Phylogenetic distribution and evolution of mycorrhizas in land plants

Mycorrhiza (2006) 16: 299–363 

DOI 10.1007/s00572-005-0033-6 

REVIEW 

B. Wang . Y.-L. Qiu 

Phylogenetic distribution and evolution of mycorrhizas 

in land plants 

Received: 22 June 2005 / Accepted: 15 December 2005 / Published online: 6 May 2006 

# Springer-Verlag 2006 

Abstract A survey of 659 papers mostly published since 

1987 was conducted to compile a checklist of mycorrhizal 

occurrence among 3,617 species (263 families) of land 

plants. A plant phylogeny was then used to map the mycorrhizal 

information to examine evolutionary patterns. Several 

findings from this survey enhance our understanding of 

the roles of mycorrhizas in the origin and subsequent diversification 

of land plants. First, 80 and 92% of surveyed land 

plant species and families are mycorrhizal. Second, arbuscular 

mycorrhiza (AM) is the predominant and ancestral type 

of mycorrhiza in land plants. Its occurrence in a vast majority 

of land plants and early-diverging lineages of liverworts 

suggests that the origin of AM probably coincided with the 

origin of land plants. Third, ectomycorrhiza (ECM) and its 

derived types independently evolved from AM many times 

through parallel evolution. Coevolution between plant and 

fungal partners in ECM and its derived types has probably 

contributed to diversification of both plant hosts and fungal 

symbionts. Fourth, mycoheterotrophy and loss of the mycorrhizal 

condition also evolved many times independently in 

land plants through parallel evolution. 

Keywords Mycorrhizas . Land plants . Fungi . 

Parallel evolution 

Introduction 

Mycorrhizas, dual organs of absorption formed when symbiotic 

fungi inhabit healthy tissues of most terrestrial plants 

(Trappe 1996), have widespread occurrence among land 

plants and are increasingly believed to have played an important 

role in the successful colonization of the land by 

B. Wang . Y.-L. Qiu (*) 

Department of Ecology and Evolutionary Biology, 

The University Herbarium, University of Michigan, 

830 N. University Avenue, Ann Arbor, MI 48109-1048, USA 

e-mail: ylqiu@umich.edu 

Tel.: +1-734-7648279 

Fax: +1-734-7630544 

plants (Pirozynski and Malloch 1975; Malloch et al. 1980; 

Harley and Harley 1987; Trappe 1987; Selosse and Le Tacon 

1998; Readetal.2000; Brundrett 2002). Since Nägeli first 

described them in 1842 (see Koide and Mosse 2004), only a 

few major surveys have been conducted on their phylogenetic 

distribution in various groups of land plants either by 

retrieving information from literature or through direct observation 

(Trappe 1987; Harley and Harley 1987; Newman 

and Reddell 1987). Trappe (1987) gathered information on 

the presence and absence of mycorrhizas in 6,507 species of 

angiosperms investigated in previous studies and mapped the 

phylogenetic distribution of mycorrhizas using the classification 

system by Cronquist (1981). He found that 82% of the 

species were mycorrhizal. From the occurrence of various 

types of mycorrhizas in different subclasses and orders of 

angiosperms, he further inferred that the ancestral type was 

glomeromycetous (the original word “zygomycotous” is not 

used because fungi forming this type of mycorrhiza are now 

placed in the new phylum Glomeromycota, see Schussler et 

al. 2001) mycorrhiza, and ascomycetous and basidiomycetous 

mycorrhizas belong to the derived types, and that evolution 

of mycorrhizas in plants had progressed from obligate 

to facultative mycorrhizas and ultimately to nonmycorrhizas. 

In the same year, Harley and Harley (1987) published a 

checklist of the mycorrhizal status of 144 families of vascular 

plants in the British flora, documenting various types of 

mycorrhizas in an entire flora for the first time. More recently, 

Gemma et al. (1992) and Zhao (2000) investigated the 

mycorrhizal status of 89 and 256 species of pteridophytes in 

Hawaii and Yunnan, China, respectively. They found that the 

percentages of mycorrhizal species in pteridophytes (74 and 

33%, respectively) were lower than in angiosperms as 

reported by Trappe (1987). Whether bryophytes have mycorrhizas 

or not is still a debatable issue, but many liverworts 

and hornworts have fungal associations (Read et al. 2000). In 

two surveys of British liverworts, Pocock and Duckett 

(1985) and Duckett et al. (1991) reported that among the 206 

and 284 examined species, respectively, 16% contained 

fungal endophytes in their rhizoids or thalli. 

Mycorrhizal research has advanced at an astonishing pace 

over the last two decades due to a recent surge of interest in

300 

the subject by botanists, mycologists, and ecologists. A large 

number of new reports on mycorrhizal occurrence in land 

plants, especially in pteridophytes and liverworts, have been 

published since the surveys by Trappe (1987) and Harley and 

Harley (1987). During the same period, our understanding of 

the land plant phylogeny was greatly improved through 

major efforts by molecular systematists. An explicit phylogenetic 

classification system of angiosperms is now available 

(Stevens 2004), and a nonflowering land plant phylogeny 

will be published soon (Qiu et al. unpublished data). Hence, 

reviewing mycorrhizal occurrence in land plants and mapping 

this information onto the newly available plant 

phylogeny will update our knowledge on mycorrhizal abundance 

in plants, facilitate examination of the origin and 

evolutionary pattern of mycorrhizal symbiosis in land plants, 

and enhance our understanding of this important biological 

interaction and its impact on evolution of both plants and 

fungi. Areas that require further study, especially the taxa that 

occupy critical positions in the land plant phylogeny and for 

which mycorrhizal information is still lacking, can also be 

identified through these efforts. 

Materials and methods 

In this review, we compiled information on mycorrhizal 

occurrence in land plants mostly from the papers published 

since 1987, as those published before were reviewed by 

Trappe (1987) and Harley and Harley (1987). First, a library 

consisting of 4,193 mycorrhiza-related references was established. 

After browsing the abstracts, 961 references that 

provided information on mycorrhizal occurrence in plant 

species from all the continents and all major habitats were 

selected. Due to time limitation and unavailability of some 

literature, 659 of these 961 references are reviewed here. 

Data on mycorrhizal status of each plant species were extracted 

from these references, based on whether the species 

was mycorrhizal or not, and if so, what type. An updated 

mycorrhizal classification system proposed by Smith and 

Read (1997) was used here. Seven types of mycorrhizas were 

all recognized (see the footnote b in Table 1), along with 

nonmycorrhiza and mycoheterotrophy. A species is considered 

to be obligately mycorrhizal if it is always found to 

form mycorrhiza, while a species is considered to be facultatively 

mycorrhizal if it is reported to form mycorrhizas in 

one habitat but not in another. If a plant species can form 

different kinds of mycorrhizas simultaneously or in different 

habitats, it is recorded as forming all relevant mycorrhizal 

types. All the data are presented in Table 1, which groups all 

the examined plant species into families that are further 

arranged according to the land plant phylogeny used in this 

study (Stevens 2004; Qiu et al. unpublished). The family 

definition for vascular plants generally follows Mabberley 

(1987), but for a small number of newly recognized families, 

the nomenclature of Stevens (2004) is used. For liverworts, 

the system by Grolle (1983) for family definition is 

followed. The data from Harley and Harley (1987), 

containing 1,101 species, were added to this table, but 

those of Trappe (1987) were not as they were not published. 

The information in Table 1 was then mapped onto a land 

plant phylogeny drawn according to a multigene analysis of 

nonflowering land plants (Qiu et al. unpublished) and the 

angiosperm classification system proposed by Stevens 

(2004). Family was used as the unit of reconstruction (i.e., 

terminal node) in this phylogeny. For each family, the 

percentages of obligate, facultative, and nonmycorrhizal 

species, and the percentage of each type of mycorrhizas 

among all mycorrhizal species were calculated and plotted 

onto the land plant phylogeny. 

One major improvement of this review over those by 

Trappe (1987) and Harley and Harley (1987) istheuseofa 

rigorously reconstructed land plant phylogeny to map the 

mycorrhizal information down to the family level for a 

worldwide coverage. Trappe (1987) covered only angiosperms, 

whereas Harley and Harley (1987)surveyedonlythe 

British vascular flora. Cronquist’s (1981) classification system 

of angiosperms used by Trappe (1987) has been significantly 

revised by molecular systematists (Soltis et al. 2000; 

Stevens 2004). Another major improvement is that information 

on mycorrhizal occurrence, more precisely fungal association 

in bryophytes, is included for the first time. Hence, 

this review should allow a thorough examination of origin 

and evolution of mycorrhizal symbiosis in land plants. 

Results and discussion 

A total of 3,617 species from 263 families of land plants were 

covered in this survey of mycorrhizal status (Tables 1 and 2). 

For land plants as a whole, 80% of the recorded species and 

92% of the families are mycorrhizal. When broken into four 

traditionally delimited groups (angiosperms, gymnosperms, 

pteridophytes, and bryophytes), these percentages vary from 

group to group. In angiosperms, the most species-rich clade 

of land plants and the dominant group in most terrestrial plant 

communities, 85 and 94% of species and families are mycorrhizal. 

These numbers are similar to those reported by Trappe 

(1987), whose study included more than twice as many 

species as this one. All of the 84 species of gymnosperms 

surveyed here are mycorrhizal, and almost all of them are 

obligately mycorrhizal. These two aspects of mycorrhizal 

status in gymnosperms highlight the essential role that mycorrhizas 

play in the life of these plants, which generally 

grow in nutrient-poor environments. For pteridophytes, 52 

and 93% of the species and families are mycorrhizal. Finally, 

46 and 71% of the bryophyte species and families have 

fungal associations. 

When this body of information is projected in a plant 

phylogenetic perspective (Fig. 1), the origin and evolutionary 

patterns of mycorrhizas in land plants become quite 

clear. Below, we discuss these various aspects of mycorrhizal 

evolution in plants. 

Origin of mycorrhizal symbiosis in land plants 

The continuous phylogenetic distribution of mycorrhizas 

throughout land plants, with the sole major exception of

301 

Table 1 An updated checklist of mycorrhizal occurrence among 

land plants 

Examined species a Mycorrhizal 

status b,c 

References 

BRYOPHYTES 

Haplomitriaceae 

Haplomitrium gibbsiae AM-like 107 

Haplomitrium ovalifolium AM-like 107 

Blasiaceae 

Blasia pusilla None 182, 328 

Lunulariaceae 

Lunularia cruciata 

Fungal association 

(G) 

182 

Marchantiaceae 

Marchantia foliacea AM-like 505 

Marchantia polymorpha Fungal association 182 

(G) 

Aytoniaceae 

Asterella wilmsii AM-like 345 

Conocephalaceae 

Conocephalum conicum AM-like 346 

Fungal association 182 

(G) 

Ricciaceae 

Riccia fluitans None 182 

Metzgeriaceae 

Apometzgeria pubescens None 466 

Metzgeria conjugata None 466 

Metzgeria fruticulosa None 466 

Metzgeria furcata None 466 

Metzgeria leptoneura None 466 

Metzgeria temperata None 466 

Aneuraceae 

Aneura pinguis ORM-like (B) 347 

Cryptothallus mirabilis Fungal association 466 

ORM-like (B) 347 

Mycoheterotrophy 77, 483 

(via ECM) 

Riccardia latifrons None 466 

Riccardia multifida None 466 

Riccardia palmate None 466 

Pelliaceae 

Pellia endiviifolia AM-like 598 

Pellia epiphylla 


(G) 

Pellia fabbroniana AM-like 483 

Codoniaceae 

Fossombronia pusilla Fungal association 182 

(G) 

Fossombronia 


wondraczekii 

Radulaceae 

Radula aquilegia None 466 

Radula complanata None 466 

Radula lindbergiana None 466 

Table 1 (continued) 

Examined species a 

Mycorrhizal 

status b,c 

Porellaceae 

Porella arboris-vitae None 466 

Porella cordaeana None 466 

Porella obtusata None 466 

Porella pinnata None 466 

Porella platyphylla None 466 

Jubulaceae 

Frullania dilatata None 466 

Frullania fragilifolia None 466 

Frullania microphylla None 466 

Frullania tamarisci None 466 

Frullania teneriffae None 466 

Jubula hutchinsiae None 466 

Lejeuneaceae 

Aphanolejeunea 

None 466 

microscopica 

Cololejeunea calcarea None 466 

Cololejeunea minutissima None 466 

Colura calyptrifolia None 466 

Drepanolejeunea None 466 

hamatifolia 

Harpalejeunea ovata None 466 

Lejeunea cavifolia None 466 

Lejeunea lamacerina None 466 

Lejeunea patens None 466 

Lejeunea ulicina None 466 

Marchesinia mackaii None 466 

Pseudolepicoleaceae 

Blepharostoma trichophyllum 

None 466 


Herbertaceae 

Herbertus borealis None 466 

Plagiochilaceae 

Pedinophyllum interruptum None 466 

Plagiochila asplenioides None 466 

Plagiochila porelloides None 466 

Plagiochila punctata Fungal association 466 

Plagiochila spinulosa None 466 

Arnelliaceae 

Southbya nigrella 

Fungal association 

(B) 

References 

182 

Geocalycaceae 

Chiloscyphus pallescens None 466 

Chiloscyphus polyanthus None 466 

Geocalyx graveolens Fungal association 466 

Harpanthus flotovianus None 466 

Harpanthus scutatus Fungal association 466 

Leptoscyphus cuneifolius None 466 

Lophocolea bidentata None 466 

Lophocolea cuspidate None 466 

Lophocolea heterophylla None 182, 466 


Saccogyna viticulosa Fungal association 466

302 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Lepidoziaceae 

Bazzania flaccida None 328 

Bazzania tricrenata None 466 

Bazzania trilobata None 328, 466 

Kurzia pauciflora Fungal association 466 

Fungal association 182, 328 

(A) 

Kurzia sylvatica Fungal association 466 

Kurzia trichoclados Fungal association 466 

Lepidozia reptans Fungal association 466 


(A) 

Telaranea murphyae Fungal association 466 

Telaranea nematodes Fungal association 466 

Scapaniaceae 

Diplophyllum albicans None 182, 466 


Diplophyllum obtusifolium Fungal association 328 

Douinia ovata None 466 

Scapania calcicola None 466 

Scapania cuspiduligera None 466 

Scapania gracilis None 466 

Scapania scandica None 466 

Scapania umbrosa None 466 

Cephaloziellaceae 

Cephaloziella 

None 466 

baumgartneri 

Cephaloziella divaricata Fungal association 466 


(A) 

Cephaloziella exiliflora Fungal association 118 

(A) 

Cephaloziella hampeana Fungal association 466 

Cephaloziella massalongi None 466 

Cephaloziella rubella Fungal association 466 

Cephaloziaceae 

Cephalozia bicuspidate Fungal association 466 


(A) 

Cephalozia catenulate None 466 


Cephalozia connivens Fungal association 466 


(A) 

Cephalozia leucantha Fungal association 466 

Cephalozia loitlesbergeri Fungal association 466 


(A) 

Cephalozia lunulifolia Fungal association 328, 466 

Cephalozia macrostachya Fungal association 466 

Cephalozia pleniceps Fungal association 466 

Cladopodiella fluitans None 466 

Cladopodiella francisci Fungal association 466 

Nowellia curvifolia Fungal association 182, 328, 466 

(A) 

Odontoschisma denudatum None 466 


(A) 

Odontoschisma elongatum None 466 

Odontoschisma sphagni Fungal association 466 


(A) 

Calypogeiaceae 

Calypogeia arguta Fungal association 466 

Calypogeia azurea Fungal association 328 

(A, B) 

Calypogeia fissa Fungal association 466 


(A) 

Calypogeia integristipula Fungal association 328 

Calypogeia muellerana Fungal association 466 


(A) 


(A, B) 

Calypogeia neesiana Fungal association 328, 466 

Calypogeia sphagnicola Fungal association 466 

Calypogeia trichomanis Fungal association 466 

Jungermanniaceae 

Anastrophyllum minutum Fungal association 328 

Barbilophozia barbata Fungal association 328 

(A, B) 

Barbilophozia hatcheri None 466 

Jamesoniella autumnalis Fungal association 466 

Jamesoniella undulifolia None 466 

Jungermannia atrovirens None 466 

Jungermannia borealis None 466 

Jungermannia exsertifolia None 466 

Jungermannia gracillima None 182, 466 

Jungermannia obovata None 466 

Jungermannia pumila None 466 

Leiocolea turbinata None 466 

Lophozia incisa 


(B) 

Lophozia sudetica Fungal association 328 

(B) 

Lophozia ventricosa Fungal association 182 

(B) 

Mylia anomala Fungal association 328, 466 

Mylia taylorii Fungal association 328 

Nardia breidleri Fungal association 466 

Nardia scalaris 


(B) 

Sphenolobus helleranus None 466

303 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Sphenolobus minutus None 466 

Gymnomitriaceae 

Gymnomitrion 

None 466 

concinnatum 

Gymnomitrion crenulatum None 466 

Gymnomitrion obtusum None 466 

Marsupella adusta None 466 

Marsupella alpina None 466 

Marsupella emarginata None 466 


Marsupella stableri None 466 

Anthocerotaceae 

Anthoceros punctatus AM-like 525 

Phaeoceros laevis AM-like 344 

PTERIDOPHYTES 

Lycopodiaceae 

Diphasiastrum alpinum AM + NM 260, 261 

Diphasiastrum 

NM 654 

complanatum 

Diphasiastrum issleri AM 260, 261 

Huperzia australiana AM 342 

Huperzia phyllantha AM 226 

Huperzia selago AM + NM 260, 261 

Huperzia serrata f. NM 654 

longipetiolata 

Lycopodiastrum 

NM 654 

casuarinoides 

Lycopodiella inundata AM 212 

AM + NM 260, 261 

Lycopodium alpinum NM 594 

Lycopodium annotinum AM 260, 261 

Lycopodium cernuum AM 181, 226 

Lycopodium clavatum AM 521, 594 

AM + NM 260, 261 

Lycopodium japonicum AM 654 

Lycopodium selago NM 594 

Palhinhaea cernua NM 654 

Phlegmariurus henryi NM 654 

Isoetaceae 

Isoetes echinospora NM 260, 261 

Isoetes histrix NM 260, 261 

Isoetes lacustris NM 260, 261 

Selaginellaceae 

Selaginella arbuscula AM 226 

Selaginella biformis AM 654 

Selaginella chrysocaulos NM 654 

Selaginella davidii AM 653, 654 

Selaginella delicatula AM 654 

Selaginella frondosa AM 654 

Selaginella helferi NM 654 

Selaginella involvens AM 654 

Selaginella kraussiana AM + NM 260, 261 

Selaginella mairei AM 343 

Selaginella moellendorfii AM 653 

Selaginella monospora NM 654 

Selaginella picta AM 654 

Selaginella pulvinata AM 343, 654 

Selaginella remotifolia AM 654 

Selaginella sanguinolenta AM 654 

Selaginella selaginoides AM 260, 261 

Equisetaceae 

Equisetum arvense NM 307, 594, 260, 

261 

AM 599 

Equisetum debile NM 654 

Equisetum diffusum AM 343 

NM 654 

Equisetum fluviatile NM 260, 261 

Equisetum hyemale AM 653, 260, 261 

Equisetum palustre NM 260, 261 

Equisetum pratense NM 260, 261 

Equisetum ramosissimum NM 368, 260, 261 

AM 653 

Equisetum sylvaticum AM + NM 260, 261 

Equisetum telmateia AM 260, 261 

Equisetum × trachydon AM 260, 261 

Equisetum variegatum NM 594 

AM 260, 261 

Marattiaceae 

Angiopteris caudatiformis AM 654, 655 

Angiopteris evecta AM 226 

Angiopteris hokouensis AM 654 

Angiopteris wangii AM 654 

Angiopteris yunnanensis AM 654 

Archangiopteris bipinnata AM 654 

Archangiopteris henryi AM 654 

Archangiopteris hokouensis AM 654 

Archangiopteris 

subrotundata AM 654 

Calathea sp. AM 510 

Christensenia assamica AM 654 

Ctenanthe sp. AM 510 

Ischnosiphon gracilis AM 510 

Marattia douglasii NM 226 

Monotagma plurispicatum AM 510 

Saranthe compositae NM 510 

Stromanthe porteana AM 510 

Psilotaceae 

Psilotum complanatum NM 226 

Psilotum nudum AM 226 

Psilotum sp. AM 483 

Ophioglossaceae 

Botrychium lanuginosum AM 653 

Botrychium lunaria AM 194, 260, 261 

Botrychium ternatum AM 653 

Botrychium virginianum AM 329

304 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Ophioglossum concinnum AM 226 

Ophioglossum lusitanicum AM 260, 261 

Ophioglossum pendulum AM 226 

Ophioglossum petiolatum AM 654 

NM 226 

Ophioglossum reticulatum AM 522, 654 

Ophioglossum themale AM 654 

Ophioglossum vulgatum AM 653, 260, 261 

Osmundaceae 

Osmunda cinnamomea AM 140 

Osmunda japonica AM 653, 654 

Osmunda regalis AM + NM 260, 261 

Hymenophyllaceae 

Crepidomanes latealatum NM 654 

Hymenophyllum 

M (endo) 260, 261 

tunbrigense 

Hymenophyllum wilsonii M (endo) 260, 261 

Mecodium badium NM 654 

Mecodium blumeanum NM 654 

Mecodium recurvum AM 226 

Sphaerocionium 

AM 226 

lanceolatum 

Trichomanes auriculatum NM 654 

Trichomanes speciosum NM 260, 261 

Vandenboscia cyrtotheca NM 226 

Vandenboscia davalliodes NM 226 

Gleicheniaceae 

Dicranopteris dichotoma AM 653 

Dicranopteris gigantea NM 654 

Dicranopteris linearis AM 226 

Dicranopteris pedata NM 654 

Dicranopteris splendida NM 654 

Diplopterygium glaucoides NM 654 

Diplopterygium glaucum NM 654 

Diplopterygium pinnatum AM 226 

Sticherus laevigatus NM 654 

Sticherus owhyhensis AM 226 

Schizaeaceae 

Anemia phyllitidis AM 31 

Lygodium conforme AM 654 

Lygodium japonicum AM 653 

NM 654 

Schizaea robusta AM 226 

Marsileaceae 

Pilularia globulifera AM + NM 260, 261 

Azollaceae 

Azolla filiculoides NM 226, 260, 261 

Plagiogyriaceae 

Plagiogyria distinctissima AM 654 

Cyatheaceae 

Alsophila constularis NM 654 

Alsophila spinulosa AM 654 

Cyathea cooperi AM 226 

Gymnosphaera gigantea NM 654 

Gymnosphaera podophylla NM 654 

Sphaeropteris brunoniana NM 654 

Dicksoniaceae 

Cibotium barometze NM 654 

Cibotium chamissoi AM 226 

Cibotium glaucum AM 226 

Cibotium st.-johnii AM 226 

Pteridaceae 

Doryopteris decipiens AM 226 

Doryopteris decora AM 226 

Histiopteris incisa AM 654 

Pellaea mairei AM 654 

Pellaea ternifolia AM 226 

Pteridium aquilinum AM 653 

Pteridium revolutum NM 654 

Pteris aspericaulis AM 653 

Pteris aspericaulis var. NM 654 

tricolor 

Pteris cretica AM 226 

Pteris cretica var. laeta NM 654 

Pteris cretica var. nervosa NM 654 

Pteris dissitifolia NM 654 

Pteris ensiformis NM 654 

Pteris esquirolii NM 654 

Pteris excelsa AM 226 

NM 654 

Pteris irregularis AM 226 

Pteris linearis NM 654 

Pteris semipinnata NM 654 

Pteris setulosocostulata AM 654 

Pteris venusta AM 655 

Pteris vittata AM 226, 653 

NM 654 

Pteris wangiana NM 654 

Vittariaceae 

Antrophyrum henryi NM 654 

Vittaria elongata AM 226 

Vittaria flexuosa NM 654 

Adiantaceae 

Adiantum bonatianum AM 654 

Adiantum capillus-veneris AM + NM 260, 261 

Adiantum edgewothii AM 654 

Adiantum flabellulatum AM 654 

Adiantum malesianum NM 654 

Adiantum philipense AM 343 

NM 654 

Aleuritopteris 

AM 654 

albomarginatata 

Aleuritopteris argentea AM 654 

Aleuritopteris duclouxii NM 654 

Aleuritopteris 

pseudofarinosa 

NM 654

305 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Anogramma leptophylla NM 260, 261 

