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Scotland’s rare tooth fungi:

Front cover image: B u x b a u m i a v i r i d i s capsules on an alder log Stewart Taylor, RSPB

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an introduction to their identification, ecology and management

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Left: Tooth fungi habitat on Deeside © Liz Holden Front cover: main image, Hydnellum peckii © Mark Gurney, RSPB Inset top: Phellodon tomentosus © Liz Holden Inset below: Bankera fuligineoalba © Mark Gurney, RSPB

Plantlife is the UK’s leading charity working to protect wild plants and their habitats. The charity has 10,500 members and owns 23 nature reserves. In 2008, Plantlife is 'Lead Partner' for 77 species under the UK Government's Biodiversity Action Plan. Conservation of these species is delivered through the charity’s Back from the Brink species recovery programme, which is jointly funded by Countryside Council for Wales, Natural England, Scottish Natural Heritage, charitable trusts, companies and individuals. It involves its members as volunteers (Flora Guardians) in delivering many aspects of this work. Plantlife’s head office is in Salisbury, Wiltshire, and the charity has national offices in Wales and Scotland. Plantlife Scotland Balallan House Allan Park Stirling FK8 2QG Tel. 01786 478509

www.plantlife.org.uk [email protected]

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Hydnellum requiring further taxonomic investigation to define exact species © Mark Gurney, RSPB

Scotland’s rare tooth fungi: an introduction to their identification, ecology and management

relationship wherein both partners gain nutrients.

What are tooth fungi? Not quite animals and certainly not plants, these fascinating organisms are members of one of the largest kingdoms on the planet, the fungi, essential to the health of all ecological systems and without which around 90% of our higher plants and trees would not survive. The parts of a fungus that we see above ground are the spore producing structures, (the ‘fruit bodies’) of a much larger organism that is mostly hidden from sight and composed of a branching network of filamentous cells.This underground network, the ‘mycelium’, enables tooth fungi to forage for nutrients and to link up with the roots of living trees in a symbiotic ‘ectomycorrhizal’

‘tooth fungi’ are a diverse group of fungi that utilise tooth like structures to produce their spores. Only four of the many genera (groups of species) that share this character are of conservation concern in Scotland and considered here: Bankera, Hydnellum, Phellodon and Sarcodon. Two other stipitate (stalked) genera could cause confusion as they have similar macroscopic structures, but these are widespread: the Wood Hedgehogs (Hydnum) and Earpick Fungus (Auriscalpium). Other tooth fungi are not stipitate and where they are of conservation concern, have not yet been found fruiting in Scotland. 1

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Hydnellum aurantiacum. © Liz Holden

Distribution in Scotland

Tooth fungi characteristics

The core concentrations of Scottish tooth fungi appear to be in the Caledonian pine forests of the Central and Eastern Highlands, with scattered records from elsewhere. Certain species, including Orange Tooth (Hydnellum aurantiacum) and Greenfoot Tooth (Sarcodon glaucopus) are rarely recorded even within the core areas.

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Rather than gills or tubes, the undersides of the caps have little tooth-like structures to support the developing spores. The four genera of conservation concern in Scotland have stalks, known as ‘stipes’ and hence known as ‘stipitate’. These fungi produce relatively long-lived fruit bodies (several weeks in some cases) between early August and October. Up to ten species, representing all four genera, have been found fruiting in close proximity to each other in ‘hot spot’ clusters. It appears that these genera do not readily colonise new sites. Once established at a site their main mode of dispersal is through vegetative growth with the mycelium moving from tree root to tree root.

Wood hedgehogs (Hydnum species) have teeth and stipes but are widespread in Scotland and not of conservation concern © Mark Gurney, RSPB

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Tooth fungi habitat ●

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Tooth fungi are woodland organisms that access nutrients in a partnership with living trees. In Scotland the host trees are thought to be mainly pine, birch and oak. Their fruit bodies appear in soil and are not on dead wood. There is evidence to suggest that the larger the area of woodland, the higher the number of species (Newton et al, 2002). As a group they seem to share a preference for fruiting in poor sandy soils, often on banks, tracksides, old quarries or borrow pits where there is a poorly developed humus layer and very little vascular plant cover.

