Collins Fungi Guide: The most complete field guide to the mushrooms and toadstools of Britain & Ireland

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INTRODUCTION

What are Fungi?

Although practically everyone realises that the mushrooms they see in the supermarket are fungi, relatively few shoppers know how they fit into the overall scheme of living things, and how they are related to the fruit and vegetables nearby on the same shelf or to the animal carcasses on the butchery and fish counters. The truth is that mushrooms are not related to either, although there has been a tradition of teaching what little is taught about fungi in plant science rather than animal science courses – largely because, like plants, they often grow from the soil and do not move. It is now recognised, however, that living things can no longer be placed in two simple Kingdoms of plants and animals. Over the past fifty years, others have been added, and for some time Fungi have been referred to as the Fifth Kingdom, although some of the latest scientific thinking recognises six or even more to accommodate the wide range of microscopic organisms now known to exist. Nonetheless plants and animals are still the major groups with which most people are familiar.

Fungi are not related to plants or animals because they differ from them in many important ways. Whereas the basic structural elements of plants and animals are cells which, en masse, form tissues, fungi are built up of uniquely different microscopic tubular thread-like bodies with multiple nuclei called hyphae; and collectively hyphae form, not tissue, but mycelium (although the word ‘cell’ is used widely in relation to small fungal structures that look like cells). The reproduction of fungi also differs greatly from plants in that they produce, not seeds, but microscopic structures called spores.

Fungi are also uniquely distinct in relation to their mode of nutrition. Plants photosynthesise, a process in which solar energy is absorbed by green chlorophyll and used to bring about the formation of nutrient substances from the raw materials of atmospheric carbon dioxide and water. No fungus can photosynthesise, and even the few species that sometimes display a green fruit body colouration or green spores do not contain chlorophyll. Fungi also have a mode of nutrition different from animals in that while animals eat, digest and then absorb the digested matter internally, fungi secrete enzymes externally from their hyphae into the environment where organic matter is broken down and then absorb the resulting chemicals from there. Like animals therefore, fungi are dependent for their nutrient source on other organisms, either living or dead. And this dictates where they grow: typically on soil, using humus and plant remains for nutrition; directly on wood or other plant matter; or sometimes parasitically on still living plants.

Fungal nutrition is a complex subject but one aspect of it is nonetheless of particular importance in relation to their occurrence in the field. Anyone who has ever collected toadstools will have noted that many, perhaps most, occur in particular types of woodland, beneath particular types of tree or consistently in company with certain types of plant. This is not mere chance, nor the result of two species requiring a similar ecological niche. It is because of an intimate association called a mycorrhiza, which means that under certain circumstances the one cannot exist, or can do so only inadequately, without the other. A mycorrhiza is a symbiotic association between fungal hyphae and the roots of higher plants, and also, to some extent, of some Pteridophytes (ferns and their allies) and the rhizoids of Bryophytes (mosses and liverworts). The fungal mycelium forms an outer sheath around the fine rootlets and this can be seen if the rootlets are examined closely with a lens. Penetration of fungal hyphae into the root is limited and occurs only between the cells of the cortex. Precisely how the mycorrhizal mycelium assists its host plant, and vice versa, is still imperfectly understood, but it seems that the fungus obtains much of its necessary supply of carbon from the roots and, in return, acts as an intermediary in the uptake of nutrients such as nitrogen, phosphate and potash from the soil. The mycorrhizal fungus seems better able to achieve this uptake, especially from poor soils, than does the plant acting on its own.

The naming and classification of fungi

Living organisms are given names and are classified into groups; fungi are no exception. The scientific names used for organisms today are based on a binomial or two-name system based on Latin and other languages and using Latin grammar derived from one devised by the Swedish biologist Linnaeus (Carl von Linné, 1707–1778). The binomial of each organism does more than signify its uniqueness, however, as it also attempts to indicate its relationship to other similar forms. Whilst each group of basically identical individuals is called a species, designated by the second of the two names, the larger groups to which similar species are considered to belong are called genera (singular genus) and it is the genera that contribute the first name. So, the fungus genus Lepiota includes species like Lepiota lilacea, Lepiota magnispora and Lepiota obscura. The specific or trivial name often attempts to give some information about the organism, for example lilacea signifies a lilac-coloured fungus and magnispora one with large spores, although sometimes, as in obscura, it does no more than recognise some undefined peculiar feature.

The category above the genus is the Family and above that is the Order. Names of families always have the ending -aceae. Names of orders end in -ales and above them in the fungal taxonomic hierarchy is the Subclass ending in -mycetidae and the Class, ending in -mycetes. A few fungal scientific names have given rise to widely used general terms. The name agaric is often used to mean any fungus with a shape roughly like most of those in the Order Agaricales – the familiar umbrella-like toadstools with gills beneath the cap. A bolete is a comparable toadstool with pores like those in the Order Boletales, especially the family Boletaceae, while a polypore is any fungus belonging to the Phylum Basidiomycota (apart from the stalked, toadstool-like boletes) that discharges its spores through pores and not from gills; most of them are bracket-shaped. Finally, a corticioid fungus is one that forms a thin, skin-like, fibrous or membranous fruit body, usually growing on wood – it takes its name from the Order Corticiales to which many of them belong.

Where to Find Fungi

Fungi are able to exploit most of the natural – and many of the artificial – raw materials of the world as nutrient sources; and to tolerate most of the environmental variables the earth can offer. Of all the natural habitats able to support life of any type, almost all are inhabited by some species of fungi. But whilst many genera of larger fungi certainly occur predominantly in one type of habitat, there are few that are wholly characteristic of individual types of woodland, grassland or other community. Nonetheless, there are certainly some fungal genera, and, more significantly, some associations of genera, that do give each habitat a characteristic mycobiota. Amanita, Lactarius and Russula for instance, which are mycorrhizal associates of trees, are predominantly woodland genera, while Hygrocybe is usually found in grassland. And a species list including Mycena capillaris, Russula fellea, Craterellus cornucopioides and Boletus satanas conjures up an image of a beech wood to a mycologist in much the same way as a list including bramble, dog’s mercury, foxglove, holly and violet helleborine might to a botanist.