Cheilosoria hancockii AM 654 

Coniogramme intermedia AM 654 

Coniogramme rosthorni NM 654 

Coniogramme simillima AM 654 

Cryptogramma crispa AM + NM 260, 261 

Gymnopteris bipinnata NM 654 

var. auriculata 

Gymnopteris vestita AM 654 

Leptolepidium subvillosum NM 654 

Onychium angustifrons NM 654 

Onychium contigium AM 654 

Onychium japonicum NM 654 

var. lucidum 

Onychium lucidum AM 654 

Sinopteris grevilleoides NM 654 

Dennstaedtiaceae 

Dennstaedtia melanostipes AM 654 

Dennstaedtia scabra AM 654 

Hypolepis punctata NM 654 

Lindsaea cultrata AM 653 

Lindsaea ensifolia NM 654 

Lindsaea javanensis NM 654 

Lindsaea odorata NM 654 

Lindsaea orbiculata AM 654 

Microlepia hookeriana AM 654 

Microlepia marginata AM 654 

Microlepia marginata AM 654 

var. calvescens 

Microlepia pilosissima AM 654 

Microlepia platyphylla NM 654 

Microlepia rhomboidea NM 654 

Microlepia strigosa AM 226 

Monachosorum henryi AM 654 

Pteridium aquilinum AM + NM 260, 261 

Pteridium decompositum AM 226 

Schizoloma heterophyllum NM 653 

Stenoloma chusanum AM 653, 654 

Aspleniaceae 

Acrophorus stipellatus NM 654 

Acystopteris tenuisecta AM 654 

Allantodia alata NM 654 

Allantodia chinensis AM 653 

Allantodia dilatata AM 654 

Allantodia doederleinii NM 654 

Allantodia laxifrons AM 654 

Allantodia megaphylla NM 654 

Allantodia spectabilis AM 654 

Allantodia stenochlamys NM 654 

Asplenium adiantumnigrum 

AM 226 

AM + NM 260, 261 

Asplenium billotii NM 260, 261 

Asplenium cheilosorum AM 654 

Asplenium excisum NM 654 

Asplenium finlaysonianum NM 654 

Asplenium florentinum AM 226 

Asplenium fuscipes NM 654 

Asplenium griffithianum NM 654 

Asplenium horridum NM 226 

Asplenium lushanense AM 654 

Asplenium macraei AM 226 

Asplenium marinum NM 260, 261 

Asplenium nidus NM 226 

Asplenium normale AM 226 

NM 654 

Asplenium onopteris NM 472 

Asplenium pekinense NM 654 

Asplenium praemosum NM 654 

Asplenium prolongatum NM 654 

Asplenium ruta-muraria AM + NM 260, 261 

Asplenium septentrionale NM 260, 261 

Asplenium sphenotomum AM 226 

Asplenium tenuicaule NM 654 

Asplenium trichomanes AM + NM 260, 261 

Asplenium unilaterale AM 226 

NM 654 

Asplenium varians NM 654 

Asplenium viride AM + NM 260, 261 

Asplenium wrightioides NM 654 

Asplenium yunnanense NM 654 

Athyriopsis longipes AM 654 

Athyriopsis ptersennii AM 654 

Athyrium anisopterum NM 654 

Athyrium biserrulatum AM 654 

Athyrium delicatulum NM 654 

Athyrium dissitifolium AM 654 

Athyrium distentifolium AM + NM 260, 261 

Athyrium esculentum AM 226 

Athyrium filix-femina AM + NM 260, 261 

Athyrium japonica AM 226 

Athyrium mackinnonii NM 654 

Athyrium macraei AM 226 

Athyrium mengtzeense AM 654 

Athyrium microphyllum AM 226 

Athyrium niponicum NM 654 

Athyrium sandwichianum AM 226 

Athyrium stigillosum AM 654 

Athyrium wardii AM 653 

Bolbitis heteroclita NM 654 

Bolbitis hokouensis NM 654 

Callipteris esculenta AM 653 

NM 654 

Ceterach officinarum NM 260, 261 

Ctenitis mariformis AM 653 

Ctenitis membranifolia NM 654 

Ctenitopsis devexa NM 654

306 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Ctenitopsis glabra NM 654 

Ctenitopsis sagenioides NM 654 

Ctenitopsis setulaosa NM 654 

Ctenitopsis subsageriacea NM 654 

Cystopteris fragilis AM + NM 260, 261 

Cystopteris montana NM 260, 261 

Cystopteris pellucida AM 653 

Diacalpe christensenae NM 654 

Diplazium donianum AM 653 

NM 654 

Diplazium lanceum AM 653 

Diplazium splendens AM 654 

Dryoathyrium boryanum NM 654 

Dryoathyrium edentulum AM 654 

Egenolfia sinensis AM 654 

Egenolfia tokinensis AM 654 

Gymnocarpium dryopteris AM + NM 260, 261 

Hypodematium crenatum AM 654 

Lomariopsis spectabilis NM 654 

Lunathyrium dolosum NM 654 

Monomelangium pullingeri AM 654 

var. daweishannicolum 

Neottopteris antrophyoides NM 654 

Neottopteris simonsiana NM 654 

Phyllitis scolopendrium AM 260, 261 

Pleocnemia winitei NM 654 

Pseudocystopteris NM 654 

atkinsonii 

Quercifilix zeylanica NM 654 

Tectaria coadunata NM 654 

Tectaria decurrens NM 654 

Tectaria dubia NM 654 

Tectaria hainanensis NM 654 

Tectaria hokouensis NM 654 

Tectaria simonii NM 654 

Tectaria variolosa NM 654 

Tectaria yunnanensis NM 654 

Woodsia alpina NM 260, 261 

Woodsia ilvensis NM 260, 261 

Thelypteridaceae 

Ampelopteris prolifera NM 654 

Dictyocline wilfordii AM 653 

Cyclogramma auriculata AM 654 

Cyclosorus acuminatus NM 654 

Cyclosorus dentatus NM 654 

Cyclosorus hokouensis AM 654 

Cyclosorus mollicesculus NM 654 

Cyclosorus parasiticus NM 654 

Cyclosorus subnigrescens NM 654 

Cyclosorus truncatus NM 654 

Dictyocline griffithii AM 654 

Glaphylopteridopsis 

erubescens 

AM 654 

Macrothelypteris toressiana AM 654 

Metathelypteris flaccida NM 654 

Oreopteris limbosperma AM 260, 261 

Parathelypteris beddomei NM 654 

Parathelypteris hirsutipes AM 654 

Parathelypteris nipponica NM 653 

Phegopteris connectilis AM + NM 260, 261 

Pronephrium 

NM 654 

gymnopteridifrons 

Pronephrium nudotum NM 654 

Pronephrium simplex NM 654 

Pseudocyclosorus AM 654 

esquirolii 

Pseudocyclosorus NM 654 

subochthodes 

Pseudophegopteris NM 654 

pyrrhorachis 

Pseudophegopteris NM 654 

yunkweiensis 

Thelypteris cyatheoides NM 226 

Thelypteris globulifera AM 226 

Thelypteris hudsoniana AM 226 

Thelypteris interrupta NM 226 

Thelypteris parasitica AM 226 

Thelypteris sandwicensis NM 226 

Thelypteris thelypteroides AM + NM 260, 261 

Blechnaceae 

Blechnum occidentale AM 226 

Blechnum orientale AM 654 

Blechnum penna-marina NM 342 

Blechnum spicant AM + NM 260, 261 

Brainea insignis NM 654 

Doodia kunthiana AM 226 

Sadleria cyatheoides AM 226 

Sadleria squarrosa AM 226 

Woodwardia japonica AM 654 

Woodwardia orientalis AM 653 

Woodwardia unigemmata NM 654 

Dryopteridaceae 

Acrorumohra diffraeta AM 654 

Arachniodes festina AM 653 

Arachniodes globisora AM 654 

Arachniodes rhomboidea AM 653 

Arachniodes simplicior AM 653 

Arachniodes sporadosora NM 654 

Cyrtomium caryotideum f. AM 654 

caryotideum 

Cyrtomium fortunei NM 654 

Dryopteris basisora NM 654 

Dryopteris caroli-hopei NM 654 

Dryopteris carthusiana AM 598 

AM + NM 260, 261 

Dryopteris chrysocoma NM 654

307 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Dryopteris cochleata NM 654 

Dryopteris cristata AM 260, 261 

Dryopteris dilatata AM 260, 261 

Dryopteris filix-mas AM + ECM + NM 260, 261 

Dryopteris fructuosa AM 654 

Dryopteris fuscipes AM 653 

Dryopteris fusco-atra AM 226 

Dryopteris glabra AM 226 

Dryopteris lepidopoda NM 654 

Dryopteris marginata AM 654 

Dryopteris odontoloma NM 654 

Dryopteris sparsa AM 654 

Dryopteris stenolepis NM 654 

Dryopteris sublacera AM 654 

Dryopteris thelypteris AM 140 

Dryopteris unidentata AM 226 

Dryopteris wallichiana AM 226 

Nothoperanema 

NM 654 

hendersonii 

Polystichum aculeatum AM + NM 260, 261 

Polystichum alteruatum NM 654 

Polystichum chingae NM 654 

Polystichum dielsii NM 654 

Polystichum eximium NM 654 

Polystichum jizhushanense AM 654 

Polystichum lonchitis AM 260, 261 

Polystichum pycnopterum NM 654 

Polystichum tsus-simense AM 654 

Polystichum vestitum NM 342 

Tectaria gaudichaudii AM 226 

Nephrolepidaceae 

Arthropteris palisotii NM 654 

Nephrolepis biserrata AM 35 

Nephrolepis faleata NM 654 

Polypodiaceae 

Arthromeris mairei NM 654 

Cheilanthes lanosa AM 449 

Colysis diversifolia NM 654 

Colysis hemitoma NM 653 

Colysis hokouensis NM 654 

Colysis pentaphylla NM 654 

Drynaria propinqua NM 654 

Lepidogrammitis rostrama NM 654 

Lepisorus contortus NM 654 

Lepisorus macrosphaerus NM 654 

Lepisorus scolopendrium NM 654 

Microsorium carinatum NM 654 

Microsorium henryi NM 654 

Microsorium 

NM 654 

membranaceum 

Microsorium punctatum NM 654 

Microsorium spectrum NM 226 

Neocheiropteris 

palmatopedata NM 654 

Neolepisorus ovatus NM 654 

Neolepisorus sinensis NM 654 

Onoclea sensibilis AM 140 

Phlebodium aureum NM 226 

Phymatopsis 

NM 654 

crenatopinnata 

Phymatopsis nigrovenia NM 654 

Phymatopsis trisecta NM 654 

Phymatosorus scolopendria NM 226 

Pleopeltis thunbergiana NM 226 

Polypodiodes amoenum NM 654 

Polypodium pellucidum NM 226 

Polypodium vulgare AM + NM 260, 261 

Pyrrosia adnaseens NM 654 

Pyrrosia gralla NM 654 

Pyrrosia lingua NM 654 

Pyrrosia mollis NM 654 

Pyrrosia subfurfuracea NM 654 

Pyrrosia tonkinensis NM 654 

Tricholepidium maculosum NM 654 

Grammitidaceae 

Adenophorus abietinus AM 226 

Adenophorus pinnatifidus NM 226 

Adenophorus tamariscinus AM 226 

Grammitis baldwinii NM 226 

Grammitis poeppigeana NM 342 

Grammitis tenella AM 226 

Loxogramme ensiformis NM 654 

Xiphopteris saffordii NM 226 

Davalliaceae 

Araiostegia perdurans NM 654 

Nephrolepis cordifolia AM 226 

Nephrolepis exaltata AM 226 

Nephrolepis multiflora AM 226 

GYMNOSPERMS 

Cycadaceae 

Cycas circinalis AM 423 

Cycas revoluta AM 423 

Zamiaceae 

Ceratozamia mexicana AM 616 

Dioon edule AM 203, 616 

Lepidozamia hopei AM 484 

Macrozamia reidlei AM 93 

Zamia pumila AM 203 

Ginkgoaceae 

Ginkgo biloba AM 206 

Pinaceae 

Abies alba ECM 189 

Abies grandis ECM 375 

Abies lasiocarpa ECM 314 

Abies spp. ECM 260, 261 

Cedrus atlantica ECM 431

308 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Larix decidua ECM 260, 261 

Larix kaempferi ECM 644, 260, 261 

Larix occidentalis ECM 156 

Picea abies ECM 8 

ECM + EEM 260, 261 

Picea engelmannii ECM 314 

Picea glauca ECM 285, 338, 414, 

470 

Picea glehnii ECM 308 

Picea mariana ECM 91 

Picea rubens ECM 232 

Picea sitchensis ECM 13, 205, 260, 

261 

Pinus banksiana ECM 91 

Pinus cembra ECM 498 

Pinus contorta ECM 98, 156, 403 

Pinus densiflora ECM 228, 309, 634, 

635 

Pinus echinata ECM 444 

Pinus edulis ECM 223, 265, 470 

Pinus elliottii ECM 470, 564 

Pinus halepensis ECM 411, 590 

Pinus jeffreyi ECM 617 

Pinus lambertiana ECM 617 

Pinus massoniana ECM 323, 564 

Pinus montezumae ECM 509 

Pinus muricata ECM 218, 280 

ECM + AM 281 

Pinus nigra ECM 260, 261 

Pinus patula ECM 487 

Pinus pinaster ECM 458, 260, 261 

Pinus pinea ECM 368, 451 

Pinus ponderosa AM 550 

ECM 156, 188, 375, 

506, 618 

Pinus radiata ECM 184 

Pinus resinosa ECM 363, 470 

Pinus rigida ECM 152 

Pinus sylvestris ECM 239, 320, 420 

ECM + EEM + 260, 261 

NM 

Pinus tabulaeformis ECM 634, 635 

Pinus taeda ECM 139, 160, 190 

Pinus thunbergii ECM 364 

Pseudotsuga menziesii ECM 70, 156, 236, 

259, 280, 375 

AM 550 

AM + ECM 116 

ECM + EEM 260, 261 

Tsuga heterophylla ECM 9, 313 

AM + ECM 116 

Ephedraceae 

Ephedra trifurca AM 137 

Welwitschiaceae 

Welwitschia mirabilis AM 296 

Gnetaceae 

Gnetum africanum ECM 445 

Gnetum buchholzianum ECM 445 

Gnetum sp. AM + ECM 445 

Araucariaceae 

Agathis robusta AM 388 

Araucaria angustifolia AM 31, 89, 650 

Araucaria cunninghamii AM 388 

Wollemia nobilis AM + EEM 388 

Wollemia sp. ECM 656 

Podocarpaceae 

Dacrycarpus dacrydioidies AM 504 

Dacrydium cupressinum AM 504 

Parasitaxus ustus Mycoheterotrophy 659 

via AM? 

Podocarpus falcatus AM 637, 638 

Prumnopitys ferruginea AM 504 

Prumnopitys taxifolia AM 504 

Taxodiaceae 

Metasequoia 

AM 559 

glyptostroboides 

Sequoia sempervirens AM 6 

Sequoiadendrom 

AM 6 

giganteum 

Cupressaceae 

Austrocedrus chilensis AM 207 

Chamaecyparis thyoides AM 106 

Juniperus communis AM + ECM 260, 261 

Juniperus communis ssp. AM + ECM 260, 261 

nana 

Juniperus monosperma AM 470 

Juniperus oxycedrusL. ssp. AM 368 

macrocarpa 

Juniperus procera AM 637, 638 

Libocedrous decurrens AM 6, 25 

Tetraclinis articulata AM 169, 412 

Thuja occidentalis AM 81, 385 

Thuja plicata AM 100 

Taxaceae 

Taxus baccata AM 637, 638, 260, 

261 

Taxus × media AM 515 

ANGIOSPERMS 

Nymphaeaceae 

Nuphar × intermedia NM 260, 261 

Nuphar lutea NM 62, 260, 261 

Nuphar pumila NM 260, 261 

Nymphaea alba NM 260, 261

309 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Nymphaea sp. AM 656 

Acoraceae 

Acorus calamus AM 531 

NM 260, 261 

Tofieldiaceae 

Tofieldia pusilla AM 260, 261 

Araceae 

Aglaonema modestum AM 655 

Alocasia longiloba AM 655 

Alocasis macrorrhiza AM 422 

Amorphophallus 

AM 422 

bannaensis 

Anthurium affine AM 510 

Anthurium pentaphyllum AM 510 

Anthurium pitteri AM 474 

Arum italicum AM 260, 261 

Arum maculatum AM 260, 261 

Arum neglectum AM 260, 261 

Caladium bicolor AM 510 

Calla palustris NM 260, 261 

Colacasia esculenta AM 531 

Dieffenbachia amoena AM 510 

Philodendron ornatum NM 510 

Philodendron undulatum NM 510 

Pistia stratiotes NM 510 

Rhaphidophora decursiva AM 422 

Hydrocharitaceae 

Egreria densa NM 510 

Elodea canadensis NM 62, 260, 261 

Hydrilla verticillata NM 260, 261 

Vallisneria americana AM 631, 632 

Butomaceae 

Butomus umbellatus NM 260, 261 

Alismataceae 

Alisma lanceolatum AM + NM 260, 261 

Alisma plantago-aquatica AM 58 

NM 62, 260, 261 

Alisma subcordatum Weak AM 626 

Alisma triviale AM 140 

Baldellia ranunculoides AM 368 

Echinodorus grandiflorus NM 510 

Sagitaria latifolia AM 626 

Limnocharitaceae 

Hydrocleys nymphoides 

Juncaginaceae 

Triglochin maritima AM + NM 260, 261 

Triglochin palustris NM 260, 261 

Potamogetonaceae 

Potamogeton crispus NM 62, 260, 261 

Potamogeton gramineus NM 62 

Potamogeton lucens NM 62 

Potamageton natans AM 58 

NM 62, 260, 261 

Potamogeton perfoliatus NM 62 

Potamogeton 

NM 260, 261 

polygonifolius 

Potamogeton praelongus NM 62 

Petrosaviaceae 

Petrosavia sakuraii Mycoheterotrophy 657, 658 

Petrosavia stellaris Mycoheterotrophy 657, 658 

Nartheciaceae 

Narthecium ossifragum AM + NM 260, 261 

Burmanniaceae 

Burmannia tenella Mycoheterotrophy 290 

(via AM) 

Thismia sp. Mycoheterotrophy 656 

Dioscoreaceae 

Tacca chantrieri NM 655 

Tamus communis AM 260, 261 

Triuridaceae 

Sciaphila polygyna Mycoheterotrophy 291 

Sciaphila tosaensis Mycoheterotrophy 642 

(via AM) 

Pandanaceae 

Freycinetia arborea AM 326 

Pandanus furcatus NM 655 

Pandanus tectorius NM 326 

Cyclanthaceae 

Asplundia gardneri NM 510 

Corsiaceae 

Arachnitis uniflora Mycoheterotrophy 79 

(via AM) 

Alstroemeriaceae 

Alstroemeria aurea AM 207 

Colchicaceae 

Colchicum autumnale AM 260, 261 

Melanthiaceae 

Paris quadrifolia AM + NM 260, 261 

Trillium flexipes AM 165 

Veratrum viride AM 140 

Smilacaceae 

Smilax aspera AM 63, 368 

Smilax corbularia AM 422 

Smilax hypoglauca AM 422 

Smilax indica AM 422 

Liliaceae 

Gagea lutea AM + NM 260, 261 

Lilium martagon AM 260, 261 

Muscari comosum AM 260, 261 

Muscari neglectum AM 260, 261 

Smilacina racemosa AM 165 

Orchidaceae 

Aceras anthropophorum ORM 260, 261 

Anacamptis pyramidalis ORM 260, 261 

Anoectochilus burmannicus ORM 422 

Anoectochilus roxburghii ORM 422

310 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Anoectochilus tortus AM 655 

Apostasia odorata AM 422 

Arundina graminifolia ORM 326 

Caladenia formosa ORM 286 

Calypso bulbosa ORM 153 

Cephalanthera austinae Mycoheterotrophy 569 

(via ECM) 

Cephalanthera 

ORM + ECM 78 

damasonium 

ORM 260, 261 

Cephalanthera longifolia ORM 260, 261 

Cephalanthera rubra ORM + ECM 78 

ORM 368, 260, 261 

Chloraea virescens ORM 207 

Coeloglossum viride ORM 260, 261 

Corallorhiza maculata Mycoheterotrophy 569, 570, 571 

(via ECM) 

Corallorhiza mertensiana Mycoheterotrophy 570 

Corallorhiza trifida Mycoheterotrophy 392, 260, 261 

Corybas dienemus ORM 342 

Cypripedium calceolus ORM 540 

ORM + NM 260, 261 

Cypripedium californicum ORM 540 

Cypripedium candidum ORM 540 

Cypripedium fasciculatum ORM 540 

Cypripedium guttatum ORM 540 

Cypripedium montanum ORM 540 

Cypripedium parviflorum ORM 540 

Dactylorhiza fuchsia ORM 260, 261 

Dactylorhiza incarnara ORM 173, 260, 261 

Dactylorhiza maculata ORM 260, 261 

Dactylorhiza maculata ssp. ORM 260, 261 

ericetorum 

Dactylorhiza majalis ORM 78, 334, 480, 

260, 261 

Dactylorhiza praetermissa ORM 260, 261 

Dactylorhiza praetermissa ORM 173 

var. junialis 

Dactylorhiza purpurella ORM 260, 261 

Epipactis atrorubens ORM + ECM 78 

ORM 304, 260, 261 

Epipactis distans ORM + ECM 78 

Epipactis helleborine ORM + ECM 78 

ORM 260, 261 

Epipactis microphylla ORM + ECM 530 

Epipactis palustris ORM 78, 481 

ORM + NM 260, 261 

Epipactis purpurata ORM 260, 261 

Epipogium aphyllum ORM 260, 261 

Goodyera repens ORM 260, 261 

Gymnadenia conopsea ORM 260, 261 

Gymnadenia odoratissima ORM 260, 261 

Haemaria discolor 

var. dawsoniana 

ORM 122 

Hammarbya paludosa ORM 260, 261 

Herminium monorchis ORM 260, 261 

Hexalectris spicata Mycoheterotrophy 572 

(via ECM) 

