Epipactis youngiana A · Figure 1 - Epipactis youngiana (© Bob Gibbons/Plantlife International)....

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Epipactis youngiana A.J.Richards & A.F. Porter Young’s Helleborine ORCHIDACEAE SYN.: Epipactis helleborine var. youngiana (Richard & Porter) Kreutz (2004) Status Schedule 8 Wildlife & Countryside Act UK BAP Priority Species since 1995 (1981) Lead partner: Plantlife International Status in Europe: British endemic 9 10km squares post 1987 UK Biodiversity Action Plan There is one current target following the 2001 Targets Review: T1- Maintain a minimum of seven populations across its range (revised). Progress on targets as reported in the UKBAP 2002 reporting round can be viewed at: http://www.ukbap.org.uk/2002OnlineReport/mainframe.htm The full Action Plan for Epipactis youngiana can be viewed online at: http://www.ukbap.org.uk/UKPlans.aspx?ID=285 Contents 1 Morphology, Identification, Taxonomy and Genetics ........................................... 2 1.1 Morphology and Identification ................................................................... 2 1.2 Taxonomic Considerations ........................................................................ 3 1.3 Genetic Implications ................................................................................ 5 2 Distribution and Current Status ....................................................................... 5 2.1 United Kingdom...................................................................................... 5 2.1.1 England .......................................................................................... 6 2.1.2 Scotland ......................................................................................... 7 2.1.3 Wales ............................................................................................. 8 3 Ecology and Life Cycle ..................................................................................... 9 4 Habitat Requirements ................................................................................... 10 4.1 History and Importance of Human Activity................................................ 10 4.2 Communities and Vegetation .................................................................. 10 4.3 Future Prospects ................................................................................... 11 5 Management Implications .............................................................................. 11 6 Threats/Factors Leading to Loss or Decline, or Limiting Recovery ........................ 12 Work on Epipactis youngiana is supported by: 1

Transcript of Epipactis youngiana A · Figure 1 - Epipactis youngiana (© Bob Gibbons/Plantlife International)....

Page 1: Epipactis youngiana A · Figure 1 - Epipactis youngiana (© Bob Gibbons/Plantlife International). 1.2 TAXONOMIC CONSIDERATIONS Epipactis youngiana is part of the E. helleborine (Broad-leaved

A.

Young’s HORCHIDSYN.: Epi

Status Schedule (1981) Status in E

UK BiodThere is o T1- Mainta Progress ohttp://ww The full Achttp://ww

Content1 Morp

1.1 M1.2 T1.3 G

2 Distri2.1 U

2.1.1 2.1.2 2.1.3

3 Ecolog4 Habita

4.1 H4.2 C4.3 F

5 Manag6 Threat

Epipactis youngiana J.Richards & A.F. Porter

elleborine ACEAE pactis helleborine var. youngiana (Richard & Porter) Kreutz (2004)

8 Wildlife & Countryside Act UK BAP Priority Species since 1995 Lead partner: Plantlife International

urope: British endemic 9 10km squares post 1987

iversity Action Plan ne current target following the 2001 Targets Review:

in a minimum of seven populations across its range (revised).

n targets as reported in the UKBAP 2002 reporting round can be viewed at: w.ukbap.org.uk/2002OnlineReport/mainframe.htm

tion Plan for Epipactis youngiana can be viewed online at: w.ukbap.org.uk/UKPlans.aspx?ID=285

s hology, Identification, Taxonomy and Genetics ...........................................2

orphology and Identification...................................................................2 axonomic Considerations........................................................................3 enetic Implications................................................................................5

bution and Current Status .......................................................................5 nited Kingdom......................................................................................5

England ..........................................................................................6 Scotland .........................................................................................7 Wales .............................................................................................8

y and Life Cycle.....................................................................................9 t Requirements ...................................................................................10 istory and Importance of Human Activity................................................10 ommunities and Vegetation ..................................................................10 uture Prospects...................................................................................11 ement Implications ..............................................................................11 s/Factors Leading to Loss or Decline, or Limiting Recovery ........................12

Work on Epipactis youngiana is supported by:

1

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7 Current Conservation Measures ......................................................................127.1 In Situ Measures...................................................................................13 7.2 Ex Situ Measures ..................................................................................13 7.3 Research Data......................................................................................13 7.4 Monitoring ...........................................................................................13 7.5 Recommendations for Future Work..........................................................13

8 References ..................................................................................................14 9 Acknowledgements .......................................................................................15 10 Contacts .................................................................................................15 11 Links ......................................................................................................15

