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Fly agaric in a spruce forest on mossy ground, an ectomycorrhizal partner in its natural setting

Mycorrhiza

Mycorrhiza: which plant goes with which fungus?

Almost every plant on Earth shares its roots with a fungus. The fungus opens up soil the plant cannot reach on its own, delivers phosphorus and water, keeps root diseases at bay, and gets paid in sugar. This partnership is more than 400 million years old, older than the root itself.1,2 The catch: not every fungus fits every plant. Once you know which system a species has, you plan trees and shrubs differently, and you also understand why porcini grow under an oak and never under an apple tree. This page starts at the very beginning (what a fungus is, how it lives, what mycorrhiza means) and works its way into the detail from there.

~85%of all vascular plant species live with a permanent fungal partner3
4 ways of lifemycorrhiza, endophyte, decomposer, pathogen: only one of them is a trade
400 millionyears old: the partnership is older than roots, leaves and seeds
4 typesof mycorrhiza exist, and two of them split almost the entire plant world between them

From the big picture to the detail

First, some sorting: fungus, way of life, mycorrhiza

Before we get to types and partners, it helps to keep three levels apart, otherwise everything later on sounds contradictory. Open whatever interests you; if you already know the basics, skip ahead.

What is a fungus, actually?

Ask a child to draw a mushroom and you get an umbrella. But that is only one shape of fruiting body, and many fungi never form one at all.

An adult would try to be more precise: a living thing that feeds through threads instead of roots, does no photosynthesis and reproduces by spores instead of seeds. Sounds reasonable. Except that none of the three points holds up:

  • Threads instead of roots? Yeasts are fungi and single-celled, with no threads at all. And some bacteria, the other way round, do form a mycelium, and even smell of forest soil while doing it.4 (Where there are threads, a single one is called a hypha and the network a mycelium. We will need both words again shortly.)
  • No photosynthesis? Yellow bird's-nest does none either: the whitish-yellow growth that pushes up from the ground in conifer forests and looks like a fungus itself. It is still a plant.
  • Spores instead of seeds? Ferns do exactly the same.

What remains is the cell wall.5

Every fungal cell is wrapped in a firm wall, and that wall contains chitin (C8H13NO5)n. Plants build their walls from cellulose (C6H10O5)n. Chemically they are almost the same molecule; chitin simply carries an extra bit of nitrogen on every building block. Animals have no cell walls at all; a beetle does carry chitin too, but as armor on the outside of its body rather than as a wall around every single cell. And bacteria use something else again, called murein.

In short: fungi are chitin beings, plants are cellulose beings.

The counter-test: downy mildew.

It forms threads, digests outside its body and in every respect lives like a fungus. It just isn't one. It belongs to the oomycetes (water molds), is more closely related to brown algae, and its cell wall is made of cellulose: a cellulose being playing at fungus.6 This is not an academic detail. Products against true fungi often have no effect on it.

You cannot tell by looking at it. People used to sort organisms by appearance and reproduction, and that is exactly why things that are not fungi sat among the fungi for decades. Only comparing their DNA set the record straight.

Not everything is what it looks like. Dolphins are not fish either, they are mammals.

And now to the point this page is about.

A fungus cannot make sugar and has to get it from outside. As we have just seen, that is not unique either: animals cannot make it, and neither can yellow bird's-nest. For this page only one question counts: where does a fungus get its sugar from?

Because that is exactly where the stories about the Wood Wide Web come from. One step at a time.

How do fungi make a living? The four ways of life

Everything hinges on where the sugar comes from. That gives four ways of life, sorted by what a fungus colonizes and whether it pays:

GroupLives on or inRole in the garden
Mycorrhizal fungiliving roots, and pays for itSupply. You want these. Porcini, chanterelle, truffle, fly agaric.
Endophytesliving tissue, without a tradeStress tolerance. A bonus: they are there anyway. You never see them.
Pathogensliving tissue, at its expenseDisease. You don't want these. Honey fungus, Phytophthora, Monilinia (brown rot), scab.
Saprotrophsdead materialDecomposition and humus. You need these. Oyster mushroom, shiitake, button mushroom.

Saprotrophs are not pests. That is the most common mix-up. They break down leaves, mulch, prunings and deadwood, and so are the basis of all humus building. Without them, mulch just sits there and never turns into anything. White rot and brown rot are simply their two ways of working on wood, and they normally happen on dead wood: in the deadwood pile, in the stump, in the hügelkultur bed. Because they need no living partner, these are exactly the fungi that can be cultivated.

Where the boundary really is fluid: between endophyte and pathogen. The same fungus can sit harmlessly in the tissue for years and turn aggressive as soon as the plant comes under stress. That is a real mechanism, covered in the very next section. And the fact that honey fungus goes on decomposing the wood after it has killed the tree does not make it a decomposer: it can do both, but it started out as a pathogen.

Endophytes: the invisible majority in every leaf

They are everywhere: in leaves, needles, bark, wood, and yes, in roots too. Practically every plant carries endophytes, often dozens of species (including the apple tree you pick from).7 You do not see them because they form no fruiting bodies as long as the tissue is alive; they lie as thin hyphae between the plant cells.