Himantoglossum hircinum ORM 260, 261 

Ionopsis utricularioides ORM 448 

Liparis loeselii ORM 260, 261 

Listera cordata ORM 260, 261 

Listera ovata ORM 260, 261 

Malaxis latifolia ORM 422 

Microtis parviflora ORM 459 

Neotinea maculata ORM 260, 261 

Neottia nidus-avis ORM + ECM 78 

ORM 260, 261 

Newiedia veratrifolia ORM 333 

Ophrys apifera ORM 260, 261 

Ophrys fuciflora ORM 260, 261 

Ophrys insectifera ORM 260, 261 

Ophrys lutea ORM 56 

Ophrys sphegodes ORM 260, 261 

Orchis mascula ORM 260, 261 

Orchis militaris ORM 260, 261 

Orchis morio ORM 75, 173, 260, 

261 

Orchis palustris ORM 195 

Orchis purpurea ORM 260, 261 

Orchis simia ORM 260, 261 

Orchis ustulata ORM 260, 261 

Phaius mishmensis AM 655 

Platanthera bifolia ORM 260, 261 

Platanthera chlorantha ORM 78, 260, 261 

Platanthera leucophaea ORM 652 

Pseudorchis albida ORM 260, 261 

Pterostylis acuminata ORM 460 

Serapias vomeracea ssp. ORM 195 

vomeracea 

Spathoglottis plicata ORM 326, 534, 565 

Spiranthes aestivalis ORM 260, 261 

Spiranthes sinensis ORM 376 

var. amoena 

Spiranthes spiralis ORM 368, 260, 261 

Stanhopea tigrina ORM 196 

Tolumnia variegata ORM 448 

Asteliaceae 

Astelia menziesiana AM 326 

Hypoxidaceae 

Curculigo capitulata AM 422 

Iridaceae 

Crocus vernus AM 260, 261 

Gladiolus segetum AM 260, 261 

Iris foetidissima AM 260, 261 

Iris germanica AM 260, 261

311 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Iris pseudacorus AM 368 

AM + NM 260, 261 

Sisyrinchium atlanticum AM 643 

Sisyrinchium sp. NM 207 

Sparaxis tricolor AM 514 

Hemerocallidaceae 

Dianella sandwicensis AM 326 

Asphodelaceae 

Asphodelus fistulosus AM 114, 115 

Bulbine alata AM 438 

Haworthia tortuosa AM 417 

Alliaceae 

Allium ampeloprasum AM 260, 261 

Allium canadense AM 140 

Allium cepa AM 7, 349, 260, 

261 

Allium oleraceum AM + NM 260, 261 

Allium porrum AM 272, 260, 261 

Allium sativum AM 260, 261 

Allium schoenoprasum AM 3, 260, 261 

Allium scorodoprasum AM 260, 261 

Allium sphaerocephalon AM 472, 260, 261 

Allium subhirsutum AM 472 

Allium triquetrum AM 260, 261 

Allium ursinum AM 260, 261 

Amaryllidaceae 

Galanthus nivalis AM 260, 261 

Leucojum aestivum AM 260, 261 

Leucojum vernum AM + NM 260, 261 

Narcissus poeticus AM 260, 261 

Narcissus pseudonarcissus AM 260, 261 

Themidaceae 

Brodiaea laxa AM 513 

Hyacinthaceae 

Hyacinthoides non-scripta AM 398, 399, 260, 

261 

Ornithogalum nutans AM 260, 261 

Ornithogalum pyrenaicum AM 260, 261 

Ornithogalum umbellatum AM 260, 261 

Scilla autumnalis AM 260, 261 

Agavaceae 

Agave americana AM 343 

Agave datylio Weak AM 112 

Agave deserti AM 151 

Agave marmorata AM 103 

Agave salmiana Weak AM 103 

Agave univittata AM 417 

Yucca elata AM 137 

Yucca periculosa Weak AM 103 

Yucca valida AM 417 

Asparagaceae 

Asparagus acutifolius AM 368, 472 

Asparagus officinalis AM 192, 383, 260, 

261 

Ruscaceae 

Convallaria majalis AM + NM 260, 261 

Maianthemum bifolium AM + NM 260, 261 

Polygonatum multiflorum AM + NM 260, 261 

Polygonatum odoratum AM + NM 260, 261 

Polygonatum verticillatum AM + NM 260, 261 

Ruscus aculeatus AM 368, 260, 261 

Arecaceae 

Astrocaryum mexicanum AM 436 

Bactris gasipaes AM 135 

Caryota monostachya AM 655 

Cocos nucifera AM 326, 580 

Euterpe edulis AM 651 

Euterpe oleracea AM 130 

Serenoa repens AM 201 

Syagrus romanzoffiana Weak AM 651 

Wallichia mooreana NM 655 

Sparganiaceae 

Sparganium angustifolium NM 62 

Sparganium chlorocarpum AM 58 

Sparganium emersum NM 62 

Sparganium erectum NM 62 

AM 260, 261 

Sparganium eurycarpum Weak AM 626 

Typhaceae 

Typha angustifolia NM 62, 368 

Weak AM 557 

AM 566, 260, 261 

Typha latifolia NM 62 

Weak AM 557 

AM 140, 185, 599 

Typha × glauca Weak AM 557 

Bromeliaceae 

Aechmea stelligera NM 510 

Ananas comosus AM 44, 246 

Hechtia aff. podantha Weak AM 103 

Vriesea limae AM 510 

Juncaceae 

Juncus acutiflorus AM + NM 260, 261 

Juncus alpinus AM + NM 260, 261 

Juncus articulatus AM 260, 261 

Juncus biglumis NM 260, 261 

Juncus bufonius AM + NM 260, 261 

Juncus bulbosus NM 62, 260, 261 

Juncus dudleyi AM 599 

Juncus effuses NM 62 

AM 140 

AM + NM 260, 261 

Juncus filiformis Weak AM 194 

NM 260, 261 

Juncus gerardi AM + NM 260, 261

312 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Juncus inflexus AM 260, 261 

Juncus maritimus NM 368, 260, 261 

Juncus nodosus AM 599 

Juncus planifolius AM 326 

Juncus roemerianus AM 278 

Juncus scheuchzerioides AM 342 

NM 560 

Juncus squarrosus AM + NM 260, 261 

Juncus torreyi AM 599 

Juncus trifidus AM 260, 261 

Luzula campestris AM + NM 260, 261 

Luzula crinita var. crinita NM 342 

Luzula hawaiiensis AM 326 

Luzula luzuloides NM 260, 261 

Luzula pallescens NM 260, 261 

Luzula pilosa AM + NM 260, 261 

Luzula spicata AM 260, 261 

Luzula sylvatica NM 260, 261 

Cyperaceae 

Bequerelia cymosa AM 426 

Bulboschoneus maritimus AM 426 

Bulbostylis barbata Facultative AM 426 

Bulbostylis capillaris NM 326 

AM 149 

Facultative AM 426 

Bulbostylis cf. conifera AM 358 

Bulbostylis densa AM 426 

Bulbostylis paradoxa AM 358 

Bulbostylis puberula NM 426 

Carex acnescens NM 426 

Carex acuta NM 62, 260, 261 

Carex acutiformis NM 260, 261 

Carex albonigra NM 426 

Carex amphibola NM 402 

Carex annectens AM 402 

Carex aphylla NM 207 

Carex aquatilis NM 426 

Carex arenaria AM + NM 260, 261 

Carex atherodes NM 402 


Carex baccans AM 426 

Carex bicknellii AM 402 

Carex bigelowii NM 501, 594, 426 

Carex blanda AM 402 

Carex boelckeiana NM 207 

Carex brevior AM 402 

Carex brizoides NM 426 

Carex brunnescens AM 194 

Carex buxbaumii AM 402 

NM 260, 261 

Carex caryophyllea NM 456, 260, 261 

Carex cephalophora AM 402 

Carex chordorrhiza NM 260, 261 

Carex crawei AM 402 

Carex cristatella AM 58, 402 

Carex curta NM 260, 261 

Carex davalliana AM 260, 261 

Carex digitata NM 260, 261 

Carex dioica NM 260, 261 

Carex distans NM 260, 261 

Carex disticha AM + NM 260, 261 

Carex divulsa AM 260, 261 

Carex ebenea NM 426 

Carex echinata NM 260, 261 

Carex elongata NM 260, 261 

Carex ericetorum NM 456, 260, 261 

Carex fillifolia NM 426 

Carex flacca NM 426 

AM + NM 260, 261 

Carex flava AM 140 

NM 426 

AM + NM 260, 261 

Carex fuscula AM 426 

Carex gayana NM 426 

Carex grannularis AM 402, 599 

Carex gravida AM 402 

Carex hallerana NM 472 

Carex hirta AM 456 

NM 260, 261 

Carex hostiana NM 260, 261 

Carex humilis NM 260, 261 

Carex hystericina NM 426 

Carex interior NM 402 

Carex lachenalii NM 426, 260, 261 

Carex lanuginosa NM 599 

Carex lasiocarpa AM 58 

Weak AM 626 


NM 426, 260, 261 

Carex lepidocarpa NM 260, 261 

Carex limosa NM 260, 261 

Carex lindleyana AM 426 

Carex lurida AM 58, 140 

Carex madoviana NM 426 

Carex maritime NM 426 

Carex membranacea NM 426 

Carex mertensii NM 584 

Carex meyenii AM 326 

Carex microchaeta NM 594 

Carex misandra NM 426 

Carex montana NM 260, 261 

Carex muricata NM 426 

AM + NM 260, 261 

Carex myosurus AM 426

313 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Carex nardina NM 426 

Carex nigra AM 140, 194 

AM + NM 260, 261 

Carex otrubae NM 260, 261 

Carex oxyandra NM 587 

Carex pallescens AM 194 

NM 260, 261 

Carex panacea AM 194 

Carex panicea AM + NM 260, 261 

Carex paniculata NM 260, 261 

Carex pellita NM 402 

Carex pendula NM 368, 260, 261 

Carex pensylvanica AM 402 

Carex pilulifera NM 260, 261 

Carex podocarpa NM 594 

Carex pulicaris NM 260, 261 

Carex remota AM + NM 260, 261 

Carex riparia NM 260, 261 

Carex rosea AM 402 

Carex rostrata NM 260, 261 

Carex scoparia AM 402 

NM 140 

Carex serotina NM 260, 261 

Carex speciosa AM 426 

Carex sprengelii NM 402 

Carex stenophylla ssp. NM 426 

eleocharis 

Carex sterilis NM 426 

Carex stipata AM 140, 402 

Carex stricta NM 140, 402 

AM 626 


Carex aff. subantarticata NM 426 

Carex subspathecea NM 426 

Carex sylvatica NM 426, 260, 261 

Carex tenera NM 402 

Carex tetanica AM 402 

Carex tribuloides AM 58, 140 

Carex trichocarpa AM 599 

Carex trifida NM 342 

Carex ursine NM 426 

Carex urticulata NM 426 

Carex vaginata NM 503, 260, 261 

Carex vesicaria AM 626 

AM + NM 260, 261 

Carex vulpina NM 260, 261 

Carex vulpinoidea AM 58, 140, 402 

Carex wahuensis AM 326 

NM 426 

Caustis dioica NM 426 

Caustis flexuosa AM 67 

Cladium amaicense AM 426 

Cladium jamaicense AM 297 

Cladium mariscus NM 368 

AM + NM 260, 261 

Cyathochaeta avenccea NM 426 

Cyperus arenarius AM 426 

Cyperus articulatus Weak AM 143 


Cyperus brevifolius AM 426 

Cyperus bulbosus NM 426 

Cyperus castaneous NM 426 

Cyperus clarkei AM 426 

Cyperus compressus Facultative AM 426 

Cyperus cyperinus AM 426 

Cyperus decompositus NM 426 

Cyperus difformis Facultative AM 426 

Cyperus distans Facultative AM 426 

Cyperus dubius AM 426 

Cyperus esculentus NM 426 

Cyperus exaltatus NM 426 

Cyperus flavescens AM 599 

Cyperus halpan Facultative AM 426 

NM 426 

Cyperus iria AM 424, 426 

Cyperus javanicus NM 426 

Cyperus kyllinga Facultative AM 355, 426 

Cyperus laevigatus AM 426 

Cyperus ligularis AM 426 


Cyperus luzulae NM 426 

Cyperus microiria NM 643 

Cyperus niveus AM 343 

Cyperus nutans AM 426 

Cyperus odoratus AM 426 

Cyperus paniceus AM 426 

Cyperus pilosus AM 426 

Cyperus platyphyllus AM 426 

Cyperus pohlii NM 426 

Cyperus pygmaeus AM 426 

Cyperus rotundus NM 326 

AM 343, 424, 425 


Cyperus squarrosus AM 426 

Cyperus stoloniferous AM 426 

Cyperus strigosus AM 599 

NM 426 

Cyperus surinamensis Facultative AM 426 

Cyperus tenuispica NM 426 

Cyperus triceps AM 426 

Eleocharis acutangula AM 426 

Eleocharis dulcis AM 426 

Eleocharis erythropoda NM 599 

Eleocharis geniculata AM 426 

NM 426 

Eleocharis multicaulis NM 260, 261

314 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Eleocharis ovata AM 140 

Eleocharis aff. pachycarpa NM 426 

Eleocharis palustris NM 62 

AM + NM 260, 261 

Eleocharis quinqueflora NM 260, 261 

Eleocharis scheuchezeri NM 426 

Eleocharis tenius NM 426 

Eleocharis triste NM 426 

Eleocharis uniglumis NM 260, 261 

Eleocharis vaginatum NM 426 

Eriophorum angustifolium AM + NM 260, 261 

Eriophorum latifolium NM 260, 261 

Eriophorum vaginatum AM + NM 260, 261 

Fimbristylis consanguinea AM 426 

Fimbristylis cymosa NM 326, 355 

Fimbristylis eragrostis AM 426 

Fimbristylis falcata Facultative AM 426 

Fimbristylis miliacea Facultative AM 426 

Fimbristylis ovata Facultative AM 426 

Fimbristylis schoenoides NM 426 

Fimbristylis trachycarya NM 426 

Fimbristylis triflora NM 426 

Frimbristyllis complanata NM 149 

Fuirena ciliaris NM 426 

Gahnia gahiniformis NM 426 

Gahnia vitiensis AM 326 

Gahnia vitiensis 

AM 326 

ssp. kauaiensis 

Hypolytrum bullatum AM 426 

Hypolytrum pulchrum AM 149, 358 

Isolepis antartica NM 426 

Isolepis aucklandica NM 342 

Isolepis nodosa AM 426 

Kobresia bellardii AM 426 

Kobresia myosuroides Facultative AM 426 

ECM 656 

Kobresia simpliciuscula NM 426, 260, 261 

Kyllinga brevifolia NM 422, 426 

Kyllinga bulbosa NM 426 

Kyllinga nemoralis NM 426 

Lagenocarpus guianensis AM 358 

Lagenocarpus sphacelata NM 426 

Lepidosperma gracile AM 397 

Machaerina augustifolia NM 326 

Machaerina mariscoides AM 326 

Mariscus dubius NM 426 

Mariscus mariscoides AM 426 

ssp. meyenii 

Mariscus meyenianus AM 326 

Mariscus paniceus NM 426 

Mariscus squarrosus NM 426 

Mesomelaena pseudostygia NM 426 

Oreobolus furcatus AM 326 

Pleurostachys cephalotes Facultative AM 426 

Pycreus flavidus AM 426 

Pycreus polystachyos Facultative AM 426 

AM 326 

Pycreus pumilus AM 426 

Pycreus puncticulatus NM 426 

Rhynchospora barbata AM 149, 358 

Rhynchospora brasiliensis AM 358 

Rhynchospora cephalotes Facultative AM 426 

Rhynchospora ciliata NM 426 

Rhynchospora cormbosa AM 426 

Rhynchospora longisetis NM 426 

Rhynchospora pubera AM 426 

Rikliella squarrosa AM 426 

Schoenoplectus grossus NM 426 

Schoenoplectus juncoides NM 426 

Schoenoplectus 

NM 426 

senegalensis 

Schoenoplectus supinus AM 426 

Schoenus ferrugineus AM + NM 260, 261 

Schoenus nigricans AM + NM 260, 261 

Scirpus acutus AM 58, 626 

Scirpus atrovirens AM 58, 140, 599, 

626 

Scirpus cespitosus NM 426 

Scirpus cyperinus AM 58, 140 

Scirpus fluviatilis AM 426 

Scirpus holoschoenus NM 368 

Scirpus maritimus AM 626 

NM 260, 261 

Scirpus pendulus AM 599 

Scirpus aff. perpusillus NM 426 

Scirpus pungens AM 599 

Scirpus robustus AM 426 

Scirpus sylvaticus NM 260, 261 

Scirpus tabernaemontani AM 599 

Scirpus validus AM 58 

Scleria latifolia Facultative AM 426 

Scleria lithosperma AM 426 

Scleria melaleuca Facultative AM 426 

Tetraria capillaris AM 397 

Trichophorum alpinum NM 260, 261 

Trichophorum cespitosum AM + NM 260, 261 

Uncinia richleriana NM 207 

Uncinia uncinata NM 326 

Restionaceae 

Alexgeorgea nitens AM 397 

Lyginia barbata AM 397 

Poaceae 

Aeluropus littoralis AM 620 

var. sinensis 

Agropyron desertorum AM 183 

Agropyron repens AM 307

315 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Agropyron smithii AM 354 

Agropyron trachycaulum AM 132 

Agropyron tsukushiense AM 643 

var. pransiens 

Agropyron tsukushiense AM 294 

var. transiens 

Agrostis alba AM 140 

Agrostis canina AM + NM 260, 261 

Agrostis capillaris AM 242 

AM + NM 260, 261 

Agrostis gigantea AM 260, 261 

Agrostis magellanica AM 560 

NM 342 

Agrostis palustris AM 126 

Agrostis scabra AM 95, 587 

Agrostis stolonifera AM 456, 467 

AM + NM 260, 261 

Agrostis stolonifera NM 307 

ssp. stolonifera 

Aira caryophyllea NM 207 

AM + NM 260, 261 

Aira praecox AM + NM 260, 261 

Alopecurus aequalis NM 260, 261 

Alopecurus geniculatus AM 260, 261 

Alopecurus myosuroides AM 260, 261 

Alopecurus pratensis AM 260, 261 

Ammocalamagrostis baltica AM + NM 260, 261 

Ammophila arenaria AM 331, 368 

AM + NM 260, 261 

Ammophila breviligulata AM 224, 225, 324, 

352 

Ampelodesmos 

AM 472 

mauritanicus 

Andropogon capillipes AM 415 

Andropogon gerardii Facultative AM 29 

AM 168, 263, 276, 

524 

Andropogon gerardii AM 88 

var. paucipilus 

Andropogon virginicus AM 326, 415 

Anthoxanthum odoratum AM 194 

AM + NM 260, 261 

Apera spica-venti AM 260, 261 

Aristida adscensionis AM 479 

Aristida contorta Weak AM 438 

Aristida holathera NM 438 

Aristida longiseta AM 604 

Aristida mutabilis AM 567 

Aristida aff. romeriana AM 143 

Aristida stricta NM 30, 415 

Arrhenatherum elatius AM + NM 260, 261 

Avena barbata AM 472 

Avena fatua AM 260, 261 

Avena sativa AM 15 

AM + NM 260, 261 

Avena strigosa AM 260, 261 

Avenula pratensis AM 260, 261 

Avenula pubescens AM 260, 261 

Axonopus pruinosus AM 358 

Beckmannia syzigachne AM 294 

Bothriochloa pertusa AM 343 

Bouteloua eripoda AM 137 

Boutelous repens AM 143 

Brachiaria decumbens AM 150, 358 

Brachiaria humidicola AM 150, 358 

Brachypodium pinnatum AM + NM 260, 261 

Brachypodium ramosum AM 472 

Brachypodium sylvaticum AM 260, 261 

Briza maxima AM 472, 260, 261 

Briza media AM + NM 260, 261 

Briza minor AM 643, 260, 261 

Bromus erectus AM 260, 261 

Bromus hordeaceus AM 493 

AM + NM 260, 261 

Bromus inermis AM 260, 261 

Bromus madritensis AM 646 

ssp. rubens 

Bromus sterilis NM 260, 261 

Bromus tectorum Facultative AM 237 

Calamagrostis canescens AM 260, 261 

Calamagrostis epigejos AM 307 

AM + NM 260, 261 

Calamagrostis stricta AM 260, 261 

Calamagrostis villosa AM 54, 366, 615 

Calamovilfa longifolia AM 88 

Catapodium rigidum AM 472 

Cenchrus ciliaris AM 20 

Cenchrus setigerus AM 567 

Chusquea culeou AM 207 

Cinna arundinacea AM 140 

Corynephorus canescens AM 81 

AM + NM 260, 261 

Cymbopogon jwarancusa AM 567 

Cymbopogon winterianus AM 327 

Cynodon dactylon AM 294, 260, 261 

Cynosurus cristatus AM 260, 261 

Dactylis glomerata AM 497 

AM + NM 260, 261 

Danthonia decumbens AM 194, 260, 261 

Dendrocalamus asper AM 608 

Dendrocalamus strictus AM 482 

Deschampsia antarctica Facultative AM 166 

NM 560 

Deschampsia caespitosa NM 342

316 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Deschampsia cespitosa AM 194, 293, 260, 

261 

Deschampsia chapmanii AM 342 

Deschampsia flexuosa AM 194, 270, 366, 

615, 260, 261 

Desmazeria rigida AM 260, 261 

Dichanthelium cynodon AM 326 

Dichanthelium 

AM 95 

lanuginosum 

Digitaria adscendens AM 294 

Digitaria chinensis AM 422 

Digitaria ciliaris Weak AM 129 

AM 643 

Digitaria eriantha AM 23 

Digitaria sanguinalis AM 260, 261 

Digitaria violascens AM 422 

Distichlis spicata AM 278 

Distichlis stricta AM 302 

Echinochloa crus-galli AM 140 

AM + NM 260, 261 

Echinolaena inflexa AM 358 

Eleusine coracana AM 576 

Eleusine indica AM 422 

Elymus canadensis AM 263, 274 

Elymus caninus AM 260, 261 

Elymus farctus AM + NM 260, 261 

Elymus pycnanthus AM + NM 260, 261 

Elymus repens AM 260, 261 

Enneapogon avenaceus AM 438 

Eragrostis dielsii Facultative AM 438 

Eragrostis ferruginea AM 643 

Eriachne aristidea Weak AM 438 

Festuca altissima NM 260, 261 

Festuca argentina AM 207 

Festuca arundinacea AM 275, 260, 261 

Festuca baffinensis AM 158 

Festuca brachyphylla AM 158 

Festuca brigantina AM 235 

Festuca contracta NM 342 

Festuca erecta AM 560 

Festuca gigantea NM 260, 261 

Festuca hyperborea NM 158 

Festuca idahoensis AM 109, 371 

Festuca juncifolia AM 260, 261 

Festuca lemanii AM 260, 261 

Festuca ovina AM 194, 456 

AM + NM 260, 261 

Festuca pallescens AM 207 

Festuca pratensis AM 194 

AM + NM 260, 261 

Festuca pumila AM 80 

Festuca rubra AM + ECM + NM 260, 261 

Festuca vivipara AM 260, 261 

Glyceria fluitans NM 260, 261 

Glyceria maxima NM 260, 261 

Glyceria plicata AM + NM 260, 261 

Heteropogon contortus AM 326, 343 

Hierochloe odorata AM 599, 260, 261 

Holcus lanatus AM 395, 586 

AM + NM 260, 261 

Holcus mollis AM 194, 260, 261 

Homolepsis aturensis AM 142 

Hordeum brachyantherum AM 293 

Hordeum comosum AM 207 

Hordeum distichon AM 260, 261 

Hordeum jubatum AM 302, 626 

Hordeum murinum AM 260, 261 

Hordeum vulgare AM 119 

AM + NM 260, 261 

Imperata cylindrica AM 422 

Ischaemum byronis AM 326 

Koeleria glauca NM 260, 261 

Koeleria macrantha AM 260, 261 

Koeleria pyramidata NM 274 

Koeleria vurilochensis NM 207 

Lagurus ovatus AM 260, 261 

Lasiurus sindicus AM 567 

Leersia hexandra AM 404 

Leersia oryzoides AM 58, 140 

NM 260, 261 

Leymus arenarius AM 240, 241 

NM 260, 261 

Lolium perenne AM 23, 140, 171, 

307 

Lolium perenne 

AM 260, 261 

ssp. multiflorum 

Lolium perenne 

AM 260, 261 

ssp. perenne 

Lolium temulentum AM + NM 260, 261 

Lygeum spartum AM 170 

Melica nutans AM + NM 260, 261 

Melica uniflora NM 260, 261 

Microstegium ciliatum AM 422 

Milium effusum AM + NM 260, 261 

Molinia caerulea AM 586, 260, 261 

Mulenbergia porteri AM 137 

Nardus stricta AM 194, 269, 260, 

261 

Nassella leucotricha AM 604 

Neyraudia reynaudiana AM 343 

Oryza sativa AM 536 

Oryzopsis hymenoides AM 11 

Palicourea rigida AM 358 

Panicum amarum AM 43

317 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Panicum 

AM 58 

clandestinum 

Panicum hemitomon AM 404 

Panicum laxum AM 149 

Panicum maximum AM 20 

Panicum micranthum AM 358 

Panicum nephelophilum AM 326 

Panicum purpurascens AM 143 

Panicum virgatum AM 83, 134 

Weak AM 626 

Panicum virginatum AM 193 

Paractaenum 

NM 438 

novae-hollandiae 

Parapholis incurva AM 260, 261 

Paspalidium flavidum AM 343 

Paspalum cf. carinatum AM 358 

Paspalum conjugatum AM 142 

Paspalum dilatatum AM 243 

Paspalum notatum AM 193, 295, 562 

Paspalum thunbergii AM 643 

Pennisetum alopecuroides AM 643 

Pennisetum glaucum AM 51, 567 

Pennisetum padicillatum AM 544 

Pennisetum parviflorum AM 545 

Pennisetum setaceum AM 326 

Phalaris arundinacea AM 58, 140 

NM 62 

AM + NM 260, 261 

Phleum alpinum AM 194, 260, 261 

Phleum arenarium NM 260, 261 

Phleum phleoides AM + NM 260, 261 

Phleum pratense AM 132, 194, 260, 

261 

Phragmites australis NM 62 

AM 140, 368 

AM + NM 260, 261 

Phragmites communis AM 620 

Piptatherum miliaceum AM 497 

Poa alpina AM + NM 260, 261 

Poa angustifolia AM 307 

Poa annua AM 560, 643 

NM 342 

AM + NM 260, 261 

Poa arctica NM 594 

Poa compressa NM 307 

AM + NM 260, 261 

Poa cookie NM 560 

AM 342 

Poa flexuosa AM + NM 260, 261 

Poa foliosa NM 342 

Poa kerguelensis AM 560 

Poa lanuginosa AM 207 

Poa literosa NM 342 

Poa nemoralis AM + NM 260, 261 

Poa palustris AM + NM 260, 261 

Poa pratensis AM 194, 467, 626 

AM + NM 260, 261 

Poa trivialis AM + NM 260, 261 

Polypogon monspeliensis AM 260, 261 

Puccinellia distans AM + NM 260, 261 

Puccinellia macquariensis AM 342 

Puccinellia maritima AM + NM 260, 261 

Puccinellia nuttalliana AM 301 

Raddiella esembeckii AM 358 

Rhynchelytrum repens AM 355 

Saccharum officinarum AM 326, 489 

Saccharum spontaneum Weak AM 439 

Schizachyrium scoparium Facultative AM 29 

AM 117, 140 

Schizachyrium stoloniferum AM 415 

Secale cereale AM 512 

AM + NM 260, 261 

Sesleria albicans AM + NM 260, 261 

Setaria glauca AM 643 

Setaria pumila AM 260, 261 

Setaria verticillata AM 260, 261 

Setaria viridis AM 643 

AM + NM 260, 261 

Sorghastrum nutans AM 264 

Sorghum bicolor AM 171 

Sorghum halpense AM 193 

Sorghum sudanense AM 311 

Spartina alterniflora NM 278 

Weak AM 391 

Spartina anglica NM 260, 261 

Spartina cynosuroides AM 278, 391 

Spartina gracilis AM 302 

Spartina patens AM 97, 278, 324 

Spartina pectinata AM 626 

Spartina × townsendii NM 260, 261 

Sphenopholis obtusa AM 625 

Sporobolus heterolepis AM 168 

Sporobolus virginicus AM 144, 326 

Sporobolus wrightii AM 491 

Stipa tenacissima AM 497 

Themeda triandra AM 23 

Thysanolaena maxima AM 422 

Trachypogon gouinii AM 144 

Trachypogon plumosus AM 149, 353, 358 

Triodia basedowii Weak AM 438 

Tripogon filiformis AM 343 

Triraphis mollis Weak AM 438 

Trisetum flavescens AM 207, 260, 261 

Triticum aestivum AM 15 

Triticum durum AM 10 

Uniola paniculata AM 563

318 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Vulpia ciliata ssp. ambigua AM 110 