1 Morphology, Identification, Taxonomy and Genetics

1.1 MORPHOLOGY AND IDENTIFICATION Epipactis youngiana is a perennial orchidaceous herb, 15-40 cm high when in flower. Epipactis is an easily identified genus, and the following characters help to distinguish this variety. Stems usually solitary, sometimes 2-3 together, typically less robust than E. helleborine (Broad-leaved Helleborine), shortly and sparsely pubescent above or glabrescent, green. Leaves more or less two-ranked, pale green, somewhat flaccid, not markedly ribbed, with an undulate margin and subacute apex. Basal leaf at least 1.2 x longer than broad, flaccid, not cucullate. Lower bracts exceeding flowers, usually less than 6 mm wide. Flowers slightly declined, rather box-shaped, 8-11 mm diameter, whitish to pale green, or in good light a clear rose-pink; hypochile purple, epichile whitish, somewhat recurved with two inconspicuous bosses. Stigma tricornute, the rostellum acute and more or less equaling the subsessile anther, producing a usually evanescent viscidium which has usually disappeared by the 4th open flower from the top; pollinia crumbling and disintegrating by day two, not often dispersed intact. Ovary pear-shaped, rather shiny, slightly ribbed, with sparse, rather prickly, stout-based hairs, becoming glabrescent. Seeds 0.9-1.3 mm in length.

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Figure 1 - Epipactis youngiana (© Bob Gibbons/Plantlife International).

1.2 TAXONOMIC CONSIDERATIONS Epipactis youngiana is part of the E. helleborine (Broad-leaved Helleborine) complex. The latter is a common and widespread outcrossing species throughout temperate Eurasia and introduced in North America (Squirrell et al 2001). E. helleborine is exceptionally variable, often varying strikingly within populations for color, robustness, indumentum and leaf shape. At least some of this variation may be attributable to developmental or environmental conditions, so that young or stressed plants often have yellowish, rather narrow, two-ranked leaves and pale flowers. This may equally affect E. youngiana plants. Nevertheless, this complex has produced many distinctive local variants, some of which have been recognized taxonomically. Some of these are outcrossing but geographically or ecologically distinct, such as E. tremolsii from the western Mediterranean, E. helleborine subsp. neerlandica from sand-dunes on Channel and North Sea coasts, E. distans from the Alps, E. latina from Italy, and E. condensata from the eastern Mediterranean (there are others; there is an account in full in Delforge 1995). E. helleborine subsp. neerlandica and its autogamous derivative E. renzii are morphologically distinct but cannot readily be separated from E. helleborine by molecular characters and are now often treated as varieties (Ehler & Pedersen 2000, Pedersen & Ehler 2000). Other taxa within the E. helleborine complex have evolved partial or complete autogamy through early pollinium disintegration while still in the anther and the absence of a viscidium on the rostellum (or its early disappearance); this syndrome is often

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accompanied by small pendant flowers which sometimes do not open fully (Richards 1982). In western and central Europe, E. dunensis, E. leptochila and E. muelleri are distinctive, invariable, local autogamous taxa of this type and there are several more in the Balkans. Harris (1993) and Harris & Abbott (1997) follow Richards & Swan (1976) in calling this inland taxon ‘E. leptochila’. Squirrell et al (2002) clearly establish that the correct name for these populations is E. dunensis. Epipactis youngiana was first detected in Northumberland by the late Tony Porter in 1975, and published in 1982 (Richards & Porter 1982). At that time it was known from four sites in that county. In the early 1980’s, very similar plants were discovered from central Scotland, associated with birch woodland colonizing bings (industrial spoil). The distinctive features of the variety are listed above, but it should be noted that it usually occurs with E. helleborine (as do other rare Epipactis species) and care must be taken not to confuse it with depauperate individuals of the latter. Table 1 summarizes the most reliable characters for E. youngiana, as compared to E. helleborine. Table 1 - Comparison of E. youngiana characteristics with E. helleborine.

EPIPACTIS YOUNGIANA EPIPACTIS HELLEBORINE Basal leaf >1.2 as long as broad Basal leaf <1.1, usually <1.0 as long as

broad Basal leaf flat, flaccid, unribbed, silky to touch, margins undulate

Basal leaf cucullate, stiff, ribbed, coarse to touch, margins not undulate

If flowers colored, clear pink Flowers with a ‘dirty’ suffusion of pink Rostellum nearly equaling anther Rostellum not more than half-way on anther Ovary shiny with sparse ‘prickly’ hairs Ovary matt, sparsely pubescent to glabrous

It was originally considered that E. youngiana was probably hybridogenous in origin, with E. helleborine as one parent and an autogamous species as the other, and that an autogamous mating system had ‘fixed’ distinctive hybridogenous features. It was noted that certain habitats in which it occurred were man-made and kept free from competition by physical and chemical features, inviting the suggestion that human activity had unwittingly brought the parental species into contact, and that the hybridogenous taxon which had thus arisen favored certain man-made conditions. Subsequent work, notably by Harris & Abbott (1997) and Hollingsworth et al (in press) has clarified certain points. E. youngiana is very closely related to E. helleborine, and English populations at least of the former variety probably originate from local populations of E. helleborine. E. phyllanthes (Green-flowered Helleborine) is definitely not a parent to E. youngiana, although E. dunensis (Dune Helleborine), which occurs in both the main districts where E. youngiana is found, could have been (as a male parent). Although observations suggest that E. youngiana must receive a substantial proportion of self pollen, it possesses the population structure of an outcrosser, so that it is certainly not always autogamous. There has been much discussion of the status of E. youngiana and those who have studied the plant in the wild maintain that in the context of this critical group it is a distinctive variety with consistent characteristics and ecology which deserves recognition (e.g. Stace 1991, Allan & Woods 1993, Delforge 1995, Preston et al 2002). P. Hollingsworth (pers. comm.) in particular points out:

‘Something is going on’ which deserves recognition.

