Research

Scotland’s native wild apple – Malus sylvestris: ecology and management

By Rick Worrell, Markus Ruhsam, James Renny, Will Jessop and Graeme Findlay

Summary Foresters and ecologists appear to have dismissed wild apple (Malus sylvestris (L.) Mill.) as unworthy of professional attention and as a result we know virtually nothing about the ecology and management of the species. This paper provides a basic account of the characteristics of wild apple based on a survey of 209 trees across Scotland and northern England. It is widely distributed, except on the most acid lithologies, and occurs infrequently as single trees or small groups, and never in stands. It grows in ancient woodland, usually near woodland edges and reaches Figure 1. Wild apple in ancient wood pasture near St Johns Town of Dalry, Galloway. its highest densities in wood pasture. It can grow to be a more substantial tree Introduction and aims paper to this (Worrell et al., 2019) points than previously reported: i.e. heights Foresters and ecologists have made out that we have been unsure even up to 14m and stem diameters up to good progress in the last few decades whether it is a native species, unclear 1m. The current generation is 60-120 on increasing our understanding of which of the apple trees growing in the years old and natural regeneration is Scotland’s rather limited palette of countryside are actually Malus sylvestris scarce, and there is a need to establish native trees and shrubs, but some species and uncertain how to identify it. The a new generation of trees. Cattle, have missed the spotlight altogether, main aim of the work described in this and presumably therefore formerly and none more so than wild apple. paper was to carry out an ecological auroch, appear to be heavily involved in Somehow both foresters and ecologists survey of wild apple in Scotland to try to promoting regeneration. It is a tree with have dismissed the tree as unworthy remedy this lack of knowledge. The main considerable potential as a spectacular of professional attention. As a result objectives were to determine: landscape tree, as a prolific producer we know virtually nothing about the • The distribution of the wild apple of a ‘non-wood forest product’ and as woodland types in which wild apple (M. sylvestris) and its ecological an important species for threatened trees grow; their site requirements and characteristics and requirements pollinator insects. Outline guidance their regeneration biology; and their • Basic information about its on conservation and management is distribution in Scotland has never been flowering, fruiting and regeneration. provided. properly described. The companion On the basis of the results we propose basic management and conservation guidance for wild apple, and propose how the species might be used in the THE AUTHORS future. Wild apple clearly needs to Rick Worrell* has been a self-employed forestry consultant since 1987, specialising in native be incorporated as a component of woodland and growing broadleaves for quality timber native woodlands and has potential as a Markus Ruhsam is a plant molecular ecologist at the Royal Botanic Gardens Edinburgh (RBGE) with spectacular landscape tree, as a prolific a particular interest in how plant populations change over time producer of a ‘non-wood forest product’, James Renny is a naturalist based in Grandtully, Perthshire and as a food source for threatened Will Jessop was an MSc student at the RBGE and is now studying for a PhD at the University of pollinator insects (Scottish Government, York 2017). In addition, our fieldwork revealed Graham Findlay is Planning and Conservation Forester at Forestry and Land Scotland, Dunkeld signs of past management and use of Office wild apples and their hybrids, and so *Corresponding author: [email protected], Upper Park, Aberfeldy, Perthshire PH15 2EH this paper briefly touches on evidence of