Zea mays AM 123 

Zoysia japonica AM 326 

Commelinaceae 

Amischotolype hispida AM 655 

Commelina communis AM 294 

NM 643 

Commelina aff. erecta AM 143 

Musaceae 

Musa acuminata AM 163 

Heliconiaceae 

Heliconia psittacorum AM 510 

Cannaceae 

Canna coccinea NM 510 

Zingiberaceae 

Alpinia conchigera AM 655 

Amomum villosum AM 422 

Costus speciosus AM 422 

Paramomum petaloideum AM 655 

Renealmia guianensis AM 510 

Costaceae 

Costus scaber AM 510 

Costus spiralis var. spiralis AM 510 

Myristicaceae 

Horsfieldia pandurifolia AM 655 

Horsfieldia tetratepala AM 422 

Myristica yunnanensis AM 655 

Pycnanthus angolensis AM 445 

Staudtia kamerunensis AM 445 

Magnoliaceae 

Liriodendron tulipifera AM 362, 470 

Magnolia henryi AM 422, 655 

Talauma ovata NM 548 

Annonaceae 

Annona cherimola AM 42 

Enantia chlorantha AM 445 

Goniothalamus griffithii AM 422, 655 

Hexalobus crispiflorus AM 445 

Mitrephora calcarea AM 655 

Pseuduvaria indochinensis NM 655 

Lauraceae 

Beilschmiedia pendula AM 356, 357 

Cryptocarya angulata AM 222 

Cryptocarya mackinnoniana AM 222 

Cryptocarya yunnanensis AM 422 

Laurus nobilis AM 368, 603 

Lindera benzoin AM 140 

Litsea dileniifolia AM 655 

Litsea liyuyingi AM 655 

Nectandra rigida AM 31 

Ocotea indecora AM 651 

Ocotea puberula AM 651 

Persea americana AM 41, 250, 326, 

609 

Phoebe lanceolata AM 422 

Aristolochiaceae 

Asarum europaeum AM + NM 260, 261 

Piperaceae 

Peperomia hesperomannii AM 326 

Peperomia membranacea AM 326 

Piper longum AM 422, 655 

Piper nigrum AM 578 

Piper sarmentosum AM 422 

Piper sp. NM 422 

Papaveraceae 

Chelidonium majus NM 260, 261 

Macleaya cordata AM 343 

Papaver rhoeas AM + NM 260, 261 

Papaver somniferum NM 260, 261 

Menispermaceae 

Cocculus trilobus AM 326 

Berberidaceae 

Berberis buxifolia AM 207 

Berberis darwinii NM 207 

Berberis vulgaris AM 260, 261 

Mahonia aquifolium AM 260, 261 

Podophyllum peltatum AM 410, 621 

Ranunculaceae 

Aconitum delphinifolium AM 594 

Aconitum napellus AM + NM 260, 261 

Actea spicata AM + NM 260, 261 

Anemone nemorosa AM + NM 260, 261 

Anemone ranunculoides AM 260, 261 

Anemone vulgaris AM 260, 261 

Batrachium circinatum NM 62 

Batrachium peltatum NM 62 

Caltha palustris NM 62 

AM + NM 260, 261 

Clematis stans AM 636 

Clematis vitalba AM 368 

AM + ECM 260, 261 

Consolida ambigua AM 260, 261 

Corydalis lutea NM 260, 261 

Corydalis solida NM 260, 261 

Fumaria officinalis AM + NM 260, 261 

Helleborus foetidus AM 260, 261 

Myosurus minimus AM 260, 261 

Pulsatilla patens AM 408 

Pulsatilla pratensis AM 408 

Pulsatilla vulgaris AM + NM 260, 261 

Ranunculus acris AM 194, 456, 586, 

260, 261 

Ranunculus acris AM 503 

ssp. pumilus 

Ranunculus adoneus AM 416, 519

319 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Ranunculus auricomus AM 586 

AM + NM 260, 261 

Ranunculus biternatus AM 560 

Ranunculus bulbosus AM 456, 260, 261 

Ranunculus crassipes AM 342 

Ranunculus ficaria AM + NM 260, 261 

Ranunculus flammula AM 62 

AM + NM 260, 261 

Ranunculus fluitans NM 260, 261 

Ranunculus lingua AM + NM 260, 261 

Ranunculus nemorosus AM 586 

Ranunculus paludosus AM 260, 261 

Ranunculus repens AM + NM 260, 261 

Ranunculus reptans × AM 260, 261 

flammula 

Ranunculus sardous AM 260, 261 

Ranunculus sceleratus AM + NM 260, 261 

Thalictrum minus AM 260, 261 

Trollius europaeus AM 194, 503, 260, 

261 

Proteaceae 

Banksia aricifolia Weak AM 454 

Conospermum longifolium AM 67 

Conospermum taxifolium AM 67 

Lomatia hirsute NM 207 

Grevillea robusta NM 326 

Telopea speciosissima AM 67 

Buxaceae 

Buxus sempervirens AM 260, 261 

Gunneraceae 

Gunnera petaloidea AM 326 

Dilleniaceae 

Hibbertia serpyllifolia AM 67 

Droseraceae 

Drosera anglica NM 326, 260, 261 

Drosera intermedia AM 212 

NM 260, 261 

Drosera rotundifolia AM 140 

AM + NM 260, 261 

Frankeniaceae 

Frankenia laevis NM 260, 261 

Frankenia plicata Weak AM 438 

Tamaricaceae 

Tamarix chinensis AM 620 

Tamarix gallica AM 368 

Plumbaginaceae 

Armeria maritime AM 277 

AM + NM 260, 261 

Armeria maritima AM 456 

ssp. halleri 

Limonium vulgare AM + NM 260, 261 

Polygonaceae 

Bistorta vivipara AM 194 

Coccoloba uvifera ECM 326 

Coccoloba warmingii AM 31 

Fagopyrum esculentum NM 260, 261 

Fallopia convolvulus NM 260, 261 

Oxyria digyna NM 260, 261 

Polygonum amphibium Facultative AM 62, 626 

NM 260, 261 

Polygonum aviculare NM 307 

AM + NM 260, 261 

Polygonum bistorta NM 594 

AM 260, 261 

Polygonum capitatum ECM + weak AM 326 

Polygonum cespitosum AM 140 

Polygonum cuspidatum AM 636 

NM 643 

Polygonum hydropiper NM 260, 261 

Polygonum lapathifolium NM 260, 261 

Polygonum lapathifolium NM 307 

ssp. pallidum 

Polygonum longisetum NM 643 

Polygonum maritimum NM 260, 261 

Polygonum mite NM 260, 261 

Polygonum pesicaria Weak AM 626 

AM + NM 260, 261 

Polygonum statice AM 343 

Polygonum viviparum ECM 594 

AM + ECM 260, 261 

Polygonum weyrichii ECM 587 

Polygonum weyrichii NM 636 

var. alpinum 

Rumex acetosa AM 194, 294, 456 

AM + NM 260, 261 

Rumex acetosella NM 207, 456 

Rumex acetosella agg. NM 260, 261 

Rumex alpinus AM + NM 260, 261 

Rumex altissimus AM 140 

Rumex conglomeratus NM 260, 261 

Rumex crispus NM 307 

AM + NM 260, 261 

Rumex hastatus AM 343 

Rumex hydrolapathum NM 62 

Rumex japonicus NM 643 

Rumex longifolius Weak AM 194 

Rumex obtusifolius AM + NM 260, 261 

Rumex palustris NM 260, 261 

Rumex scutatus NM 260, 261 

Ruprechtia laxiflora NM 651 

Caryophyllaceae 

Agrostemma githago NM 260, 261 

Arenaria serpyllifolia NM 260, 261 

Cerastium alpinum NM 260, 261 

Cerastium arcticum NM 260, 261

320 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Cerastium arvense AM 456 

NM 260, 261 

Cerastium cerastoides NM 260, 261 

Cerastium fontanum AM 194 

NM 456 

Cerastium fontanum AM 342 

ssp. fontanum 

Cerastium fontanum AM + NM 260, 261 

ssp. glabrescens 

Cerastium glomeratum NM 560 

Cerastium holosteoides NM 643 

var. hallaisanense 

Cerastium semidecandrum NM 260, 261 

Colobanthus apetalus AM 342 

var. alpinus 

Colobanthus kerguelensis NM 560 

Colobanthus muscoides NM 342 

Dianthus carthusianorum AM 456 

NM 260, 261 

Dianthus caryophyllus NM 554 

Dianthus deltoides NM 260, 261 

Dianthus gratianopolitanus NM 260, 261 

Gypsophila fastigiata NM 456 

Honkenya peploides NM 260, 261 

Lychnis flos-cuculi AM + NM 260, 261 

Lychnis viscaria NM 260, 261 

Minuartia hybrida AM 260, 261 

Minuartia verna NM 260, 261 

Moehringia trinervia NM 260, 261 

Myosoton aquaticum NM 260, 261 

Polycarpaea corymbosa AM 336 

Sagina apetala NM 260, 261 

Sagina procumbens NM 260, 261 

Sagina subulata NM 260, 261 

Saponaria officinalis NM 260, 261 

Scleranthus annuus NM 260, 261 

Silene acaulis AM 594 

AM + ECM + NM 260, 261 

Silene alba NM 260, 261 

Silene colorata 

AM 368 

ssp. canescens 

Silene dioica Weak AM 194 

NM 260, 261 

Silene maritima NM 260, 261 

Silene nutans NM 260, 261 

Silene otites AM 260, 261 

Silene vulgaris NM 456, 260, 261 

Schiedea spergulina AM 326 

Spergula arvensis AM + NM 260, 261 

Spergularia marina AM or NM 260, 261 

Spergularia media AM + NM 260, 261 

Stellaria alsine NM 260, 261 

Stellaria graminea NM 260, 261 

Stellaria holostea AM + NM 260, 261 

Stellaria media AM 294 

AM + NM 260, 261 

Stellaria media ssp. media AM 342 

Stellaria nemorum AM + NM 260, 261 

Stellaria pallida AM + NM 260, 261 

Stellaria palustris NM 260, 261 

Stellaria parviflora NM 342 

Amaranthaceae 

Achyranthes aspera NM 422 

AM 343 

Achyranthes fauriei NM 643 

Achyranthes splendens NM 326 

Amaranthus caudatus AM 33 

Amaranthus lividus AM 294 

Amaranthus retroflexus NM 260, 261 

Amaranthus spinosus NM 422 

AM 343 

Amaranthus tricolor AM 33 

Amaranthus viridis AM 343 

Atriplex barclayana Weak AM 112 

Atriplex canescens AM 57 

Atriplex glabriuscula NM 260, 261 

Atriplex halimus NM 368 

Atriplex hortensis NM 260, 261 

Atriplex julacea AM 543 

Atriplex limbata NM 438 

Atriplex littoralis AM 260, 261 

Atriplex patula AM 260, 261 

Atriplex prostrata NM 260, 261 

Atriplex semibaccata NM 326 

Beta vulgaris AM 260, 261 

Celosia cristata AM 33 

Chenopodium album NM 307 

AM + NM 260, 261 

Chenopodium 

AM 343 

ambrosioides 

Chenopodium bonushenricus 

NM 260, 261 

Chenopodium glaucum NM 260, 261 

Chenopodium murale NM 326 

Chenopodium rubrum NM 260, 261 

Enchylaena tomentosa NM 438 

Gomphrena globosa AM 33 

Halimione portulacoides AM + NM 260, 261 

Nototrichium sandwicense NM 326 

Ptilotus latifolius NM 438 

Ptilotus obovatus var. obovatus 

NM 438 

Ptilotus sessilifolius NM 438 

Salicornia europaea AM 277 

AM + NM 260, 261

321 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Salsola kali NM 19, 300, 260, 

261 

AM + NM 438 

Sclerolaena diacantha AM 438 

Sclerolaena holtiana NM 438 

Suaeda glauca AM 620 

Suaeda maritima AM 533 

AM + NM 260, 261 

Suaeda vera AM + NM 260, 261 

Aizoaceae 

Cheredospsis purpurea NM 417 

Delosperma herbium Falcutative AM 417 

Sesuvium portulacastrum NM 326 

AM 143 

Tetragonia tetragonioides AM 326 

Phytolaccaceae 

Stegnosperma halimifolia AM 112 

Nyctaginaceae 

Abronia maritima Falcutative AM 543 

Abronia umbellata AM 543 

Boerhavia repens NM 326 

Bougainvillea spectabilis NM 326, 651 

Mirabilis jalapa NM 326 

Neea sp. ECM 656 

Pisonia sp. ECM 656 

Pisonia umbellifera NM 326 

Cactaceae 

Chamacereus sylvestris NM 417 

Cochemia poselgeri AM 112 

Coryphanta radians Weak AM 103 

Escontria chiotilla Weak AM 103 

Ferocactus acanthodes AM 151 

Ferocactus flavovirens Weak AM 103 

Ferocactus latispinus Weak AM 103 

Ferocactus peninsulae AM 112 

Lemaireocereus thurberi Weak AM 112 

Lophocereus schottii AM 112 

Machaerocereus gummosus Weak AM 112 

Mammilaria carnea Weak AM 103 

Mammilaria dioica Weak AM 112 

Mamillaria elongata NM 417 

Myrtillocactus 

Weak AM 103 

geometrizans 

Neobuxbaumia tetetzo Weak AM 103 

Nopalea karwinskiana AM 20 

Opuntia cholla AM 112 

Opuntia excelsa AM 20 

Opuntia ficus-indica AM 151 

Opuntia lindsayi AM 112 

Opuntia pilifera AM 103 

Opuntia puberula AM 20 

Opuntia streptacantha AM 103 

Opuntia stricta var. dillenii AM 143 

Opuntia tuna NM 417 

Pachycereus 

AM 494 

pectin-aboriginum 

Pachycereus pringlei Weak AM 112 

Selenicereus macdonaldiae AM 417 

Stenocereus queretaroensis AM 463, 464 

Stenocereus stellatus Weak AM 103 

Portulacaceae 

Lyallia kerguelensis NM 560 

Montia fontana 

NM 342 

ssp. fontana 

Montia perfoliata NM 207 

Montia sibirica AM 260, 261 

Portulaca intraterranea NM 438 

Portulaca lutea NM 326 

Portulaca oleracea AM + NM 260, 261 

Portulaca sclerocarpa NM 326 

Olacaceae 

Coula edulis AM 445 

Ongokea gore AM 445 

Strombosis grandifolia AM 445 

Loranthaceae 

Viscum album NM 260, 261 

Santalaceae 

Santalum ellipticum NM 326 

Santalum paniculatum AM 326 

Thesium alpinum NM 456 

Thesium humifusum NM 260, 261 

Loasaceae 

Caiophora sylvestris NM 207 

Loasa berghii NM 207 

Cornaceae 

Cornus sanguinea AM + NM 260, 261 

Balsaminaceae 

Impatiens capensis AM 140 

Impatiens glandulifera AM + NM 260, 261 

Impatiens noli-tangere AM + NM 260, 261 

Impatiens parviflora AM + NM 260, 261 

Impatiens walleriana AM 322 

Tetrameristaceae 

Tetramerista glabra AM 568 

Polemoniaceae 

Polemonium caeruleum AM 260, 261 

Fouquieriaceae 

Fouquieria digueti AM 112 

Theaceae 

Pyrenaria cheliensis AM 422 

Ebenaceae 

Diospyros nigrocartex AM 422, 655 

Theophrastaceae 

Jacquinia pungens AM 20 

Primulaceae 

Anagallis arvensis AM 472, 260, 261

322 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Anagallis tenella AM 260, 261 

Asterolinum stellatum AM 472 

Cyclamen hederifolium AM 260, 261 

Glaux maritima AM 260, 261 

Hottonia palustris AM + NM 260, 261 

Lysimachia japonica AM 643 

Lysimachia nemorum AM + NM 260, 261 

Lysimachia nummularia AM + NM 260, 261 

Lysimachia terrestris AM 140 

AM + NM 260, 261 

Lysimachia thyrsiflora NM 62 

AM + NM 260, 261 

Lysimachia vulgaris AM 368 

AM + NM 260, 261 

Primula elatior AM 260, 261 

Primula farinosa AM 260, 261 

Primula veris AM + NM 260, 261 

Primula vulgaris AM 260, 261 

Samolus valerandi AM 368, 260, 261 

Trientalis europaea AM 501 

AM + NM 260, 261 

Myrsinaceae 

Ardisia crenata AM 87 

Ardisia tenera AM 422, 655 

Measa indica AM 422 

Myrsine alyxifolia AM 326 

Rapanea ferruginea AM 31 

Styracaceae 

Strichinus brasiliensis AM 651 

Diapensiaceae 

Diapensia lapponica ERM 260, 261 

Lecythidaceae 

Barringtonia macrostachya AM 655 

Barringtonia racemosa AM 655 

Cariniana estrellensis AM 651 

Sapotaceae 

Aningeria adolfi-friedericii AM 637, 638 

Aningeria robusta AM 445 

Argania spinosa AM 434 

Autranella congolensis AM 445 

Baillonella toxisperma AM 445 

Gambeya africana AM 445 

Omphalocarpum procerum AM 445 

Palaquium gutta AM 568 

Actinidiaceae 

Actinidia deliciosa AM 523 

Clethraceae 

Clethra barbinervis AM 335 

Ericaceae 

Andromeda polifolia ERM 260, 261 

Arbutus menziesii ABM 374 

ECM 375 

Arbutus unedo ABM 200, 238, 421, 

368, 472, 260, 

261 

Arctostaphylos alpinus ABM 594 

ERM 260, 261 

Arctostaphylos uva-ursi ABM 259, 507 

M(endo) + ERM + 260, 261 

ABM + ECM + 

EEM 

Calluna vulgaris ERM 72, 238, 299, 

260, 261 

Cassiope tetragona ERM 594 

Cavendishia capitulata ERM 474 

Cavendishia melastomoides ERM 474 

Daboecia cantabrica ERM 260, 261 

Disterigma humboldtii ERM 474 

Empetrum nigrum ERM 594, 260, 261 

Erica arborea ERM 368 

Erica cinerea ERM 99, 260, 261 

Erica erigena ERM 260, 261 

Erica mackaiana ERM 260, 261 

Erica multiflora ERM 472 

Erica tetralix ERM 260, 261 

Erica vagans ERM 260, 261 

Gaultheria erecta ERM 474 

Gaultheria procumbens ERM 238 

Gaultheria shallon ERM 22, 639 

Gonocalyx costaricense ERM 474 

Kalmia latifolia ERM 238 

Ledum palustre ERM 260, 261 

Ledum palustre 

ERM 594 

ssp. decumbens 

Ledum palustre 

ERM 594 

ssp. groenlandicum 

Leucopogon parviflorus ERM 401 

Leucothoe fontanesiana ERM 238 

Loiseleuria procumbens ERM 594, 260, 261 

Oxydendrum arboreum ERM 238 

Pernettya mucronata ERM 207 

Phyllodoce caerulea ERM 260, 261 

Pieris floribunda ERM 238, 555 

Rhododendron 

ERM 154 

brachycarpum 

Rhododendron 

ERM 238 

calendulaceum 

Rhododendron 

ERM 238 

carolinianum 

Rhododendron catawbiense ERM 238 

Rhododendron maximum ERM 238 

Rhododendron 

ERM 238 

mucronulatum 

Rhododendron obtusum ERM 601

323 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Rhododendron ponticum ERM 260, 261 

Sphyospermum buxifolium ERM 474 

Sphyospermum cordifolium ERM 474 

Vaccinium angustifolium ERM 157 

Vaccinium calycinum ERM + AM 325, 326 

Vaccinium corymbosum ERM 238, 645 

Vaccinium dentatum ERM + AM 325, 326 

Vaccinium macrocarpon ERM 99, 238, 260, 

261 

Vaccinium myrtillus ERM 72, 260, 261 

Vaccinium oxycoccos ERM 260, 261 

Vaccinium reticulatum ERM + weak AM 326 

ERM + AM 325 

Vaccinium uliginosum ERM 594, 260, 261 

Vaccinium vitis-idaea ERM 594, 260, 261 

Woollsia pungens ERM 401 

Ericaceae “Pyrolaceae” 

Moneses uniflora EEM + M(endo) 260, 261 

Orthilia secunda EEM + M(endo) 260, 261 

Pyrola media M(endo) 260, 261 

Pyrola minor EEM + M(endo) 260, 261 

Pyrola rotundifolia EEM + M(endo) 260, 261 

Ericaceae (Monotropoideae) 

Monotropa hypopitys MTM 260, 261 

Monotropa uniflora MTM 647 

Monotropastrum humile MTM 384 

Sarcodes sanguinea ERM + MTM 332 

Ericaceae (Epacridoideae) 