The status of E. youngiana is very similar to that of E. subsp. neerlandica and E. var. renzii: there are some morphological differences to indicate divergence, which with the absence of genetic differences have led Ehler & Pedersen (2000) and Pedersen & Ehler (2000) to describe renzii as a variety of E. helleborine. The lack of taxon specific alleles and shared polymorphism means E. youngiana is also

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more appropriately described as a variety. Lang (2004) describes E. youngiana as E. helleborine var. youngiana.

Disproving a hybrid origin for E. youngiana from E. helleborine x dunensis is difficult, given that E. dunensis is itself a recent derivative of E. he

lleborine as

ne but can be distinguished morphologically, it is best described as a variety bbott 1997, Bateman in press, Hollingsworth et al in press).

zym:

g and both studies identified high levels of genetic variation in

it and E. helleborine may indicate

that E. youngiana is a type of population from which future autogenous lines

Hollingsworth et al (in press) hypothesize that E. youngiana may be undergoing “detour”, which is illustrated by the presence of distinctive , reflecting possible recent hybridization between E. helleborine,

nt Status

.1 UNITED KINGDOM There is no suggestion that any plants resembling Epipactis youngiana have yet been found outside the UK, so this variety must be considered endemic. It has been recorded in a total of at least 11 localities in nine 10 km squares and in 2003 it survived in at least eight of these in seven squares.

well. It would therefore be difficult to find conclusive evidence that E. youngiana is correctly interpreted as a new hybrid from two closely related taxa.

The taxonomic status of E. youngiana has, and continues to be, queried by several influential authors including Harris & Abbott (1997) and Bateman (in press). The merging consensus seems to be that because E. youngiana is genetically indistinct from e

E. helleboriof E. helleborine (Harris & A

1.3 GENETIC IMPLICATIONS There have been two major studies, Harris & Abbott (1997) the data and interpretations in which are reported in more detail in Harris (1993); and Hollingsworth et al (in press), which reached similar but more detailed conclusions. Harris’ work was based on iso es alone, while Hollingsworth et al also examined cpDNA. The essential findings are

Epipactis youngiana is closely related to E. helleborine and cannot be distinguished from it by any reliable marker.

E. phyllanthes is not a parent of E. youngiana, but E. dunensis could be.

E. youngiana has a level of genetic diversity and heterozygosity typical of an outbreeder, but unlike the Northumberland populations of E. phyllanthes and E. dunensis, these do not differ from E. helleborine, or from panmixis.

This has caused Harris & Abbott to doubt the hybridogenous origin of E. youngiana.

Hollingsworth et al (in press) found no clear evidence that E. youngiana plants are self pollinatinplants studied.

Given that within the Epipactis group there is evidence of self pollination to produce “new” varieties, the morphological distinctiveness of E. youngiana alongside the genetic indistinctiveness between

could originate.

an evolutionarygenetic markersE. youngiana, E. dunensis and E. phyllanthes.

2 Distribution and Curre

2

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2.1.1 ENGLAND

Figure 2 – Distribution of Epipactis youngiana in England (Derived from Miniscale by

rdnance Survey. Reproduced from Ordnance Survey digital map data. © Crown

he site on Magnesian limestone. However, it has not been relocated since (Abbott, pers.

comm. January 2005). Locations of E. youngiana in England are listed in Table 2.

OCopyright 2003. All rights reserved). Attention was first drawn to Epipactis youngiana by the late Tony Porter in the mid 1970s. He was actively surveying Northumberland Epipactis sites after the discovery of what is now called E. dunensis (Dune Helleborine) on inland River Tyne sites and lead mines, and of E. phyllanthes (Green-flowered Helleborine) associated with a lead mine (Richards & Swan 1976). In a mature oak plantation at Dissington (the type locality) he noted that amongst a large population of typical E. helleborine (Broad-leaved Helleborine), about half the individuals were very distinct, with the suite of characters noted above. Porter later discovered an almost pure population of what was undoubtedly the same plant some 35 km to the west, associated with a metal mine near Settlingstones. Three other small oakwood populations were discovered subsequently in Northumberland (VC 67) and one associated with mining trackways in Durham (VC 66), leading to the formal publication of the new species (Richards & Porter 1982). Two of these additional sites definitely survive (2003), but after the felling of the Dissington site in 1984 and the subsequent disappearance of the orchids there, the Settlingstones population remains the most important English site of three known to survive. In 1988, Dr Frank Horsman discovered a small population in mid west Yorkshire (SE44). Tis

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Table 2 – Site details for locations of Epipactis youngiana in England. (VC=Vice-County) VC SITE / LOCALITY GRID