Woodlands

More kinds of macro-fungi are to be found in woodlands than anywhere else and every organised fungal foray will include a visit to at least one. Woodlands offer a rich and continuing nutrient source and a wide range of microhabitats (from the twigs and leaves of tree tops to roots many centimetres below the soil surface), a soil environment reasonably well buffered against extremes of temperature and a fairly moist environment all year. They are conveniently divided into the major categories of coniferous, broadleaved and mixed, with additional habitats provided by the woodland edges and by open, grassy areas or glades within the wood itself.

Within these broad divisions, however, there is a wide range of environments. Broad-leaved woodland for instance embraces such disparate habitats as the almost pure stands of beech on the chalk downs with their extremely sparse ground flora, the rich oak wood with a ground flora dominated by early season species like blue-bells that flower before the canopy closes, and the carr, dominated by alder and willows and with almost permanent standing water. Copses and hedgerows are among several other more specialised types of woodland habitat. Each woodland type will be considered briefly in turn.

In Britain, there are only three native conifers: Scots pine, juniper and yew, and natural woodland dominated by any of them is now extremely rare, although the remnants of old Caledonian pine forest in Scotland are highly important habitats, not only for many rare fungi, but for flowering plants and bryophytes too. The coniferous woodland and forest that occurs so widely today has largely been planted over the past two or three centuries, and although it includes some Scots pine, it is dominated by exotics – especially other species of pine, larches, Norway and Sitka spruces, Douglas fir, hemlock, western red cedar and firs. Whilst native woodland and forest of some of these species occurs extensively in other parts of western Europe, most of the remainder are North American introductions. The dense shade cast by many conifers that inhibits much plant life on the ground has no direct effect on fungi and they are often highly conspicuous on the more or less bare needle litter of the forest floor. Conversely, the general paucity of tree species in conifer plantations means there is likely to be a paucity of fungal species too. The fungi of British coniferous woodland and plantation forest comprise those native species that have adapted to the new environment and those that have presumably been introduced along with their associated tree species. The Larch Bolete Suillus grevillei for instance is an obligate mycorrhizal associate of larches and could not have existed in Britain before the introduction of larch trees from Europe in the mid 17th century.

No large genus of fungi is exclusively associated with conifers but among terrestrial Basidiomycetes, Cortinarius and Suillus are well represented, either in coniferous woods or associated with conifers in mixed stands. Notable and common individual agaric species include Tricholomopsis rutilans on conifer stumps, Auriscalpium vulgare and other small species emerging from buried pine cones, and Chroogomphus rutilus under pines. Wood-destroying bracket-forming fungi are more frequently specific to particular types of tree, and probably the most important example affecting a wide range of conifers is Heterobasidion annosum. One of the few fungi that destroys yew wood, although it occurs on broad-leaved trees also, is the yellow bracket Laetiporus sulphureus.

Over much of Britain, mixed broad-leaved woodland, containing greater or lesser numbers of oaks, beech, limes, hornbeam, elms, birches, ash and alder, is the native vegetation type. Many fungi are associated with particular types of broad-leaved tree, and some tend to occur only in more or less pure woodland of individual species – Russula fellea, for example, occurs almost exclusively in old beech woods. Others, like Leccinum scabrum, which is always found with birch, noticeably goes wherever the tree goes, whether in pure or mixed woodland, in copses or as isolated individuals in parkland. Birch woodland and birch trees in general have quite characteristic fungal associates and there are perhaps more common species invariably associated with birches than with any other single tree genus or species. Other fungi by contrast are found in company with any of a range of broad-leaved trees. Many kinds, for instance, occur commonly with both beech and oaks; old broad-leaved woodland, such as that in the New Forest, Hampshire, which contains both these types of tree, probably supports the richest diversity of fungal species to be found in any British habitat.

Among more local and restricted types of woodland with interesting mycobiota are those found mainly on wetlands, particularly copses and riverside plantations of willow or poplar, and especially alder carr. A surprisingly large number of fungal species are alder associates: Pholiota alnicola and Gyrodon lividus are among the most striking, while almost the entire genus Naucoria is found with alders, hence the English name aldercaps.

Hedgerows are in effect long narrow woods with rigorous management, and although most of the broad-leaved tree species found in woodland also occur in hedgerows, along with at least some of their associated fungi, certain trees have come to be thought of as predominantly hedgerow species. The various species of elm came into this category before elm disease removed so many of them from the landscape after the 1970s and there has been a consequent decline in the numbers of elm-associated fungi like Rigidoporus ulmarius and Entoloma aprile, although fallen elm trunks and stacks of logs led to a temporary increase in the occurrence of such species as Pleurotus cornucopiae. Ash is also very much a hedgerow tree over much of lowland Britain and two fungi in particular are very commonly but not obligatorily associated with it: the wood-rotting Basidiomycete Inonotus hispidus and the ascomycete Daldinia concentrica.

Grassland

Grasslands in Britain today are entirely artificial habitats, created largely in historical times on areas cleared of the ancient forest and used for grazing animals. Some grasslands are essentially short-term environments, lasting for one or a few years as breaks from other crops being grown in rotation on arable farmland. As habitats for macro-fungi, they are almost barren because no large basidiomycete has enough time to build up a mature mycelium in the soil before it is disturbed by ploughing. Such grasslands also tend to be subject, either directly or indirectly, to the fertilisers and crop-protection chemicals that form such an integral part of much modern farming and which do little to create a favourable environment for fungi. Apart from a few species such as Volvariella gloiocephala and Cyathus olla that grow on and among stubble, the modern farm field environment, with the exception of old set-aside land, will not repay much mycological study.