What they live on: whatever is lying around between the cells anyway, sugars and amino acids that seep through the cell walls. Very little, lost in the noise. That is why the plant does not notice. And that is precisely the difference from mycorrhiza, which receives large amounts of sugar, but through a real organ and in return for something. The endophyte only skims off.

What they do, three things that matter in the garden:

  • They occupy the space. Where an endophyte sits, a pathogen can no longer get in.
  • They produce defensive compounds against plant-eating insects and fungal diseases.8
  • They train the defenses of the plant, which then reacts faster to real attacks.

This is even put to use: pasture grasses with endophytes are tougher, and become toxic to grazing animals.9 It is the best-known case of this double nature.

The point about stress. It is not the endophyte that wears the tree down. Rather, many of these fungi are opportunistic pathogens. As long as the tree is vigorous, its defenses keep them in check and they stay harmless. When drought knocks out those defenses, they switch strategy and start to grow. That is why branches suddenly die back in drought years without any disease having "arrived": the pathogen had been sitting inside for years. Diplodia on pines and Botryosphaeria on fruit trees work exactly like this.10

And one more thing that closes the circle: when the leaf falls, many of these fungi are the first to start decomposing it. They are already inside and can get going right away. That does not make them decomposers; endophyte is a way of life of its own. It just means that nobody has to arrive from outside before a freshly fallen leaf starts to break down.11

Aren't pathogens bacteria? What really makes plants sick

Bacteria are just one group among several, and for trees and shrubs the smallest:

  • Fungi, by far the largest share for trees: honey fungus, Monilinia, scab, powdery mildew, rusts, root rot (Heterobasidion).
  • Oomycetes look like fungi and live like fungi, but they are not fungi; they are more closely related to brown algae. They include Phytophthora, the cause of many root and collar rots and of late blight in potatoes. For control this is not an academic detail: products against true fungi often do not work on them.
  • Bacteria, for example fire blight, feared in orchards, but the exception.
  • Viruses, usually spread by aphids or pruning tools, and incurable.
  • Nematodes, which are animals, not microbes, and often underestimated in the soil.

And the distinction that is worth most in the garden: a fungus on dead wood is not a pathogen. Whatever grows in the deadwood pile, in the stump or in the hügelkultur bed is decomposing; it is not attacking anything. Only when fruiting bodies appear on a living trunk is it worth a second look.

For a tree, then: whoever talks about disease means a fungus in the vast majority of cases. And whoever hears about a giant fungus in the forest is not necessarily hearing about a partner: the record holder among fungi is a tree killer. More on that further down.

So what does "mycorrhiza" mean?

Literally "fungus root", from Greek: mykes fungus, rhiza root. And that already says the most important thing:

Mycorrhiza is not a species of fungus or a group of fungi, it is a connection. It means the shared organ of fungus and living root, not the fungus on its own. You cannot buy it or hold it in your hand; it only comes into being where both sides meet.

Three features define it: close, regulated contact between fungus and living root; an exchange from which both sides benefit; and a distinct structure at the contact point that can be recognized under the microscope.

A fungus that can form such a structure is called a mycorrhizal fungus. As a rule it does not maintain just one such connection but many at once, to several roots and often to several plants.

Why do some types seem to show up twice later on? Three levels to keep apart

This is where things almost always get muddled, so here it is before the types come up:

  • Level 1, the fungus's way of life: mycorrhiza, endophyte, saprotroph, pathogen. Four of them, see above.
  • Level 2, the type of connection, within the "mycorrhiza" way of life: arbuscular, ecto, ericoid, orchid. Each type has its own group of fungi, and the groups are not interchangeable.
  • Level 3, variants within one type. This is where arbutoid and monotropoid belong: they are formed by the same ectomycorrhizal fungi as ordinary ectomycorrhiza, just on different plants and therefore built differently.

The key point: which variant forms depends on the plant, not on the fungus. One and the same ectomycorrhizal fungus forms ordinary ectomycorrhiza with a pine, an arbutoid one with a strawberry tree and a monotropoid one with yellow bird's-nest: one individual fungus, three variants.12,13

An analogy: charging cables.

  • Level 2, the types, are different connector families: USB and Lightning. An apple tree has one socket, an oak the other, and a fungus with the wrong plug simply cannot connect, however close it lies. Blueberries and orchids each have yet another port of their own.
  • Level 3, the variants, are plug shapes within one family: USB-A, USB-C, Micro-USB. All USB, all the same fungi; only the socket looks different on a pine, a strawberry tree and yellow bird's-nest.
  • Willow and poplar are the devices with two different sockets, so they can bridge two families.
  • Yellow bird's-nest is the device that only ever charges and never gives anything back.

Where the analogy breaks down: a plug is a question of fit, a mycorrhiza is a trade. And the socket is not there beforehand: fungus and root build it together, from scratch, every time.

When this page talks about "the two systems", it means the two big families: arbuscular and ecto. That is the distinction that matters when you plant.

Are all fungi mycorrhizal fungi?

No, quite the opposite: they are a minority. The vast majority of fungi are decomposers. Only certain groups can form mycorrhiza, and those that can usually depend on it.