Astroloma conostephioides ERM 393 

Astroloma humifusum ERM 393 

Astroloma pinifolium ERM 393, 394 

Brachyloma daphnoides AM 67 

ERM 393 

Epacris impressa ERM 393, 394 

Epacris impressa ERM 393 

var. grandiflora 

Epacris microphylla AM 67 

ERM 128 

Leucopogon ericoides ERM 393 

Leucopogon juniperinus AM 67 

Leucopogon parviflorus ERM 393, 556 

Lysinema ciliatum ERM 36 

Styphelia adscendens ERM 393 

Styphelia tameiameiae ERM + AM 325, 326 

Woollsia pungens ERM 127, 312 

Icacinaceae 

Pittosporopsis kerrii AM 422 

Boraginaceae 

Arnebia hispidisima AM 567 

Borago officinalis NM 260, 261 

Bourreria sonorae AM 112 

Cerastium arvense NM 207 

Cordia alliodora AM 20 

Cordia curassavica Weak AM 103 

Cordia ecalyculata AM 651 

Cordia trichotoma AM 651 

Cynoglossum creticum AM 207 

Cynoglossum lanceolatum AM 343 

Echium vulgare AM 260, 261 

Heliotropium anomalum AM 326 

Heliotropium curassavicum AM 326 

Heliotropium sp. AM 567 

Lithospermum arvense NM 260, 261 

Lithospermum officinale NM 260, 261 

Lithospermum 

AM 260, 261 

purpurocaeruleum 

Myosotis alpestris AM + NM 260, 261 

Myosotis arvensis NM 260, 261 

Myosotis decumbens Facultative AM 194 

Myosotis discolor AM 260, 261 

Myosotis laxa 

AM 260, 261 

ssp. caespitosa 

Myosotis palustris Facultative AM 62 

Myosotis ramosissima AM + NM 260, 261 

Myosotis scorpioides AM + NM 260, 261 

Myosotis sylvatica AM + NM 260, 261 

Omphalalappula concava NM 438 

Plagiobothrys figuratus AM 293 

Pulmonaria officinalis AM + NM 260, 261 

Silene andicola NM 207 

Stellaria graminea AM 194 

Stellaria media AM 207 

Symphytum officinale AM 260, 261 

Symphytum tuberosum AM + NM 260, 261 

Tournefortia argentea AM 326 

Trichodesma zeylanicum AM 438 

Rubiaceae 

Asperula cynanchica AM 260, 261 

Borreria articularis AM 336 

Borreria pusilla AM 336 

Canthium parvifoliam AM 422 

Cephalanthus occidentalis AM 140 

Chesalia curviflora AM 422, 655 

Coffea arabica 

AM 602 

cv. guatemala 

Coprosma ernodeoides AM 326 

Coprosma kauensis AM 326 

Coprosma perpusilla NM 342 

ssp. subantarctica 

Declieuxia fruticosa AM 149 

Duperrea pavettaefolia AM 655 

Galium album AM 194, 260, 261 

Galium antarcticum AM 560 

Galium aparine NM 207 

AM + NM 260, 261 

Galium boreale AM 260, 261

324 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Galium cruciata AM 260, 261 

Galium mollugo AM 456 

AM + NM 260, 261 

Galium odoratum AM + NM 260, 261 

Galium palustre AM + NM 260, 261 

Galium saxatile AM + NM 260, 261 

Galium sp. AM 140 

Galium uliginosum NM 260, 261 

Galium verum AM 260, 261 

Geophila herbacea AM 422, 655 

Hedyotis costata AM 422 

Hedyotis elatior AM 326 

Hedyotis foggiana AM 326 

Hedyotis terminalis AM 326 

Lasianthus hookeri AM 422 

Lasianthus sikkimensis AM 655 

Lasianthus verticillatus AM 655 

Metadina trichotoma AM 655 

Mitragina ciliata AM 445 

Morinda lucida AM 422 

Mycetia hirta AM 655 

Nauclea diederrichii AM 445 

Neonauclea tsaiana AM 655 

Nertera granadensis AM 326 

Oldenlandia aspera AM 336 

Ophiorrhiza 

AM 422 

austro-yunnanensis 

Paederia scandens AM 643 

Pausynistalia johimbe AM 445 

Prismatomeria tetrandra AM 422 

Psychotria calocarpa AM 655 

Psychotria henryi AM 422 

Psychotria siamica NM 655 

Psychotria sp. AM 326 

Randia ruglosa AM 315 

Relbuneum hypocarpium AM 207 

Relbuneum richardianum AM 207 

Rubia peregrina AM 368 

AM + ECM 472 

ECM 260, 261 

Sherardia arvensis AM 472 

Vangueria infausta AM 84, 85 

Vangueria infausta AM 221 

ssp. infausta 

Gentianaceae 

Blackstonia perfoliata AM 368, 472 

AM + NM 260, 261 

Centaurium erythraea AM 472, 260, 261 

Gentiana algida NM 594 

Gentiana nivalis AM + NM 260, 261 

Gentiana pneumonanthe AM + NM 260, 261 

Gentiana verna AM 561, 260, 261 

Gentianella amarella AM 260, 261 

Gentianella campestris AM 194, 260, 261 

Gentianella germanica AM 456, 260, 261 

Voyria aurantiaca Mycoheterotrophy 79 

(via AM) 

Voyria caerulea 

Mycoheterotrophy 79 

(via AM) 

Voyria corymbosa Mycoheterotrophy 79 

(via AM) 

Voyria obconica 


(via AM) 

Voyria rosea 


(via AM) 

Voyria tenuiflora Mycoheterotrophy 79 

(via AM) 

Voyriella parviflora Mycoheterotrophy 79 

(via AM) 

Loganiaceae 

Anthocleista schweinfurthii AM 445 

Buddleia asiatica AM 326 

Apocynaceae 

Adenium obesum AM 623 

Adenium somalense AM 623 

Allamanda cathartica AM 623 

var. schotti 

Allamanda violacea AM 623 

Alstonia boonei AM 445 

Alyxia oliviaeformis AM 326 

Amsonia tabernaemontana AM 623 

Apocynum cannabinum AM 623 

Apocynum medium AM 623 

Aspidosperma parvifolium NM 548 

Aspidosperma polyneuron NM 651 

Catharanthus roseus AM 623 

Dyera costulata NM 568 

Landolphia heudelottii NM 47 

Mandevilla sanderei AM 623 

Nerium oleander AM 623 

Pachypodium lamerei AM 623 

Picralima nitida AM 445 

Plumeria obtuse AM 623 

Plumeria rubra AM 623 

Rauwolfia serpentina AM 623 

Rauwolfia verticillata AM 623 

Saba senegalensis NM 47 

Stemmadenia 

AM 244 

donnell-smithii 

Strophanthus capensis AM 623 

Tabernaemontana australis AM 651 

Thevetia peruviana AM 623

325 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Trachelospermum AM 623 

jasminoides 

Vinca major AM 260, 261 

Vinca minor AM 623, 260, 261 

Apocynaceae 

(Asclepiadoideae) 

Asclepias incarnata AM 58, 140 

Calotropis gigantea AM 343 

Cynanchum 

AM 207 

nummularifolium 

Periploca graeca NM 368 

Stapelia glabricaulis AM 417 

Oleaceae 

Fraxinus excelsior AM 465, 622 

AM + ECM 260, 261 

Fraxinus ornus AM 368 

Fraxinus oxycarpa AM 368 

Fraxinus pennsylvanica AM 28 

Jasminum laurifolium AM 655 

Jasminum wangii AM 422 

Ligustrum vulgare AM 368, 260, 261 

Olea europaea AM 17, 101 

Olea europaea 

AM 637, 638 

ssp. cuspidate 

Olea europaea 

AM 108, 368 

ssp. sylvestris 

Olea europaea var. oleaster AM 472 

Phillyrea angustifolia AM 368, 472 

Phillyrea latifolia AM 368 

Gesneriaceae 

Corallodiscus flabellatus AM 343 

Cytrandra longifolia AM 326 

Pedaliaceae 

Sesamum indicum AM 612 

Plantaginaceae 

Callitriche hamulata AM 260, 261 

Littorella uniflora AM 62, 432, 260, 

261 

Plantago coronopus AM 260, 261 

Plantago drummondii AM 438 

Plantago lanceolata AM 194, 307, 446, 

456, 260, 261 

Plantago major AM 49, 368, 260, 

261 

Plantago major 

AM 260, 261 

ssp. intermedia 

Plantago maritima AM 260, 261 

Plantago media AM 48, 194, 260, 

261 

Plantago princeps AM 326 

Bignoniaceae 

Incarvillea arguta AM 343 

Jacaranda mimosaefolia AM 548, 651 

Jacaranda puberula AM 651 

Oroxylum indicum AM 655 

Stenolobium stans AM 548 

Tabebuia chrysotricha AM 651 

Tabebuia roseo-alba AM 651 

Tecoma stans Weak AM 103 

AM 112 

Zeyheria tuberculosa NM 651 

Verbenaceae 

Aegiphila sellowiana AM 651 

Citharexylum myrianthum AM 651 

Clerodendron japonicum AM 422 

Lantana camara AM 326 

Lippia graveolens Weak AM 103 

Lippia nodiflora AM 143 

Tectona grandis AM 476 

Teucrium flavum AM 368 

Verbena hastata AM 140 

Verbena officinalis AM 260, 261 

Vitex agnus-castus AM 368 

Vitex montevidensis AM 651 

Vitex negundo AM 343 

Vitex rotundifolia AM 326 

Vitex vestita AM 655 

Lamiaceae 

Ajuga genevensis AM 260, 261 

Ajuga pyramidalis AM 194 

Ajuga reptans AM 260, 261 

Ballota nigra NM 260, 261 

Betonica officinalis AM 212 

Clinopodium gracile AM 643 

Clinopodium vulgare AM + NM 260, 261 

Colebrookea oppositifolia AM 655 

Coleus parviflorus AM 469 

Coleus × hybridus AM 322 

Collinsonia canadensis AM 140 

Elsholtzia blanda AM 422 

Elsholtzia cypriani AM 343 

Galeopsis segetum NM 260, 261 

Galeopsis tetrahit NM 260, 261 

Glechoma hederacea AM + NM 260, 261 

Gomphostemma microdon AM 655 

Hyptis lantiflora AM 112 

Lamiastrum galeobdolon AM + NM 260, 261 

Lamium album AM 260, 261 

Lamium amplexicaule AM 294, 260, 261 

Lamium purpureum AM 260, 261 

Lavandula augustifolia AM 348 

Lavandula officinalis AM 104 

Lavandula pedunculata AM 339 

Lavandula spica AM 39 

Leucas aspera AM 336 

Lycopus europaeus AM + NM 260, 261

326 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Melittis melissophyllum AM + NM 260, 261 

Mentha aquatica AM 368 

AM + NM 260, 261 

Mentha arvensis AM 248 

AM + NM 260, 261 

Mentha arvensis 

AM 1 

ssp. haplocalyx 

Mentha cardiaca AM 1 

Mentha citrata AM 1 

Mentha piperita AM 1 

AM + NM 260, 261 

Mentha pulegium AM 260, 261 

Mentha spicata AM 1 

Mentha viridis AM 1 

Nepeta cataria AM 260, 261 

Ocimum basilicum AM 105, 171, 343 

Origanum vulgare AM 260, 261 

Paraphlomis javanica AM 655 

Prunella vulgaris AM 194, 586 

AM + NM 260, 261 

Pycnanthemum tenuifolium AM 140, 599 

Pycnanthemum 

AM 140 

virginianum 

Rosmarinus officinalis AM 472 

Salvia azurea AM 633 

Salvia officinalis AM 105 

Salvia pratensis AM 260, 261 

Salvia splendens AM 322 

Salvia verbenaca AM 260, 261 

Scutellaria galericulata AM + NM 260, 261 

Stachys × ambigua AM 260, 261 

Stachys maritima AM 368 

Stachys officinalis AM 260, 261 

Stachys palustris AM 260, 261 

Stachys recta 

NM 368 

var. psammophila 

Stachys sylvatica AM + NM 260, 261 

Stenogyne purpurea AM 326 

Teucrium chamaedrys AM 260, 261 

Teucrium fruticans AM 348, 472 

Teucrium scordium AM 260, 261 

Teucrium scordonia AM 260, 261 

Thymus mastichina AM 339 

Thymus polytrichus AM 630 

ssp. britannicus 

Thymus pulegioides AM 456 

Thymus serpyllum AM 456, 260, 261 

Thymus vulgaris AM 105, 583 

Thymus zygis AM 339 

Acanthaceae 

Barleria cristata AM 343 

Carlowrightia californica AM 112 

Justicia procumbens var. AM 643 

leucantha 

Phlogacanthus curviflorus AM 655 

Pseudoranthemum AM 422, 655 

palatiferum 

Ruellia peninsularis AM 112 

Scrophulariaceae 

Antirrhinum majus AM 74 

Bartsia alpina AM + NM 260, 261 

Buddleja davidii NM 260, 261 

Calceolaria crenatiflora AM 207 

Calceolaria polyrrhiza AM 207 

Castilleja arvensis NM 326 

Chaenorhinum minus NM 260, 261 

Cymbalaria muralis AM 260, 261 

Digitalis purpurea AM + NM 260, 261 

Eremophila longifolia Weak AM 438 

Eremophila macdonnellii Weak AM 438 

Euphrasia minima NM 260, 261 

Euphrasia officinalis s. l. NM 260, 261 

Euphrasia stricta NM 456 

Kickxia spuria AM 260, 261 

Lamourouxia rhinanthifolia NM 103 

Linaria repens AM + NM 260, 261 

Linaria supina AM + NM 260, 261 

Linaria vulgaris AM + NM 260, 261 

Melampyrum arvense NM 260, 261 

Melampyrum cristatum NM 260, 261 

Melampyrum pretense NM 194 

AM + NM 260, 261 

Melampyrum sylvaticum NM 194, 260, 261 

Mimulus guttatus NM 62 

AM 95 

Mimulus ringens AM 140 

Odontites lutea AM 368 

Odontites verna NM 260, 261 

Pedicularis capitata NM 594 

Pedicularis kanei NM 594 

Pedicularis langsdorffii NM 594 

Pedicularis palustris NM 260, 261 

Pedicularis sylvatica NM 260, 261 

Pedicularis verticillata NM 594 

Penstemon cardwellii Facultative AM 585 

Penstemon frutescens Facultative AM 587 

Plantago asiatica AM 643 

Rhinanthus angustifolius AM 456 

NM 260, 261 

Rhinanthus minor Weak AM 194 

Scrophularia auriculata AM 260, 261 

Scrophularia nodosa AM + NM 260, 261 

Scrophularia umbrosa NM 260, 261 

Scrophularia vernalis NM 260, 261

327 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Verbascum lychnitis AM 260, 261 

Verbascum nigrum AM 260, 261 

Verbascum thapsus AM 207, 260, 261 

Veronica agrestis AM + NM 260, 261 

Veronica alpina AM 260, 261 

Veronica 


anagallis-aquatica AM 260, 261 

Veronica arvensis AM 643, 260, 261 

Veronica beccabunga NM 62, 260, 261 

Veronica chamaedrys AM 194, 456 

AM + NM 260, 261 

Veronica filiformis AM 260, 261 

Veronica fruticans AM 260, 261 

Veronica officinalis AM 194 

AM + NM 260, 261 

Veronica peregrina AM 260, 261 

Veronica persica AM 643, 260, 261 

Veronica serpyllifolia AM 194, 260, 261 

Veronica spicata AM 260, 261 

Lentibulariaceae 

Pinguicula alpine NM 260, 261 

Pinguicula vulgaris NM 260, 261 

Convolvulaceae 

Calystegia sepium AM 260, 261 

Calystegia soldanella NM 368 

Convolvulus arvensis AM 260, 261 

Convolvulus elegantissimus AM 472 

Convolvulus eyreanus AM 438 

Ipomoea arborescens Weak AM 103 

Ipomoea batatas AM 455 

Ipomoea imperati AM 326 

Ipomoea pes-caprae AM 64, 65, 144, 

326, 336 

Ipomoea stolonifera Weak AM 143 

Jacquemontia ovalifolia AM 326 

Jacquemontia reclinata AM 202 

Solanaceae 

Atropa belladonna AM 260, 261 

Capsicum annuum AM 50 

Cestrum intermedium AM 651 

Datura metal AM 216 

Datura stramonium AM 343 

Lycium barbarum AM 260, 261 

Lycium fremontii AM 112 

Lycium sandwicense AM 326 

Lycopersicon esculentum AM 407 

Nicotiana velutina AM 438 

Petunia × hybrida AM 322 

Solanum argenteum AM 651 

Solanum dulcamara NM 62 

AM 260, 261 

Solanum elipticum AM 438 

Solanum khasianum AM 343 

Solanum hindsianum AM 112 

Solanum nigrum AM + NM 260, 261 

Solanum surattense AM 567 

Solanum torvum NM 422 

Solanum tuberosum AM 386 

Aquifoliaceae 

Ilex aquifolium AM + ECM 260, 261 

Ilex paraguariensis AM 31 

Ilex verticallata AM 140 

Araliaceae 

Didymopanax 

AM 31 

angustissimum 

Hedera helix AM 368, 472, 260, 

261 

Macropanax dispermus AM 655 

Panax ginseng AM 178 

Panax quinquefolius AM 390 

Pseudopanax laetevirens AM 207 

Stilbocarpa polaris NM 342 

Trevesia palmata NM 422 

Trevesia palmata NM 655 

var. costata 

Pittosporaceae 

Pittosporum gayanum AM 326 

Apiaceae 

Actinotis helianthi AM 67 

Aegopodium podograria AM + NM 260, 261 

Aethusa cynapium AM 260, 261 

Angelica archangelica AM 260, 261 

Angelica sylvestris AM 260, 261 

Anthriscus sylvestris NM 260, 261 

Apium graveolens AM 490, 260, 261 

Apium nodiflorum NM 260, 261 

Astrantia major AM + NM 260, 261 

Azorella macquariensis AM 342 

Berula erecta Facultative AM 62 

NM 260, 261 

Bupleurum baldense AM 472 

Bupleurum falcatum AM 260, 261 

Carum carvi AM 194, 260, 261 

Chaerophyllum temulentum AM + NM 260, 261 

Conium maculatum AM 260, 261 

Conopodium majus AM 260, 261 

Coriandrum sativum AM 220 

Daucus carota AM 171 

AM + NM 260, 261 

Echinophora spinosa AM 368 

Eryngium campestre AM 260, 261 

Eryngium maritimum AM 368 

AM + NM 260, 261 

Eryngium paniculatum AM 207 

Foeniculum vulgare AM 305 

Heracleum sphondylium AM + NM 260, 261

328 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Hydrocotyle bonariensis AM 143 

Hydrocotyle 

AM 342 

novae-zeelandiae 

Hydrocotyle vulgaris AM 368 

AM + NM 260, 261 

Myrrhis odorata AM 260, 261 

Oenanthe aquatica NM 260, 261 

Oenanthe crocata NM 260, 261 

Osmorhiza chilensis AM 207 

Pastinaca sativa AM 307, 260, 261 

Petroselium crispum AM 171 

NM 260, 261 

Peucedanum ostruthium AM 260, 261 

Peucedanum palustre AM 260, 261 

Pimpinella major AM + NM 260, 261 

Pimpinella saxifraga AM 194, 456 

AM + NM 260, 261 

Platysace linearifolia ECM 67 

Sanicula europaea AM 260, 261 

Scandix pecten-veneris AM 260, 261 

Seseli libanotis AM 260, 261 

Seseli tortuosum AM 368 

Silaum silaus AM 260, 261 

Sium latifolium NM 260, 261 

Trachymene glaucifolia AM 438 

Campanulaceae 

Brighamia insignis AM 326 

Campanula glomerata AM 260, 261 

Campanula lasiocarpa AM 587 

Campanula patula AM 260, 261 

Campanula persicifolia AM 194 

AM + NM 260, 261 

Campanula punctata AM 636 

ssp.hondoensis 

Campanula rapunculoides AM 260, 261 

Campanula rotundifolia AM 194, 437, 456, 

260, 261 

Campanula trachelium NM 260, 261 

Clermontia fauriei AM 326 

Cyanea leptostegia AM 326 

Downingia elegans AM 293 

Jasione montana AM + NM 260, 261 

Lobelia cardinalis AM 140 

Lobelia dortmanna AM 62, 432, 260, 

261 

Lobelia sp. AM + ECM 656 

Lobelia yuccoides AM 326 

Phyteuma orbiculare AM 260, 261 

Phyteuma spicatum AM + NM 260, 261 

Pratia nummularia AM 422 

Trematolobelia kauaiensis AM 326 

Menyanthaceae 

Menyanthes trifoliata NM 260, 261 

Nymphoides peltata NM 260, 261 

Goodeniaceae 

Calogyne sp. AM + ECM 656 

Dampiera stricta AM + ECM 67 

Goodenia cycloptera AM 438 

Goodenia lunata AM 438 

Goodenia sp. AM + ECM 656 

Scaevola chamissoniana AM 326 

Scaevola gaudichaudii AM 326 

Scaevola parvibarbata AM 438 

Scaevola procera AM 326 

Scaevola sericea AM 326 

Asteraceae 

Achillea millefolium AM 194 

AM + NM 260, 261 

Achillea ptarmica AM 194 

NM 260, 261 

Ageratina espinosarum Weak AM 103 

Ageratum conyzoides AM 336, 422 

Ambrosia artemisifolia AM 143 

Anaphalis margaritacea Facultative AM 585 

AM 260, 261 

Angianthus sp. AM + ECM 656 

Antennaria dioica AM 194, 260, 261 

Anthemis arvensis AM 260, 261 

Anthemis tinctoria AM 260, 261 

Arctium lappa AM + NM 260, 261 

Arctium minus s. l. AM 260, 261 

Arnica angustifolia AM 443 

Arnica montana AM 194, 269 

Artemisia absinthium NM 260, 261 

Artemisia californica AM 646 

Artemisia campestris AM 260, 261 

Artemisia codonocephala AM 343 

Artemisia dracunculus AM 105 

Artemisia ludoviciana AM 633 

Artemisia maritima AM + NM 260, 261 

Artemisia princeps AM 643 

Artemisia tridentata AM 183 

Artemisia tridentata AM 552 

ssp. wyomingensis 

Artemisia vulgaris AM 307, 260, 261 

Aster novi belgii AM 260, 261 

Aster sericeus AM 633 

Aster subulatus AM 326 

Aster tripolium AM 113, 518, 260, 

261 

Baccharis racemosa AM 207 

Balduina angustifolia AM 30 

Bebbia juncea AM 112 

Bellis perennis AM + NM 260, 261 

Berkheya coddii AM 597

329 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Bidens asymmetrica × AM 227 

sandvicensis 

Bidens bipinnata AM 422 

Bidens cosmoides AM 326 

Bidens frondosa AM 140, 368 

Bidens pilosa AM 143, 343, 422 

Bidens sandwicensis AM 227, 326 

Bidens tripartita AM 140 

AM + NM 260, 261 

Blainvillea acmella AM 343 

Calendula officinalis AM 260, 261 

Callistephus chinensis AM 428 

Carduus acanthoides AM 307, 260, 261 

Carlina vulgaris AM 456, 260, 261 

Carthamus tinctorius AM 94 

Centaurea cyanus AM 260, 261 

Centaurea jacea AM 194, 260, 261 

Centaurea maculosa AM 109, 371 

Centaurea nigra AM 260, 261 

Centaurea pratensis AM 260, 261 

Centaurea scabiosa AM 260, 261 

Centaurea subciliata AM 368 

Chrysanthemum 

AM 551 

morifolium 

Chrysanthemum segetum AM 260, 261 

Cicerbita alpina AM + NM 260, 261 

Cichorium intybus AM 260, 261 

Cirsium acaule AM 260, 261 

Cirsium arvense AM 307, 260, 261 

Cirsium helenioides AM 260, 261 

Cirsium oleraceum AM 260, 261 

Cirsium palustre AM + NM 260, 261 

Cirsium purpuratum AM 636 

Cirsium setosum AM 620 

Cirsium vulgare AM 626 

AM + NM 260, 261 

Conyza blinii AM 343 

Conyza canadensis AM 326, 343 

Cotula plumose NM 342, 560 

Crassocephalum 

AM 422 

crepidioides 

Crepis biennis AM 260, 261 

Crepis capillaris AM 260, 261 

Crepis paludosa AM + NM 260, 261 

Crepis praemorsa AM 194 

Crepis setosa AM 207 

Cynara cardunculus AM 369 

Dittrichia viscosa AM 497 

Dubautia ciliolata AM 326 

Dubautia knudseni AM 326 

Dubautia plantaginea AM 326 

Dubautia scabra AM 326 

Eclipta alba AM 577 

Emilia sonchifolia AM 326, 336 

Erechtites hieracifolia AM 326 

Ericameria diffusa AM 112 

Erigeron acer NM 260, 261 

Erigeron annuus AM 643 

Erigeron bonariensis AM 643 

Erigeron borealis AM 260, 261 

Erigeron canadensis AM 294 

AM + NM 260, 261 

Eriophyllum lanatum AM 293 

Eupatorium cannabinum AM 260, 261 

Eupatorium coelesticum AM 422 

Eupatorium maculatum AM 140 

Eupatorium odoratum AM 422 

Eupatorium perfoliatum AM 140 

Eupatorium serotinum AM 599 

Filaginella uliginosa AM 194 

Florestina tripteris AM 143 

Flourensia cernua AM 137 

Galinsoga ciliata AM 260, 261 

Galinsoga parviflora AM 260, 261 

Galinsoga supinum AM 260, 261 

Galinsoga sylvaticum AM 260, 261 

Galinsoga uliginosum AM 260, 261 

Gnaphalium norvegicum AM 502, 503, 260, 

261 

Gnephosis eriocarpa Weak AM 438 

Gutierrezia sarothrae AM 492, 493 

Gymnosperma glutinosum Weak AM 103 

Haplopappus venetus AM 543 

Helianthus annuus AM 121, 260, 261 

Helianthus niveus AM 543 

Helichrysum italicum AM 368 

Helichrysum sp. AM + ECM 656 

Helichrysum stoechas AM 368 

Helipterum sp. AM + ECM 656 

Hieracium albiflorum Facultative AM 585 

Hieracium alpinum AM 260, 261 

Hieracium bifidum AM 260, 261 

Hieracium lachenalii AM 260, 261 

Hieracium lactucella AM 194 

Hieracium laevegatum AM + NM 260, 261 

Hieracium murorum AM + NM 260, 261 

Hieracium pilosella AM 270, 456, 260, 

261 

Hieracium sagittatum AM 260, 261 

Hieracium subaundum AM 260, 261 

Hieracium umbellatum AM 194, 260, 261 

Hieracium vulgatum AM + NM 260, 261 

Homogyne alpina AM + ECM 260, 261 

Hypochaeris radicata Facultative AM 584, 585, 260, 

261 

Hypochoeris achyrophorus AM 472

330 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Inula conyza AM 260, 261 