Volume 74 No. 1 Spring 2020 | 21 Research

the common history of people and wild trees). Ecological information was form stands. It occurs at highest density apples in Scotland. recorded for up to 209 of these covering: in wood pastures (up to ten mature Rather more is known about the • Tree size and structure: Height trees/ha), and makes a very startling ecology of the species in continental (m), crown diameter (m), stem contribution to the landscape in spring Europe, i.e. that it is a tree of woodland diameter (cm) at ground level, and (Figure 1). edges, growing as single individuals number of stems (183 trees). Trees This project produced the first or small groups, widely distributed thought to have been pollarded were map which unambiguously shows but never common, tolerating a range noted occurrences of wild apple (Figure 2 of soil types, often found on river • Site characteristics: Elevation, left). However this is an incomplete floodplains, and relatively intolerant of topographic position, habitat picture due to the limited number of shade (Stephan et al., 2003, Schnitzler (ancient woodland, native samples and varying sampling effort et al., 2014). Interestingly it is listed in woodland, other woodland, wood in different parts of the country. The Europe as a noble hardwood, capable pasture, hedgerow, or field); Botanical Society of Britain and Ireland of small-scale decorative timber National Vegetation Classification (BSBI) map for Malus sylvestris (Figure 2 production (Karen Russel, pers comm.). (NVC) woodland community middle) and a map we produced (Figure The ultimate aim of this paper is to (Rodwell, 1991); associated ground 2 right) using data from the Native encourage forestry and conservation flora species (209 trees) Woodland Survey of Scotland (Forestry professionals in Scotland to take the • Location: National Grid Reference Commission Scotland, 2014) give better species seriously and address some of and other information describing coverage, but do not distinguish between the symptoms of our collective neglect. the location. wild, hybrid and domestic apple. Whilst none of these maps is wholly adequate Methods Germination studies on its own, together they build up a A total of 342 apple trees were located in Apple seeds were collected from 24 wild fairly clear overall picture. Details of Scotland and parts of northern England apple trees and sown in the autumn in identification using both morphological (as described in Worrell et al., 2019), potting compost. Their germination and characteristics and DNA analysis are focusing primarily on wild apples (M. the growth of seedlings were recorded. given in the companion paper (Worrell sylvestris) and probable hybrids (M. et al., 2019). sylvestris × domestica), but also including Results Wild apple has strongholds in Loch a few feral domestic apple trees. These Species distribution Lomond and the Trossachs, Highland trees were identified to species, initially Wild apple occurs as single trees or Perthshire, and Dumfries and Galloway. in the field based on morphology, with occasionally in small groups, scattered It is apparently largely absent in many emphasis on leaf hairiness, crown through native woodland, wood pastures, parts of Deeside and Strathspey. Oddly, structure and, if available, fruit size as hedgerows, pastures and low-lying it also appears scarce in much of the described in Worrell et al. (2019), and moorland. As with other tree species of Western Highlands, but not wholly confirmed later via DNA analysis (342 the Rosaceae (cherries, rowan) it does not absent. Possibly the most interesting

Figure 2. Distribution of Malus sylvestris: Left – wild apple verified by DNA in this study, with darker red dots reflecting greater numbers of samples at a location. The distribution partly reflects natural factors and partly surveying effort.Middle – according to the BSBI New Atlas but including M. domestica and hybrids (www.brc.ac.uk/plantatlas/plant/malus-sylvestris-sl). Right – woods containing ‘crab apple’ from the Native Woodland Survey of Scotland, though actually the data points include domestic and hybrid apple and some that were misidentified as apple.

22 | SCOTTISH FORESTRY Research

human dispersal; these are mainly hybrids and domestic apples.

Elevation The highest elevation trees were at about 300m asl in both Scotland and northern England. The highest tree in the Lake District is at 305m in Patterdale, on an Above: Figure 3. The most north-western wild apple tree recorded in Scotland, at 200m asl, near exposed ridge (see Figure 4); and this Dundonnell, south of Ullapool. Below: Figure 4. The highest apple (a hybrid) located at 300m in hybrid tree had grown to 10m tall and Patterdale, Lake District. It had its top 8m blown out (lying to right), but is regrowing vigorously. over 70cm stem diameter, had its top 8m blown out, but is now re-growing vigorously – suggesting that apple may be capable of growing at higher elevations than this. Anecdotally, a wild apple seedling planted accidentally in the southern uplands at 650m appears to be growing adequately (Philip Ashmole, pers comm.), which is rather startling, though its long term survival is obviously uncertain.