REF. DATE NO. RECORDER SOURCE

66 Derwent Walk NZ1557 1997> 2 Richards, Davison, Kergon

sight

67 Killingworth NZ2770 1995 2 Davison, Kergon, Richards

sight, photos

67 Gosforth Park NZ2570 1996> 5-20 Davison, Kergon, Richards

Sight, photos

67 Dissington NZ1372 1975-84

40-100

Porter, Richards, Swan

type locality, hb, photos

67 Hexham, Oakerlands crossroad

NY9462 1976-82

2-6 Richards, Furness sight, photo

67 Stonecroft Mine Settlingstones

NY8568 1975> 50-150

Porter, Richards Sight, photo, hb

2.1.2 SCOTLAND In about 1987, attention was drawn to certain Scottish bing populations of Epipactis by Professor Jim Dickson (e.g. Dickson et al 2000), notably populations at Bardykes and Bothwell Bings. These did not appear to differ morphologically from Northumbrian E. youngiana, although they did not perhaps have exactly the same overall appearance (‘gestalt’), and rather more intermediate types with E. helleborine seemed to occur. Nevertheless, they were distinctive and have now been referred to as E. youngiana for nearly two decades. There are five definite localities in four 10 km squares, and as far as it is known all of these survive. Another two localities in new separate squares have not been confirmed, but from the habitat and general area seem likely. Locations of E. youngiana in Scotland are listed in Table 3. Table 3 – Site details for locations of Epipactis youngiana in Scotland. (VC=Vice-County) VC SITE / LOCALITY GRID

REF. DATE NO. RECORDER SOURCE

84 Philipstoun bing NT0576 1996 5 K. Watson SNH database 2002

83 Gorebridge bing NT3362 1996 18 K. Watson SNH database 2002

77 Bothwell Priory bing

NS6859 1996 54 K. Watson SNH database 2002

77 Bardykes bing NS6758 1996 43 K. Watson SNH database 2002

86 Almond bing NS9676 1995 13 K. Watson SNH database 2002

77 Newton (West) bing

NS6661 2002 ID not confirmed

J. McIntosh K. Watson pers. comm.

77 Ross bing, Ferniegair, near Motherwell

NS7455 11/07/2000

ID not confirmed

Gill Smart K. Watson pers. comm.

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Figure 3 – Distribution of Epipactis youngiana in Scotland (Derived from Miniscale by Ordnance Survey. Reproduced from Ordnance Survey digital map data. © Crown Copyright 2003. All rights reserved).

2.1.3 WALES A strange population of Epipactis occurs on sand dunes at Kenfig in South Wales, which has some features of E. youngiana. An unpublished review of this taxon (Richards and Lewis, February 2003) concluded that this plant represented another local derivative from E. helleborine (Broad-leaved Helleborine), possibly associated with metal pollution from Margam steelworks, but could not be conclusively identified as E. youngiana. This site (Margam) also has populations of E. cambrensis C. Thomas, which is almost certainly a variety of E. phyllanthes (Green-flowered Helleborine), and a dune form of E. helleborine which is very similar to E. neerlandica. Both of these differ from the E. youngiana-like plant there.

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3 Ecology and Life Cycle Epipactis youngiana is a perennial, but is inconstant and relatively short-lived. About 40-80% of the flowers set capsules. Abundant self-pollination occurs and reproductive limitation is certainly under maternal control; proportional set is greater on big plants and in warm wet autumns when plants stay green longer. Capsules are estimated to release >1000 seeds each from late October onwards by wind. Many seeds remain in capsules but many plants must release >10,000 seeds annually. About 70-80% seeds contain embryos. Observations by Davison and Kergon (pers. comm.) of germination on buried glass slides shows that seeds germinate in May, produce an internal swelling first which develops into a 1-2 mm diameter subterranean protocorm by that autumn (year 1). A single leaf is produced in years 2-3, an aerial shoot in years 4-5 and plants probably first flower years 5 to 6. First year flowering plants typically produce 4-6 flowers; mature plants up to 30 flowers. Typically, 50-80% of above-ground plants in a population are immature and non-flowering, and many flowering shoots are subsequently eaten by deer. Leaves and shoots appear above ground in late May, buds are set late June and flowering occurs July 20th-August 20th in most years. Plants disappear by early December. Some mature plants definitely fail to appear some years, to reappear in subsequent years, but the proportion is not known. Plants with multiple shoots are uncommon, less so than in E. helleborine (Broad-leaved Helleborine) and especially E. purpurata (Violet Helleborine). Mature flowering plants rarely appear for more than two subsequent seasons, and it is likely that a major reproductive effort kills many, so that ongoing reproduction by seed is essential. Long-term study of the Stonecroft population shows that even locations are inconstant between years. Within a metapopulation, areas of local abundance rarely last for more than five years, suggesting a high rate of population turn over as subsites become locally favorable and then unfavorable again. Plants can reappear in ‘old’ locations where they have not been seen for 20 years, but are almost certainly not the same plants. This pattern of population cycling is also observed in E. phyllanthes (Green-flowered Helleborine) and E. dunensis (Dune Helleborine). Flowers are visited by wasps (Vespa-spp.) that drink nectar from the hypochile. There is little evidence that pollinia are removed intact and then placed on the viscidium, as happens in E. helleborine, but the flowers are relatively showy and rewarding and the stigma is accessible compared with autogamous species such as E. phyllanthes. Loose pollen massulae can be observed on wasps and the genetic results indicate that considerable cross-pollination ensues, although some fail-safe autogamy obviously occurs. Even in allogamous species, much pollination of intact pollinia is probably geitonogamous. As far as is known all Epipactis are self-compatible, and all except the most obligately autogamous probably have mixed-mating systems. In common with all Epipactis, E. youngiana is a poor competitor. It emerges very late in the season and is easily smothered, and the small seedlings appear to need almost open sites in which to succeed. It seems likely that various factors which have kept otherwise mesic sites free from vegetation, for example pollution by metals such as zinc and lead, pollutants in coal waste, limited trampling and path maintenance, and weeding of young trees in commercial woodland, have all promoted its survival. It appears to resent both full sunlight and deep shade, and both waterlogging and desiccation. Soils tend to be well-drained, deep and often rather stony brown earths, typically with a pH of 5.8-6.5 and a balanced and plentiful supply of essential elements. In common with other orchids, E. youngiana adopts heterobasidioid fungi for its own nutrition. Watling (pers. comm.) identified the presence of a rather specialized group of jelly fungus, which includes species of Sebacina, Tulasnella or Ceratobasidium, associated with E. youngiana in Glasgow in 1992. These symbionts are not all