Established permanent grasslands, meadows, commons, lawns and parkland are very different however. These are relatively undisturbed habitats, although the grass itself may be grazed or mown. The soil temperature is relatively uniform, protected by the grass swards, but may be prone to drying out in summer. Among major agaric genera, Agaricus, Clitocybe and Hygrocybe are especially frequent on these grasslands, and representatives of all groups that do not have an obligate mycorrhizal association with trees or that depend on wood as nutrient source may be expected. The sub-division of grasslands is less straightforward than that of woodlands, and although certain well-defined types can be recognised, there are less obviously characteristic mycobiotas in each.

The rich lowland pastures of water meadows and valley bottoms, the rough unimproved and mainly upland grasslands where no fertiliser has been used, and the fine downland pastures overlying chalk all include species of the major grassland genera. In the first, Agaricus arvensis, Agaricus urinascens and Agaricus campestris occur typically with Psilocybe semilanceata and many of the related common dung fungi. On the unimproved upland areas will be found many of the brightly coloured species of Hygrocybe and such rarities as Tulostoma niveum, while the short downland turf supports more Agaricus and Hygrocybe species and very frequently Calocybe gambosa, St George’s Mushroom. The Fairy Ring Mushroom Marasmius oreades is also common on short grazed or mown turf and it has been determined from growth rates that some of the fairy rings on undisturbed downland may be a hundred or more years old. Marasmius oreades also occurs on established lawns along with several small species of relatively unremarkable-looking brown fungi, including species of Coprinellus, Panaeolus and Psathyrella.

Gardens and urban environments

The garden, like the farm, is not generally a good environment for fungi as the soil, other than under lawns, is too frequently disturbed. Nonetheless, there are a few species that are tolerant of the moderate disturbance in herbaceous borders and shrubberies, including Coprinus comatus and the Stinkhorn Phallus impudicus. Agaricus bitorquis is one of the commonest culprits in raising tarmac from pavements and paths and is so much a species of disturbed ground that its presence can be predicted with reasonable certainty within two or three years of new roadside verges being laid.

One increasingly important garden habitat however is the wood chip mulch laid on flower beds and shrubberies. This has been responsible for a number of alien agarics like Psilocybe cyanescens and Leratiomyces ceres appearing and spreading in Britain, presumably with imported wood, but it has also meant that many native woodland species now appear in gardens and even Morels, Morchella spp., are no longer as rare there as before. Many introduced agarics must come, albeit not always noticed, attached to their mycorrhizal hosts. One that is particularly widely distributed in parks and gardens with Eucalyptus is Laccaria fraterna, which has been found at many sites, especially in South West England. Greenhouses are sometimes the habitats for aliens that have arrived with plants or compost material. Among them are the distinctive brown-capped Gymnopilus dilepis (rather like a small Tricholomopsis rutilans), the pale brown Agrocybe putaminum and perhaps most striking of all, the beautiful yellow Leucocoprinus birnbaumii. Compost heaps, especially when neglected for sufficiently long periods, can be a fruitful environment for some interesting agarics, including species of Coprinellus, Coprinopsis and Psilocybe.

Fungi may also sometimes be found in houses. Fortunately for the owners, they are usually no more significant than microscopic moulds, but many old houses, especially with cellars, will support some growth of Coniophora puteana, the cause of wet rot or the serious wood-destroying alien Serpula lacrymans, the cause of dry rot. Other Basidiomycetes, including Antrodia vaillantii and even the agaric Agrocybe arvalis, may also be found occasionally in cellars.

Heathland, moorland and mountains

Heathland and moorland are habitats where, even more than coniferous woodland, the environment is shaped by an acidic soil. The dominant plants are not trees or grasses but usually members of the heather family, the Ericaceae. More than in any other British habitats, fungi, as one component of lichens, may approach a dominant role in some areas. As habitats for fungi, the high soil acidity is challenging but moisture content remains generally high and the plant life relatively undisturbed for long spells, interrupted only by the limited grazing of sheep or deer and by periodic burning, either accidental or deliberate. Among the common small Basidiomycetes is Marasmius androsaceus, the Horsehair Parachute, which causes a die-back of heather stems and then grows on the dead remains, while Lichenomphalia umbellifera is among a group of closely related agarics that grow on and among mosses and form a lichenised association with algae growing in the same habitat. Hypholoma and Hygrocybe species are other common mountain and moorland fungi, while in communities containing dwarf willows, mycorrhizal species of such familiar woodland genera as Russula and Lactarius occur commonly at high altitude.

Fire sites

Much moorland is periodically burned, either accidentally as a result of lightning strikes or by deliberate actions designed to stimulate the growth of the heather. Fire can sometimes stimulate spore germination and can clear the soil of competing microorganisms and fire sites have their own characteristic fungi. In woodland, the ascomycete tree pathogen Rhizina undulata presents such a serious threat that even the lighting of fires by forest workers to brew tea is prohibited. Among the commoner agaric fungi of burned ground are Pholiota highlandensis, Hebeloma anthracophila, Tephrocybe anthracophila and Tephrocybe atrata.

Bogs, marshes and other wetland areas

Some of the woodland habitats discussed already, like birch woodland and alder carr, are essentially wetlands. Other habitats dominated by trees may present fungi with a moist, fairly uniform environment but they must adapt to poor aeration and the possibility of high concentrations of sulphurous gases. These are the conditions of bogs on acidic peaty soils, fens on less acidic peats and marshes on silty soils. Bogs may occur at both high and low altitudes, grading from moorland in the former where rainfall is high and drainage impeded. Fens occur typically in the upper parts of some old river estuaries but in all these habitats, mosses, especially species of Sphagnum, make up important components and it tends to be fungi that grow in close association with mosses that are the most characteristic species. Galerina, Pholiota and Hypholoma are among the genera of small, brownish and often inconspicuous Basidiomycetes found most frequently. Some of them have greatly elongated stems to facilitate spore dispersal clear of the Sphagnum or other vegetation. A very few Basidiomycetes like Arrhenia lobata that grow on and among the bases of rushes, sedges or other herbaceous plants may even be almost entirely aquatic.