Climatic conditions Wild apples occur in a very wide range of climatic conditions (see Figure 5) i.e. in areas with: • annual rainfall ranging between 600mm (Lothians) to over 3000mm (Lake District) location at which wild apple was found (Argyll), to apparent absence just a little • annual temperature ranging from in the west was on the valley slopes further west. It is common in northern over 1500 degree days (above above Dundonnell, near Ullapool, where Perthshire but apparently absent in 6oC) in lowland Dumfries and the seven trees occur in ancient woodland nearby Strathspey and Deeside. The Galloway coast, to 1100 degree days and wood pasture. These were only incidence of wild apple reduces markedly in Shetland, and down to 700–800 recorded because the estate owner and on acid lithology such as Moine schist degree days on some upland sites. gardener had a special interest in the and granites, in the same way as for Some of the finest woods for wild trees; they would otherwise not have elm and ash. However, occasionally apple, with the highest densities of apple been picked up in this survey. There had wild apple trees are recorded on acid trees, were at Thorneythwaite, near been botanical surveys on this estate geology on favourable sites, e.g. several Seatoller in the Lake District, one of the that had recorded these trees as early large trees at the western end of Loch UK’s wettest locations (annual rainfall as 1923 (D Rice, pers comm.). They give Rannoch. 3550mm). The climatic envelope shown every impression of being of natural Figure 2 (middle and right) show in Figure 5 encompasses conditions origin (e.g. Figure 3); and if wild apple high densities of records in the Central found over virtually all of Scotland occurs naturally as far west and north Belt between Glasgow and Edinburgh, below about 200-300m and it can be as Dundonnell, it is probably present including many peri-urban areas. DNA concluded that climatic conditions are elsewhere in the Western Highlands. So analysis and field survey (Worrell et al., unlikely to limit the occurrence of wild the apparent lack of it here is probably 2019) showed that most of these are in apples anywhere in Scotland below due to insufficient surveying as well fact planted hybrid and domestic apples. this elevation. The tree from Shetland as rarity. There is also a cluster of The records in lowland agricultural areas suggests particularly impressive site wild apple located around Contin and in Angus, Aberdeenshire and Moray are tolerance. Torrachilty, which are currently the in all likelihood mainly planted trees and most northern records on the mainland. include wild apple, hybrids and domestic Habitat and site characteristics Remarkably there is one record for a wild apple trees. We found planted trees of Table 1 shows the breakdown of broad apple confirmed by DNA analysis from all sizes in lowland areas, suggesting habitats types in which wild and hybrid/ a sea-cliff of an uninhabited island in planting had happened continuously feral domestic apples were recorded. Shetland. over the last hundred years or more, and Over the whole study area, wild apple is There are some abrupt transitions, was ongoing. There is also a scattering well represented in woodland (45% of for example between high densities of trees beside roads, tracks and railways occurrences), and wood pasture (37%); around Loch Lomond and Faslane which are clearly the result of accidental with only 16% occurring in hedges

Volume 74 No. 1 Spring 2020 | 23 Research

and pastures. Of those trees recorded in woodland, 75% were in ancient woodland. The proportion of wild apples in ancient woodland was highest in Highland Scotland (44%), and lowest in Lowland Scotland (8%). In Lowland Scotland, where ancient woodland is scarce, most wild apples appear in hedges and pasture (58% of occurrences). Most of the wild apples in northern England and the Southern Uplands were located in ancient wood pasture (75% and 59% respectively). The picture is rather different for hybrid and feral domestic apples (Table 1). While most of these were still found in woodland (53% of occurrences), they were more commonly found in Figure 5. The climatic envelope of wild apple in northern Britain in terms of annual temperature and precipitation. Each point represents a sampled wild apple tree. The arrows indicate the climatic hedges, pasture and moorlands (33%) conditions in different locations where wild apples were located in this study. than wild apples. It is interesting that we still found a few hybrid apples in ancient woodland (though only 11 trees suggests that the pasturage of cattle in species in Scotland might be more shade in total). These have all the appearances these woods, which continues to this day tolerant than the literature suggests. of having arrived by natural means and in some, is beneficial to recruitment of hence are likely to be a result of natural apple regeneration (see Figure 10). This Woodland communities and associated hybridisation (rather than being planted probably explains the striking frequency tree and plant species stock), but we will never know their of wild apple in places like Galloway Wild apples occurred in oak woods, origins for certain. which have a strong history of cattle birch woods, ash woods and wet alder A considerable proportion of wild farming. woodland; therefore showing quite apples were found in upland gullies a high range of ecological tolerances and riparian woodlands, which may Shade tolerance (see Table 2). They are however most simply reflect that this is where remnant Wild apples are reported in the European common in oak-birch woodland. native woodland is most common, or literature as growing only in open The tree and shrub species growing in may indicate an ecological preference. conditions such as woodland edges, the immediate proximity of wild apple There are interesting and quite large glades and hedges, and this is interpreted trees reflect the main woodland types, populations at high water mark on the as the species being intolerant of shade i.e. oak, birch, ash and alder (Table islands and shoreline of Loch Lomond (e.g. Stephan, 2003). In this survey 3). The high incidence of hawthorn is and Loch Ken (Dumfries and Galloway) most trees were found on woodland eye-catching; the species being one of and we assume these have spread via edges, but we found several trees in far the most common associates in wood floating apples. These loch-shore trees more shaded conditions including a few pasture, and suggests that the two seem to be able to thrive in a surprisingly growing, and fruiting, under a partial species might have similar regeneration wide variety of soils including wet sand understory of oak. This suggests that the requirements. and gravel, rock crevices, bog myrtle mires and deep alluvial gleys (Figure 6). Figure 6. A small wild apple tree on the shoreline on the Loch Lomond islands. We have also observed apples floating down watercourses, so water may be one dispersal method.