You can see one consequence of this in the supermarket: button mushrooms, oyster mushrooms and shiitake are for sale, porcini and chanterelles are not (or only wild-picked). The first are decomposers and grow on straw, manure or wood. The second need a living tree and therefore cannot be grown in a shed; you can only gather them or plant a tree whose roots have been inoculated beforehand. That is exactly what is done with truffles.

The flip side: a fungus growing in your compost is not a partner for your tree. And a mycorrhizal fungus cannot live on compost.

The words that keep coming up on this page
Hypha
A single fungal thread, twenty to fifty times thinner than a fine root. Plural: hyphae.
Mycelium
The network of many hyphae: the actual fungus.
Fruiting body
What you gather and eat. Only the spore-producing organ, not the organism.
Spore
The fungus's unit of dispersal: single-celled, fine as dust, spread by the wind. This is how a fungus reaches new ground. Ferns, mosses and bacteria have spores too, so for fungi there is nothing special about it.
Rhizomorph
A cord-like strand of bundled hyphae. Some fungi (above all honey fungus) use these to bridge open stretches without roots.
Symbiont
A partner in a relationship from which both sides benefit.
Saprotroph
Decomposer: lives on dead material.
Endophyte
Lives inside living plant tissue without the plant noticing.
Pathogen
A disease-causing organism.
Myco-heterotrophic
A plant that feeds wholly or partly on a fungus instead of on photosynthesis.
Genet
One individual fungus: everything that has grown from a single spore, however widely spread out.

The basics

What a fungal partner actually does

A fine root is about half a millimeter thick. A fungal hypha is twenty to fifty times thinner, which lets it into soil pores that stay closed to the root. A single gram of soil often holds tens of meters of hyphae.14,15 This multiplies the surface through which the plant takes things up, without it growing a single extra millimeter of root.

Four services lie behind this:

🧪

Phosphorus

The most important point. Phosphorus is usually plentiful in soil, but tightly bound and barely mobile. Around a root it is used up after a short time, and the plant cannot reach the rest. The fungus fetches it from areas well beyond that zone.15

💧

Water and drought tolerance

The same reach applies to water. Colonized plants often come through dry spells better16, and on free-draining soils that is often the difference between establishing and struggling.

🛡️

Protection from root diseases

The fungal sheath occupies the root surface and makes it inaccessible to harmful fungi; on top of that come defensive compounds and a primed immune response in the plant.17,18 Citrus is considered especially dependent on its fungal partner.19

🔗

Exchange with the neighbors

Everything attached to the same fungus is connected. It is proven that carbon, nitrogen and phosphorus move from one plant to another along the way; how much and how far is what the argument is about. More on that further down.

The forms

The four types of mycorrhiza (and the two that cover almost everything)

Now the second level: within the "mycorrhiza" way of life there are four types. They differ in how closely fungus and root grow together: does the fungus grow into the root cell or only wrap around it? In practice it is enough to tell the first two apart; the other two concern blueberries and orchids. If the levels are not clear yet, look them up above.

FormWhoWhat defines it
Arbuscular mycorrhizaAbout 72% of all vascular plant species: almost all fruit, vegetables, herbs, grasses, palms, legumes and nearly all tropical crop treesThe fungus grows into the root cell and forms little tree-shaped exchange organs there, the arbuscules. It forms no visible fruiting bodies, so you never get to see it. Only a few hundred fungal species serve this huge group of plants (244 had been described by 2015, DNA analyses point to 300 to 1,600)20, and accordingly they are not very picky.
EctomycorrhizaAbout 2% of species, but they include the dominant forest trees of the temperate and boreal zones plus some tropical familiesThe fungus wraps the root tip in a sheath and does not enter the cells. Around 20,000 fungal species20, many of them tied to just a few tree species, and almost all well-known edible wild mushrooms belong here: porcini, chanterelle, truffle, bay bolete.
Ericoid mycorrhizaThe heath family: blueberry, lingonberry, cranberry, rhododendron, heatherSpecialized in acidic, wet, humus-rich soils. The fungus unlocks nitrogen directly from undecomposed raw humus, which is why blueberries grow where there is nothing to be had for other plants.21
Orchid mycorrhizaAll orchids, about 10% of all vascular plant speciesAn orchid seed is fine as dust and has no reserves. Without the right fungus to feed it during germination, it does not come up. That is why you cannot simply sow orchids.22

Two variants, and why they are not types of their own

Books often list two more types. But they are not types: both are formed by perfectly ordinary ectomycorrhizal fungi and therefore belong in the same drawer as row two.

Arbutoid mycorrhiza: strawberry tree and bearberry. A sheath as in ectomycorrhiza, but the fungus also enters the cell.

Monotropoid mycorrhiza: yellow bird's-nest and its relatives, plants without chlorophyll. One and the same individual fungus is attached to a spruce on one side and to yellow bird's-nest on the other, which lives entirely on the tree's sugar.13,23 The best evidence that carbon flows from plant to plant through a fungus.

In short: which variant forms depends on the plant, not on the fungus. In the analogy above they are USB-A, USB-C and Micro-USB, but all of them USB. One individual fungus can serve a pine, a strawberry tree and yellow bird's-nest at the same time. When this page talks about "the two systems", it means the two big connector families, arbuscular and ecto, and these two variants count as ecto.