Inula salicina AM 260, 261 

Ixeris denticulate AM 129 

Jaumea carnosa AM 90 

Lactuca hirsuta AM 140 

Lactuca sativa AM 40, 171, 500 

Laggera pterodonta AM 343 

Lapsana communis AM 260, 261 

Launaea sarmentosa AM 336 

Leontodon autumnalis AM 194, 260, 261 

Leontodon hispidus AM 260, 261 

Leontodon hispidus AM 456 

ssp. danubialis 

Leontodon hispidus AM 456 

ssp. hispidus 

Leucanthemum vulgare AM 194, 260, 261 

Leuceria achillaeifolia NM 207 

Liatris tenuifolia 

Weak AM 30 

var. laevigata 

Lipochaeta connata AM 326 

Matricaria maritime AM 307 

ssp. inodora 

Matricaria matricarioides AM 260, 261 

Matricaria recutita AM + NM 260, 261 

Microseris laciniata AM 293 

Mutisia decurrens AM 207 

Mutisia spinosa AM 207 

Mycelis muralis AM + ECM + NM 260, 261 

Omalotheca norvegica AM 194 

Onopordum acanthium AM + NM 260, 261 

Otanthus maritimus AM 368 

Othonna gregorii AM + NM 438 

Palafoxia lindenii AM 144 

Parthenium argentatum AM 461 

Parthenium hysterophorus AM 343 

Pectis saturejoides AM 143 

Petasites albus AM + NM 260, 261 

Petasites frigidus NM 594 

Petasites hybridus AM + NM 260, 261 

Phagnalon rupestre AM 472 

Picris echioides AM 260, 261 

Picris hieracioides AM 260, 261 

Piptocarpha axillaris AM 651 

Pityopsis graminifolia Weak AM 30 

Pleurophyllum hookeri AM 342 

Podolepis sp. AM + ECM 656 

Polycalymma stuartii AM 438 

Porophyllum numularium AM 143 

Pulicaria angustifolia AM 567 

Pulicaria dysenterica AM 260, 261 

Remya kauaiensis AM 326 

Rhodanthe floribunda AM 438 

Rhodanthe moschata AM 438 

Sagittaria latifolia AM 58, 626 

Sanvitalia procumbens AM 260, 261 

Saussurea alpina AM 260, 261 

Scorzonera humilis AM 194 

Senecio articulatus AM 417 

Senecio bicolor AM 368, 260, 261 

Senecio bracteolatus AM 207 

Senecio erucifolius AM 260, 261 

Senecio jacobaea AM + NM 260, 261 

Senecio pendulus AM 417 

Senecio praecox AM 208 

NM 103 

Senecio squalidus AM 260, 261 

Senecio sylvaticus AM + NM 260, 261 

Senecio viscosus AM 307, 260, 261 

Senecio vulgaris AM 625 

AM + NM 260, 261 

Serratula tinctoria AM 212, 260, 261 

Siegesbeckia orientalis AM 343 

Solidago altissima AM 643 

Solidago canadensis AM 298, 626 

Solidago chilenesis AM 207 

Solidago gigantea AM 140 

Solidago graminifolia AM 140 

Solidago litoralis AM 368 

Solidago sp. AM 58 

Solidago virgaurea AM 194, 260, 261 

Sonchus arvensis AM 302 

AM + NM 260, 261 

Sonchus asper AM + NM 260, 261 

Sonchus oleraceus AM 343, 643, 260, 

261 

Spilanthes callimorpha AM 422 

Synedrella nudiflora AM 422 

Tagetes erecta AM 2, 102, 350 

Tagetes patula AM 322, 350, 554 

Tagetes tenuifolia AM 350 

Tanacetum parthenium NM 260, 261 

Tanacetum vulgare AM + NM 260, 261 

Taraxacum japonicum AM 643 

Taraxacum officinale AM 307 

Tithonia diversifolia AM 422, 539 

Tragopogon pratensis AM 260, 261 

Tridax procumbens AM 336 

Tripleurospermum AM 260, 261 

inodorum 

Tripleurospermum AM 260, 261 

maritimum 

Tussilago farfara AM 307 

AM + NM 260, 261 

Verbesina encelioides AM 326 

Vernonia noveboracensis AM 140 

Viguiera eriophora AM 103

331 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Waitzia sp. AM + ECM 656 

Wedelia trilobata AM 326 

Wilkesia gymnoxiphium AM 326 

Xanthium sibiricum AM 343 

Xanthocephalum 

AM 137 

microcephala 

Zinnia acerosa AM 137 

Zinnia elegans AM 2, 343 

Adoxaceae 

Adoxa moschatellina NM 260, 261 

Caprifoliaceae 

Carlemannia tetragona AM 655 

Linnaea borealis AM 260, 261 

Lonicera caprifolium AM 260, 261 

Lonicera implexa AM 368 

Lonicera pericylmenum AM 260, 261 

Lonicera xylosteum AM + NM 260, 261 

Sambucus nigra AM + NM 260, 261 

Sambucus racemosa AM + ECM+NM 260, 261 

Symphoricarpos rivularis AM + NM 260, 261 

Viburnum lantana AM 260, 261 

Viburnum lentago AM 140 

Viburnum opulus AM 260, 261 

Viburnum tinus AM 368 

Dipsacaceae 

Dipsacus fullonum AM + NM 260, 261 

Knautia arvensis AM 194, 260, 261 

Pycnocomon rutifolium AM 368 

Scabiosa columbaria AM 260, 261 

Scabiosa ochroleuca AM 456 

Succisa pratensis AM 194, 260, 261 

Valerianaceae 

Centranthus ruber NM 260, 261 

Valeriana dioica NM 260, 261 

Valeriana laxiflorac AM 207 

Valeriana officinalis AM + NM 260, 261 

Hamamelidaceae 

Liquidambar styraciflua AM 139 

Crassulaceae 

Aeonium decorum AM 417 

Crassula aquatica NM 260, 261 

Crassula moschata NM 342, 560 

Crassula phyturus AM 417 

Crassula rupestris AM 417 

Echeveria sedoides NM 417 

Kalanchoe daigremontana AM 417 

Sedum acre AM + NM 260, 261 

Sedum adolfii AM 417 

Sedum album NM 260, 261 

Sedum dasyphyllum NM 260, 261 

Sedum forsteranum NM 260, 261 

Sedum reflexum NM 260, 261 

Sedum rosea AM + NM 260, 261 

Sedum telephium NM 260, 261 

Sempervivum tectorum M(endo) + NM 260, 261 

Haloragaceae 

Myriophyllum alterniflorum NM 260, 261 

Proserpinaca palustris AM 140 

Grossulariaceae 

Ribes alpinum AM + ECM + NM 260, 261 

Ribes magellanicum AM 207 

Ribes nigrum AM 260, 261 

Ribes rubrum AM 260, 261 

Ribes uva-crispa AM 260, 261 

Saxifragaceae 

Chrysosplenium 

AM + NM 260, 261 

alternifolium 

Chrysosplenium 

NM 260, 261 

oppositifolium 

Saxifraga aizoides AM + NM 260, 261 

Saxifraga granulate AM 194 

AM + NM 260, 261 

Saxifraga hieracifolia NM 594 

Saxifraga hirculus NM 594 

Saxifraga oppositifolia AM + NM 260, 261 

Saxifraga stellaris NM 260, 261 

Vitaceae 

Cayratia japonica AM 643 

Leea compactiflora AM 655 

Leea indica AM 655 

Leea marcophylla AM 655 

Vitis vinifera AM 306, 517 

Geraniaceae 

Erodium cicutarium AM + NM 260, 261 

Erodium crinitum AM 438 

Geranium carolinianum AM 643 

Geranium dissectum AM 260, 261 

Geranium lucidum AM + NM 260, 261 

Geranium molle AM + NM 260, 261 

Geranium nodosum AM 260, 261 

Geranium pratense AM 260, 261 

Geranium robertianum AM 262, 472 

AM + NM 260, 261 

Geranium sanguineum NM 260, 261 

Geranium sessiliflorum AM 207 

Geranium sylvaticum AM 194, 260, 261 

Pelargonium hortorum AM 435 

Combretaceae 

Terminalia catappa AM 326 

Terminalia superba AM 445 

Onagraceae 

Camissonia californica AM 543 

Chamerion angustifolium AM + NM 260, 261 

Circaea alpina AM 260, 261 

Circaea lutetiana AM 260, 261

332 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Epilobium alsinifolium AM + NM 260, 261 

Epilobium anagallidifolium AM 260, 261 

Epilobium angustifolium Facultative AM 585 

Epilobium brunnescens NM 342 

ssp. brunnescens 

Epilobium ciliatum AM 260, 261 

Epilobium coloratum AM 140 

Epilobium hirsutum Facultative AM 62 

AM 260, 261 

Epilobium lanceolatum AM 260, 261 

Epilobium montanum AM + NM 260, 261 

Epilobium obscurum AM 260, 261 

Epilobium palustre AM 260, 261 

Epilobium parviflorum AM 260, 261 

Epilobium pendunculare AM 342 

Epilobium roseum AM 260, 261 

Epilobium tetragonum AM 260, 261 

Ludwigia palustris AM 140 

Ludwigia prostrata AM 422 

Oenothera biennis AM 260, 261 

Oenothera erythrosepala AM 260, 261 

Lythraceae 

Cuphea carthagenensis AM 326 

Lafoensia pacari AM 651 

Lythrum alatum AM 599 

Lythrum hyssopifolia NM 260, 261 

Lythrum maritimum AM 326 

Lythrum salicaria AM 140, 462, 558, 

629 

AM + NM 260, 261 

Myrtaceae 

Acca sellowiana AM 31 

Acmena resa AM 222 

Angophora hispida ECM 67 

Baeckea ramosissima ECM 67 

Campomanesia 

AM 651 

xanthocarpa 

Eucalyptus bosistoana AM 5 

Eucalyptus camaldulensis AM + ECM 179 

Eucalyptus citriodora AM 343 

AM + ECM 179 

Eucalyptus cloeziana AM + ECM 179 

Eucalyptus delegatensis AM 5 

Eucalyptus dives AM 5 

Eucalyptus dumosa Weak AM 5 

Eucalyptus dunnii AM + ECM 442 

Eucalyptus globoidea ECM 133 

Eucalyptus globulus AM 5 

ECM 359 

Eucalyptus globulus ECM 12 

ssp. bicostata 

Eucalyptus gomphocephala AM 5 

Eucalyptus grandis AM + ECM 179 

ECM 96 

Eucalyptus haemostoma ECM 67 

Eucalyptus largiflorens AM 5 

Eucalyptus macarthurii AM 5 

Eucalyptus marginata ECM 365 

Eucalyptus microcorys ECM 120 

Eucalyptus racemosa ECM 67 

Eucalyptus regnans ECM 341 

Eucalyptus sieberi ECM 133 

Eucalyptus stricta ECM 67 

Eucalyptus torelliana ECM 120 

Eucalyptus urophylla AM 5 

AM + ECM 179 

Eucalyptus viminalis AM 5 

AM + ECM 66 

Eucalyptus viridis AM 5 

Eugenia uniflora AM 651 

Gomidesia spectabilis AM 31 

Kunzea capitata ECM 67 

Leptospermum flavescens ECM 67 

Leptospermum lanigerum ECM 67 

Luma apiculata AM 207 

Melaleuca quinquenervia AM + ECM 316 

Metrosideros polymorpha AM 326 

Myrtus communis AM 368, 382, 472 

Plinia rivularis Weak AM 651 

Psidium cattleianum AM 326 

Psidium guajava AM 31, 198, 337 

Rhodomyrtus tomentosa AM 326 

Syzygium firmum AM 215 

Syzygium guineense AM 637, 638 

Syzygium jambolanum AM 548 

Syzygium latilimbum AM 422 

Syzygium makul AM 215 

Syzygium operculatum AM 215 

Syzygium rubicundum AM 215 

Tristaniopsis whiteana NM 568 

Melastomataceae 

Clidemia hirta AM 326 

Graffenrieda emarginata AM + ECM 266 

Leandra dasytricha AM 31 

Melastoma affine NM 422 

Melastoma melabathricum AM 568 

Krameriaceae 

Krameria paucifolia AM 112 

Zygophyllaceae 

Balanites aegyptiaca AM 47 

Fagonia cretica AM 567 

Larrea tridentata AM 112, 137 

Morkillia mexicana Weak AM 103 

Tribulus cistoides NM 326 

Tribulus terrester AM 343

333 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Viscainoa geniculata AM 112 

Zygophyllum dumosum AM 268 

Zygophyllum howittii NM 438 

Parnassiaceae 

Parnassia palustris AM + NM 260, 261 

Celastraceae 

Euonymus europaeus AM 260, 261 

Maytenus boaria AM 207 

Maytenus chubutensis AM 207 

Maytenus magellanica AM 207 

Maytenus phyllantoides AM 112 

Humiriaceae 

Sacoglottis gabonensis AM 445 

Pandaceae 

Panda oleosa AM 445 

Rhizophoraceae 

Combretocapus rotundatus NM 568 

Poga oleosa AM 445 

Linaceae 

Linum bienne AM 472 

Linum catharticum AM 456 

AM + NM 260, 261 

Linum perenne 

NM 260, 261 

ssp. anglicum 

Linum strictum AM 472 

Linum trigynum AM 472 

Linum usitatissimum AM 171, 180 

AM + NM 260, 261 

Erythroxylaceae 

Erythroxylon compactum NM 103 

Euphorbiaceae 

Acalypha acmophylla AM 343 

Actinostemon concolor AM 651 

Adelia virgata AM 112 

Alchornea cordifolia AM 445 

Alchornea tiliaefolia AM 422, 655 

Aleurites moluccana AM 326 

Amperea xiphoclada ECM 67 

Baccaurea ramiflora AM 422, 655 

Breynia fruticosa AM 422 

Celaenodendron 

AM 20 

mexicanum 

Chamaesyce atrococca AM 326 

Chamaesyce celastroides AM 326 

Chamaesyce degeneri AM 326 

Chamaesyce hypericifolia AM 326 

Cleidion bracteosum AM 422 

Cleidion brevipetiolatum AM 655 

Cleistanthus sumatranus AM 422 

NM 655 

Croton ciliatoglanduliferus 

Weak AM 103 

Croton floribundus AM 651 

Croton kongensis AM 422 

Croton machrostachyus AM 637, 638 

Croton punctatus AM 143 

Croton urucurana AM 651 

Ditaxis lanceolata AM 112 

Drypetes gossweileri AM 445 

Drypetes hoaensis NM 655 

Epiprinus silhetianus NM 655 

Euphorbia amygdaloides AM 260, 261 

Euphorbia cyparissias AM 260, 261 

Euphorbia dulcis AM 260, 261 

Euphorbia esula AM 260, 261 

Euphorbia gatbergensis AM 417 

Euphorbia helioscopia AM + NM 260, 261 

Euphorbia heterophylla AM 343 

Euphorbia hirta AM 216, 343 


Euphorbia lathyrus AM 260, 261 

Euphorbia paralias AM 260, 261 

Euphorbia parliramulosa AM 417 

Euphorbia peplis AM 368 

Euphorbia peplus AM 472 

AM + NM 260, 261 

Euphorbia royleana AM 343 

Euphorbia tannensis AM 438 

Euphorbia thymifolia AM 343 

Glochidion assamicum NM 655 

Hevea brasiliensis AM 288, 526, 527 

Hieronyma alchorneoides AM 31 

Jatropha cinerea AM 112 

Jatropha cuneata AM 112 

Macaranga denticulata AM 422, 648 

Macaranga indica AM 655 

Macaranga peltata AM 315 

Manihot esculanta AM 252 

Mercurialis annua AM 260, 261 

Mercurialis perennis AM + NM 260, 261 

Pedilanthus cymbiferus NM 103 

AM 112 

Pedilanthus macrocarpus AM 112 

Phyllanthus asteranthus AM 422 

Phyllanthus lacunarius AM 438 

Phyllanthus niruri AM 577 

Phyllanthus urinaria AM 343 

Ricinodendron heudelotii AM 445 

Ricinus communis AM 318 

Sauropus androgynus AM 253 

Sebastiania 

AM 651 

commersoniana 

Uapaca acuminata AM + ECM 445 

Uapaca guineense ECM 52, 445 

Uapaca heudelotti AM 52

334 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Uapaca staudtii ECM+AM 413 

ECM 445 

Uapaca vanhouttei ECM 445 

Violaceae 

Viola arvensis AM + NM 260, 261 

Viola calaminaria AM 588 

Viola canina AM 269 

AM + NM 260, 261 

Viola hirta AM 260, 261 

Viola kitaibeliana AM + NM 260, 261 

Viola lutea AM 260, 261 

Viola maculata AM 207 

Viola odorata AM 260, 261 

Viola palustris AM + NM 260, 261 

Viola reichenbachiana AM + NM 260, 261 

Viola riviniana AM 260, 261 

Viola rupestris AM 260, 261 

Viola tricolor AM 194, 456 

AM + NM 260, 261 

Viola wailenalenae AM 326 

Viola × wittrockiana AM 322 

Salicaceae 

Casearia sylvestris AM 31, 651 

Populus alba ECM+AM 368 

ECM+NM 260, 261 

Populus balsamifera ECM 470 

Populus balsamifera ECM 271 

ssp. trichocarpa 

Populus × canadensis AM + ECM 260, 261 

Populus canescens ECM 260, 261 

Populus deltoides AM 140 

Populus euroamericana AM + ECM 317 

Populus nigra AM + ECM 260, 261 

Populus nigra ‘Italica’ AM + ECM 260, 261 

Populus serotina AM + ECM 260, 261 

Populus tremula AM + ECM 260, 261 

Populus tremula × ECM 59 

tremuloides 

Populus tremuloides ECM 145, 338 

ECM+AM 430 

Populus trichocarpa ECM 59, 260, 261 

Salix alba ECM 260, 261 

Salix alba vitellina ECM 260, 261 

Salix arbuscula ECM 260, 261 

Salix arctica ECM 594 

Salix aurita ECM+NM 260, 261 

Salix babylonica AM + ECM 316, 260, 261 

Salix caprea AM + ECM 260, 261 

Salix cinerea AM + ECM 260, 261 

Salix daphnoides ECM 260, 261 

Salix fragilis ECM 260, 261 

Salix gracilistyla AM 511 

Salix herbacea ECM+EEM 260, 261 

Salix lapponum ECM 260, 261 

Salix myrsinites ECM 260, 261 

Salix nigra AM 140 

Salix nigricans ECM+NM 260, 261 

Salix phylicifolia ECM 260, 261 

Salix polaris ECM 594 

Salix purpurea AM + ECM 260, 261 

Salix reinii ECM 587 

Salix repens AM 605 

AM + ECM 260, 261 

Salix reticulata ECM 594, 260, 261 

Salix triandra ECM 260, 261 

Salix viminalis AM 60 

AM + ECM 260, 261 

Xylosma ciliatifolium AM 651 

Xylosma pseudosalzmannii AM 651 

Turneraceae 

Turnera ulmifolia AM 143 

Clusiaceae 

Calophyllum sclerophyllum AM 568 

Calophyllum soulattri AM 568 

Clusia minor AM 474 

Clusia rotundata AM 474 

Cratoxylum arborescens AM 568 

Garcinia cowa NM 655 

Garcinia kola AM 445 

Garcinia lucida AM 445 

Garcinia xanthochymus Facultative AM 422 

Hypericaceae 

Hypericum calycinum NM 260, 261 

Hypericum hirsutum NM 260, 261 

Hypericum humifusum NM 260, 261 

Hypericum maculatum AM 194, 260, 261 

Hypericum montanum NM 260, 261 

Hypericum perforatum AM 409, 260, 261 

Hypericum pulchrum AM 260, 261 

Hypericum tetrapterum AM + NM 260, 261 

Elatinaceae 

Elatine hexandra AM 62 

Ochnaceae 

Lophira alata AM 445 

Chrysobalanaceae 

Licania platypus AM 321 

Oxalidaceae 

Oxalis acetosella AM 598 

AM + NM 260, 261 

Oxalis corniculata AM 294, 643, 260, 

261 

Oxalis corymbosa AM 643 

Oxalis exilis AM 260, 261 

Oxalis europaea AM 260, 261 

Oxalis stricta AM 260, 261 

Oxalis valdiviensis AM 207

335 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Cunoniaceae 

Ceratopetalum apetalum AM 387 

ECM 389 

Elaeocarpaceae 

Aristotelia chilensis AM 207 

Elaeocarpus austroyunnanensis 

AM 422 

Sloanea guianensis AM 31 

Polygalaceae 

Polygala nicaeensis AM 368 

Polygala amarella AM + NM 260, 261 

Polygala calcarea AM 260, 261 

Polygala serpyllifolia AM 260, 261 

Polygala vulgaris AM 194 

AM + ECM 260, 261 

Salomonia sp. 