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necessarily associated with trees, although Ceratobasidium has very recently been proven to be ectomycorrhizal (Oberwinkler, via Watling pers. comm.). While sebacinoid fungi are not dependent on trees, they could occur in the habitats where E. youngiana grows.

4 Habitat Requirements

4.1 HISTORY AND IMPORTANCE OF HUMAN ACTIVITY Epipactis youngiana has only been recognized for 30 years. However, none of the segregates of the E. helleborine (Broad-leaved Helleborine) complex were differentiated until the 1920s, so all were doubtless overlooked in earlier years. The close association of E. youngiana with mine spoil and coal-bing habitats has led to suggestions that its evolutionary origin must be recent and favoured by these new, anthropogenic habitats. However, the only factors which link the English populations (apart from the obvious ones of lowland woodlands of north-east England) appear to be the open nature of the ground vegetation, and good drainage. In general, the populations of the rarer Epipactis in north-east England have been linked to heavy-metal polluted ground. Whether these are ex-mine sites such as Stonecroft, or polluted river shingles like most of the E. dunensis (Dune Helleborine) sites, it is likely that they have only been in their present condition for 150 years or less. Equally, carefully managed, even-aged oak stands such as are (were) typical of the Dinnington and Gosforth Park sites are also a relatively recent, post nineteenth century, habitat. It can also be argued that if E. youngiana is a hybridogenous species (and there is no firm evidence either way), then one of the putative parents, E. dunensis, which is virtually restricted to metal-rich habitats inland, has been brought into contact with the widespread E. helleborine by the relatively recent activities of man, which are likely to have provided a suitable habitat away from its coastal sites. The evidence does indeed tend to suggest that E. youngiana may have arisen within the last 150 years or so. This would help to explain the lack of distinctive genetic markers, and as Hollingsworth (pers. comm.) has pointed out, E. youngiana may represent, as do several endemic Euphrasia (Eyebrights), ‘active evolution (with) multiple origins of allogamy-autogamy transitions’ and ‘(a) product of hybridization between allogamous and autogamous taxa’.

4.2 COMMUNITIES AND VEGETATION In England, Epipactis youngiana typically grows in lowland habitats, to 150 m above sea level, in mature deciduous woodland, often with oaks Quercus spp.. It usually occurs in glades or openings, with herb-rich vegetation and sparse grass, and at least 50% open ground, typically with Rumex conglomeratus (Clustered Dock), R. acetosa (Common Sorrel), Silene dioica (Red Campion), Epilobium montanum (Broad-leaved Willowherb), Arrhenatherum elatius (False Oat-grass), Poa nemoralis (Wood Meadow-grass), Senecio jacobaea (Common Ragwort), Filipendula ulmaria (Meadowsweet), Fragaria vesca (Wild Strawberry) and Solidago virgaurea (Goldenrod). Pyrola minor (Common Wintergreen) is a characteristic rare associate. These are essentially modifications of NVC W10 Quercus-Pteridium woodland on skeletal soils, with Dryopteris filix-mas (Male-fern) characteristically replacing Pteridium aquilinum (Bracken). Urtica (Nettle spp.) and Rubus (Bramble spp.) are absent except when very depauperate. Sites are often by paths or tracksides, or on steeply sloping ground, streambanks and wooded mineral tip slopes. Soils are invariably brown earths but usually well-drained, with a low clay fraction, and often with a highly skeletal base from tailings, shingles or ballast which has been infiltrated by leaf-mould.