Two things do not belong on this list even though they often turn up here. First, the dark septate endophytes: fungi that live in roots without forming any of the classic structures. They are widespread on extreme sites and their effect is unclear, but they are endophytes, not mycorrhiza. Second, the plants with no fungal partner at all, about eight percent of species (they get a section of their own in a moment).

Worldwide guide

Which plant has which mycorrhiza? From apple to coconut palm

The rule of thumb holds on every continent: arbuscular is the norm, ectomycorrhiza the exception.3,24 But that exception is concentrated in a few families, and those very families make up the great forests of the northern hemisphere, Australia and Southeast Asia.

Species or groupSystemNote
Apple, pear, quince, cherry, plum, peach, almondarbuscularAll pome and stone fruit, plus strawberry, raspberry, blackberry, currants and gooseberry.
WalnutarbuscularOften mixed up: despite its size, the walnut is not an ectomycorrhizal tree. Its close relative the pecan, on the other hand, is.25,26
Hazel, oak, beech, birch, sweet chestnut, hornbeam, pecanectoThe edible-mushroom trees. Truffles grow under hazel and oak, birch boletes under birch, porcini and bay boletes under spruce and pine.
Pine, spruce, larch, fir, Douglas fir, cedarectoConifers are almost without exception ectomycorrhizal partners. Without colonization, pine seedlings practically do not grow on raw soil.
AlderectoDoubly equipped: ectomycorrhiza for phosphorus, plus Frankia nodules for nitrogen from the air. That is why it copes with raw soils where nothing else will establish.
Oleaster (Russian olive), sea buckthornarbuscularAlso with Frankia nodules, but with the other fungal system than alder27, a difference that is easy to overlook when putting a group together.
Willow, poplarbothThe most important exception of all. They can form arbuscular and ectomycorrhiza side by side and so bridge two groups. Ectomycorrhiza usually dominates; the arbuscular form joins in mainly in young plants and on wet sites.3,28
Eucalyptus, many acaciasecto (acacias arbuscular depending on the species, some also ecto)Australia is the great ectomycorrhiza continent outside the north.
Mango, avocado, papaya, banana, citrus, coffee, cocoa, olive, fig, date and coconut palmarbuscularTropical and subtropical crops practically without exception. Citrus is considered especially dependent, above all on rootstocks with coarse roots.19,29
Corn (maize), wheat, millet, rice, potato, tomato, onion, leek, carrot, squash, bean, pea, clover, alfalfaarbuscularAlmost all field crops and vegetables. Leek and onion are considered especially heavily colonized and are often used as indicator plants in trials.
Blueberry, lingonberry, cranberry, rhododendronericoidA system of its own, with fungi of its own. A blueberry bed hardly benefits from an ectomycorrhizal neighbor and not at all from an arbuscular one. The well-known exception: a few fungi from the group around Rhizoscyphus ericae colonize heather and pine at the same time, but only over a few centimeters. No large shared network comes out of it.30,31
Cabbage, rapeseed, mustard, radish, arugula, cress, horseradishnoneAll brassicas. What that means in practice.
Beetroot, chard, spinach, quinoa, amaranthnoneGoosefoot and amaranth family.
Rhubarb, sorrel, buckwheat, knotweednoneKnotweed family, mostly not colonized or only weakly.
LupinnoneInstead it forms dense root clusters that dissolve phosphorus out of the soil with acids on their own32, its own solution to the same problem.
Colonized root: white fungal mycelium on fine roots, next to it a sheathed ectomycorrhizal root tip and a stained microscope image showing arbuscules in root tissue
The same root, magnified twice. Top right: ectomycorrhiza, where the fungus wraps a sheath around the root tip. Bottom right: the arbuscular form, where it grows into the cell and forms the fine branchings that give it its name.

The exceptions

Plants without a fungal partner, and why that matters in the garden

About eight percent of vascular plants manage entirely without mycorrhiza, and another seven percent are colonized sometimes and sometimes not.3,33,34 They gave up the partnership again over the course of evolution and unlock phosphorus themselves: with acids that release it from its bonds, and with strikingly dense root clusters. It works, but it costs the plant more energy, which is why these species are typically at home on disturbed, nutrient-rich sites.

And the kitchen garden, of all places, is full of them: cabbage in all its forms, radishes, mustard, arugula, cress, horseradish. Plus beetroot, chard, spinach, rhubarb and most lupins.

The green manure trap

This leads to a practical point that you rarely find written down anywhere: mustard and oilseed radish as a cover crop do not feed the fungi. Where only brassicas grow for a year, the amount of arbuscular fungi in the soil drops noticeably because they lack a host; in one trial it fell by a good half after brown mustard.35,36,37 Brassicas probably also release compounds that inhibit colonization.38

In the vegetable garden this usually has no consequences. Before planting trees and shrubs, before corn or before strongly dependent crops such as citrus, though, it is a real drawback. If you choose green manure with the fungi in mind, go for phacelia, clover, vetch, rye or simply a grass mix, and leave out mustard and oilseed radish, which form no partnerships, as well as buckwheat, which is only weakly colonized.39

In practice

Mycorrhiza in the garden: the most common questions

The questions that actually come up when you plant, from "do I need to do anything?" to "how do I connect two groups?".

When does mycorrhiza arrive on its own, and when not?

It depends on the system, and the difference is big.