Wild apple in wood pasture Wild apple is also strongly represented in wood pasture (37% of records); this being particularly obvious in southern Scotland and northern England (mainly the Lake District); but also occurring elsewhere, as far north as Dundonnell, Ross-shire. It is in ancient wood pasture that we found wild apples at their highest densities. The pattern of occurrence

24 | SCOTTISH FORESTRY Research

Table 1. The percentage of apple trees in different habitats and regions. About 40 ground flora species were PERCENTAGE OF TREES IN DIFFERENT HABITATS AND REGIONS recorded growing with wild apple trees. The most common associated species Wild apple were bracken, bluebell, wood sorrel, Highland Southern Lowland Northern Total Total no. of broom and wood sage (indicators of HABITAT Scotland Uplands Scotland England samples oak-birch (dry acidic) woodland); dog's Ancient Woodland 44% 28% 8% 0% 34% 56 mercury (indicator of ash (dry neutral) woodland); and creeping buttercup and Other native woodland 5% 3% 8% 0% 5% 9 rushes (indicators of alder (neutral) wet Non-native (mixed) wood 6% 3% 25% 0% 6% 10 woodland). Ancient woodland indicator species were well represented. Epiphytic ALL WOODLAND 55% 34% 41% 0% 45% 75 lichen and moss communities on wild apple bark do not appear particularly Wood pasture 29% 59% 0% 75% 37% 61 diverse; though domestic apples in the Hedges, pasture, 14% 7% 58% 25% 16% 28 western Highlands and Inner Hebrides moorland often have spectacular Lobarion Other 3% 0% 0% 0% 2% 3 communities (Alison Averis, pers TOTAL 100% 100% 100% 100% 100% 167 comm.). Old horizontal branches of wild apple sometimes host luxurious moss carpets, sometimes including common Hybrid and feral domestic apples polypody ferns. Highland Southern Lowland Northern Total Total no. of HABITAT Scotland Uplands Scotland England samples Tree dimensions and structure Ancient Woodland 49% 20% 0% 20% 26% 11 Wild apples are rather more substantial trees than suggested in the literature, Other native woodland 6% 0% 21% 0% 10% 4 with an average stem diameter of 52cm Non-native woodland 12% 0% 36% 0% 17% 7 (at ground level), and a maximum ALL WOODLAND 67% 20% 57% 20% 53% 22 diameter of 120cm. Eleven of the wild apple trees sampled for structure had Wood pasture 0% 60% 0% 60% 14% 6 stem diameters over 90cm. This is very different to some published accounts, Hedges, pasture, 33% 20% 43% 20% 33% 14 e.g. Stephan et al. (2003) suggest a moorland maximum stem diameter of 45cm. Other 0% 0% 0% 0% 0% 0 Mean height was 8m, with a maximum TOTAL 100% 100% 100% 100% 100% 42 of 14m; well in excess of the maximum value of 10m which is generally quoted. Notes: for hybrid/feral domestic apples, sample numbers for individual regions were too low to draw Average crown diameter was 8m, with a conclusions. ‘Other’ habitats were adjacent to railway tracks and farm tracks. Samples in ‘Northern England’ came mainly from the Lake District, Cumbria, with a few from Northumberland. maximum of 17m (the tree in Figure 1 of Worrell et al., 2019). So we need to revise our mental picture of mature wild apple Table 2. Occurrence of wild apple in different woodland types trees from small trees, to something Woodland type NVC types Proportion of records more the size of wild cherry trees. The hybrid apple trees in this survey Oak-birch woodland Mainly W11, some W10 59% were rather smaller than the wild Ash woodland W9 23% apple trees, but we suspect this is not Alder wet woodland W7 18% a biological reality but an outcome of Total 100% sampling. A higher proportion of hybrid trees were younger, smaller planted trees, whereas wild apples were larger, Table 3. Occurrence of native trees and shrubs growing in the vicinity of wild apple older, self-set trees. Apple trees coppice and pollard very easily, and our sample Tree species Proportion of locations Shrub species Proportion of locations included 14 apparent pollard trees (see with species (%) with species (%) Figure 7). In the Lake District we found Oak 17 Hawthorn 18 signs of historic pollard cutting. Birch 16 Hazel 9 Ash 12 Willow 3 Flowering and fruiting Alder 9 Blackthorn 2 Both wild and hybrid apple trees flower and fruit regularly and profusely. In the Rowan 5 Dog Rose 2 two fieldwork seasons of this project, Holly 5 Juniper 1