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In Scotland, E. youngiana and E. dunensis (Dune Helleborine) are most commonly associated with Betula (Birch), although Salix caprea (Goat Willow) is frequently present. The canopy is usually intermediate in age and ranges from spindly, dense scrub to more widely spaced, older, often multi-stemmed individuals. Typical plant associates include Fragaria vesca (Wild Strawberry), Viola riviniana (Common Dog-violet), Pyrola minor (Common Wintergreen), Hieracium spp. (Hawkweeds), Circaea lutetiana (Enchanter’s-nightshade), Epilobium montanum (Broad-leaved Willowherb), Geranium robertianum (Herb-Robert), Geum urbanum (Wood Avens), Urtica dioica (Common Nettle), sparse grasses and frequent pleurocarp mosses (e.g. Eurhynchium sp.). Substrate is usually freely draining, coarse shale, base-rich with high pH, at least initially.

4.3 FUTURE PROSPECTS Sites rapidly become unsuitable in the absence of management, through increased shade and ground cover, although other nearby sites may become suitable as colonization proceeds on spoil, or through forest management. It is easy to prescribe management which favors Epipactis youngiana (see sections 5 and 6), but there are many difficult philosophical problems with respect to its conservation, and some basic principles need to be addressed first:

This plant may have essentially evolved to an industrial age which has now passed. Has the time for this variety also passed? If we are to keep it, is it as a relic of that industrial age, or does it have another niche?

Should man-made post-in / preserved

e this variety?

ld

If so, should these only be to sites in the immediate vicinity of existing ones?

nization is to be encouraged, should this be done by habitat modification?

ve answers are given to at least ome of these questions.

what appears to be an actively evolving

ures and kills young seedlings. All brambles, gorse and

dustrial sites such as bings be maintained on behalf of this variety?

Should such sites be managed to preserv

This is an essentially colonizing variety with short-lived habitat occupancy; shouit be encouraged to colonies new sites?

If so, should introductions to new sites of seed and plants be made artificially?

How does one define a suitable site for introduction?

If autonomous colo

As the mining of coal and heavy metals is virtually extinct, it seems likely that E. youngiana may disappear of its own accord unless positis The taxonomic status of E. youngiana, as variant, also leads to important questions on adopting conservation strategies capable of conserving endemic variety in Britain (see section 7.0).

5 Management Implications See section 4.3. Short-term management can often be successfully achieved by the local removal of branches and trees to create glades and admit some light. This should be done in winter (after the New Year). No bonfires should be lit anywhere near sites (one Epipactis dunensis location was destroyed this way). Opening canopies will encourage ground vegetation, which should be strimmed, preferably before mid May, or very carefully after that, taking care not to disturb seedlings. Scriefing is probably not successful as it spoils soil struct

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other scrub must be removed, preferably manually, in late winter or early spring. Brambles, nettles and Rosebay Willowherb (Chamerion angustifolium) outcompete E. youngiana. This has been illustrated at Dissington where, even before it was felled, the site was essentially lost when woodland husbandry ceased after the gamekeeper retired nd the brambles encroached.

no circumstances should any fertilization of the ground take place, and nitrogen fixers (Fabaceae, Pea family, and Alnus, Alders) should be discouraged. Tension zones, e.g.

e

Threats/Factors Leading to Loss or Decline, or Limiting Recovery

n from any tall herb, but especially brambles, gorse, etc, or any coarse

Loss of new colonizing habitat on, for example, woodland seres on mining spoil.

Excessive grazing of flowering shoots by deer and rabbits.

Hollingsworth et al (in press) suggest that, given the lack of a distinct genetic identity stent identification in the field, E. youngiana is not yet a

demic species. This requires a re-design of species-

a In

between woodland paths and encroaching vegetation, are very important and should bully strimming. caref maintained, e.g. by

Numbers of roe deer and rabbits should be kept down.

6

Excessive shading.

Competitiograss.

Lack of woodland management.

Forest clearance.

Drought.

Fertilization of soil, increased levels of nitrogen.

Flooding.

7 Current Conservation Measures Epipactis youngiana clearly illustrates the problems that taxonomic complexity bequeaths to conservation and in particular to the UK BAP process, which is currently rigorously focused on a typological, species-based approach. Brochmann et al (2003), in a review of the status of British species, reflected that all British endemic putative species are in taxonomically difficult groups. A solution is therefore to ensure that the BAP process is able to conserve taxonomically difficult groups of plants that seem to be actively evolving as endemics.

and the difficulties of consicohesive, distinct and reproductively isolated species. However, the British E. youngiana populations represent complex populations that have not yet achieved separate evolutionary trajectories from E. helleborine, although they may be in the process of doing so. Given that evidence indicates that E. youngiana is in the process of diverging in evolutionary terms from E. helleborine, strategies to conserve this process, instead of a fraction of the species diversity currently present are urgently required, not just for his variant but for all British ent

focused conservation programs towards a process that is capable of flexibility in conserving groups of species, some of which are rarer than others.

