Waxcap grassland is short-sward, nutrient-poor grassland that supports a rich assemblage of larger

fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...

, particularly waxcaps (species of ''

Hygrocybe ''Hygrocybe'' is a genus of agarics (gilled fungi) in the family Hygrophoraceae. Called waxcaps in English (sometimes waxy caps in North America), basidiocarps (fruit bodies) are often brightly coloured and have dry to waxy caps, white spores, an ...

'' and related genera), characteristic of such habitats. Waxcap grasslands occur principally in Europe, where they are declining as a result of agricultural practices. The fungal species are consequently of conservation concern and efforts have been made in the United Kingdom and elsewhere to protect both the grasslands and their characteristic fungi. Over 20 species of European waxcap grassland fungi are assessed as globally "vulnerable" or "endangered" on the IUCN Red List of Threatened Species.

Background

The association of waxcaps with unimproved (nutrient-poor) grasslands was first noted in 1949 in the Netherlands, but current interest was stimulated by a series of papers published by Dutch

mycologist Mycology is the branch of biology concerned with the study of fungus, fungi, including their genetics, genetic and biochemistry, biochemical properties, their Taxonomy (biology), taxonomy and ethnomycology, their use to humans, including as a so ...

Eef Arnolds in the 1980s. Arnolds not only confirmed the association of waxcaps with unimproved grasslands, but also noted the rapid decline in such habitats in the Netherlands. Similar studies were subsequently undertaken elsewhere in Europe, initially in Denmark and the United Kingdom.

Definition and description

Waxcap grasslands are characterized by being unimproved (unfertilized and nutrient-poor), short-sward (through grazing or mowing), moss-rich, and long-established (not recently sown). They occur in both upland and lowland areas and may be on acidic, neutral, or calcareous soil. They support a wide range of characteristic larger fungi, but may not be equally species-rich in plants.

Characteristic species

Larger fungi characteristic of waxcap grasslands include

agarics An agaric () is a type of fungus fruiting body characterized by the presence of a pileus (cap) that is clearly differentiated from the stipe (stalk), with lamellae (gills) on the underside of the pileus. In the UK, agarics are called "mushrooms ...

(gilled mushrooms) belonging to the genera '' Cuphophyllus'', '' Gliophorus'', ''

Gloioxanthomyces ''Gloioxanthomyces'' is a genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species ...

'', ''

Neohygrocybe ''Neohygrocybe'' is a genus of agaric fungi in the family Hygrophoraceae. ''Neohygrocybe'' species belong to a group known as waxcaps in English, sometimes also waxy caps in North America or waxgills in New Zealand. In Europe, ''Neohygrocybe'' sp ...

'', and ''

Porpolomopsis ''Porpolomopsis'' is a genus of fungi in the family Hygrophoraceae. It was circumscribed in 2008 by Andreas Bresinsky to contain '' P. calyptriformis''. Bresinsky separated it from the genus ''Hygrocybe'' based on its color and the absence ...

'' (waxcaps), '' Entoloma'' (pinkgills), '' Dermoloma'', '' Pseudotricholoma'', ''

Camarophyllopsis ''Camarophyllopsis'' is a genus of agarics (gilled fungi) in the family Clavariaceae. Basidiocarps (fruit bodies) are dull-coloured and have dry caps, rather distant, decurrent lamellae, white spores, and smooth, ringless stems. In Europe specie ...

'', and ''

Hodophilus ''Hodophilus'' is a genus of agarics (gilled fungi) in the family Clavariaceae. Basidiocarps (fruit bodies) are dull-coloured and have dry caps, rather distant, decurrent lamellae, white spores, and smooth, ringless stems. In Europe species are ...

'';

clavarioid fungi The clavarioid fungi are a group of fungi in the ''Basidiomycota'' typically having erect, simple or branched basidiocarps (fruit bodies) that are formed on the ground, on decaying vegetation, or on dead wood. They are colloquially called club fun ...

(club and coral fungi) belonging to the genera '' Clavaria'', ''

Clavulinopsis ''Clavulinopsis'' is a genus of coral fungi in the family Clavariaceae. The genus, first described scientifically by Casper van Overeem in 1923, has a widespread distribution. The name means "having the appearance of ''Clavulina''". Species , ...

