Climax' woodland vegetation of these islands (Tansley 1949; Newbold and Goldsmith 1981). The notion of a single dominant form of vegetation covering the whole climatic region (ie the concept of a 'Climatic Climax' vegetation of a region or large geographical area) has tended to go out of fashion as we learn more about the requirements of plants and of the dynamic and sometimes cyclical nature of vegetation change (Watt 1947; Gimingham 1972). However, while we should approach the ecological concept of climax vegetation with caution, it is still a useful background scheme to assist the interpretation of much that we observe in vegetation dynamics, and it also remains a tool for the prediction and direction of habitat management and active conservation.

Beech in competition with Oaks

The potential of beech to outgrow oak is partially due to the fact that the former is essentially both shallow-rooting and mycorrhizal, and thus it makes better use of, and more fully occupies, the most fertile upper horizons of the soil. This feature also allows beech to colonise and thrive on shallower, more rocky substrates than those preferred by oak, or required by the latter if it is to compete successfully with other tall tree species, including beech. The competitive benefit of shallow rooting allows beech trees to restrict the amount of water and mineral nutrients available in the soil to other species. This is also the case with Tilia spp. (Limes), another tree genus that did not reach Ireland in the post-glacial 'plant steeplechase' (Mitchell 1986, p. 72).

F. sylvativa casts such a heavy shade and its leaf litter is so resistant and slow to decay, that it restricts to a minimum the number of species that can maintain themselves in the field- and ground-layers of the woodland structure beneath its canopy (Watt & Fraser 1933). Another aspect of shallow rooting, however, is that it makes the beech tree vulnerable to summer drought, since it may not put its roots down deep enough to tap ground water sufficiently under conditions of prolonged dry weather. Whenever (or in Fermanagh, if ever!) drought occurs, large trees and their seedlings must compete for scarce water in the upper soil layers, which restricts the species ability to regenerate (Rackham 1980).

Invasion by F. sylvatica poses a threat to the purity of native or long-established plantation oak woodland in places like Killarney, where it likewise threatens the even rarer Yew woodland (Reynolds 2002).

Sexual reproduction, mast years, fruit predation and dispersal

Beech is not all that well equipped for seed dispersal. The tree reaches sexual maturity when it is somewhere between 30 and 80 years old, depending on the shade environment it inhabits. It then produces its tiny, green, wind-pollinated, monoecious flowers each April, at around the same time as the leaves are opening. Heavy seed crops of beech (and of Q. petraea (Sessile Oak) and Q. robur (Pedunculate Oak)) are sporadic, the so-called 'mast years' occurring roughly every five to 15 years. The timing of mast years, in beech at least, is dependent upon a good warm summer the year before fruiting, involving high July temperatures, low rainfall and plenty of sunshine. This must then be followed by a mild, frost-free spring. These special seasonal conditions together induce a heavy beech fruit crop. However, even within the best mast years, the quantities of fruit produced on individual beech trees varies greatly (Matthews 1955; C.D. Pigott, in: Milner 1992, p. 93).

The one or two seeds (triangular nuts), in their woody pericarp husk covered with coarse bristles fall beneath the tree, and birds such as rooks, jays, pigeons and pheasants, plus squirrels, mice and voles, collect them as food. The irregular, sporadic, heavy fruiting mast years, it is now realised, are the trees' evolved response to limit its losses due to this heavy seed predation by birds and mammals. In the intervening years between 'masts', beech trees produce very few or no fruits. It follows that the populations of birds and rodents decline due to

food shortage in these lean years. Then, when a mast arrives, the quantity of fruit is suddenly so massive that the diminished animal population cannot eat it all, and a high proportion of the trees' seed crop escapes predation. Many trees in the world with large edible fruit use this strategy, but in beech it is absolutely vital for survival, for only the seed released in mast years contribute to species regeneration. The few fruit and seedlings produced in the intervening years are all eaten (C.D. Pigott, in: Milner 1992, p. 92).

While a high proportion of the nuts collected by wild animals are eaten and digested, it is the few nuts that are transported but which the animals fail to eat that reproduce the species. Apart from the possibility of some degree of wind dispersal of the triangular nut, which might roll on a suitable surface, this is the only means of dispersal open to the tree (Ridley 1930; Hadfield 1957).