Arbuscular mycorrhiza: already there. Every meadow and every field is full of it, because grasses and herbs are partners of these fungi themselves. An apple, a walnut or an elder that you plant in a meadow is colonized within a few weeks to months. You do not need to do anything.

Ectomycorrhiza: this is the problem case. In a meadow without host trees there are hardly any spores of these fungi. An oak, beech, hazel or sweet chestnut on open ground finds little at first. Spores arrive by wind and animals, and colonization takes months to years. In a trial with oak seedlings on fallow land, all seedlings up to eight meters from the forest edge were colonized; from about 15 meters on, colonization dropped off markedly.52

And there is a sequence: the pioneer fungi come first (deceivers, earthfans, poisonpies). The late colonizers, meaning brittlegills, webcaps and amanitas, need decades and an established stand.53,54 So you can easily have half a forest for ten years that is still thinly equipped on the fungal side.

What does it actually achieve to plant the same types next to each other?

It saves you exactly that waiting time, and that is the real benefit of planning a group in the first place.

If you plant an ectomycorrhizal shrub next to an established tree of the same system, the existing mycelium grows into the new roots. That happens within one season, instead of waiting years for spores to drift in by chance.52,55 A hazel planted next to an established oak is therefore something completely different from the same hazel alone in the corner of a meadow.

That is exactly why it pays to group trees and shrubs of the same system instead of scattering them across the site.

Do the types cross? Do they interfere with each other?

No, they do not get in each other's way. Hyphae of both systems live side by side in the same soil, each on its own roots. A mixed planting is no disadvantage.

What you have to let go of is the idea of one big network. Many individual fungi are only a few meters across, the largest known ones a few dozen; any piece of land is a patchwork of many such individuals, not one continuous cable. The "road" mentioned on the plant guilds page is therefore more of a chain of handover points than a cable. Covered in detail further up.

How do I connect two groups?

There is an elegant solution for this: willow and poplar can form both types, and so can eucalyptus and alder. Ectomycorrhiza usually dominates; the arbuscular form joins in mainly in young plants and on wet or extreme sites. Worldwide, just under 90 plant genera with this kind of dual mycorrhiza are known28; in the garden these four are the obvious choices.

A row of basket willows between an oak and hazel group and a fruit group is therefore the connecting element, and on the side it gives you prunings, early-spring forage for bees and weaving material. If you want to put the idea into practice on your own land, this is where to start.

Do I need to buy mycorrhiza and inoculate?

For fruit trees and bushes it is almost always money down the drain: they need arbuscular fungi, and those are already present in established garden or meadow soil.

It makes sense for oak, beech, hazel and sweet chestnut on open ground, exactly where the right fungi are missing and colonization would otherwise take years. And on land where soil life is fundamentally lacking: long-term arable fields, raw subsoil after construction, stripped topsoil, land reclamation.

Even then, though: a wheelbarrow of soil from the root zone of an old, healthy tree of the same species is usually more effective and cheaper than any product. You get exactly the fungi that occur locally and cope with the site. Many products you can buy contain only arbuscular fungi anyway and are therefore useless for an oak or a birch from the start.

What helps without inoculating: keep the soil covered, turn it as little as possible, leave roots in the ground over winter and avoid long periods without plants. Fungi live on living roots; bare soil starves them.

Then why are there so many different products, even special ones for conifers?

Because the two groups differ fundamentally in one respect: how picky they are.

Arbuscular fungi are generalists. There are only a few hundred species for around 200,000 partner plants, so they simply cannot afford to be picky. One and the same fungus colonizes apple, tomato, corn and lavender. That is why a single universal product really is enough here, and most contain only a handful of species.

Ectomycorrhizal fungi are often specialists. Over twenty thousand species, and many of them are tied to one tree genus or even a single species: the birch bolete grows only with birch, the larch bolete only with larch, the black truffle only with oak and hazel.56 An all-purpose product cannot exist here even in principle, hence the split into conifer, broadleaf, oak and so on. Then there is the site: conifer fungi are mostly adapted to acidic soil, others to chalky soil.

Still, the choice is smaller than the labels suggest. Only what can be propagated can be sold, and that means mainly pioneer species, the generalists among the specialists: dyeballs (Pisolithus), common earthballs, deceivers, false truffles (Rhizopogon), poisonpies. Exactly the fungi that would find a young tree first anyway. The late colonizers that characterize an old stand cannot be raised in a bag.

And what most people actually want does not exist. Porcini and chanterelles still cannot be grown commercially (for the chanterelle it worked once in the lab, in 1997)57,58, so there is no product, however high the demand. The one exception is the truffle, and that is not sold as powder but as a ready-inoculated, certified young tree from the nursery.

Part of the variety is simply product range. The market is barely regulated, and tests of commercial products regularly find discrepancies between label and contents, right down to packs with no detectable viable fungi. In a worldwide test of 28 products, 84% led to no colonization at all in sterile soil.59,60,61

What this means in practice is exactly what the previous answer says: where inoculation is needed at all, a wheelbarrow of soil from the root zone of an old, healthy tree of the same species is the more reliable route. The right species are guaranteed to be in there, alive and used to the site.

Are non-native trees more susceptible to disease?