'', and '' Ramariopsis''; and earthtongues belonging to the genera ''

Geoglossum ''Geoglossum'' is a genus of fungi in the family Geoglossaceae. They are commonly called earth tongues. The type species is ''Geoglossum glabrum''. ''Geoglossum'' species are distinguished from the related genus ''Trichoglossum'' by the lack of ...

'', ''

Glutinoglossum ''Glutinoglossum'' is a genus of six species of earth-tongue fungi in the family Geoglossaceae. The widespread type species, '' G. glutinosum'', is commonly known as the "glutinous earth tongue". '' G. heptaseptatum'' is known only fro ...

'', '' Microglossum'', and ''

Trichoglossum ''Trichoglossum'' is a genus of fungi in the family Geoglossaceae. They are commonly called hairy earth tongues. The type species is ''Trichoglossum hirsutum''. Members of the genus ''Trichoglossum'' have tiny hairs known as setae on the spor ...

''. File:Hygrocybe intermedia.jpg, File:2009-06-30 Entoloma mougeotii.jpg, File:Dermoloma cuneifolium 73323.jpg, File:Clavaria rosea - Goździeniec różowy-1.jpg, File:Velvety Earth Tongues.jpg,

The "CHEG" or "CHEGD" assessment system

In 1995, Rald proposed a simple count of the number of waxcap species present at a given site in order to assess its value as a waxcap grassland. He suggested that the presence of 17 or more species meant the site was of national importance, 9-16 species of regional importance, 4-8 species of local importance, and 3 or fewer of no importance. This system was modified by Rotheroe and others to include all the characteristic macrofungi and not just waxcaps. Known as the "CHEG" system, this is widely used in survey work today. The acronym "CHEG" stands for the main groups of relevant fungi: C - the

clavarioid The clavarioid fungi are a group of fungi in the ''Basidiomycota'' typically having erect, simple or branched basidiocarps (fruit bodies) that are formed on the ground, on decaying vegetation, or on dead wood. They are colloquially called club fun ...

species (club and coral fungi); H - species of ''Hygrocybe'' and related genera (waxcaps); E - ''Entoloma'' species (pinkgills); and G - the geoglossoid fungi (earthtongues). More recently the modified term "CHEGD" has been used to include species of '' Dermoloma'', '' Pseudotricholoma'', ''

'', and ''

'' which also inhabit these grasslands.https://cieem.net/grasslands-and-chegd-fungi/

Conservation

The past 75 years have witnessed a loss of more than 90% of unimproved grasslands in western Europe, mainly due to agricultural intensification through ploughing and reseeding, manuring, and the application of fertilizers and other chemicals. The maintenance of a short sward by grazing or mowing (and removal or cuttings) has also been shown to be important for fruiting of waxcaps, with haycutting in July, followed by aftermath grazing/mowing to 3 cm having found to be optimal. In 1988, Arnolds estimated that only some 200 ha of unimproved waxcap grasslands remained in the Netherlands. As a result, both the unimproved, nutrient-poor grasslands and the larger fungi typical of such grasslands are of conservation concern, with 23 waxcap-grassland species currently assessed as globally "vulnerable" or "endangered" on the IUCN Red List of Threatened Species.

Globally threatened waxcap-grassland fungi

Conservation in the United Kingdom

In the United Kingdom, survey work has shown that surviving waxcap grasslands are more extensive than in many other European countries, thanks mainly to large areas of unimproved upland sheep pastures and also to many unimproved lawns (especially in churchyards and country houses) and amenity grasslands. Nonetheless, five species characteristic of waxcap grasslands - '' Entoloma bloxamii'', ''

Porpolomopsis calyptriformis ''Porpolomopsis calyptriformis'' is a species of agaric (gilled mushroom) in the family Hygrophoraceae. It has been given the recommended English name of Pink Waxcap. The species has a European distribution, occurring mainly in agriculturally un ...

'', ''

Hygrocybe spadicea ''Hygrocybe spadicea'' is a species of agaric (gilled mushroom) in the family Hygrophoraceae. It has been given the recommended English name of Date Waxcap. The species has a European distribution, occurring mainly in agriculturally unimproved gr ...