via AM 

Fabaceae 

Acacia albida AM 174 

Acacia ampliceps AM 477 

Acacia arabica AM 317 

Acacia auriculiformis AM 52, 229 

Acacia bonariensis AM + ECM 211 

Acacia caven AM 211 

Acacia cochliacantha Weak AM 103 

Acacia constricta Weak AM 103 

Acacia eriopoda AM 477 

Acacia farnesiana AM 343, 600 

Weak AM 103 

Acacia holosericea ECM 186 

Acacia koa AM 326, 406 

Acacia leptocarpa AM 52 

Acacia linifolia AM 68 

ECM+AM 67 

Acacia mangium ECM 186, 187 

AM 52, 257, 520 

Acacia nilotica AM 473, 538 

Acacia obtusifolia ECM+AM 67 

Acacia pennatula AM 18 

Acacia polyphylla AM 651 

Acacia suaveolens ECM+AM 67 

Acacia tortilis AM 567 

Acacia ulicifolia ECM+AM 67 

Adenanthera pavonina AM 52 

Aeschynomene indica AM 546 

Afzelia africana NM 47 

ECM 45, 52 

Afzelia bipidensis ECM+AM 413, 445 

Afzelia pachyloba AM + ECM 445 

Afzelia quanzensis ECM 418 

Albizia corniculata AM 655 

Albizia falcataria AM 14 

Albizia ferruginea AM 52, 256 

Albizia gummifera AM 637, 638 

Albizia hassleri AM 651 

Albizia kalkora AM 343 

Albizia lebbeck AM 52, 216, 477, 

538, 548 

Albizia schimperiana AM 637, 638 

Alysicarpus regosus AM 336 

Alysicarpus vaginalis AM 343 

Amorpha crenulata AM 202 

Amphimas ferrugineus AM 445 

Amphimas pterocarpoides AM 52, 445 

Anadenanthera colubrina AM 452, 651 

Anadenanthera falcata AM 548 

Anadenanthera 

AM 651 

macrocarpa 

Anadenanthera peregrina AM 528, 593 

Anthonotha fragrans ECM 52 

ECM+AM 413, 445 

Anthonotha macrophylla ECM 52, 413, 445 

Anthyllis cytisoides AM 170, 234, 583 

Anthyllis tetraphylla AM 472 

Anthyllis vulneraria AM 456 

AM + NM 260, 261 

Aphanocalyx 

ECM 413 

cynometroides 

Arachis hypogaea AM 111 

Astragalus alpinus NM 594 

Astragalus applegatei AM 55 

Astragalus glycyphyllos NM 260, 261 

Atylosia scarabaeoides AM 343 

Baphia nitida AM 52 

Bauhinia forficata AM 211, 651 

Berlinia bracteosa ECM 445 

Berlinia confusa ECM 52, 445 

Bossiaea obcordata AM 67 

Brachystegia 

ECM 445 

cynometroides 

Brachystegia eurycoma ECM 445 

Brachystegia leonensis ECM 52 

Brachystegia zenkeri ECM 445 

Bussea occidentalis AM 52 

Caesalpinia eriostachys AM 20 

Caesalpinia ferrea AM 548 

Caesalpinia gilliesii AM 211 

Caesalpinia pannosa AM 112 

Caesalpinia peltophoroides AM 548 

Cajanus cajan AM 172, 624 

Calliandra eriophylla Weak AM 103 

Calliandra parvifolia Weak AM + ECM 211 

Calliandra selloi AM 211 

Calliandra tweedii AM + ECM 211 

Calopogonium caeruleum AM 289 

Calopogonium mucunoides AM 439

336 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Canavalia rosea AM 144, 336 

Cassia grandis AM 548 

Cassia siamea AM 52, 230, 251 

Cassia spectabilis AM 258 

Castanospermum australe AM 4 

Cathormion polyanthum AM 211 

Centrolobium tomentosum AM 372, 651 

Ceratonia siliqua AM 147 

Cercidium praecox AM 112 

Chamaecrista 

AM 143 

chamaecristoides 

Cicer arietinum AM 124, 125 

Colophospermum mopane AM 477 

Copaifera langsdorffii AM 548, 651 

Cordyla pinnata AM 47 

Coronilla emerus AM 368 

ssp. emeroides 

Coronilla scorpioides AM 472 

Coronilla varia AM 159 

NM 260, 261 

Crotalaria burhia AM 567 

Crotalaria incana AM 143 

Crotalaria medicaginea AM 343, 567 

Cullen discolor AM 438 

Cullen pallida AM 438 

Cylicodiscus gabunensis AM 445 

Cynometra alexandri AM 591 

Cynometra leonensis AM 52 

Cynometra sanagaensis AM 445 

Cytisus scoparius AM 260, 261 

Dalbergia obtusifolia AM 655 

Dalbergia sissoo AM 317, 473 

Daniella ogea AM 445 

Daviesia corymbosa AM 67 

Desmanthus illinoensis AM 541 

Desmodium heterophyllum AM 355 

Desmodium microphyllum AM 343 

Desmodium oxyphyllum AM 643 

Desmodium paniculatum AM 541 

Detarium macrocarpum AM 445 

Dialium guineense AM 47, 52 

Dicorynia guianensis AM 71, 161 

Didelotia africana ECM + Weak AM 413 

ECM 445 

Didelotia letouzeyi ECM 413, 445 

Dillwyinia parvifolia ECM+AM 67 

var. tricopoda 

Dillwynia retorta ECM 67 

AM 589 

Diphysa robinoides AM 143 

Dipteryx panamensis AM 321 

Distemonanthus 

AM 445 

benthamianus 

Dorycnium hirsutum NM 368 

Enterolobium 

AM 211, 651 

contortisiliquum 

Enterolobium cyclocarpum AM 143, 256 

Eperua falcata AM 161 

Erythrina berteroana AM 142 

Erythrina crista-galli AM 211 

Erythrina poeppigiana AM 148 

Erythrophleum ivorense AM 52, 445 

Eysenhardtia polystachya Weak AM 103 

Faidherbia albida AM 46 

Genista hirsute AM 339 

Genista tinctoria AM 260, 261 

Geoffroea decorticians AM 211 

Gilbertiodendron ECM 445 

brachystegioides 

Gilbertiodendron dewevrei ECM+AM 591 

ECM 445 

Gilbertiodendron preussi ECM 52 

Gleditsia amorphoides Weak AM + ECM 211 

Gliricidia sepium AM 52, 199, 258 

Glyceria maxima NM 62 

Glycine max AM 37 

Gompholobium 

AM 67 

grandiflorum 

Gossweilerodendron AM 445 

balsamiferum 

Guibourtia tessmannii AM 445 

Havardia albicans AM 18 

Hedysarum confertum AM 496 

Hedysarum coronarium AM 351 

Hedysarum spinosissimum AM 496 

Hippocrepis comosa AM 260, 261 

Hippocrepis unisiliquosa AM 472 

Holocalyx balansae Weak AM 651 

Hymenaea courbaril NM 548 

Indigofera brevidens Weak AM 438 

Indigofera cordifolia AM 567 

Indigofera linifolia AM 567 

Inga leiocalycina AM 627 

Inga marginata AM 31 

Inga sessilis AM 651 

Inga striata AM 651 

Inga urguuensis AM 211 

Intsia palembanica AM + ECM 14 

Julbernardia seretii ECM+AM 591 

ECM 445 

Koompassia malacensis NM 568 

Kummerowia striata AM 213 

Lablab purpureusus AM 640 

Laburnum anagyroides AM 260, 261

337 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Lathyrus montanus AM 260, 261 

Lathyrus japonicus AM 260, 261 

Lathyrus niger AM + NM 260, 261 

Lathyus odoratus AM 260, 261 

Lathyrus pratensis AM 194 

AM + NM 260, 261 

Lens culinaris AM 319 

Leucaena leucocephala AM 18, 52, 254, 

326 

Lonchocarpus campestris AM 651 

Lonchocarpus 

Weak AM 651 

muehlbergianus 

Lonchocarpus nitidus AM + ECM 211 

Lotus corniculatus AM 140, 159, 194, 

307, 456 

AM + NM 260, 261 

Lotus glaber AM 508 

Lotus tenuis AM 396 

Lotus uliginosus AM 260, 261 

Lupinus arcticus NM 594 

Lysiloma candida AM 112 

Machaerium minutiflorum AM 651 

Machaerium stipitatum Weak AM 651 

Macrotyloma uniflorum AM 453 

Medicago littoralis AM 368 

Medicago lupulina AM 260, 261 

Medicago marina AM 368 

Medicago polymorpha AM 294, 260, 261 

Medicago sativa AM 38, 159, 351, 

260, 261 

Melilotus alba AM 260, 261 

Melilotus officinalis AM 159, 260, 261 

Microberlinia bisulcata ECM 413 

Mimosa adenantheroides Weak AM 103 

Mimosa bimucronata AM 452 

Mimosa biuncifera Weak AM 103 

AM 82 

Mimosa calcicola Weak AM 103 

Mimosa lacerata Weak AM 103 

Mimosa luisana AM 103 

Mimosa polyantha Weak AM 103 

Mimosa pudic AM 326 

Mimosa purpusii Weak AM 103 

Mimosa scabrella AM 31, 651 

Mimosa texana Weak AM 103 

Monopetalanthus le-testui ECM 445 

Monopetalanthus ECM 445 

microphyllus 

Mora excelsa AM 592 

Olneya tesota AM 112 

Onobrychis viciifolia AM 260, 261 

Onobrychis sativa AM 260, 261 

Ononis repens AM 260, 261 

Ononis spinosa AM 260, 261 

Ormosia arborea NM 548 

AM 651 

Oxystigma buchholzii AM 445 

Oxystigma mannii AM 445 

Oxytropis scammaniana AM 594 

Pachyelasma tessmannii AM 445 

Paraberlinia bifoliolata ECM 445 

Parapiptadenia rigida AM 211, 452, 651 

Parkia bicolor AM 52 

Parkia biglobosa AM 47, 247 

Parkia speciosa AM 14 

Parkinsonia aculeata AM 211 

Peltophorum dubium AM 211 

NM 548, 651 

Pentaclethra macrophyll AM 52, 445 

Phaseolus lunatus AM 20 

Phaseolus mungo AM 478 

Phaseolus radiatus AM 478 

Phaseolus vulgaris AM 37 

Piptadeniastrum africanum AM 52, 445 

Plagiosiphon longitubus AM 445 

Plagiosiphon multijugus AM 445 

Platycyamus regnellii AM 548 

Poecilanthe parviflora AM 211, 651 

Prioria copaifera AM 592 

Prosopis articulata AM 112 

Prosopis cineraria AM 379 

Prosopis glandulose AM 137 

Prosopis juliflora AM 76, 175, 567 

Prosopis laevigata AM 495 

Psoralea bituminosa AM 472 

Pterocarpus angolensis AM 419 

Pterocarpus mildbraedii AM 445 

Pterocarpus soyauxii AM 445 

Pterogyne nitens Weak AM 651 

Pueraria phaseoloides AM 150 

Pultenaea elliptica ECM + AM 67 

Retama sphaerocarpa AM 16, 17, 497 

Robinia pseudoacacia ECM 330 

AM + ECM 582, 260, 261 

ECM + EEM 86 

Samanea saman AM 52 

Schizolobium parahyba AM 548 

Schrankia quadrivalvis AM 143 

Scorodophloeus zenkeri AM 445 

Scorpiurus muricatus AM 472 

Senna corymbosa AM 211 

Senna macranthera AM 548, 651 

Senna multijuga AM 548 

Senna siamea AM 441 

Senna spectabilis AM 548 

Sesbania aegyptiaca AM 231

338 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Sesbania grandiflora AM 231 

Sesbania punicea AM 211 

Sesbania tomentosa AM 326, 227 

Sophora chrysophylla AM 326, 406 

Spartium junceum AM 368 

Strophostyles helvola AM 596 

Strophostyles umbellata AM 532 

Stryphnodendron AM 36 

microstachyum 

Stylosanthes guvanensis AM 545 

Stylosanthes 

AM 545 

scarbaeoidesand 

Swainsona phacoides AM 438 

Tamarindus indica AM 47, 247, 343 

Tephrosia purpurea AM 216, 343 

Tephrosia sphaerospora AM 438 

Tetraberlinia bifoliolata ECM + Weak AM 413 

ECM 445 

Tetraberlinia moreliana ECM 413 

Tipuana tipu AM 548 

Toubaouate 

ECM 445 

brevipaniculata 

Trifolium arvense AM 307 

Trifolium aureum AM 194 

Trifolium dubium AM 643, 260, 261 

Trifolium fragiferum NM 260, 261 

Trifolium glomeratum AM 260, 261 

Trifolium hybridum AM 307 

AM + NM 260, 261 

Trifolium medium AM 260, 261 

Trifolium pratense AM 140, 159, 194, 

204, 456 

AM + NM 260, 261 

Trifolium repens AM 140, 194, 283, 

643, 260, 261 

Trifolium spadiceum AM 194 

Trifolium subterraneum AM 171 

AM + NM 260, 261 

Trigonella foenumgraecum AM 220 

Ulex europaeus AM 368, 260, 261 

Ulex gallii NM 260, 261 

Ulex parviflorus AM 497 

Vicia angustifolia AM 643 

var. segetalis 

Vicia cracca AM 194, 294, 260, 

261 

Vicia faba AM 191 

Vicia nigricans AM 207 

Vicia hirsute AM 643, 260, 261 

Vicia sativa AM 260, 261 

Vicia sepium AM 194, 260, 261 

Vicia sylvatica AM + ECM 260, 261 

Vicia tetrasperma AM 307, 260, 261 

Vicia villosa AM 140 

Vigna aconitifolius AM 343 

Vigna luteola AM 273 

Vigna marina AM 326, 547 

Vigna mungo AM 319 

Vigna parkeri AM 440 

Vigna radiate AM 377 

Vigna unguiculata AM 37, 287 

Xylia xylocarpa AM 315 

Zornia sp. Weak AM 103 

Rosaceae 

Acaena magellanica AM 342 

Acaena minor NM 560 

AM 342 

Acaena novae-zelandiae AM 260, 261 

Acaena ovalifolia AM 207 

Acaena pinnatifida AM 207 

Adenostoma fasciculatum AM + ECM 21 

Agrimonia eupatoria AM 260, 261 

Alchemilla alpina AM 80, 194, 260, 

261 

Alchemilla glabra AM 260, 261 

Alchemilla glaucescens AM 194 

Alchemilla glomerulans AM 503 

Alchemilla vulagris agg. AM + NM 260, 261 

Alchemilla xanthochlora AM 260, 261 

Aphanes arvensis AM 260, 261 

Cotoneaster integerrimus AM 260, 261 

Crataegus laevigata AM + ECM 260, 261 

Crataegus monogyna ECM 368 

AM + ECM 260, 261 

Dryas octopetala ECM 80, 594 

AM + ECM 260, 261 

Duchesnea chrysantha AM 643 

Filipendula ulmaria AM + NM 260, 261 

Fragaria chiloensis AM 207 

Fragaria vesca AM + NM 260, 261 

Fragaria xananassa AM 537 

Geum rivale AM 194 

AM + NM 260, 261 

Geum urbanum AM 260, 261 

Geum virginianum AM 140 

Hagenia abyssinica AM 637, 638 

Malus communis AM 260, 261 

Malus domestica AM 368 

Malus micromalus AM 26 

Malus pumila 

AM 233 

var. domestica 

Malus sylvestris AM + ECM 260, 261 

Osteomeles anthyllidifolia AM 326 

Ovidia andina AM 207 

Potentilla anglica AM 260, 261

339 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Potentilla anserine AM 307, 626, 260, 

261 

Potentilla argentea AM 260, 261 

Potentilla cinerea AM 456 

Potentilla crantzii AM 194, 260, 261 

Potentilla erecta AM 194, 586 

AM + NM 260, 261 

Potentilla fruticosa AM 260, 261 

Potentilla hookeriana AM 443 

Potentilla palustris AM + NM 260, 261 

Potentilla pulchella AM 443 

Potentilla reptans AM 260, 261 

Potentilla sterilis AM 260, 261 

Potentilla tabernaemontani AM 260, 261 

Prunus africana AM 637, 638 

Prunus avium AM 465 

AM + ECM 260, 261 

Prunus cerasifera AM 73 

Prunus cerasus ECM 260, 261 

Prunus domestica AM + NM 260, 261 

Prunus maritima AM 324 

Prunus padus AM + ECM 260, 261 

Prunus sellowii Weak AM 651 

Prunus spinosa AM + NM 260, 261 

Pyrus communis AM 219 

Pyrus pyraster AM + ECM 260, 261 

Rosa arvensis AM 260, 261 

Rosa canina AM 81, 260, 261 

Rosa hybrida AM 160 

Rosa rubiginosa AM 207 

Rosa rugosa AM 324 

Rosa sempervirens AM 368 

Rubus caesius AM 260, 261 

Rubus chamaemorus NM 260, 261 

Rubus fruticosus s. l. AM 260, 261 

Rubus idaeus AM 573, 574 

AM + NM 260, 261 

Rubus nessensis AM + ECM 260, 261 

Rubus rufus AM 422 

Rubus saxatilis AM 260, 261 

Rubus ulmifolius AM 368 

Sanguisorba minor AM 260, 261 

Sanguisorba officinalis AM 260, 261 

Sibbaldia procumbens AM 594 

NM 260, 261 

Sorbus aria AM + ECM 260, 261 

Sorbus aucuparia AM + ECM + NM 260, 261 

Sorbus torminalis ECM 260, 261 

Spiraea latifolia AM 140 

Spiraea tomentosa AM 140 

Rhamnaceae 

Colletia hystrix AM 207 

Colubrina glabra AM 112 

Colubrina glandulosa AM 651 

Colubrina oppositifolia AM 227 

Condalia globosa AM 112 

Discaria articulata AM 207 

Discaria chacaye AM 207 

Frangula alnus AM + ECM + NM 260, 261 

Hovenia dulcis AM 548 

Phylica ericoides AM 24 

Rhamnus catharticus AM + NM 260, 261 

Rhamnus lycioides AM 17, 108 

Ventilago calyculata AM 655 

Zizyphus mauritiana AM 47, 343, 378, 

381, 655 

Zizyphus nummularia AM 567 

Zizyphus obtusifolia AM 112 

Ziziphus xylopyrus AM 380 

Elaeagnaceae 

Elaeagnus commutata AM 613 

Hippophae rhamnoides AM 405, 260, 261 

Hippophae tibetana AM 581 

Shepherdia canadensis AM 581 

Ulmaceae 

Celtis wightii AM 422 

Gironniera subaequalis NM 422 

Trema micrantha AM 548, 651 

Trema orientalis AM 445 

Ulmus glabra AM 260, 261 

Ulmus procera AM + ECM 260, 261 

Cannabaceae 

Humulus lupulus AM + NM 260, 261 

Moraceae 

Artocarpus altilis AM 136 

Brosimum alicastrum AM 18 

Brosimum lactescens AM 31 

Broussonetia papyrifera AM 343 

Ficus cyrtophylla NM 655 

Ficus guaranitica AM 651 

Ficus hirta AM 422 

Ficus hispida AM 655 

Ficus langkokensis AM 655 

Milicia excelsa AM 445 

Milicia regia AM 279 

Morus alba AM 475, 535 

Musanga cecropioides AM 445 

Sorocea bonplandii Weak AM 651 

Urticaceae 

Boehmeria zollingeriana AM 422 

Cecropia glaziovii AM 651 

Cecropia pachystachya AM 651 

Elatostema parvum AM 422 

Parietaria judaica AM + NM 260, 261 

Pipturus albidus AM 326

340 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Touchardia latifolia NM 326 

Urera glabra AM 326 

Urtica dioica NM 611 

AM + NM 260, 261 

Urtica urens NM 260, 261 

Cucurbitaceae 

Bryonia dioica AM 260, 261 

Citrullus lanatus AM 310, 438 

Ctenolepis cerasiformis AM 567 

Cucumis callosus AM 567 

Cucumis sativus AM 499 

Cucurbita foetidissima AM 396 

Cucurbita pepo AM 340 

Mukia maderaspatana AM 216 

Begoniaceae 

Begonia angustinei AM 422 

Begonia cathayana AM 422 

Hillebrandia sandwicensis AM 326 

Tetramelaceae 

Tetrameles nudiflora AM 422 

Fagaceae 

Castanea sativa ECM 260, 261 

Castanopsis borneensis ECM 267 

Castanopsis indica NM 422 

AM 655 

Fagus sylvatica ECM 529, 260, 261 

Lithocarpus densiflora ECM 375 

Lithocarpus leucostachyus NM 422 

Nothofagus dombeyi ECM 207 

Quercus alba ECM 444, 470 

Quercus cerris ECM 260, 261 

Quercus ilex ECM 368, 472, 260, 

261 

Quercus petraea ECM 53, 260, 261 

Quercus pubescens ECM 138 

Quercus robur ECM 53, 368, 260, 

261 

Quercus rubra ECM 214 

Myricaceae 

Comptonia peregrina Facultative AM 284 

Myrica cerifera AM 468 

Myrica faya AM 326 

Myrica gale Facultative AM 146 

AM + NM 260, 261 

Myrica pensylvanica NM 324 


Juglandaceae 

Engelhardtia roxburghiana ECM 267 

Juglans nigra AM 465 

Juglans regia AM 177 

AM + ECM + NM 260, 261 

Betulaceae 

Alnus acuminata ECM 61 

Alnus cordata AM 361 

Alnus glutinosa AM 210 

ECM + AM 368 

ECM 471 

AM + ECM + 260, 261 

EEM + NM 

Alnus incana AM 34 

AM + ECM 260, 261 

Alnus sinuate ECM 271 

Alnus tenuifolia ECM 641 

Betula alleghaniensis ECM 162, 516 

Betula lenta ECM 619 

Betula nana ECM 594, 260, 261 

Betula papyrifera ECM 303 

Betula pendula ECM 92, 155 

ECM+EEM 260, 261 

Betula pubescens ECM 370 

ECM+EEM 260, 261 

Betula verrucosa ECM 217 

Carpinus betulus ECM 529, 260, 261 

Corylus avellana ECM 367, 529, 260, 

261 

Casuarinaceae 

Allocasuarina littoralis ECM 579 

Casuarina 

Weak AM 485 

cunninghamiana AM 316 

Weak ECM 579 

Casuarina equisetifolia AM 447, 326, 542, 

606 

Casuarina equisetifolia ECM 579 

ssp. equisetifolia 

Tropaeolaceae 

Tropaeolum majus AM 360, 610 

Moringaceae 

Moringa concanensis AM 450 

Caricaceae 

Carica papaya AM 360, 595 

Bataceae 

Batis maritime NM 326 

Resedaceae 

Reseda lutea AM + NM 260, 261 

Reseda luteola AM + NM 260, 261 

Capparidaceae 

Atamisquea emarginata AM 112 

Capparis sandwichiana NM 326 

Brassicaceae 

Alliaria petiolata NM 260, 261 

Alyssum montanum NM 456 

Arabidopsis thaliana Weak AM + NM 260, 261 

Arabis alpina NM 260, 261 

Arabis hirsuta NM 472, 260, 261

341 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Armoracia rusticana AM + NM 260, 261 

Barbarea vulgaris NM 260, 261 

Biscutella laevigata NM 456 

AM 446, 260, 261 

Blennodia pterosperma NM 438 

Brassica napus AM + NM 260, 261 

Brassica nigra NM 83, 260, 261 

Brassica oleracea AM + NM 260, 261 

Brassica rapa NM 260, 261 

Cakile maritima NM 368 

AM + NM 260, 261 

Capsella bursa-pastoris Facultative AM 167 

AM 164 

AM + NM 260, 261 

Cardamine amara NM 260, 261 

Cardamine bulbifera AM + NM 260, 261 

Cardamine concatenata NM 165 

Cardamine corymbosa NM 342 

Cardamine flexuosa AM + NM 260, 261 

Cardamine hirsuta AM + NM 260, 261 

Cardamine impatiens NM 260, 261 

Cardamine pratensis NM 260, 261 

Cardamine trifolia NM 260, 261 

Cardaminopsis arenosa AM 456 

Cochlearia anglica AM + NM 260, 261 

Cochlearia danica AM 260, 261 

Cochlearia officinalis AM + NM 260, 261 

Diplotaxis erucoides AM + NM 260, 261 

Diplotaxis muralis AM + NM 260, 261 

Erophila verna NM 260, 261 

Erysimum cheiranthoides NM 260, 261 

Hesperis matronalis Facultative AM 167 

Iberis amara AM 260, 261 

Isatis tinctoria NM 260, 261 

Lepidium bidentatum AM 326 

Lepidium phlebopetalum NM 438 

Lepidium ruderale NM 260, 261 

Lepidium serra NM 326 

Matthiola incana Facultative AM 167 

Nasturtium officinale NM 260, 261 

Pringlea antiscorbutica NM 560 

Raphanus raphanistrum NM 260, 261 

Raphanus sativus AM + NM 260, 261 

Rorippa amphibia NM 62 

Rorippa islandica AM 140 

Rorippa palustris NM 260, 261 

Sinapis alba NM 260, 261 

Sisymbrium loeselii NM 260, 261 

Sisymbrium officinale AM + NM 260, 261 

Sisymbrium orientale NM 260, 261 

Thlaspi arvense NM 260, 261 

Thlaspi caerulescens AM 488 

Thlaspi montanum AM 488 

Thlaspi praecox AM 488, 614 

Thymelaeaceae 

Daphne laureola AM 260, 261 

Daphne gnidium AM 368 

Daphne mezereum AM + NM 260, 261 

Gonystylus bancanus AM 568 

Pimelea linifolia AM 67 

Wikstroemia furcata AM 326 

Bixaceae 

Cochlospermum vitifolium AM 20 

Cistaceae 

Cistus creticus ECM 368 

Cistus ladanifer ECM 339 

Cistus incanus ECM 368 

Cistus monspeliensis ECM 368 

ECM + AM 472 

Fumana thymifolia AM 472 

Helianthemum almeriense EEM + ECM 249 

Helianthemum apenninum AM + ECM + EEM 260, 261 

Helianthemum canum AM + ECM + EEM 260, 261 

Helianthemum guttatum ECM + EEM 209 

Helianthemum 

ECM 80 

nummularium 

AM + ECM + EEM 260, 261 

Helianthemum ovatum ECM 330 

Tuberaria guttata ECM 260, 261 

Dipterocarpaceae 

Hopea mengarawan NM 568 

Shorea balangeran AM 568 

Shorea leprosula ECM 14 

Shorea teysmanniana AM 568 

Shorea uliginosa AM 568 

Malvaceae 

Abelmoschus esculentus AM 69 

Abutilon californicum AM 112 

Abutilon indicum AM 343 

Abutilon otocarpum AM 438 

Abutilon theophrasti AM 553 

Althaea officinalis AM 260, 261 

Bastardiopsis densiflora AM 651 

Gossypium hirsutum AM 7, 649 

Gossypium tomentosum AM 326 

Kokia kauaiensis AM 326 

Malva sylvestris AM 260, 261

342 



Mycorrhizal 

status b,c 

References 



Mycorrhizal 

status b,c 

References 

Thespesia populnea AM 326 

Sida ammophila NM + Weak AM 438 

Sida cordifolia AM 567 

Sida fallax AM 326 

Sida szechuensis AM 343 

Malvaceae (Bombacoideae) 