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Page 13: Epipactis youngiana A · Figure 1 - Epipactis youngiana (© Bob Gibbons/Plantlife International). 1.2 TAXONOMIC CONSIDERATIONS Epipactis youngiana is part of the E. helleborine (Broad-leaved

7.1 IN SITU MEASURES There are currently few or no conservation measures in operation in England. Stonecroft Mine is a Site of Special Scientific Interest (SSSI), and Gosforth Park, a Local Nature Reserve (LNR), is owned and managed by the Natural History Society of Northumbria. Derwent Walks is a Countryside Park managed by Durham County Council. The Gosforth

vant LBAPs, its genetic status, nd the possibility that it is still evolving into a separate species, means that its statutory

c n obust. Several sites, including Bardykes bing, Bothwell Priory bing has been used as a source of bing

Park population is part of the Newcastle LBAP (Local Biodiversity Action Plan), but little active management is occurring. Although Epipactis youngiana is a Red Data Book variety, is listed on Schedule 8 of the Wildlife & Countryside Act (1981) and appears on all releaprote tio is not very rbing and Almond bing, are SSSIs. Bardykes material for hardcore, but the Epipactis woods seem to have been left alone. A monitoring strategy has been set up at Almond bing in conjunction with the LBAP and local ranger service. Experimental ground vegetation thinning took place early in 2005. The UKBAP (UK Biodiversity Action Plan) Steering Group for this species has agreed in principle that E. youngiana should be treated and conserved as one of the Epipactis roup. This provides a mechanism for conservation action for the group as a whole and

safeguards the evolutionary process that may be in train. g

7.2 EX SITU MEASURES The Hardy Orchid Society is involved in a project to investigate the possibility of seed germination and the raising of young plants in artificial media. This was previously ttempted unsuccessfully by Hardy Orchid Society member Steve Davison, who

nevertheless had some success in raising young plants between buried glass microscope Epipactis has a reputation for being difficult with respect to the raising

ESEARCH ATA

Essentially no research projects are active at present; see sections 1.3 and 7.2. s of the Epipactis leptochila (Narrow-lipped

further publications

ONITORING is now in place with Falkirk LBAP and ranger service (see section

mes to the

a

slides. In general, of seeds in artificial media (Rasmussen 1995), and strongly dislikes inorganic nitrogen, although limited success with E. helleborine has been achieved with Curtis’ medium.

.3 R D 7

However, genetic research on the statuHelleborine) complex has been published (Squirrell et al 2002) andare expected.

7.4 MA monitoring scheme7.1). Otherwise, monitoring occurs without formal notification except sometiNorthumberland Wildlife Trust in England by A. J. Richards. Surveys have been carried out at Bardykes bing and Almond bing (Watson 1993, 1994, 1995).

lity of habitats should be considered.

7.5 RECOMMENDATIONS FOR FUTURE WORK

Support local monitoring schemes to track population levels.

Where the variety is listed on the Local Biodiversity Action Plan, some management to ensure continued availabi

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Page 14: Epipactis youngiana A · Figure 1 - Epipactis youngiana (© Bob Gibbons/Plantlife International). 1.2 TAXONOMIC CONSIDERATIONS Epipactis youngiana is part of the E. helleborine (Broad-leaved

8 References . & Woods, P. (1993). Wild Orchids of Scotland. H.M.S.O., Edinburgh. Allan, B

Dickson

of Epipactis

Harris,

Preston

Richard

Richard

urnal of Botany, 88: 1409-1418. rrell, J., Hollingsworth, P.M., Bateman, R.M., Tebbitt, M.C. & Hollingsworth, M.L.

lexity and breeding system transitions: conservation ptochila complex (Orchidaceae). Molecular Ecology, 11:

Brochmann, C., Gabrielsen, T.M., Nordal, I., Landvik, J.Y. & Elven, R. (2003). Glacial survival or tabula rasa? The history of North Atlantic biota revisited. Taxon, 52: 417-450.

Bateman, R. M. (in press). How many orchid species are currently native to the British Isles? In: Bailey, J. (ed.) Contributions to taxonomic research on the British and European Flora. BSBI, London.

Delforge, P. (1995). Orchids of Britain and Europe. HarperCollins, London. , J.H., Macpherson, P. & Watson, K.J. (2000). The changing flora of Glasgow. Edinburgh University Press, Edinburgh.

ies Ehler, B.K. & Pederson, H.A.E. (2000). Genetic variation in three spec(Orchidaceae): geographic scale and evolutionary inferences. Biological Journal of the Linnaean Society, 69: 411-430. S.A. (1993). The status of Epipactis Zinn. (Orchidaceae) on Bings in mid Strathclyde. Report SS/F2B/459A) to Scottish Natural Heritage.

Harris, S.A. & Abbott, R.J. (1997). Isozyme analysis of the reported origin of a new hybrid orchid species Epipactis youngiana (Young’s helleborine) in the British Isles. Heredity, 79: 402-407.