The finding is clear, the cause is not. A study in 48 Czech castle parks examined the soil under 73 non-native tree species: there, pathogens made up twice as large a share of fungal species and three times as large a share of fungal biomass as under native trees, while ectomycorrhizal fungi made up a smaller share. The longer a tree species had been growing locally, the lower the share of pathogens.62

What causes this, though, the study does not say. It could be poorer supply with suitable fungal partners; it could just as well be which pathogens were imported together with the plants. Both are plausible, neither is proven.

Put honestly, the finding is: fewer fungal partners, more pathogens, cause unknown. It still works as a rule of thumb for planting, as an observation rather than an explanation.

And edible mushrooms: can I plan for those too?

Yes, but it is a different route from the inoculation above, even though the same fungal groups are involved. If you want to harvest truffles, porcini or chanterelles, you buy specifically inoculated planting stock (a hazel or oak whose roots were colonized with a particular fungal species at the nursery) and you need the right soil to go with it, which for truffles above all means a high pH. This has nothing to do with an all-purpose mycorrhiza powder.

Mushrooms such as oyster mushroom or shiitake, on the other hand, are not mycorrhizal fungi at all but decomposers: they are grown on logs or substrate, independently of any living tree. That is why you can buy them and not porcini.

A cluster of porcini at the foot of an old broadleaf tree in the grass
Porcini do not stand at the foot of the trunk by chance: they live off this tree and it lives off them. That is exactly why you cannot grow them on straw like an oyster mushroom. Without a living partner there is no porcini.

Next steps

What this means for your planting plan

Three sentences are enough for everyday use. First: a fungus only colonizes plants of its own system, so if you put a forest tree among nothing but fruit trees, you give it no partner. Second: neighborhood matters, not area; place trees and shrubs of the same system so that their root zones touch. Third: a plant guild should not be locked into a single system, otherwise it has no connection to whatever grows next to it.

Permaplans garden map with the mycorrhizal root zones layer: colored root circles per tree or shrub for ecto, endo, dual system and heath family over the aerial photo, plus plus signs as planting suggestions for gaps
This is how it looks in Permaplans: the "Mycorrhiza root zones" layer colors the estimated root zone of every tree and shrub by its system (blue ecto, yellow endo, green both, purple the heath family such as blueberry and rhododendron). Where areas of the same color overlap, a shared fungus is likely; the plus signs suggest plants for the gaps.

How to turn this into a complete planting, with an anchor plant, roles and a timeline, is covered on the sister page: Plant guilds: what belongs together.

Plan your site in the browser Browse species

Evidence

Sources

The superscript numbers in the text point here, and ↩ takes you back to the text. Where textbook knowledge is concerned, the standard work is cited; where experts disagree (above all on the Wood Wide Web), both sides are listed. Planning conclusions such as grouping trees and shrubs are our own inferences from these findings, not study results.