'', '' Microglossum atropurpureum'', and ''

Microglossum olivaceum ''Microglossum olivaceum'' (vernacular name: olive earthtongue) is a species of fungus belonging to the family Geoglossaceae. Synonym: * ''Geoglossum olivaceum'' Pers. (= basionym In the scientific name of organisms, basionym or basyonym means ...

'' - were formerly the subject of national Biodiversity Action Plans and waxcap grasslands as a specific habitat were the subject of several local Biodiversity Action Plans. Four species - ''E. bloxamii'', ''H. spadicea'', ''M. atropurpureum'', and ''M. olivaceum'' - are currently listed as "Priority Species" under Section 41 of the Natural Environment and Rural Communities Act (in England) and Section 7 of the Environment (Wales) Act (in Wales). JNCC ( Joint Nature Conservation Committee) has issued "Guidelines for the Selection of Biological SSSIs" ( Sites of Special Scientific Interest) that offer a measure of protection to waxcap grasslands. Using the "CHEGD" system, sites should be considered for notification as SSSIs if the total number of waxcap species reaches or exceeds 19. Thresholds are also stipulated for other CHEGD species. As a result, several waxcap-grassland sites, such as the banks of Llanishen and Lisvane Reservoirs in

Cardiff Cardiff (; cy, Caerdydd ) is the capital and largest city of Wales. It forms a principal area, officially known as the City and County of Cardiff ( cy, Dinas a Sir Caerdydd, links=no), and the city is the eleventh-largest in the United Kingd ...

Down Farm in

Dorset Dorset ( ; archaically: Dorsetshire , ) is a county in South West England on the English Channel coast. The ceremonial county comprises the unitary authority areas of Bournemouth, Christchurch and Poole and Dorset (unitary authority), Dors ...

The Leasowes The Leasowes is a 57-hectare (around 141 acre) estate in Halesowen, historically in the county of Shropshire, England, comprising house and gardens. The parkland is now listed Grade I on English Heritage's Register of Parks and Gardens and the h ...

Shropshire Shropshire (; alternatively Salop; abbreviated in print only as Shrops; demonym Salopian ) is a landlocked historic county in the West Midlands region of England. It is bordered by Wales to the west and the English counties of Cheshire to th ...

, and the lawns of Roecliffe Manor in

Leicestershire Leicestershire ( ; postal abbreviation Leics.) is a ceremonial and non-metropolitan county in the East Midlands, England. The county borders Nottinghamshire to the north, Lincolnshire to the north-east, Rutland to the east, Northamptonshire t ...

, have been designated Sites of Special Scientific Interest, Waxcap grassland surveys have been undertaken by the British Mycological Society, the

National Trust The National Trust, formally the National Trust for Places of Historic Interest or Natural Beauty, is a charity and membership organisation for heritage conservation in England, Wales and Northern Ireland. In Scotland, there is a separate and ...

, Plantlife, and the various national conservation bodies. A leaflet on managing waxcap grasslands in Britain and Ireland has been published by Plantlife and the Fungus Conservation Forum.