Dunes and salt marshes

Sand dunes might appear an improbable habitat for fungi, apparently lacking nutrients and being in a state of permanent movement. Nonetheless, in the more stable areas of dunes there is a small but characteristic range of species but they are often difficult to find and collect because some of the most interesting, like Agaricus devoniensis, barely reveal their caps above the surface of the sand. Many of the species, like Psathyrella ammophila, have a very long stem that enables them either to reach moisture supplies at depth or to grow in association with such plants as marram. Among other Basidiomycetes likely to be found are Conocybe dunensis, Inocybe dunensis, Hohenbuehelia culmicola and Phallus hadriani. Agaricus bernardii is perhaps the most striking of those dune fungi that also occur on salt marshes and other coastal habitats subject to salt spray.

Collecting, Studying and Identifying Fungi

To naturalists who are used to identifying birds and mammals, the study of fungi (rather like the study of some insects and other small creatures, and to a certain extent some plants) requires a rather different approach. This is because relatively few species can be named accurately in the field, other than by the most experienced collectors, and even they almost always find it necessary to collect fruit bodies for close inspection. Most fungi, moreover, require microscopic examination and sometimes simple chemical tests too, either to verify an identification based on superficial characters or, sometimes, to identify them at all.

The legality of collecting

Rightly, many kinds of plant and animal now enjoy some form of legal protection. It is illegal for instance to uproot any plant without the permission of the owner of the land on which it is growing; and some may not be dug up or picked under any circumstances. However, in being neither plant nor animal, fungi slip through some of the protection legislation. Largely because of this, a group of organisations with a particular interest in fungal conservation has published guidelines in The Wild Mushroom Pickers’ Code of Conduct and its provisions should be closely followed whether collecting for study or for the kitchen. The following guidelines are based on those in the Code of Conduct:

Always obtain the landowner’s or site manager’s permission before you enter land and explain the purpose of your visit.

Follow The Countryside Code.

Try to minimise damage to vegetation, leaf litter, soil and other habitat features.

Avoid removing dead wood unless this is necessary to identify a fungus.

Take a reputable field guide with you and try to identify as many fungi as you can in situ.

Ancient woodlands usually contain a rich variety of different types of fungi and may include some rare species. Particular care should be taken when collecting from these sites.

If collecting fungi for study:

Collect the minimum amount of material or number of specimens required for a proper description and reliable identification.

Fungi are enjoyed by many people because of their beauty and intrigue. For this reason you should take care to minimise the visual effect of collecting samples.

Record accurately the localities and habitat data for rare species.

Do not collect legally protected species.

Always offer the results of your survey to the landowner or manager and explain the significance of what you have found.

If you have permission to collect for scientific purposes do not abuse it by also collecting to eat.

Supply information to local and national databases and retain ‘voucher specimens’ for deposit in museum collections.

And if collecting fungi to eat:

Be aware that some fungi are very poisonous and many others may make you unwell. Some people have allergic reactions after eating certain species. Make quite sure you can identify fungi by attending a field course or foray led by an expert before you start collecting.

Do not collect species you don’t intend to eat.

Respect and protect other species, including poisonous ones.

Do not collect uncommon, rare, Red List or legally protected species.

Only collect from plentiful populations and take no more than what you want for your personal consumption. Pick no more than 1.5 kilograms per visit or no more than half of the fruit bodies of any single species present, whichever is the lower amount. Larger quantities taken for profit or other use should be agreed beforehand with the landowner or site manager.

Do not collect ‘buttons’ (mushrooms that have not expanded). Giving buttons time to expand will allow spores to be discharged and will give you a bigger mushroom to eat.

On some Sites of Special Scientific Interest (SSSIs), most nature reserves and on other protected areas it is unlikely that culinary collecting is allowed. Always consult the site owner or manager before collecting.

Class A drugs in fungi

There is another dimension to the legality of collecting fungi however – not a conservation but a chemical one. Some fungal species, most of which are popularly and collectively called magic mushrooms, contain one of the two related indole hallucinogens psilocin and psilocybin (chemicals that block the action of the nerve impulse transmitter serotonin in brain tissue). The result of eating them is a relatively short-lived hallucinogenic effect, similar to that produced by LSD. These indole compounds are Class A drugs and as such are embraced by Section 21 of the Drugs Act 2005 and it is an offence to import, export, produce, supply, possess or possess with intent to supply magic mushrooms in whatever form, whether prepared or fresh.

Although the Law and its interpretation is rather complex, and no prosecutions have yet been brought, it is clear that deliberately and knowingly collecting magic mushrooms is illegal unless you intend immediately to destroy them or pass them to someone who is licensed to hold them. Licences are held by such bodies as the two Royal Botanic Gardens at Kew and Edinburgh but are unlikely to be issued to individuals. If you collect magic mushrooms without knowing what they are, it appears you will only commit an offence if you retain them once they have been identified. You will not be committing an offence of possession if magic mushrooms are growing uncultivated on your premises.

The list of those species that are classed as magic mushrooms is only provisional and there is uncertainty about the validity of some of the chemistry on which it is based. Nonetheless, it does have legal standing. The relevant species are indicated in the text of this Guide.

Equipment

For collecting

To remove specimens from their natural growing position, a range of appliances will be helpful, most of which can be satisfied by a visit to a garden centre. A hand trowel, a smaller trowel-like tool often called a widger such as those sold for use with house plants, a pair of secateurs (preferably anvil pattern which are better at cutting through hard woody material), a sharp knife and ideally a small folding saw should enable most situations to be dealt with. Immediately after collection, each specimen should be laid out separately (bracken or leaves can make useful partitions) in a suitable carrier – a traditional pattern Sussex trug is ideal.