Hollingsworth, P.M., Squirrell, J., Hollingsworth, M.L., Richards, A.J. & Bateman, R. (in press) Taxonomic complexity, conservation and recurrent origins of self-pollination in Epipactis (Orchidaceae). In: Bailey, J. (ed.) Contributions to taxonomic research on the British and European Flora. BSBI, London.

Lang, D.C. (2004). Britain’s Orchids. WildGuides Ltd., Old Basing, Hants. Pedersen, H.A.E. & Ehler, B.K. (2000). Local evolution of autogamy in Epipactis

helleborine ssp. neerlandica (Orchidaceae). Plant Systematics and Evolution, 223: 173-183. , C.D., Pearman, D.A. & Dines, T.D. (2002). New Atlas of the British Flora. Oxford University Press, Oxford.

Rasmussen, H.N. (1995). Terrestrial orchids, from seed to mycotrophic plant. Cambridge University Press, Cambridge. s, A.J. (1982). The influence of minor structural changes in the flower on the breeding systems and speciation in Epipactis Zinn. (Orchidaceae). In: Armstrong, J.A., Powell, J.M. & Richards, A.J. (eds.) Pollination and Evolution. Royal Botanic Garden, Sydney. pp. 47-53. s, A.J. & Porter, A.F. (1982). On the identity of a Northumberland Epipactis. Watsonia, 14: 121-128.

Richards, A.J. & Swan, G.A. (1976). Epipactis leptochila (Godfery) Godfery and E. phyllanthes G E Sm. occurring in south Northumberland on lead and zinc soils. Watsonia, 11: 1-5.

Squirrell, J., Hollingsworth, P.M., Bateman, R.M., Dickson, J.H., Light M.H.S., MacConaill, M. & Tebbitt, M.C. (2001). Partitioning and diversity of nuclear and organelle markers in native and introduced populations of Epipactis helleborine (Orchidaceae). American Jo

Squi(2002). Taxonomic compgenetics of the Epipactis le1957-1964.

Stace, C.D. (1991). New Flora of the British Isles. Cambridge University Press, Cambridge.

Watson, K. (1993). A survey of Young’s helleborine (Epipactis youngiana) at Bardykes bing, Glasgow. Back from the Brink report, Plantlife.

Watson, K. (1994). Almond bing: survey of helleborine orchids. Report to Falkirk Council. Watson, K. (1995). Almond bing: monitoring of helleborine orchid populations. Report to

Falkirk Council.

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Page 15: Epipactis youngiana A · Figure 1 - Epipactis youngiana (© Bob Gibbons/Plantlife International). 1.2 TAXONOMIC CONSIDERATIONS Epipactis youngiana is part of the E. helleborine (Broad-leaved

9 Acknowledgements pecial thanks are due to the two teams S of molecular geneticists that have taken a

rve particular mention in this regard, the ter for commissioning this dossier. Quite early in the history of this controversial

s were made (for instance by Martin Wigginton) that it should be ed Data Book, and be given Schedule 8 status in the Wildlife and

se have been largely ignored.

P to thank Mrs. P Abbott, Prof Richard Bateman, Prof Roy Watling and K additional information. We would also like to thank the Epipactis y g group for comments on the text, and Plantlife International volunteer D at for his editorial work.

10 Contacts he lead partner in the conservation of this Schedule 8 and national

78509

special interest in this critical group; led by Peter Hollingsworth at Royal Botanic Garden Edinburgh, and by Stephen Harris, now the School of Plant Sciences, Oxford. Scottish Natural Heritage and Plantlife International have a long-standing interest in Scottish populations of E. youngiana which goes back at least 14 years, and Jim Dickson, Chris ydes, Keith Watson and Deborah Long deseS

latspecies, decisionincluded in the RCountryside 1981 Act. These decisions have led to the current interest in a variety which might otherwi lantlife would like eith Watson for oungiana steerinr James Pe

Plantlife Scotland is tBAP species. Dr Deborah Long Plantlife Scotland Balallan House Allan Park Stirling FK8 2QG

Or contact us by e-mail: [email protected]

Tel: 01786 4 Other key addresses are: Professor A J Richards, High Trees, South Park, Hexham NE46 1BT 01434 602403. [email protected] Peter Hollingsworth, Royal Botanic Garden, Edinburgh EH3 5LR. [email protected] of Falkirk Area LBAP and Gosforth Park - Cramlington Wildlife Corridor LBAP can be found on www.ukbap.org.uk.

11 Links The full Action Plan for Epipactis youngianahttp://www.ukbap.org.uk/UKPlans.aspx?ID=285

can be viewed at

ARKive species web page for Epipactis youngiana: http://www.arkive.org/species/ARK/plants_and_algae/Epipactis_youngiana/

ISBN: 1 904749-21-3

Original draft by Prof. A. J. Richards Edited by Plantlife Scotland/Plantlife International

First draft dated May 2004 Last revised 13 February 2007

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