Toxicity of the fruit

The beech tree does attempt to protect its fruit in another way, but obviously this is not effective enough. The nuts and the pericarp contain saponins plus a poisonous uncharacterised substance called 'fagin', which in sufficiently high dosage is toxic enough to kill horses and cattle. Previously oil was extracted from beech nuts and the residue made into animal feed cake. This cake fodder, or the beech nuts themselves, have been known to poison both horses and cattle, the former being much more susceptible. Humans have also suffered after eating up to 50 or more nuts, a diet which produced symptoms of headache, soreness of the mouth and throat, vomiting and other unpleasant and dangerous effects. THE NUTS SHOULD THEREFORE BE REGARDED AS POISONOUS AND BE COMPLETELY AVOIDED (Cooper & Johnson 1998). Despite this, in his popular guide book to wild flowers and trees, Mabey (1972) gives instructions for extracting beech oil from the nuts, and also mentions the difficult labour of shelling them to eat.

Seed survival, germination and establishment

The seed may survive a year or so, but there is no evidence of prolonged viability or of a soil seed bank. Seed germinates in spring but seedlings are very susceptible to late frost, which may indeed be the limiting factor determining the northern and eastern boundaries of the indigenous distribution of the species in Europe, if not in England. The seedlings are very shade tolerant and are often found under bushes or other plants which undoubtedly provide them shelter from frost. For this same reason, beech seedlings and saplings never establish in open conditions (Jonsell et al. 2000). Nevertheless, saplings do not long survive under the dense shade cast by the beech itself (Grime et al. 1988), and their success in replacing existing trees is heavily dependent upon the frequency of light gaps appearing in the woodland canopy.

Variation

Although as a species F. sylvatica in Britain and Ireland does not vary greatly, and from around 1955 onward much of the seed of broadleaved trees planted here has been imported from continental sources (Gordon & Fraser 1982), a number of distinct forms of horticultural interest have arisen over the years. The most notable varieties probably are the purple 'Copper Beech', several pendulous forms, an erect fastigate form often used as a street tree ('fastigata'), and most interesting of all, the 'Fern-leaved Beech' (= 'Cut-leaved Beech'), named 'heterophylla' (= 'laciniata'). This is an example of a 'chimaera', an unusual type of graft having inner tissues of the ordinary beech overlain by tissue of the cut-leaved form (Mitchell 1974).

Fermanagh occurrence

It may seem very odd that the first record of this species in Fermanagh dates from 1934! However 19th century Irish botanists knew that Beech was alien and widely planted so they did not deem it worth recording (eg Cybele Hibernica 1866, 1898). In Fermanagh, Beech has now been recorded in 276 tetrads, 52.3% of those in the VC. F. sylvatica is very commonly found throughout the lowlands: it is more frequent in the east of the county and especially so on the larger demesnes. Although we do not have any pure beech woodland as such, merely small copses in mixed species plantations, the often linear representation Beech displays on the tetrad map reflects the fact that it is commonly planted in roadside hedgerows and along other linear geographic features. These include slopes and river banks (eg the Tempo, Colebrooke and Swanlinbar Rivers), plus along estate avenues (eg Castle Archdale and Castle Coole). Frequently on such estate lands it is possible to come across fine individual specimen trees, occasionally up to 30 m in height and some possibly over 200 years in age.

British and Irish occurrence

The New Atlas hexad map shows F. sylvatica common and widespread throughout most of both islands, but the distribution thinning to the north and west in both Scotland and Ireland. As discussed above, the information available does not allow accurate discrimination of minority native occurrences that possibly occur in SE England and SE Wales, so the map shows all sites as native.

European occurrence

The distribution pattern of F. sylvatica in Britain and Ireland suggests that exposure to wind and to frost are significant factors limiting the species occurrence (Preston et al. 2002). However, from what we know of the requirements for successful regeneration, it is clear that we really need to consider the map of the indigenous European distribution of the species (Jalas & Suominen 1976, Map 284). This shows that beech is widespread throughout W and C Europe and extends northwards to 60^oN on coastal Norway. Southwards, the presence of F. sylvatica thins and fades away in the Mediterranean basin, although the tree is recorded in S France and down the length of Italy to Sicily, the only other Mediterranean island it reaches is Corsica. In the east, the species reaches Moldavia and the Crimea (Jonsell et al. 2000).