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Soils supporting tooth fungi are usually well drained although humidity levels are often relatively high. Humidity might be maintained for, example, by proximity to water or overhanging branches. The fruiting success of these fungi appears to be linked to low nitrogen levels in the soil as well as climatic variables. Fruiting is not limited to old growth forests and fruit bodies do sometimes occur with young trees or in plantation woodland on suitable ground.There is however, usually a link to old growth forest either through proximity to that habitat or through scattered old growth forest trees.

Ideal tooth fungi habitat © Stewart Taylor

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Bankera fuligineoalba © Liz Holden

The tooth fungi genera Bankera species The flesh can be quite easily broken and generally the caps do not fuse together.The fruit bodies have white to buff teeth and white spores.The cap is fleshy and often has debris sticking to it. It is white at first becoming tinged brown with age. Dried material usually smells strongly of fenugreek or curry powder. Sarcodon species The flesh can be quite easily broken and generally the caps do not fuse together. Fruit bodies have greyish teeth and brown spores. The cap is fleshy, a brown colour and covered in distinct scales.There is no strong spicy smell when dried. 4

Sarcodon squamosus © Liz Holden

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Sarcodon glaucopus © Liz Holden Bankera fuligineoalba © Mark Gurney

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Hydnellum species The flesh is tough and the caps often fuse together incorporating surrounding vegetation and debris; there are multiple stipes beneath. Caps have brownish teeth and brown spores. The cap can be thick or thin and whilst starting very pale, becomes some shade of brown, with blue, orange or pink tints in some species.The caps of thin-fleshed species are often concentrically zoned.The cut flesh itself is often zoned and in some species contains bright blue or orange colours. In damp weather fresh fruit bodies of some species produce blood red droplets (guttules).There is no strong spicy smell when dried.

Hydnellum peckii © Liz Holden



Hydnellum requiring further taxonomic investigation to define exact species © Liz Holden

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Hydnellum caeruleum © Liz Holden

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Hydnellum peckii © David Genney



Hydnellum ferrugineum © Mark Gurney RSPB

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Phellodon tomentosus © Liz Holden

Phellodon melaleucus © Mark Gurney RSPB

Phellodon species The flesh is tough and the caps often fuse together incorporating surrounding vegetation and debris; there are multiple stipes beneath. Caps have white teeth and white spores. Caps are often concentrically zoned and colours vary from brown to blue black. Dried material usually smells strongly of fenugreek or curry powder.

Look alike non-target species

Phellodon niger © Liz Holden

The Earpick Fungus (Auriscalpium) – is a small, dark brown fungus that grows throughout the year on old pinecones.The relatively long, thin, hairy stipe is often set to one side of the cap. The Wood Hedgehogs (Hydnum) – have teeth more or less the same colour as the uniformly coloured cap ie pale cinnamon to buff or terracotta.The spores are whitish.The cap is fleshy and easily broken. Tiger’s Eye (Coltricia perennis) – is common in similar habitats, has a stipe, tough flesh and the top of the cap has concentric brown zones like several of the tooth fungi. Look carefully to find pores rather than teeth below the cap.

Why are tooth fungi of conservation concern? Reports produced during the 1980s and 1990s in parts of western and central Europe indicated that fruiting of these fungi was in significant decline and in 1999, concern about their status in the UK led to a grouped Biodiversity Action Plan (BAP) being established.The revised 2007 BAP list includes one additional species bringing the total number of species in the plan to 15, of which 13 occur in Scotland. 8