Volume 74 No. 1 Spring 2020 | 25 Research

84% of wild apples and 92% of hybrid/ in cattle droppings on such sites (MA feral domestic apples were recorded as Smyth, pers comm. – see Figure 10). flowering or fruiting, though it is not Older wild apple seedlings look rather possible to say how typical these years like blackthorn, which adds to the were. In this respect apples are like other challenge of finding them, and are regularly flowering trees in the Rosaceae, similarly defended by thorns (Figure 11). such as cherries and rowan, and different from many wind-pollinated trees where Tree age flowering is typically more variable. Fruit By counting rings on cut sections of is produced in significant quantities on lower branches of a small sample of most trees and represents a considerable trees, we estimate that most trees, wild annual investment of resources. The and hybrid, were about 60–120 years fall of apples is variable, with most trees old. This would date their establishment dropping large quantities of fruit during to the late-19th to mid-20th centuries, the autumn (see Figure 8), but others when many native woods expanded as retaining apples in the crown during a result of agricultural abandonment winter, and some very occasionally during economic depressions, or of persisting into the following spring (see deliberate protection as forestry practice Figure 9). Apples are insect-pollinated, developed. A large proportion of larger and much insect activity was apparent trees had many veteran characteristics, to the main stem by the thinnest strip around trees on still, sunny days during and the Woodland Trust Veteran tree of bark. Only two trees were found flowering in mid-to-late May, with bees inventory has a number of records for apparently affected by disease. prominent, but hoverflies, flies, and ‘crab apple’ (https://ati.woodlandtrust. occasionally, beetles were also present. org.uk/). However, large trees in our Discussion and recommendations sample, that appeared to be quite old, This paper sets out the main ecological Germination and regeneration turned out to be younger than they first attributes of wild apple, which should Seeds germinate readily. Seed-lots from appeared. enable foresters and ecologists to better 24 parent trees were planted in compost understand its ecological place and value in a seed tray and left outdoors over A hardy tree in our ancient woods and wood pastures, winter; 42% of the seed-lots germinated Nearly all the trees surveyed were in and to integrate it into native woodland in the first spring and 54% germinated very good health. They appear to be management. Key aspects are: the following spring. Two seed-lots incredibly resilient. They blow over 1. It is widely distributed, except on produced seedlings both years and only and re-sprout freely as ‘phoenix’ trees. the most acid lithologies, and occurs two seed-lots (8%) failed to germinate. About 20% of mature trees were phoenix infrequently as single trees or small Naturally-regenerated saplings were trees, making wild apple one of the most groups, and never in stands. It grows found only very occasionally in this adept species at this growth form. A in ancient woodland, usually near survey, though we did not specifically sizeable proportion of the oldest trees woodland edges and reaches its set out to find them. The best examples grow hollow then collapse, but then get highest densities in wood pasture. were in areas of wood pasture where a second wind and start growing again as 2. It can grow to be a more substantial there is a history of pasturage of cattle. phoenix trees. Broken limbs will live on tree than previously reported Wild apple seeds and seedlings appear and recover provided they are connected (height up to at least 14m and stem diameters in excess of 1m). 3. The current generation of wild apple trees is quite old. Natural regeneration is apparently scarce and so we need to establish a new generation in our woods and wood pastures. 4. We only partly understand its reproduction and regeneration, i.e. pollination, seed dispersal, germination and early growth; this limits our ability to achieve natural regeneration. 5. It is a tree with considerable potential as a spectacular landscape tree, as a prolific producer of a non-wood forest product, and as an important species for threatened Figure 7. Pollard wild apple tree at Loch Katrine. pollinator insects.