References

{{reflist, 2, refs= {{cite web , url=https://www.iucnredlist.org/search?taxonomies=100001&searchType=species , title=Fungi: The IUCN Red List of Threatened Species , access-date=2022-06-01{{clarify, reason=this url returns a vague search results page that may or may not support the claims of this article;, date=June 2022 {{cite journal , author=Schweers ACS. , year=1949 , title=Hygrophorusweide, een associatie , journal=Fungus , volume=19 , pages=17–18 {{cite journal , author=Arnolds E. , year=1988 , title=The changing macromycete flora in the Netherlands , journal=Transactions of the British Mycological Society , volume=90 , issue=3 , pages=391–406 , doi=10.1016/s0007-1536(88)80148-7 {{cite journal , author=Arnolds E. , year=1989 , title=The influence of increased fertilization on the macrofungi of a sheep meadow , journal=Opera Botanica , volume=100 , pages=7–21 {{cite journal , vauthors=Griffith GW, Easton GL, Jones AW , year=2002 , title=Ecology and diversity of waxcap (''Hygrocybe'' spp) fungi , journal=Botanical Journal of Scotland , volume=54 , pages=7–22 , url=http://www.aber.ac.uk/waxcap/downloads/ecology-diversity.doc , format=DOC , doi=10.1080/03746600208685025, s2cid=84829857 {{cite journal, author1=Griffith, G.W. , author2=Gamarra, J.G.P. , author3=Holden, E.M. , author4=Mitchel, D. , author5=Graham, A. , author6=Evans, D.A. , author7=Evans, S.E. , author8=Aron, C. , author9=Noordeloos, M.E. , author10=Kirk, P.M. , author11=Smith, S.L.N. , author12=Woods, R.G. , author13=Hale, A.D. , author14=Easton, G.L. , author15=Ratkowsky, D.A. , author16=Stevens, D.P. , author17=Halbwachs, H. , date=30 August 2014 , title=The international conservation importance of Welsh 'waxcap' grasslands, journal=Mycosphere, volume=4 , issue=5, pages=969–984, doi=10.5943/mycosphere/4/5/10 {{cite journal , author=Rald E. , year=1995 , title=Vokshatte som indikatorarter for mykologisk verifulde overdrevslokaliteter , journal=Svampe , volume=11 , pages=57–65 {{cite journal , vauthors=Rotheroe M, Newton A, Evans S, Feehan J , year=1996 , title=Waxcap-grassland survey , journal=The Mycologist , volume=10 , pages=23–25 , doi=10.1016/s0269-915x(96)80046-2 UK BAP priority fungi species http://jncc.defra.gov.uk/page-5165 {{cite web, title=Section 41 Species - Priority Actions Needed (B2020-008), url=http://publications.naturalengland.org.uk/publication/4958719460769792, website=publications.naturalengland.org.uk, publisher=Natural England, access-date=17 February 2017, date=October 2013 Section 7 Priority Species https://www.biodiversitywales.org.uk/environment-wales-act Provisional atlas of waxcap fungi ''Hygrocybe'' in Cardiff https://www.aber.ac.uk/waxcap/downloads/Grantham-CardiffWaxcapAtlas2009.pdf waxcap grassland action plan https://neenp.org.uk/natural-environment/durham-priority-habitats/waxcap-grassland-action-plan/ Guidelines for the Selection of Biological SSSIs: Chapter 14: Non-lichenised Fungi https://data.jncc.gov.uk/data/d1fcb171-8086-4f5b-ade5-a34c5edc78c5/SSSI-Guidelines-14-Non-lichenisedfungi-2018a.pdf Llanishen and Lisvane Reservoirs: Ecology http://www.friendsofcardiffreservoirs.org/ecology/ Down Farm SSSI https://consult.defra.gov.uk/natural-england/down-farm/supporting_documents/Down_Farm_Supporting_Info_final%20notified%207%20Feb%202019.pdf The Leasowes https://www.dudley.gov.uk/news/rare-mushrooms-to-get-special-protection-at-halesowen-beauty-spot/ Roecliffe Manor SSSI https://pap.charnwood.gov.uk/AnitePublicDocs/00135127.pdf Grassland Fungi Project https://www.nationaltrust.org.uk/hardcastle-crags/features/grassland-fungi-project Waxcap Watch//https://www.plantlife.org.uk/uk/discover-wild-plants-nature/habitats/grassland/waxcaps-fungi/waxcapp-survey {{cite book , author=Anon , title=Grassland gems: managing lawns and pastures for fungi (leaflet) , publisher=Plantlife , pages=4 , url=https://www.plantlife.org.uk/application/files/3414/8232/2984/Grassland_gems-_managing_lawns_and_pastures_for_fungi.pdf {{cite journal , author=Evans S , year=2003 , title=Waxcap-grasslands - an assessment of English sites , journal=English Nature Research Reports , volume=555 , pages=1–49 , url=http://publications.naturalengland.org.uk/publication/131003#:~:text=Waxcap%2Dgrasslands%20%E2%80%93%20an%20assessment%20of%20English%20sites%20(ENRR555),England%20on%2012%20May%202004%20.&text=Waxcap%2Dgrassland%20fungi%20are%20of,throughout%20the%20UK%20and%20Europe. Scottish Fungi - grassland fungi project http://sites.google.com/site/scottishfungi/conservation/grassland-fungi

External links

WaxCap Website
'' University of Wales, Aberystwyth''. Grasslands Fungus ecology Habitats