  1. Remy, W., Taylor, T. N., Hass, H. & Kerp, H. (1994): Four hundred-million-year-old vesicular arbuscular mycorrhizae. PNAS 91: 11841–11843. doi:10.1073/pnas.91.25.11841 ↩
  2. Brundrett, M. C. (2002): Coevolution of roots and mycorrhizas of land plants. New Phytologist 154: 275–304. doi:10.1046/j.1469-8137.2002.00397.x ↩
  3. Brundrett, M. C. & Tedersoo, L. (2018): Evolutionary history of mycorrhizal symbioses and global host plant diversity. New Phytologist 220: 1108–1115. doi:10.1111/nph.14976 ↩ ↩ ↩ ↩
  4. Jiang, J., He, X. & Cane, D. E. (2007): Biosynthesis of the earthy odorant geosmin by a bifunctional Streptomyces coelicolor enzyme. Nature Chemical Biology 3: 711–715. doi:10.1038/nchembio.2007.29 ↩
  5. Gow, N. A. R., Latgé, J.-P. & Munro, C. A. (2017): The fungal cell wall: structure, biosynthesis, and function. Microbiology Spectrum 5(3): FUNK-0035-2016. doi:10.1128/microbiolspec.FUNK-0035-2016 ↩
  6. Fawke, S., Doumane, M. & Schornack, S. (2015): Oomycete interactions with plants: infection strategies and resistance principles. Microbiology and Molecular Biology Reviews 79: 263–280. doi:10.1128/MMBR.00010-15 ↩
  7. Rodriguez, R. J., White Jr, J. F., Arnold, A. E. & Redman, R. S. (2009): Fungal endophytes: diversity and functional roles. New Phytologist 182: 314–330. doi:10.1111/j.1469-8137.2009.02773.x ↩
  8. Arnold, A. E., Mejía, L. C., Kyllo, D., Rojas, E. I., Maynard, Z., Robbins, N. & Herre, E. A. (2003): Fungal endophytes limit pathogen damage in a tropical tree. PNAS 100: 15649–15654. doi:10.1073/pnas.2533483100 ↩
  9. Schardl, C. L., Leuchtmann, A. & Spiering, M. J. (2004): Symbioses of grasses with seedborne fungal endophytes. Annual Review of Plant Biology 55: 315–340. doi:10.1146/annurev.arplant.55.031903.141735 ↩
  10. Slippers, B. & Wingfield, M. J. (2007): Botryosphaeriaceae as endophytes and latent pathogens of woody plants: diversity, ecology and impact. Fungal Biology Reviews 21: 90–106. doi:10.1016/j.fbr.2007.06.002 ↩
  11. Osono, T. (2006): Role of phyllosphere fungi of forest trees in the development of decomposer fungal communities and decomposition processes of leaf litter. Canadian Journal of Microbiology 52: 701–716. doi:10.1139/w06-023 ↩
  12. Molina, R. & Trappe, J. M. (1982): Lack of mycorrhizal specificity by the ericaceous hosts Arbutus menziesii and Arctostaphylos uva-ursi. New Phytologist 90: 495–509. doi:10.1111/j.1469-8137.1982.tb04482.x ↩
  13. Bidartondo, M. I. & Bruns, T. D. (2001): Extreme specificity in epiparasitic Monotropoideae (Ericaceae): widespread phylogenetic and geographical structure. Molecular Ecology 10: 2285–2295. doi:10.1046/j.1365-294X.2001.01358.x ↩ ↩ ↩
  14. Leake, J., Johnson, D., Donnelly, D., Muckle, G., Boddy, L. & Read, D. (2004): Networks of power and influence: the role of mycorrhizal mycelium in controlling plant communities and agroecosystem functioning. Canadian Journal of Botany 82: 1016–1045. doi:10.1139/b04-060 ↩
  15. Smith, S. E. & Read, D. J. (2008): Mycorrhizal Symbiosis. 3rd ed., Academic Press, London. ISBN 978-0-12-370526-6. doi:10.1016/B978-0-12-370526-6.X5001-6 ↩ ↩
  16. Augé, R. M. (2001): Water relations, drought and vesicular-arbuscular mycorrhizal symbiosis. Mycorrhiza 11: 3–42. doi:10.1007/s005720100097 ↩
  17. Marx, D. H. (1972): Ectomycorrhizae as biological deterrents to pathogenic root infections. Annual Review of Phytopathology 10: 429–454. doi:10.1146/annurev.py.10.090172.002241 ↩
  18. Jung, S. C., Martinez-Medina, A., Lopez-Raez, J. A. & Pozo, M. J. (2012): Mycorrhiza-induced resistance and priming of plant defenses. Journal of Chemical Ecology 38: 651–664. doi:10.1007/s10886-012-0134-6 ↩
  19. Menge, J. A., Johnson, E. L. V. & Platt, R. G. (1978): Mycorrhizal dependency of several citrus cultivars under three nutrient regimes. New Phytologist 81: 553–559. doi:10.1111/j.1469-8137.1978.tb01628.x ↩ ↩
  20. van der Heijden, M. G. A., Martin, F. M., Selosse, M.-A. & Sanders, I. R. (2015): Mycorrhizal ecology and evolution: the past, the present, and the future. New Phytologist 205: 1406–1423. doi:10.1111/nph.13288 ↩ ↩
  21. Read, D. J., Leake, J. R. & Perez-Moreno, J. (2004): Mycorrhizal fungi as drivers of ecosystem processes in heathland and boreal forest biomes. Canadian Journal of Botany 82: 1243–1263. doi:10.1139/b04-123 ↩
  22. Rasmussen, H. N. & Rasmussen, F. N. (2009): Orchid mycorrhiza: implications of a mycophagous life style. Oikos 118: 334–345. doi:10.1111/j.1600-0706.2008.17116.x ↩
  23. Leake, J. R. (1994): The biology of myco-heterotrophic (‘saprophytic’) plants. New Phytologist 127: 171–216. doi:10.1111/j.1469-8137.1994.tb04272.x ↩ ↩
  24. Wang, B. & Qiu, Y.-L. (2006): Phylogenetic distribution and evolution of mycorrhizas in land plants. Mycorrhiza 16: 299–363. doi:10.1007/s00572-005-0033-6 ↩
  25. Huang, G.-M., Zou, Y.-N., Wu, Q.-S., Xu, Y.-J. & Kuča, K. (2020): Mycorrhizal roles in plant growth, gas exchange, root morphology, and nutrient uptake of walnuts. Plant, Soil and Environment 66: 295–302. doi:10.17221/240/2020-PSE ↩
  26. Bonito, G., Brenneman, T. & Vilgalys, R. (2011): Ectomycorrhizal fungal diversity in orchards of cultivated pecan (Carya illinoinensis; Juglandaceae). Mycorrhiza 21: 601–612. doi:10.1007/s00572-011-0368-0 ↩