26 | SCOTTISH FORESTRY Research

Left: Figure 8. A heavy crop of fallen apples, Bridge of Earn. © Andy Fairbairn

established in cow pats (Buttenschøn conditions. The other key message is and Buttenschøn, 1998), suggesting an that efforts to manage wood pastures almost obligate relationship. Wild boar, and establish new ones, should include badger and bears are reported eating and wild apple. dispersing wild apples, as are birds such The dispersal of apples by water is as blackbirds and thrushes. This points implicated by the population of trees to important ecological associations at high water mark on the shores and between wild apple and some of our islands of Loch Lomond and Loch Ken, now extinct large mammals, especially which is an interesting observation aurochs, which might partly explain not encountered in the literature the contemporary lack of regeneration. from the Continent and worthy of Fortunately, it appears that modern further investigation. We also observed cattle are helping to fulfill the role as apples dispersing down watercourses, regeneration agent in upland wood and wonder if this is linked to their pastures; though the historic switch occurrence in upland gully woodlands. from cattle to sheep rearing has almost certainly been detrimental to wild Conservation status and ecological value apple. The status of apples as food for Wild apple is regarded as threatened in animals points mainly in the direction several European countries (Kik et al., A new generation of wild apple trees of animals now extinct in Scotland such 2011), though IUCN’s overall assessment The current generation of wild apple in as bears and aurochs, and highlights the for Europe is that the species is ‘data native woodland appears to be mainly difficulties of properly managing wild deficient’. The lack of understanding of 60–120 years old, and regeneration apple with these animals missing. The its distribution in Scotland, reported in appears to be scarce. Foresters and thorny early growth of wild apple also this paper, is a good example of that data ecologists need to learn to recognise suggests adaptation to co-existence with deficiency. The main threat reported by young wild apple and identify likely grazing animals and indicates an element IUCN is that of hybridisation, though places to search for seedlings and of co-evolution, which is significant in lack of regeneration owing to changes saplings. More basic research is the context of the debate about the role in woodland management is also required to evaluate the occurrence of of large herbivores in shaping native mentioned. IUCN’s main conservation regeneration, starting in wood pasture forests in the past (Vera, 2000). The requirement is that research is necessary with frequent wild apple trees. We association between cattle and wild to determine the effect of hybridisation suggest owners and agents should make apple suggests that controlled cattle and efforts made to establish ex situ a special effort to protect naturally- grazing may be a key for establishing conservation collections; this project regenerated seedlings and saplings, and regeneration in the future, i.e. wild has gone a long way towards fulfilling otherwise plant a few to perpetuate the apple may be a beneficiary of efforts to that in Scotland. It would also be useful species. Planting could be carried out as introduce controlled grazing in native to bring some of the best examples of a priority into existing native woodland/ woodland and wood pasture. The fact wood pasture with wild apple into formal wood pasture where the species is that hawthorn is such a frequent co- conservation protection. known to occur locally. This can then be associate of wild apple suggests that both Wild apple is a relatively rare complemented by including wild apple species benefit from similar regeneration tree in most (but not all) woods we in native woodland creation schemes, planted in small numbers (say up to 10–20 in a wood) in oak-birch woods (NVC W11), ash woodland (W8/9) and on drier alder woodland sites (W7).

Seed dispersal and an association between apples and animals Association between cattle and wild apple regeneration has been reported from Denmark, with cattle and horses both eating and dispersing the seeds and providing germination sites in dung (Figure 10, and Buttenschøn and Buttenschøn, 1998). Seeds pass unscathed through the animals and the poaching of ground can create good regeneration conditions. In a study in Denmark, over 90% of wild apple seedlings were found to have Figure 9. Previous years apples retained until next flowering season, Patterdale, Lake District.