For taking specimens home

To carry fungi safely, rigid containers are needed. Never put specimens in a plastic bag as they will disintegrate rapidly and be useless for further study. Small tubes and small plastic boxes with tight-fitting lids such as those used for storing food in deep freezers are good because several will fit into a haversack-style bag. On returning home, the tubes and boxes may be placed in a fridge (not a deep freeze) to keep your specimens fresh for a few days, although it is important to prepare spore prints as soon as possible (see page 00), and remember to dry collections promptly for long-term storage before they deteriorate. Several features important for identification can change rapidly after fungi have been picked.

For recording information

A notebook with several pencils is essential and a loose-leaved pattern with a waterproof cover will enable the information to be filed permanently later. An assortment of lined and plain pages will facilitate the making of both notes and sketches. It is essential to record transient characters such as colour, smell, shape and size as soon as possible. An alternative or an accompaniment to a notebook – and very useful in the rain – is a small hand-held voice recorder although the information will have to be transcribed later. Watercolour pencils are useful for making notes of the colour of fresh specimens and although a digital camera is a very useful aid it should not replace field sketches. Each collection of specimens should be accorded an individual number and ideally this should be written on a jewellers’ tag which can usefully be attached to the specimens themselves. If the fungi are to form part of a permanent collection, the tags need never be removed but can be dried with the specimens.

For detailed examination

Although a hand lens magnifying around 10 times is highly useful, especially in the field, a microscope is essential for the certain naming of most fungi, but this need not be sophisticated or expensive if its only purpose is as an aid to identification rather than detailed academic study. A basic ‘student’ pattern of microscope giving a magnification of about 150 times will be adequate for inspecting the shape and overall form of spores and spore-bearing structures, although for examination of the surface details of spores, an oil immersion objective is an enormous help. As size of spores and other bodies is such an important criterion, however, a micrometer eyepiece (and calibration slide) permitting measurements to be made quickly and accurately is essential. A camera lucida attachment which projects an image of the field of view onto a sheet of paper at the side of the microscope is a valuable aid to accurate drawing of spores and other microscopic features but by no means essential. A camera attachment to enable digital images to be made of microscopic preparations is desirable but really a luxury for all except the most serious collector. Gummed labels, a few simple dissecting needles, scalpels, microscope slides and cover glasses, an accurate ruler and dividers for measuring complete the basic requirements.

For permanent storage of specimens

For long-term storage of fungal specimens, careful preparation is needed and this is very difficult to achieve satisfactorily without a well-ventilated drying oven, preferably incorporating a circulating fan. Within the oven, fresh specimens should be laid out carefully and separately on wire shelves or racks. A suitable purpose-made oven can be purchased from companies supplying science equipment to schools. A domestic fan oven can be used but the time over which it is required makes this less practicable. A temperature of 75°C for about 48 hours will dry most types of small- or medium-sized toadstool, although it may be necessary to cut large fruit bodies into sections and dry them for a week or so. It is important to dry, not cook, them.

Dried fungi are best kept in stout paper packets or, for large specimens, cardboard boxes and stored in a dry, fairly warm place with crystals of naphthalene scattered among the specimens to deter insects and moulds.

Chemical tests

A small range of chemical reagents will be necessary for use with particular groups of fungi, which cause characteristic colour changes when the chemicals are applied to them (either as macroscopic reactions, visible to the naked eye, or as microscopic effects, to be seen under the microscope). These reagents are shown here, along with details of the various tests.

Collection and examination in the field

First, use a camera and take plenty of photographs. Digital photography means that large numbers of pictures, either as memory joggers or permanent records, can be taken at no cost. Then make as many notes and collect as much information as possible because many features can change in a few hours and it is usually impossible to return to the same spot to record some previously overlooked but important fact, such as the type of tree beneath which the toadstool was growing. The relevant pages of your notebook should therefore contain: the date and weather conditions; the precise locality; the nature of the habitat (broad-leaved woodland, wet ditch, garden lawn, recently disturbed roadside verge or stable sand dune for instance); the substrate on which the fungus is found (the soil, living branches, rotting timber or even another old toadstool for example); and the nature of the local vegetation, especially the species of tree or plant on which, or close to which, the specimen is growing (it makes sense to take leaves or another sample if the species cannot be identified on the spot). Even the presence of particular kinds of moss or other fungi nearby can be significant. There is however a caveat to this. When recording closely associated vegetation, how close is close? At best, people often simply record the species nearest to their specimen, although the fungus may not in reality be associated with it but with one 10 or even 30 metres away. Tree roots spread considerable distances, especially the fine rootlets with which mycorrhizal species will be associated. Moreover, a little learning can be a dangerous thing. For example, most collectors, if they see Amanita muscaria, look for a birch and record it, if Suillus grevillei they look for larch or Russula atropurpurea for oak. And having found them, they enter the information in their notes and look no further. Existing beliefs are thus perpetuated. It is potentially much more meaningful and scientifically useful to record as much as possible of the local vegetation – all trees for example within, say, 15 metres. Of course, it will take longer but it is far better to produce a body of information from which real, rather than imagined, relationships can be deduced. And it may teach mycologists something about higher plants, pteridophytes, bryophytes, ecology and biodiversity, of which many are more ignorant than they are prepared either to admit or even realise!

Note the relationship of the fruit bodies one to the other. This is especially important with agarics which may occur as solitary individuals, attached to each other in various ways in tufts or clumps, or trooping (groups of isolated individuals growing close together but not obviously attached).