Perceived threats ● Habitat loss at both macro and micro scales e.g. clear felling, alteration of site conditions by invasive plants such as bracken or rhododendron or loss of suitable, sandy soil microhabitats ● Eutrophication of soils through airborne pollution or agricultural run-off ● The application of fungicides or substances containing nitrogen and / or phosphate ● Liming alters the soil pH dramatically.Work in Sweden and Germany has found that liming was detrimental to established ectomycorrhizal species (Taylor & Finlay, 2003) ● Compaction or disturbance of soil by trampling or machine ● Lack of awareness of the habitat requirements of tooth fungi amongst land managers ● Lack of understanding of the ecology and taxonomy of the group What you can do if you find tooth fungi ● These are species of conservation concern and collection should be kept to the minimum necessary to establish identification. ● Tooth fungi, some of which are listed on the UK BAP (2007) list and in the preliminary Red List of Threatened Fungi (Evans, 2007), are not currently listed on Schedule 8 of the Wildlife and Countryside Act (1981).Although a licence from the government is not required to pick fruit bodies, permission of the landowner must be sought. ● These fungi are not poisonous and working with the fruit bodies will not require action beyond normal health and safety procedures. Most species are either too

A survey plot for tooth fungi © Liz Holden

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tough or bitter to be edible although some are collected for dying craft materials. Check whether their presence is already recorded at the site using your own land management records, by checking the Fungal Records Database of Britain and Ireland through the British Mycological Society website or by contacting Plantlife Scotland (contact details on back page). If they are already well recorded then there is no need to disturb them further; if not then advice can be given on how to proceed. Refer to the management guidelines below if interventions are proposed for the site. Consider undertaking a simple survey, as there is very little base line data available from which to assess the impact of management interventions. Surveys should take place over at least 5 years, as some tooth fungi do not fruit regularly. Monthly visits from early August to October each year are also recommended as some species fruit early and others later. Temporarily mark each fruiting site to avoid confusion in successive monthly visits. GPS readings, good field notes and photographs can help to accurately map your tooth fungi and inform future site management. Contact Plantlife if assistance is required. 9

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© Liz Holden

Management guidelines If you are in woodland with suitable host trees and poor sandy soils it is possible that you either already have tooth fungi, or if they do not appear to be present, that you could encourage their presence through appropriate management. Current research (Van der Linde, 2008) suggests that the mycelium of tooth fungi can be successfully transferred to new sites on the inoculated roots of seedling trees. This work is at an early stage but may be a future consideration. General guidelines in areas where tooth fungi are known to fruit: ● The exact ecological requirements of these fungi are not yet fully understood, so great care should be taken when changing the management of sites known to support tooth fungi. ● Ensure contractors and land managers are aware of the presence of tooth fungi. 10

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Get to know species specialists and local fungus recording groups who may be able to help inform site management and provide regular and relevant advice including how to monitor any management interventions. In the vicinity of known fruit bodies, maintain a wide age structure of trees to increase the likelihood of continuity of host trees.The exact stocking densities to enable the inoculation of neighbouring young trees with fungal mycelium have not been determined in this habitat but will relate to the extent of the host tree’s own root system. Opening up the canopy beyond 1.5 times the radius of the canopy is not recommended close to the known area colonised by tooth fungi. It can be difficult to ascertain which trees are actually hosting the target mycelium; the further development of molecular tools will assist with this process in the future. Meanwhile the maintenance of a sustainable buffer zone (a minimum of 50m is suggested) of potential host trees around known fruiting sites is recommended to ensure host continuity. Extended rotations, continuous cover forestry and small coupe felling are more likely to sustain tooth fungi populations than clear fell.When carefully applied these forms of management can benefit tooth fungi by increasing the longevity of the woodland site, maintaining continuity of hosts (selective thinning can be used to reduce the likelihood of felling host trees) and maintaining appropriate environmental conditions such as humidity. Increase woodland size where possible by extending existing woodlands and incorporating existing woodland fragments. Further research is required to determine which areas of the forest are likely to be most suitable for the mycelia of these fungi.

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It is not yet possible to offer specific management recommendations to favour the development of these species in the wider forest. Where natural regeneration is not possible then restocking should be with native host species. Minimise heavy disturbance and compaction from vehicle access or foot traffic. Ploughing and scarification should be avoided in areas where tooth fungi are known to be fruiting or where there is a plan to encourage the presence of tooth fungi. Even light disturbance should be avoided during the fruiting season. Maintain areas around fruiting tooth fungi free from the development of humus and dense vascular plant cover. Verge cutting with a bar can be appropriate as long as the dead plant material does not accumulate to enrich the soil.The effect of herbicides on tooth fungi is unknown. At sites threatened by, for example, Rhododendron ponticum or Snowberry (Symphoricarpus albus) take measures to control, contain or eradicate invasive nonnative plant species using Forestry Commission guidelines on suitable techniques. Bracken, particularly in riverside locations, can be difficult to access by machinery but could be mown or managed by volunteers. Avoid the use of fungicides, lime or the application or accidental run off of any nitrogen or phosphorus rich substance in or adjacent to target areas. Monitor the effects of management interventions. Remember that it is possible to manage for a range of conservation interests by creating a mosaic of habitats.