Volume 74 No. 1 Spring 2020 | 27 Research

encountered. It hits the sweet spot of fertility and good luck. between being common enough to be The apple fruit as an image is ecologically meaningful and rare enough universally known and obviously to be notable when you find one (rather appealing: think of Newton’s apple, New like aspen in Scotland (Worrell et al., York as ‘The Big Apple’, and nowadays of 1995). In addition to the problem of course, the Apple iPhone. regeneration, there may be issues with Foresters and ecologists need to pollination resulting from the scarcity Figure 10. A naturally regenerated wild apple reinstate wild apple in our collective of trees. When collecting seed for the seedling in an old cow pat, southern Scotland. professional consciousness as a tree © Mary Anne Smyth. germination studies there was some of considerable value, and one to evidence that isolated trees produced be brought into active woodland fewer apples, and those apples had fewer Viking ship in Norway dating from 820, management. seeds in them, suggesting that isolation and there is a reference in the medieval inhibits pollination and therefore period, from the town of Egremont in Recommendations seeding. This could be a mechanism Cumbria, of a ‘crab fair’ involving apples • Professional and popular literature contributing to the apparent scarcity in (Egremont, Crab Fair (no date)). There on native woodland should feature areas like the Western Highlands, where are written accounts from Scotland and wild apple in the same way as for it is possible that land-use pressures Norway from 1500–1780 of wild apples other native tree and shrub species led to the reduction of woodland to being used for juice and vinegar (Åsen, • The main populations of pure wild the point where pollination distances no date). In this survey we looked for apple should be catalogued, and became so large that this contributed to evidence of wild apples planted close owners, managers and agencies regeneration failure. to old farms, as has historically been made aware of their existence and Their most obvious ecological the case with wild cherry, rowan and value. These areas include Loch values would appear to be the prolific sycamore; it was conspicuously absent. Lomond/Trossachs, Galloway, flowering as a source of pollen and Presumably its place was gradually taken Highland Perthshire and the Lake nectar for pollinating insects, and fruit by domestic apple grown in orchards District, and smaller areas further for mammals and birds, so this is a and gardens; with orchards known north and west. It would be useful species to promote in the context of our to have been present in Scotland as to continue searching for wild apple threatened pollinators. Wild apple is early as 1100 (Robertson, 2007; Hayes, in western Scotland to complete our recorded as having 118 associated insects 2016). In modern times crab apple inventory of the species and mite species, and ranks tenth out has maintained a tenuous place in the • Some of the best examples of wood of 28 tree and shrub species listed by public consciousness because of its use pasture with frequent wild apple Kennedy and Southwood (1984). as for setting jams and making apple need to be brought into formal jellies; this makes apple one of the conservation protection. Landscape value more useful non-timber forest products • In areas with frequent wild apple, Mature wild apple trees are almost (Martynoga, 2015). It is still used as an owners should be encouraged to without exception characterful and effective pollinator of domestic apples establish a new generation of trees impressive; a veteran wild apple at in both orchards and gardens, and the flowering time is as spectacular as it myriad cultivars of foreign crab apple are comes (Figure 1). This is equally true popular as decorative garden trees. of many of the older hybrid apples we Wild apple suffers from an image found too. Planted singly, or in small problem. The common name crab apple clumps, at the edges of woods or near and its colloquial equivalent ‘scroggie’, clearings, or in wood pasture, wild apple derive from words meaning small, untidy gives outstanding value. and scrubby. It genus name, Malus, means ‘evil-one’ – think of mal-words History, uses and cultural associations like malformed, malice, malaria and Wild apple appears to have been used dismal – apparently because the apple by people for millennia, but evidence was the fruit of the tree of ‘knowledge is scarce. It is listed from sediments in of goodness and evil’ that tempted Eve Yorkshire from 10,500 years ago, and and led to mankind’s expulsion from is recorded as burnt timber or food the Garden of Eden, with the apple remains from the late Neolithic to the appearing in some Christian art as a early Bronze Age from a few sites in symbol for sin. On the other hand, Scotland, and rather more in England apples have several positive cultural (Dickson and Dickson, 2000). It was connotations, e.g. the term ‘apple of protected as pannage for pigs in Europe my eye’ is of biblical origin; the apple as early as the 6th and 7th centuries is sacred in Norse mythology and has Figure 11. Naturally regenerated wild apple (Vera, 2000). It has been found in a featured in several cultures as a symbol saplings and young trees in cattle pasture, Cree Valley, Galloway.