  27. Gardner, I. C., Clelland, D. M. & Scott, A. (1984): Mycorrhizal improvement in non-leguminous nitrogen fixing associations with particular reference to Hippophaë rhamnoides L. Plant and Soil 78: 189–199. doi:10.1007/BF02277850 ↩
  28. Teste, F. P., Jones, M. D. & Dickie, I. A. (2020): Dual-mycorrhizal plants: their ecology and relevance. New Phytologist 225: 1835–1851. doi:10.1111/nph.16190 ↩ ↩
  29. Graham, J. H. & Syvertsen, J. P. (1985): Host determinants of mycorrhizal dependency of citrus rootstock seedlings. New Phytologist 101: 667–676. doi:10.1111/j.1469-8137.1985.tb02872.x ↩
  30. Villarreal-Ruiz, L., Anderson, I. C. & Alexander, I. J. (2004): Interaction between an isolate from the Hymenoscyphus ericae aggregate and roots of Pinus and Vaccinium. New Phytologist 164: 183–192. doi:10.1111/j.1469-8137.2004.01167.x ↩
  31. Grelet, G.-A., Johnson, D., Vrålstad, T., Alexander, I. J. & Anderson, I. C. (2010): New insights into the mycorrhizal Rhizoscyphus ericae aggregate: spatial structure and co-colonization of ectomycorrhizal and ericoid roots. New Phytologist 188: 210–222. doi:10.1111/j.1469-8137.2010.03353.x ↩
  32. Lambers, H., Shane, M. W., Cramer, M. D., Pearse, S. J. & Veneklaas, E. J. (2006): Root structure and functioning for efficient acquisition of phosphorus: matching morphological and physiological traits. Annals of Botany 98: 693–713. doi:10.1093/aob/mcl114 ↩
  33. Cosme, M., Fernández, I., Van der Heijden, M. G. A. & Pieterse, C. M. J. (2018): Non-mycorrhizal plants: the exceptions that prove the rule. Trends in Plant Science 23: 577–587. doi:10.1016/j.tplants.2018.04.004 ↩
  34. Brundrett, M. C. (2009): Mycorrhizal associations and other means of nutrition of vascular plants: understanding the global diversity of host plants by resolving conflicting information and developing reliable means of diagnosis. Plant and Soil 320: 37–77. doi:10.1007/s11104-008-9877-9 ↩
  35. Njeru, E. M., Avio, L., Sbrana, C., Turrini, A., Bocci, G., Bàrberi, P. & Giovannetti, M. (2014): First evidence for a major cover crop effect on arbuscular mycorrhizal fungi and organic maize growth. Agronomy for Sustainable Development 34: 841–848. doi:10.1007/s13593-013-0197-y ↩
  36. Gavito, M. E. & Miller, M. H. (1998): Changes in mycorrhiza development in maize induced by crop management practices. Plant and Soil 198: 185–192. doi:10.1023/A:1004314406653 ↩
  37. Lekberg, Y. & Koide, R. T. (2005): Is plant performance limited by abundance of arbuscular mycorrhizal fungi? A meta-analysis of studies published between 1988 and 2003. New Phytologist 168: 189–204. doi:10.1111/j.1469-8137.2005.01490.x ↩
  38. Vierheilig, H., Bennett, R., Kiddle, G., Kaldorf, M. & Ludwig-Müller, J. (2000): Differences in glucosinolate patterns and arbuscular mycorrhizal status of glucosinolate-containing plant species. New Phytologist 146: 343–352. doi:10.1046/j.1469-8137.2000.00642.x ↩
  39. Likar, M., Bukovnik, U., Kreft, I., Chrungoo, N. K. & Regvar, M. (2008): Mycorrhizal status and diversity of fungal endophytes in roots of common buckwheat (Fagopyrum esculentum) and tartary buckwheat (F. tataricum). Mycorrhiza 18: 309–315. doi:10.1007/s00572-008-0181-6 ↩
  40. Ferguson, B. A., Dreisbach, T. A., Parks, C. G., Filip, G. M. & Schmitt, C. L. (2003): Coarse-scale population structure of pathogenic Armillaria species in a mixed-conifer forest in the Blue Mountains of northeast Oregon. Canadian Journal of Forest Research 33: 612–623. doi:10.1139/x03-065 ↩
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  44. Beiler, K. J., Durall, D. M., Simard, S. W., Maxwell, S. A. & Kretzer, A. M. (2010): Architecture of the wood-wide web: Rhizopogon spp. genets link multiple Douglas-fir cohorts. New Phytologist 185: 543–553. doi:10.1111/j.1469-8137.2009.03069.x ↩
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  46. Karst, J., Jones, M. D. & Hoeksema, J. D. (2023): Positive citation bias and overinterpreted results lead to misinformation on common mycorrhizal networks in forests. Nature Ecology & Evolution 7: 501–511. doi:10.1038/s41559-023-01986-1 ↩ ↩
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  48. Simard, S. W., Perry, D. A., Jones, M. D., Myrold, D. D., Durall, D. M. & Molina, R. (1997): Net transfer of carbon between ectomycorrhizal tree species in the field. Nature 388: 579–582. doi:10.1038/41557 ↩ ↩
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  52. Dickie, I. A. & Reich, P. B. (2005): Ectomycorrhizal fungal communities at forest edges. Journal of Ecology 93: 244–255. doi:10.1111/j.1365-2745.2005.00977.x ↩ ↩
  53. Last, F. T., Dighton, J. & Mason, P. A. (1987): Successions of sheathing mycorrhizal fungi. Trends in Ecology & Evolution 2: 157–161. doi:10.1016/0169-5347(87)90066-8 ↩
  54. Twieg, B. D., Durall, D. M. & Simard, S. W. (2007): Ectomycorrhizal fungal succession in mixed temperate forests. New Phytologist 176: 437–447. doi:10.1111/j.1469-8137.2007.02173.x ↩
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Sources last checked: October 2026. Spotted a mistake? Let us know.

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