Remove the specimens carefully from the substrate, noting especially if the stem base is rooting (inserted deeply into the ground, rather like a carrot root). Make a record of any features that are immediately obvious and striking (a hairy cap, a smell of almonds or a bright red colour for example) and also those characters that change on handling (such as a colour change similar to bruising where the tissues have been touched, or an aroma that becomes suddenly apparent). Whenever possible, try to take at least one immature fruit body but remember that without some fully mature individuals identification will usually be impossible. It is much better to place only one collection in each container for bringing home; many toadstools look remarkably similar, especially when seen apart from their habitat, and trying to relate a mixed collection to one species can cause endless frustration. The need not to mix collections is even more important when collecting fungi to eat because an inedible or poisonous species can so easily be tossed into the pan along with the edible ones.

Two or three fruit bodies of each species should be adequate for identification, and perhaps five or six if the material is to be preserved as part of a permanent collection. It is unlikely that the time available to the average amateur collector will permit the examination and identification of more than about six or, at the most, ten unknown species from each collecting expedition. To take home more than this is wasteful therefore – especially as the main autumn collecting period lasts only a few weeks during which time perhaps ten trips may be made.

Always remember that it is far better to identify with certainty ten species and so enrich your fund of knowledge and experience than return home wastefully with fifty or more unknowns, most of which there will never be any prospect of naming.

Patterns of growth

Examination at home

At home, it is possible to begin a systematic examination of your specimens, but whilst numerous macroscopic features of the fruit bodies and microscopic details of their structure can be recorded, not all are needed to identify every fungus and each individual species description in this book uses only some of them therefore. The basis of this account is the identification of agarics which will comprise the bulk of the harvest from most collecting expeditions. Additional and different features needed for other groups are described here.

Spores and most other features of microscopic interest do retain their form more or less indefinitely and can be examined at leisure after small pieces of dried tissue have been macerated in warm water, Melzer’s reagent (see here), methylene blue or cotton blue (see here). It should be remembered nonetheless, that for some fungi, such as those in the genera Lactarius and Russula, features like spore print colour, taste and smell are of such importance for identification that accurate naming of dried specimens without detailed notes on the fresh material is often impossible.

A few types of Basidiomycete such as the jelly-like members of the family Dacrymycetaceae do not dry satisfactorily, and although they can be preserved in alcohol, formaldehyde or other liquid, the results are not very satisfactory. Most types of Ascomycete, apart from a few terrestrial species, are much less fleshy than most large Basidiomycetes and will dry very adequately in the air at room temperature.

Identification

The identification of fungi is not inherently any more difficult than the identification of other organisms but it requires a degree of patience and there are a number of unique factors. One major problem lies with the ephemeral nature of most fungal fruit bodies and the fairly irregular appearance of some from season to season. Thus, in many instances, there are only a few weeks each season in which to practise identification and some species may only be seen once every few years, even in the same locality. Beginners should take comfort therefore from the fact that even experienced field mycologists have to re-learn a great deal each autumn. Only the professional or very serious amateur with access to a fungal herbarium has year-round availability of material for study.

Criteria used to describe and identify agarics

Macroscopic features of the fruit body

The Cap

Dimensions. The diameter to be used is the average measured in centimetres across at least two diameters of a mature cap. The height, which is used normally only with markedly conical or bell-shaped caps, is the average in centimetres of at least two heights measured from the apex to the cap edge. Cap diameter and height vary considerably with overall growing conditions, and whilst there are obviously large and obviously small types of agaric, size alone is almost never a criterion for determining the limits of a species. Moreover, the size ranges given in the species descriptions should never be considered as excluding the existence of some particularly large or small individuals.

Colour. This can vary greatly as the cap matures and opens and is also sometimes variable from one part of the cap to another. It also changes in some species when the cap is wet and this phenomenon can aid identification in certain groups. The commonest cap colours among fungi are shades of brown, yellow and orange, although they can range from white and greys through pink and red to black. Blue and mauve are uncommon and green very rare. Although, technically, and for the greatest accuracy, it is best to describe colours in relation to standard colour charts, these are often expensive and difficult to obtain and time-consuming to use. Coloured sketches or colour descriptions related to everyday objects are more practical and have largely been adopted in this book.

Shape. Cap shape shows the greatest difference between mature and immature specimens and this difference should be noted. The description of cap shape used normally in species descriptions relates to the mature form and five main types are recognised: bell-shaped, conical, convex (or in its extreme form, domed) with the appearance of an upturned bowl, flat and, finally, uplifted, where the entire cap is depressed into a bowl-like form with uplifted margins. In describing shape, the presence and form of any central bump or depression should also be noted. A cap with some form of central bump is called umbonate or, if the bump is very small and pimple-like, papillate; one with a depression is called depressed although a very deep depression rendering the cap funnel-shaped is found quite commonly and is characteristic of some genera. A depression in the cap with a small bump or pimple in the centre is also fairly common and is called umbilicate.

Cap shapes

Surface characters. There are many different cap surface features, often highly characteristic of particular groups, but only the commonest and most important need be considered. Note if the surface is shiny, dull or silky, if it is dry or moist, sticky, glutinous, oily, greasy or gelatinous. The surface texture varies widely too, but the most important types to note are smooth and split or cracked, with a covering that may range from glistening, powdery, granular and scaly to more or less hairy or woolly. The presence of faint lines on the cap extending from the margin partway to the centre may be important; sometimes these are only apparent or enhanced when the cap is wet and their extent from the margin should be recorded. In some genera, the entire cap and fruit body may darken when wet (hygrophanous). In a few genera, most especially Russula, the extent if any to which the cap cuticle (skin) may be peeled away from the edge is an important charter, ranging from peeling at the edge only to a proportion of the cap radius or fully to the centre.

Marginal characters. The margin of the cap may differ significantly from the remainder in colour or surface texture, and the edge may be either perfectly smooth (entire) or more or less wavy, split or eroded. There are often small radial lines around the circumference associated with the gill attachment but some caps are lined radially as a separate marginal feature.