Uses

The usually white, close grained wood is hard, smooth and strong and is widely used for furniture, tool handles, sports equipment and kitchen utensils. Milner (1992) contains interesting information on both the uses of the timber and other parts of the tree, and the extremely limited folklore that is associated with it.

Names

The genus name 'Fagus' is the ancient Roman name for the tree (Gilbert-Carter 1964), but it may be derived from the Greek verb 'phagein' meaning 'to eat', since in famine times beech mast was eaten by starving people, and must have made many of them ill (Milner 1992). Grigson (1974) has however pointed out that in Greek 'phegos' refers to oak not beech, and the local Greek oak, Q. macrolepis (Valonia Oak) does have very large acorns that were sometimes eaten by man. The Latin specific epithet 'sylvatica' means 'growing in woods' from 'silva', 'woodland' (Gilbert-Carter 1964).

The English common name 'Beech' is derived from the Old English 'bece', 'boc', 'beoce', words that mean, with a difference in gender only, 'a book' and 'a beech tree'. The connection between the two is that the Runic tablets on which people once wrote were made of beechwood (Prior 1879). 'Bece' and its associated dialect forms are word elements in many place names, including some well beyond the indigenous area of the tree, eg Beckwith in W Yorkshire (previously 'Becwudu', meaning 'beech wood', around 972 AD), and Bitchfield in Northumberland (Hadfield 1957; Grigson 1987).

Alternative English common names include 'Buck', transferred directly from the Old English. Another name is a reference to a rough-barked variety of the tree, 'Hay Beech'. The fruit of the tree is well known by the name 'Mast', and other forms exist as 'Buck Mast' and 'Buck's Mast'. In Hampshire, when pigs are turned out into the beechwoods in autumn to feed on the beech nuts, they are said to be turned out to mast (Britten & Holland 1886).

Research literature

There is a huge wealth of published research on beechwoods in both the British Isles and in Europe. Rodwell (1991a) provides a comprehensive bibliography and is a suitable entry point to the literature on this and other forms of British Isles woodland and scrub for anyone who requires further reading.

Threats

As mentioned above, F. sylvatica invades native Irish oak woodland, but it is under no conservation threat itself since it is an introduction. However, in Fermanagh and other areas of Ireland at least, the beech population is old and very definitely living on borrowed time due to frequent wind throw and a general lack of regeneration. The frequency of wind throw appears to be increasing as global warming makes our climate more extreme. Being a familiar and attractive tree, additional beech planting is surely desirable. The invasive fungal pathogen Phytophthora ramosum that recently arrived in N Ireland is known to have infected and severely attacked beech trees in England, but this has not yet occurred here.

Castanea sativa Mill., Sweet Chestnut

Introduced, neophyte, deliberately planted and rare or very occasional. European temperate, also cultivated and widely naturalised in Europe.

17 September 1986; McMullin, A.S. & Corbett, P.; Mullynacoagh Td shore, Upper Lough Erne.

January to November.

Growth form and preferred habitats

This is potentially a very large deciduous tree, up to 30 m with a wide, broad crown. Older trees have deeply fissured bark on the trunk, the longitudinal fissures often spirally curved, allowing the species to be easily recognised at any time of year. Ecologically, C. sativa can tolerate most soils, although it thrives best on moist, acidic, sandy ones (T.D. Dines, in: Preston et al. 2002). In season, the leaf canopy casts a heavy shade, while the deep, persistent leaf-litter creates a mor humus, often with a pH of 4 or under (Rackham 1980, p. 332). In Fermanagh, conspicuous, distinctive and often very large old trees of this species are almost always associated with parkland or tree collections (arboreta) in landed estates such as Castle Coole, Crom or Colebrooke. Unlike the quasi-native chestnut woods in Kent, E Essex and SE Suffolk described by Rackham (1980, pp. 332-9) which are or were coppiced every 12-14 years for small timber, in Ireland C. sativa never formed woodland stands. A possible reason for this is the more recent date of introduction and plantation of the tree in Ireland, probably sometime before the 17th century – although nobody really knows a definite date (Nelson & Walsh 1993, p. 105).