Maintenance of paths, roads and car parks in areas known to support tooth fungi: ● Aim for minimal interference with the established soil profile e.g. avoid excavation to create foundations or deep layers of new surfacing material to level the paths. ● Endeavour to retain existing path widths and edges where tooth fungi are known to fruit ● New construction materials should be locally sourced if possible and mineral in origin.Wood chips would not be appropriate. ● A few carefully sited borrow pits, away from the locations of the rarest tooth fungi, could provide additional suitable habitat for these species to fruit in. For this purpose and where appropriate, it is recommended that topsoil is not replaced. Excess topsoil should be placed several metres into the woodland and away from tooth fungus fruiting sites. ● Path drainage should be established with care and in consultation with species specialists.The maintenance of ditches and banks can be beneficial in some circumstances. ● Material derived from any clearance of trees should be piled up several metres into the surrounding woodland if staying on site. ● Minimise disturbance and compaction from machinery involved in the project, or the storage of materials. ● It is important to maintain the host trees but it may also be important to have a clear margin between the path edge and the trees, if that does not compromise humidity levels.

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Recommended texts and references UKBAP species lists and action plans are available at www.ukbap.org.uk The preliminary assessment for the Red Data List of Threatened British Fungi (Evans 2007) can be found through the British Mycological Society’s website (www.britmycolsoc.org.uk) D.N. Pegler, P.J. Roberts and B.M. Spooner. Royal Botanic Gardens, Kew 1997 British Chanterelles and tooth fungi J. Breitenbach and F. Kränzlin.Verlag Mykologia, 1986 Fungi of Switzerland Vol 2 Non gilled fungi

Contacts for advice and further information For help with reporting or identifying tooth fungi contact, Plantlife Scotland Balallan House Allan Park Stirling FK8 2QG Tel. 01786 478509 [email protected] www.plantlife.org.uk For help with contacting mycological contractors and volunteer groups: contact Plantlife Scotland

Newton, A.C., Holden, E., Davy, L.M.,Ward, S.D., Fleming, L.V. & Watling, R. (2002). Status and distribution of stipitate hydnoid fungi in Scottish coniferous forests. Biological Conservation (107) 181-192

For help with contacting fungal recording groups contact the British Mycological Society at www.britmycolsoc.org.uk or Plantlife Scotland

Parfitt, D., Ainsworth, A.M., Simpson, D., Rogers,H.J. & Boddy, L. (2007). Molecular and morphological discrimination of stipitate hydnoids in the genera Hydnellum and Phellodon. Mycological Research, 111, 761-777.

For information on dealing with invasive nonnative plant species see the Forestry Commission Practice Guides for example ‘Managing and controlling invasive rhododendron’ by Colin Edwards (2006) at www.forestry.gov.uk

Taylor, A.F.S. & Finlay, R.D. (2003). Effects of liming and ash application on below ground ectomycorrhizal community structure in tow Norway Spruce forests. Water, Air and Soil Pollution: Focus 3: 63-76, Netherlands

Guidance provided in this leaflet is integrated into HaRPPS, the Forestry Commission's webbased information and decision support system, providing quick and easy access to information about a range of Priority and protected woodland species and habitat management: http://www.forestresearch.gov.uk/fr/INFD758CCA

Van der Linde, S. (2008).‘Ecology and conservation of stipitate hydnoid fungi associated with Scots Pine’. PhD thesis, University of Aberdeen 12

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Sarcodon squamosus © Mark Gurney RSPB

This leaflet was written for Plantlife Scotland by Liz Holden, Field Mycologist

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Phellodon niger © Mark Gurney, RSPB