Cap margins

Flesh characters. These are very important for the identification of many species and should be noted carefully. Thickness (from cap upper surface to under surface where the gills emerge) both in the centre and at the margin may be measured in millimetres for record purposes but is usually mentioned in the species descriptions only if it is significantly thick or thin. Colour is an important feature and especially any changes in it when the flesh is cut or broken; some of these occur almost immediately, others take some minutes to become apparent. Smell is important too and is best experienced by holding a piece of broken cap to the face in cupped hands. It can be described most readily by reference to familiar everyday objects such as flour, aniseed or radish. Sometimes it nonetheless defies precise categorising and can only be said to be pleasant, unpleasant, nauseous, mushroomy or earthy. For some groups of agarics, flesh taste is an important feature in identification, but it is important to stress that tasting a fungus is not the same as eating it. A small piece of the cap flesh should be chewed on the tip of the tongue, the sensation (hot and peppery for instance) noted and the fragments then politely spat out. It should be stressed that taste is never the first criterion to use when identifying an unknown species; it must be employed only when the fungus has been fairly positively placed in a particular group or in a genus like Russula within which individual species descriptions mention taste as an important diagnostic feature. In a few genera such as Lactarius and Mycena, the presence of fluid exuding from the cut or broken flesh is an important feature. The colour of this fluid should be noted, together with any changes it makes on exposure to the air; this exposure and colour change is best achieved by touching the fluid drops onto a white tissue or handkerchief. The consistency of the flesh should also be recorded, noting for instance if it is crumbly, rubbery or brittle.

The Gills

Certain features of the gills are important in the identification of all gilled fungi. Of greatest significance is the manner by which the gills are attached to the stem and there are six major types of attachment, with a degree of inter-grading between some of them. Free gills do not actually make contact with the stem, and, when viewed from below, a circular gap is apparent around the top of the stem where the cap underside can be seen. When this gap is very wide and the gills end a considerable distance from the stem, they are said to be remote. Adnate gills are attached to the stem squarely, their entire width making contact, whereas adnexed and emarginate gills are only attached to the stem for a small or a large proportion of their width respectively. Emarginate gills appear to be notched at the lower edge where they meet the stem, while decurrent gills arch downwards at the inner edge and appear to run down, gradually merging with the stem. Gill width is sometimes used as a distinguishing feature as is the thickness (from face to face) of each individual gill. The spacing of gills is an expression of their appearance when seen from beneath the cap and ranges from crowded (where the gills are very close together) to distant (where they are very wide apart).

The appearance of the gill margin (its lower edge) is especially important in some genera. It may be described as undulating or wavy in the same way as the edge of the cap itself, but quite commonly gills are found that appear toothed or minutely ragged. Sometimes the gill edge is of a different colour to the remainder of the gill whereas overall gill colour, especially the difference between immature and mature gills, is very important in some groups. Surface features of gills are not as important in identification as the surface features of the cap, although in a few groups the relative waxiness or pubescence (minute hairiness, like the surface of a peach) is used as a diagnostic criterion.

Finally, it should be noted if the gills are deliquescing or self-liquefying and turning inky, a characteristic of many species in the genera Coprinus, Coprinopsis, Coprinellus and Parasola.

Gill attachments

Gill margins

Spacing the gills

Stages in colour change of gills

The Stem

Attachment to the cap. Although most agarics have the stem attached centrally under the cap – like an umbrella handle – some are slightly excentric and a few, which appear superficially bracket-like, have it attached laterally. A very few are stem-less and have a fruit body form more commonly associated with other groups of fungi (see here).

Attachment of stem to cap

Dimensions. The length of the stem is measured from its point of emergence from the substrate to its attachment to the gills (which may be some distance from the cap surface if the gills are markedly decurrent). The diameter of the stem is a fairly variable character and difficult to measure accurately; but make note of a markedly stout or markedly thin stem (taking width in relation to length).

Shape. Most stems are equal (more or less parallel sided) but some taper either upwards or downwards; most species with rooting stems taper downwards into the ‘root’, while others are markedly club-shaped (clavate) with a highly pronounced taper upwards or even bulbous (with a pronounced swelling at the base like an onion). Some bulbous forms have a marked and rather sharp margin around the upper edge of the bulb; these are called marginate bulbous while those that swell very abruptly at the base are called abruptly bulbous. Stem shape in transverse section should be noted if it is other than circular; some species have markedly flattened or grooved stems for instance, while it is sometimes important to observe if it is solid in section, hollow or stuffed (apparently with a central hollow filled with cottony tissue).

Stem shapes

Sections through stems

Colour and texture

Many of the cap flesh characters like colour, taste and smell and also some of surface textural features are applicable to the stem too, although it is important to note any differences in colour, texture or surface features that are restricted to the extreme apex or base. Additional features to note on the stem surface are the various degrees of wrinkling or grooving mentioned above, while the flesh is often much tougher than that of the cap and can be cartilaginous, chalky, corky, fibrous, leathery or even woody, in addition to the possibilities referred to already.

Other stem features

Other stem features have manifestations on the stem and, to some extent, on the cap too; these are the presence or absence of veils – more or less filmy sheets of tissue produced on the fruit bodies of some agaric species. They are of two types, partial veils and universal veils. A partial veil covers the gills of an agaric fruit body as it emerges from the substrate, stretching from the cap edge to the stem. As the cap grows and expands, this veil tears in one of two main ways. An arachnoid (spider or cobweb-like) veil splits radially leaving fibrils on the cap edge and sometimes adhering across the gills too. By contrast, a membranous veil tears concentrically around the cap, sometimes leaving flaps of tissue conspicuously on the cap edge and more importantly a ring of tissue (called the ring) on the stem itself. In many fungi the ring is barely detectable but in others it forms a very obvious feature, important in identification. The ring may take several forms; if it joins only loosely to the stem, it is called movable, if more tightly adhering, attached. If it is on the upper part of the stem, a ring is called superior; if on the lower half, inferior; a ring with a cotton wool-like roll of tissue on the underside is referred to as double. Sometimes there is no proper ring, only a ring-zone or slightly rougher tissue and contrasting colour on the stem.