28 | SCOTTISH FORESTRY Research

by protecting existing saplings wherever they can be Vol. 53, 2, 455-478. found, and by planting small numbers of genuine wild Martynoga, F (Ed) (2015) A handbook of Scotland’s wild harvest. Saraband. apple Mitchell, A (1992) Trees of Britain and Northern Europe. Collins. • Elsewhere across Scotland, small admixtures of wild Robertson, FW (2007) A History of Apples in Scottish Orchards. Garden apple should be planted as part of new native broadleaved History Vol 35 pp37-50. woodland on suitable sites, with trees numbering in the Stace, CA (2010) New flora of the British Isles. Third edition. Cambridge tens at most, and planted as a component of shrub areas, University Press. mainly in clearings and on woodland edges Rodwell, JS (Ed) (1991) British Plant Communities. Volume 1: Woodlands and • Controlled grazing of cattle should be encouraged in Scrub. Cambridge University Press. native woodland and wood pasture with mature wild Ruhsam, M, Jessop, W, Cornille, A, Renny, J and Worrell, R (2018) Crop- apple trees; new wood pasture should include planting of to-wild introgression in the European wild apple Malus sylvestris in Northern wild apple Britain. Forestry. 92 (1), 85-96. • Seed collections from known pure populations of wild Scottish Government (2017) Pollinator Strategy for Scotland 2017–2027. apple should be encouraged Available from: https://www.nature.scot/sites/default/files/2018-04/ • Research should be undertaken to determine conditions Pollinator%20Strategy%20for%20Scotland%202017-2027.pdf [Accessed that favour natural regeneration and to clarify the Schnitzler A, Arnold C, Cornille A, Bachmann O, & Schnitzler C (2014) Wild invertebrate, plant and fungal associates of wild apple, its European apple (Malus sylvestris (L.) Mill.) Population dynamics: Insight from genetics and ecology in the Rhine Valley. Priorities for a future conservation regeneration biology and its adaptive genetic variation. programme. PLoS ONE, 9, 1–11. Stephan BR, Wagner, I & Kleinschmit, J 2003. EUFORGEN Technical Acknowledgements: Guidelines for genetic conservation and use for wild apple and pear (Malus sylvestris This project was funded by the Patsy Wood Trust, Scottish and Pyrus pyraster). International Plant Genetic Resources Inst. Rome. Forestry Trust, Sustainable Forestry SCIO and Forest Vera, FWM (2000) Grazing Ecology and Forest History. CABI publishing. Enterprise Scotland. The report is dedicated to the memory of Worrell, R (1995) European Aspen (Populus Tremula L.): a review with Patsy Wood. We are grateful to the many people who located particular reference to Scotland 1: Distribution, ecology and genetic variation. trees for us or sent in leaves for sampling. Our work was also Forestry 68, 94-105. supported by the Falkland Stewardship Trust, who handled the Worrell, R, Ruhsam, M, Renny, J, Jessop, W, & Findlay, G (2019) Scotland’s finances, and Forestry Commission Scotland who sent us data Native Wild Apple – Malus sylvestris: Genetic issues and conservation. Scottish from the Native Woodland Survey of Scotland. The genetic Forestry 72 (2) 33-41. analysis at the Royal Botanic Garden Edinburgh was supported by the Scottish Government’s Rural and Environment Science and Analytical Services Division. I especially thank Neil Mackenzie and Alison Averis for their thorough refereeing of the manuscript. ERERICIC BBOYDOYD FORESTFORESTRRYY LLTTDD References Åsen, PA (no date) Vileppler in Norge (Wild apples in Norway). Available at: http://consideratecandidum.com/villepler/historikk_og_bruk.html [accessed 10/02/2020]. Buttenschøn, RM & Buttenschøn, J (1998) Population dynamics of Malus sylvestris stands in grazed and ungrazed, semi-natural grasslands and fragmented woodlands in Mols Bjerge, Denmark. Ann. Bot. Fennici 35: 233–246. Decourtye, L (1977) “Evereste”, un nouveau pommier d’ornement. L’Hortic Fr., 54, 309–311. Dickson, C.A. and Dickson, JH (2000) Plants and People in Ancient Scotland. NPI Media Group. Egremont Crab Fair (no date) Available at: http://www.egremontcrabfair.com/ history/index.html (accessed on 10/02/2020) Forestry Establishment and Forestry Commission (2017) Forestry Facts and Figures. https://www. Maintenance Contractors forestresearch.gov.uk/tools-and-resources/statistics/forestry-statistics Forestry Commission Scotland (2014) Scotland’s Native Woodlands: Results Contractors based throughout from the Native Woodland Survey of Scotland. Forestry Commission Scotland. most of Scotland Edinburgh. Hayes, C (2016) Orchards of the Forth Valley: A Programme for Revival. The Mob: Forth Naturalist and Historian Vol 39, 97-112. 07788 923820 Kik, C, Korpelainen, H, Vögel, R, Asdal, Å, Eliáš, P, Draper, D & Magos Tel: 01369 703316 Brehm, J (2011) Malus sylvestris. The IUCN Red List of Threatened Species: e.T172170A6841688. Available from: https://dx.doi.org/10.2305/IUCN. E-mail: eboeboydyd10099775@[email protected] UK.2011-1. RLTS.T172170A6841688.en. [accessed 10/02/2020] Kennedy, CEJ and Southwood, TRE (1984) The Number of Species of wwwww.b.boydforestroydforestry.co.uk Insects Associated with British Trees: A Re-Analysis. Journal of Animal Ecology

Volume 74 No. 1 Spring 2020 | 29