Flowering reproduction

Trees flowering in June and July, the minute flowers being borne on long catkins held erect at the tips of shoots. Individual catkins are 10-20 cm long, with female flowers at their base and more numerous male flowers on the remainder of the catkin length. Various insects pollinate the flowers and the characteristic prickle-covered green chestnuts develop until the autumn. In most of Ireland, the vast majority of ovules abort and it is doubtful if any of the chestnuts produced in northern counties are fertile and capable of germination as self-sown trees are very rare here. The survey of the Belfast urban area in the 1990s produced a solitary seedling at the edge of playing fields (Beesley & Wilde 1997; Reynolds 2002).

European region of origin

C. sativa is a native species of the more humid areas of eastern Mediterranean countries. However, selected cultivated forms, some of them grafted with superior fruiting scions, have long since been introduced into more northerly regions of Europe, and the tree and its edible nuts and other products have been present in Britain since Roman times (Zohary & Hopf 2000, p. 189). Despite the familiar 18th century English common name 'Spanish Chestnut' and the present day frequency of the tree in the Pyrenees, the species is unlikely to be native as far west as Spain (Rackham 1980, p. 329). The native range of C. sativa is controversial. The editors of Flora Europaea took a conservative view, regarding it as indigenous in the Balkans (Albania, Greece, Jugoslavia) and Turkey, and less probably in SC Europe (Tutin et al. 1993). It is common in Italy as well as in southern France and Calabria, where it is extensively coppiced (Rackham 1980, p. 332).

British and Irish occurrence

The New Atlas hectad map demonstrates that C. sativa is much more thinly scattered in Ireland than in Britain, and especially so when compared to S England and Wales. The tree flowers and fruits readily, and in suitable, warmer conditions in S England, if not elsewhere at present, the nuts germinate in the spring after their production. Unlike Beech and Oaks, losses due to seed predation appear rare in England. As it cannot regenerate under its own shade, transport into gaps, or into suitable sites under other species is necessary, the likely vectors being rooks (Rackham 1980, pp. 334-5).

Fermanagh occurrence

This is a rare or very occasional tree in Fermanagh, having only been recorded in 15 tetrads, 2.8% of those in the VC. It is confined to estate parks and woodland plantations, plus a few outliers nearby. Only one or two of the records occur on the margins of the demesnes mentioned above and, although I greatly doubt it, these might just be self-sown trees. However, in all our years of field recording RHN and I have never found either seedlings or saplings of Sweet Chestnut anywhere in Fermanagh.

In truth, the reproductive biology and local behaviour of the exotic (ie non-native) trees and shrubs in N Ireland have had almost no attention paid to them. In many cases we do not know whether they can successfully set seed, disperse and establish in our part of the world, as many of them undoubtedly do in S England (T.D. Dines, in: Preston et al. 2002).

A well-known English tree expert, Alan Mitchell, suggested that only a few exotic, introduced trees "plant themselves" in Britain and Ireland. He highlighted one very obvious exception "that proves the rule", as Acer pseudoplatanus (Sycamore). Mitchell went on to list Sweet Chestnut along with several introduced conifers that he believed could produce natural seedlings, although he reckoned these were likely to be "very local" (A. Mitchell, in: Milner 1992, p. 158). In Fermanagh, as elsewhere, in the autumn we can certainly find beneath the trees shiny chestnuts inside the leathery, green or brown, heavily-spined husks which split into four valves. However, it is very doubtful if they contain viable seed in Ireland as far north as Fermanagh. This contrasts with the situation in Essex and Kent where Rackham (1980) found C. sativa regenerating successfully, and probably even capable of invading woodland of other species. A study of the ability of exotic, non-native trees to regenerate or not, would be another example of a feasible school project, where a very little study could fill a gap in our knowledge regarding the performance and the local behaviour of long-introduced plants.

Uses

The tree is valued for its nut crop. For many generations it formed an important part of the traditional diet of farming communities, and it was also fed to stock animals.

Names

In S England, Sweet Chestnut survived throughout the Dark Ages, and acquired an Anglo-Saxon name, 'cyst' or 'cisten', the latter form (pronounced 'chisten'), being a derived survival from the Classical Latin name, 'Castanea'. The tree name then made the transition to a word element in at least a few English place names, for instance, 'Cystewde' of 1272, which is today's 'Chest Wood' near Colchester (Rackham 1980, p. 330).

Threats

None.

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