The manifestations of a universal veil are somewhat different. Universal veils are not formed by all agarics but when present they envelop the entire developing ‘button’ and as the fruit body expands and matures, fragments of the veil remain as flakes on the cap surface, often in contrasting colour to the remainder of the cap. More significantly, however, the veil remains enveloping the base of the stem as a structure called a volva, the presence of which is characteristic of certain genera. The volva takes one of two main forms: when its tissues adhere closely to the stem base, it is called adherent and its surface may be loose and scaly, powdery or, sometimes, marked with characteristic concentric zones; when the volva envelops the stem base like a loose bag, it is said to be free and its tissues may then vary from very fragile to fairly tough and membranous.

Ring types

Adherent volva

Free volva

Development of the arachnoid veil

Remains of universal veil as cap flecks

Remnants of partial veil

Spore Print

A spore print is the coloured pattern produced when spores are discharged from the gills onto white paper. It is one of the basic features employed in the identification and, to some degree (although less now than formerly), in the classification of agarics. It is fairly consistent within genera, with all species tending to produce spore prints in one main colour type: dark brown to purple-brown; olive-brown through mid-brown to red-brown; black; pink; and white-cream. However, a few genera such as Flammulaster do span colour boundaries. But remember that the colour of the mature gills may not always give an indication of the colour of the spore print; some fungi with dark coloured gills produce white or colourless spores and consequently a white print. Careful attention must be paid to the way in which a spore print is obtained and to the manner in which its colour is assessed. In some agaric genera, most notably Russula, subtle variations in spore print colour are very important in the identification of individual species, and, to facilitate this, carefully produced colour charts are provided below with which the spore print colours should be matched.

Chart of Russula spore print colours

To produce a spore print, use a fresh, mature but not over-ripe fruit body. The freshness is important because it can be difficult to obtain a print from specimens that have been kept in a fridge or allowed to dry out slightly. It is an operation that should be started immediately on returning home after a collecting expedition. Cut the cap from the stem at the apex with a sharp knife and place it gills downwards onto a piece of smooth, stiff white paper or card. Cover the cap with an inverted jar or similar cover and leave it undisturbed. Depending on the state of cap maturity, it may need to be left for anything between half an hour and 24 hours. The spores will then be discharged from the gills as described later and produce a pattern on the card; this is the spore print, and for most identification purposes the colour of the pattern on the paper, when dry, can be used. For critical genera like Russula, however, scrape the spores into a small heap with a microscope slide cover glass, flatten the heap gently with the cover glass and assess the colour of this mass of spores in daylight (take the card to a window during the daytime, because evening light in autumn and artificial light can give confusing colours).

Chemical tests on the fruit body

The identification of some species is aided by a colour change that occurs when a chemical reagent is applied to the tissues. The following are the most useful of these tests and reference is made to them in the species descriptions but it should be appreciated that each is only of value for particular groups and must only be used to aid diagnosis when they are referred to specifically. All of the reagents should be prepared freshly; certainly each season, although sulphovanillin is best made up each time it is needed. Some of the chemicals, especially chloral hydrate and the concentrated mineral acids, are very dangerous. Special permission may be needed before they can be purchased and all should be stored and used with respect and none allowed to be used by children.

Iron salts test

Apply a few drops of a fresh aqueous solution of 10 per cent (weight: volume) ferric chloride or ferric sulphate to the stem and observe any colour change. If it is possible to obtain a large crystal of ferric sulphate, this can be rubbed directly on the stem and so used for rapid identification of certain species of Russula in the field. It can be kept fresh when not in use by placing it in a corked tube containing a wad of cotton wool moistened with dilute ammonium sulphate.

Melzer’s test

(the same reagent is also used for microscopic tests)

Prepare Melzer’s reagent by adding 1.5 g iodine, 5 g potassium iodide and 100 g chloral hydrate to 100 ml of water. Warm carefully (do not boil) and stir thoroughly. When added to tissues or spores (especially white spores) en masse, a colour change to dark blue is called an amyloid response; a change to red-brown is dextrinoid; no change is non-amyloid and non-dextrinoid.

Schaeffer’s test

Apply a streak across the cap of Agaricus species with a glass rod dipped in pure aniline and cross this with another rod dipped in concentrated nitric acid. A flame red colour at the point of intersection gives Schaeffer +; no red colour gives Schaeffer -. This test has been included in the Guide because it is useful for advanced study in the genus Agaricus, but concentrated nitric acid is an extremely dangerous chemical and the procedure should not be used by inexperienced collectors or those unfamiliar with the practices of laboratory chemistry.

Sulphovanillin test

Add 8 ml concentrated sulphuric acid to 3 ml water and dissolve 1 g pure vanillin in this. A characteristic colour change occurs in some groups when this is applied to the stem. Concentrated sulphuric acid is an extremely dangerous chemical and the reagent must be prepared with the greatest care and only by those experienced in its use.

Miscellaneous reagent tests

In certain groups, application of one of the following reagents to the cap, stem or hyphae gives a characteristic colour reaction:

Aqueous 75 per cent ammonia

Aqueous 40 per cent formaldehyde (formalin)

Aqueous 2 per cent phenol

Aqueous 10–40 per cent potassium or sodium hydroxide (KOH or NaOH)

Congo Red aqueous solution

Dilute or concentrated solutions of hydrochloric, nitric or sulphuric acids. These acids are extremely dangerous chemicals and should only be used by those experienced in their handling.

Microscopic features

Spores

Spore shape and