Adansonia digitata AM 47 

Bombax malabaricum AM 343 

Ceiba pentandra AM 18 

Ceiba speciosa AM 548 

Chorisia speciosa AM 651 

Ochroma pyramidale AM 321 

Pseudobombax 

AM 651 

grandiflorum 

Malvaceae (Tilioideae) 

Corchorus tridens AM 567 

Grewia biloba AM 343 

Heliocarpus americanus AM 651 

Heliocarpus 

AM 244, 245 

appendiculatus 

Heliocarpus pallidus AM 20 

Luehea candicans AM 651 

Luehea divaricata AM 651 

Luehea grandiflora AM 548 

Luehea seemannii AM 321 

Tilia cordata ECM 529 

AM + ECM + NM 260, 261 

Tilia platyphyllos ECM 549, 260, 261 

Tilia × vulgaris ECM 260, 261 

Malvaceae (Sterculioideae) 

Eribroma oblonga AM 445 

Guazuma ulmifolia AM 18, 32, 651 

Pterigota macrocarpa AM 445 

Pterospermum 

AM 422 

menglunense 

Pterospermum yunnanensis AM 655 

Theobroma cacao AM 131, 321 

Waltheria indica AM 143, 326 

Rutaceae 

Acronychia pedunculata AM 655 

Agathosma ovata AM 24 

Boronia thujona AM 67 

Citrus aurantifolia AM 400 

Citrus aurantium AM 429 

Citrus jambhiri AM 176 

Eriostemon australasius AM 67 

Fagara heitzii AM 445 

Flindersia brayleana AM 222 

Melicope barbigera AM 326 

Paramignya retispina AM 655 

Zanthoxylum rhetsa AM 315 

Meliaceae 

Azadirachta excelsa AM 282 

Azadirachta indica AM 255, 427, 477 

Cabralea canjerana AM 31 

Cedrella fissilis AM 548, 593, 651 

Chisocheton siamensis Facultative AM 422 

Ekebergia capensis AM 637, 638 

Entandrophragma AM 445 

angolense 


candollei 


cylindricum 

Entandrophragma utile AM 445 

Guarea cedrata AM 445 

Guarea kunthiana Weak AM 651 

Khaya ivorensis AM 445 

Lansium domesticum AM 422 

Lovoa trichilioides AM 445 

Melia azedarach AM 422 

Melia toosanden NM 422 

Swietenia macrophylla AM 433 

Trichilia casaretti Weak AM 651 

Trichilia claussenii Weak AM 651 

Trichilia elegans Weak AM 651 

Simaroubaceae 

Castella peninsularis AM 112 

Irvingia gabonensis AM 445 

Anacardiaceae 

Anacardium excelsum AM 321 

Anacardium occidentale AM 27, 47 

Antrocaryon klaineanum AM 445 

Astronium graveolens AM 651 

Campnosperma 

AM 568 

auriculatum 

Cyrtocarpa edulis AM 112 

Mangifera sp. AM 486 

Pistacia atlantica AM 197 

Pistacia lentiscus AM 368, 472 

Pistacia palestina AM 197 

Pistacia terebinthus AM 104 

Pistacia vera AM 197 

Schinus patagonicus AM 207 

Schinus terebinthifolius AM 548, 651 

Sclerocarya birrea AM 47 

Trychoscypha acuminata AM 445 

Burseraceae 

Bursera fagaroides NM 103 

AM 112 

Bursera hindsiana Weak AM 112 

Bursera microphylla Weak AM 112 

Canarium schweinfurthii AM 445 

Sapindaceae 

Acer campestre AM + ECM + NM 260, 261

343 



Mycorrhizal 

status b,c 

References 

Acer platanoides AM 607 

AM + ECM + NM 260, 261 

Acer pseudoplatanus AM 465, 622 

AM + ECM + NM 260, 261 

Acer rubrum AM 140 

Acer saccharum AM 141, 162, 470 

Aesculus hippocastanum AM 260, 261 

Allophylus schweinfurthii AM 445 

Aphania senegalensis AM 47 

Blighia welwitschii AM 445 

Cardiospermum corindum AM 112 

Deinbollia pycnophylla AM 445 

Dodonaea viscosa AM 326, 343, 479 

Eriocoelum macrocarpum AM 445 

Harpullia cupanioides AM 655 

Lepisanthes senegalensis Facultative AM 422 

Litchi chinensis AM 326 

Matayba guianensis AM 31 

Pometia tomentosa AM 422 

NM 655 

Sapindus saponaria AM 548 

Some unplaced taxa 

Bruniaceae 

Staavia radiata AM 24 

Escalloniaceae 

Escallonia rubra AM 207 

a The families in column 1 are divided into four groups: bryophytes, 

pteridophytes, gymnosperms and angiosperms. For each group, the 

order of families is consistent with the phylogeny used in Fig. 1 

b The mycorrhizal types shown in column 2 are consistent to the 

current widely used system of Smith and Read (1997): AM 

arbuscular mycorrhiza, ECM ectomycorrhiza, EEM ectendomycorrhiza, 

ORM orchid mycorrhiza, ERM ericoid mycorrhiza, 

MTM monotropoid mycorrhiza, ABM arbutoid mycorrhiza, 

as well as NM nonmycorrhiza, and mycoheterotrophy 

c For the bryophytes, fungal associations were used instead of 

mycorrhiza; A, B, and G indicate ascomycetous, basidiomycetous, 

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mosses (Fig. 1; but see Boullard 1988 for mycorrhizoids in 

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Furthermore, liverworts are clearly favored as the earliest 

divergent lineage of extant land plants according to emerging 

molecular evidence (Qiu et al. 1998; Dombrovska and Qiu 

2004; Groth-Malonek et al. 2005; Qiu et al. unpublished), 

despite some lingering controversy on relationships among 

three bryophyte lineages: liverworts, mosses, and hornworts 

(see Qiu and Lee 2000; Renzaglia et al. 2000). Hence, the 

widespread occurrence of fungal association in liverworts 

deserves some attention here. 

Mycorrhizas, or more precisely mycorrhizoids and 

mycothalli, are found in many of the liverworts that have 

been investigated (Pocock and Duckett 1985; Boullard 

1988; Duckett et al. 1991). Several aspects of these 

symbioses revealed by recent studies may help clarify their 

evolutionary relationship to mycorrhizas in vascular plants. 

First, in cases where detailed ultrastructural analysis has 

been done, structures similar to those in the endomycorrhizas 

of vascular plants, i.e. arbuscules, have been 

observed in some liverworts (Read et al. 2000; Carafa et 

al. 2003; Russell and Bulman 2005). Second, crossinoculation 

experiments have convincingly demonstrated 

that the same fungi can infect both liverworts and 

angiosperms and produce similar structures for either 

glomeromycetous or ascomycetous mycorrhizas (Read et 

al. 2000). Third, although isotopic tracing of nutrient, 

water, and carbon exchange has not been done for most 

liverwort–fungus symbioses, the carbon flow from Betula 

through the ectomycorrhizal fungus Tulasnella to the 

mycoheterotrophic liverwort Cryptothallus has been confirmed 

by a 14 C feeding experiment (Bidartondo et al. 

2003). Finally, the evolutionary pattern of plant–fungus 

symbiosis observed in angiosperms by Trappe (1987) and 

in vascular plants (this study), i.e., glomeromycetous 

mycorrhizas being the common type and occurring in 

early-diverging plant lineages and ascomycetous and 

basidiomycetous mycorrhizas being the rare type and 

found in late-evolving plant lineages, is also seen in 

liverworts (Fig. 1). These four lines of evidence indicate 

that even though the plant structures involved in symbiosis 

are probably structurally and functionally analogous 

between liverworts and vascular plants, the nature of the 

biological interaction is almost certain to be evolutionarily 

homologous. Besides further investigations of the functional 

aspects of mycorrhizas in liverworts and other 

bryophytes as suggested by Read et al. (2000), one line of 

evidence that will provide conclusive support for evolutionary 

homology between the mycorrhizal symbioses in 

bryophytes and vascular plants could come from studies of 

plant genes that are involved in the processes, some of

Table 2 Mycorrhizal investigation of four major groups of land plants at both family and species levels 

Group Total number of 

families/species 

surveyed 

No. (%) of mycorrhizal 

families (obligate and 

facultative) 

No. (%) of 

facultatively 

mycorrhizal species 


obligately 

mycorrhizal 

species 


nonmycorrhizal 

families 


nonmycorrhizal 

species 

Bryophytes 28/143 20 (71%) 60 (42%) 6 (4%) 8 (29%) 77 (54%) 

Pteriophytes 28/426 26 (93%) 185 (43%) 39 (9%) 2 (7%) 202 (47%) 

Gymnosperms 12/84 12 (100%) 83 (99%) 1 (1%) 0 (0%) 0 (0%) 

Angiosperms 195/2,964 184 (94%) 2,141 (72%) 396 (13%) 11 (6%) 427 (14%) 

Sum 263/3,617 242 (92%) 2,469 (68%) 442 (12%) 21 (8%) 706 (20%) 

353 

which have been identified in recent studies (Stracke et al. 

2002; Liu et al. 2003; Ane et al. 2004; Demchenko et al. 

2004; Levy et al. 2004). If the same genes are found to 

control development of mycorrhizas in all groups regardless 

of their expression in the gametophytes of bryophytes 

or the sporophytes of vascular plants, and if the gene 

phylogenies are congruent to the plant phylogeny, the final 

doubt about homology of the two types of symbioses will 

be dispelled. 

If the mycorrhizal symbiosis in liverworts, other bryophytes, 

such as the moss Takakia (see Boullard 1988), and 

hornworts proves to be homologous to that in vascular 

plants, it becomes obvious that mycorrhiza was indeed an 

important means of water and nutrient uptake adopted by the 

earliest land plants (Pirozynski and Malloch 1975). Fossil 

evidence is in agreement with such a scenario. Both 

glomeromycetous fungi (Redecker et al. 2000) andliverworts 

(Wellman et al. 2003) evolved by the Ordovician, 

when plants colonized the land (Kenrick and Crane 1997). 

Fossil mycorrhizas formed between nonseptate fungi and the 

protracheophyte Aglaophyton major have been uncovered 

from the Rhynie chert, confirming that this type of plant– 

fungus interaction existed at least 400 million years ago 

(Remy et al. 1994;Kerpetal.2004; Taylor et al. 2005). It is 

possible that evolution of mycorrhiza might even have 

predated the origin of land plants, if fungal association in the 

extinct charophyte Palaeonitella (Taylor et al. 1992) is 

mycorrhizal. We conclude that although more work is 

needed to solidify the relationship between mycorrhizal 

symbioses in bryophytes and vascular plants, including 

morphological, physiological, and molecular characterization 

of the plant–fungus interactions in both bryophytes and 

early vascular plants (see Read et al. 2000), all available 

evidence seems to point to an origin of mycorrhizas at the 

beginning of land plant evolution. 

Arbuscular mycorrhiza is the ancestral type 

of mycorrhiza 

When the different types of mycorrhizas are mapped onto 

the land plant phylogeny, it becomes clear that arbuscular 

mycorrhiza (AM), the endomycorrhiza formed by glomeromycetous 

fungi and plants, is the ancestral type, as it 

occurs in a vast majority of plants and in all early-diverging 

lineages of major clades of land plants (Fig. 1). Even in 

bryophytes where the plant structures involved in symbiosis 

are rhizoids and thalli of the gametophyte, glomeromycetous 

fungi-mediated symbiosis results in formation 

of AM-like structures and is found in most early-diverging 

lineages of liverworts and in hornworts (Ligrone and Lopes 

1989; Read et al. 2000; Russell and Bulman 2005). These 

two aspects of AM symbiosis suggest that the genetic 

mechanism underlying interaction between the plants and 

glomeromycetous fungi might have been established in the 

common ancestor of all land plants. Future molecular 

studies should be able to test this hypothesis to determine if 

the same genes are involved in plants–glomeromycetous 

fungi symbioses throughout land plants. 

Trappe (1987) reported that AM is the ancestral type of 

mycorrhiza in angiosperms. Our survey, which broadens 

the scope to include all land plants, extends this conclusion 

to all land plants. This result is not likely to change even if 

relationships among three bryophyte lineages and vascular 

plants are altered (see Qiu and Lee 2000) because all of 

their early-diverging lineages possess AM when they are 

mycorrhizal. Furthermore, fossil evidence clearly indicates 

that AM is the most ancient type of mycorrhiza. The 

Devonian protracheophyte Aglaophyton major provides 

the best fossil record for existence of AM-like symbiosis as 

early as 400 million years ago; the arbuscules are visible in 

the protosteles of this early Devonian plant (Remy et al. 

1994; Taylor et al. 2005). Fossil roots of younger age from 

the Triassic show that this type of symbiosis has been 

present continuously in early land plant evolution 

(Stubblefield et al. 1987). On the other hand, the oldest 

ectomycorrhizal fossils have so far been found only in the 

middle Eocene (50 million years ago) (LePage et al. 

1997). The fossil record of glomeromycetous fungi, found 

in the Ordovician (Redecker et al. 2000), also clearly 

predates that of ascomycetous fungi unearthed from the 

Lower Devonian (Taylor et al. 1999). Given all these lines 

of evidence, it seems safe to conclude that AM represents 

the ancestral type of mycorrhiza in land plants. 

Many independent conversions of arbuscular 

mycorrhiza to other types of mycorrhizas 

If the ancestral state of AM in land plants is accepted, and 

further, if it can be agreed upon that the glomeromycetous 

fungi–plant interaction is underpinned by a genetic mech-

354 

anism that has been stably inherited since its establishment in 

the common ancestor of all land plants, the phylogenetic 

distribution of AM and other types of mycorrhiza shown in 

Fig. 1 leads to the conclusion that there have been many 

independent conversions of AM to these other types of 

mycorrhizas. Brundrett (2002) previously proposed these 

evolutionary changes. In this review, we find overwhelming 

evidence in support of this hypothesis. Many plant lineages, 

ranging from pteridophytes through gymnosperms to angiosperms, 

contain some species that can form both AM and 

other types of mycorrhizas, e.g., Dryopteridaceae, Pinaceae, 

Gnetaceae, Araucariaceae, Cupressaceae, Orchidaceae, Cyperaceae, 

Poaceae, Ranunculaceae, Polygonaceae, Caryophyllaceae, 

Ericaceae, Rubiaceae, Oleaceae, Aquifoliaceae, 

Campanulaceae, Goodeniaceae, Asteraceae, Caprifoliaceae, 

Grossulariaceae, Myrtaceae, Melastomataceae, Euphorbiaceae, 

Salicaceae, Polygalaceae, Fabaceae, Rosaceae, 

Rhamnaceae, Juglandaceae, Betulaceae, Casuarinaceae, 

Cistaceae, Malvaceae, and Sapindaceae (Table 1; Fig. 1). 

These species might be in a transitional state from AM to 

other types of mycorrhizas. These other types of mycorrhizas 

have probably evolved as a consequence of 

emergence of new lineages in fungi on one side (e.g., 

certain groups of Ascomycetes and Basidiomycetes) and in 

plants on the other side (e.g., Pinaceae, Orchidaceae, 

Ericaceae, Fagales, and Malvales), and the associations 

formed by these new lineages that were able to colonize 

habitats that were previously not very successfully 

occupied by AM plants (Malloch et al. 1980). Because 

these other types of mycorrhizas have been classified using 

both morphological and plant partner criteria (Smith and 

Read 1997), we follow this classification in our discussion 

below. 

Ectomycorrhiza 

Ectomycorrhiza (ECM) is a partnership between mostly 

basidiomycetous fungi and various groups of land plants. It 

primarily occurs in Pinaceae, Araucariaceae, Cupressaceae, 

Gnetaceae, Polygonaceae, Nyctaginaceae, Myrtaceae, 

Salicaceae, Fabaceae, Fagales, and Malvales. In addition, 

many other families of vascular plants contain a small 

percentage of species that form this type of mycorrhiza. 

Most leafy liverworts also have ascomycetous and basidiomycetous 

fungal associations (Table 1; Fig. 1). Although 

ECM is much rarer than AM in land plants, among all types 

of mycorrhizas formed by ascomycetous and basidiomycetous 

fungi, it is the most common one. When viewed in a 

plant phylogenetic perspective, the distribution of ECM 

clearly suggests many independent origins of this type of 

mycorrhiza because its occurrence is sporadic throughout 

land plants and it is mostly found in derived lineages in 

major plant clades. Previously, Bruns and Shefferson 

(2004) estimated at least 12 independent origins of ECM 

within angiosperms by mapping ectomycorrhizal information 

provided in Smith and Read (1997) onto an angiosperm 

phylogeny (Soltis et al. 2000). Fitter and Moyersoen 

(1996) conducted a similar survey and reached a similar 

conclusion. Our study revealed many more events of 

evolution of this type of mycorrhiza in land plants (Fig. 1). 

Examination of the evolutionary histories of both plants 

and fungi may provide some ideas to understand how this 

type of mycorrhiza evolved repeatedly. The plants that 

form ECM typically grow in nutrient-poor environments, 

e.g., temperate and timberline forests of both Southern and 

Northern Hemispheres (Malloch et al. 1980), the spruce– 

hemlock–redwood forest of the coastal fog belt in the 

Pacific Northwestern USA, the wet eucalyptus forest in 

Australia, and the wet dipterocarp forest of Australasia (J. 

Trappe, personal communication). Phylogenetically, these 

plants represent derived lineages of some major land plant 

clades that once lived in a less nutrient-deficient environment, 

e.g., Pinaceae and Cupressaceae, as opposed to other 

conifers such as Cephalotaxaceae, Taxaceae, and Taxodiaceae, 

and Fagales and some Malvales vs other rosids. 

Meanwhile, the fungal species that participate in ECM 

symbiosis evolved repeatedly from saprotrophic homobasidiomycetes 

and also reverted to free-living condition 

many times independently (Hibbett et al. 2000), and they 

also represent derived lineages of fungi as a whole (Lutzoni 

et al. 2004). Thus, evolution of ECM could be viewed as an 

adaptation by those major plant clades to the change of 

environment when the climate on earth became more 

seasonal and arid (Hickey and Doyle 1977; Malloch et al. 

1980), which immobilized nutrients, or simply as an 

adaptation to the environments that were limited in nutrient 

availability. From these considerations, it may be argued 

that the plasticity exhibited by ECM evolution reflects the 

opportunistic nature of a collective response by both plant 

and fungal partners to environmental challenges. This 

seems to be a short-term evolutionary strategy in comparison 

to AM, which clearly represents a major evolutionary 

innovation to solve a permanent water-and-nutrients deficiency 

problem faced by plants when they first made the 

virtually irreversible invasion onto the land in the 

Ordovician. 

Fig. 1 A phylogenetic tree of 263 land plant families mapped with" 

mycorrhizal information. a Non-angiosperms (including bryophytes, 

pteridophytes, and gymnosperms), b angiosperms except core 

eudicots, c core eudicots except rosids, and d rosids. For each 

family, the number in the column Total shows the total number of 

examined species; the M% column shows the percentage of 

obligately mycorrhizal species. The FM% column shows the 

percentage of facultatively mycorrhizal species, and the NM% 

column shows the percentage of nonmycorrhizal species. Thicker 

terminal branches were used to indicate 21 nonmycorrhizal 

families. For each of the 242 mycorrhizal families, the percentages 

of different types of mycorrhizae among all mycorrhizal species are 

shown in the M type column. The mycorrhizal types are consistent to 

the current widely used system of Smith and Read (1997): AM 

arbuscular mycorrhiza, ECM ectomycorrhiza, EEM ectendomycorrhiza, 

ORM orchid mycorrhiza, ERM ericoid mycorrhiza, MTM 

monotropoid mycorrhiza, ABM arbutoid mycorrhiza, as well as 

mycoheterotrophy. For the bryophytes, fungal associations were 

used instead of mycorrhiza: A, B, and G indicate ascomycetous, 

basidiomycetous, and glomeromycetous fungal associations, respectively. 

1 Families in which ectomycorrhizal species have been 

reported. 2 Families in which mycoheterotrophic species have been 

reported

a 

angiosperms 

Total 

M% FM% NM% 

Taxaceae 

1 

Cupressaceae 

Taxodiaceae 

2 

Podocarpaceae 

1 

Araucariaceae 

1 

Gnetaceae 

2 

11 

3 

6 

5 

3 

100 

100 

100 

100 

100 

100 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

AM only 

82% AM,18% AM+ECM 

AM only 

83% AM, 17% mycoheterotrophy? 

60% AM, 20% ECM, 20% AM+EEM 

67% ECM, 33% AM+ECM 

Welwitschiaceae 

Ephedraceae 

Pinaceae 

1 

Ginkgoaceae 

1 

2 

43 

1 

100 

100 

98 

100 

0 

0 

2 

0 

0 

0 

0 

0 

AM only 

AM only 

86% ECM, 7% AM+ECM, 5% ECM+EEM, 2% AM+ECM+EEM 

AM only 

Zamiaceae 

Cycadaceae 

Davalliaceae 

Grammitidaceae 

Polypodiaceae 

Nephrolepidaceae 

1 

Dryopteridaceae 

Blechnaceae 

Thelypteridaceae 

Aspleniaceae 

Dennstaedtiaceae 

Adiantaceae 

5 

2 

4 

8 

37 

3 

40 

11 

33 

99 

20 

25 

100 

100 

75 

38 

8 

33 

53 

64 

36 

32 

60 

48 

0 

0 

0 

0 

3 

0 

8 

9 

6 

12 

5 

12 

0 

0 

25 

63 

89 

67 

40 

27 

58 

56 

35 

40 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

96 %AM, 4% AM+ECM 

AM only 

AM only 

AM only 

AM only 

AM only 

Vittariaceae 

Pteridaceae 

Dicksoniaceae 

Cyatheaceae 

Plagiogyriaceae 

3 

22 

4 

6 

1 

33 

45 

75 

33 

100 

0 

9 

0 

0 

0 

67 

45 

25 

67 

0 

AM only 

AM only 

AM only 

AM only 

AM only 

Azollaceae 

Marsileaceae 

Schizaeaceae 

Gleicheniaceae 

Hymenophyllaceae 

Osmundaceae 

Ophioglossaceae 

Psilotaceae 

Marattiaceae 

Equisetaceae 

Selaginellaceae 

Isoetaceae 

Lycopodiaceae 

Anthocerotaceae 

Gymnomitriaceae 

Jungermanniaceae 

Calypogeiaceae 

1 

1 

4 

10 

11 

3 

11 

3 

17 

12 

17 

3 

17 

2 

7 

21 

8 

0 

0 

75 

40 

36 

67 

91 

67 

88 

25 

76 

0 

35 

100 

0 

48 

100 

0 

100 

25 

0 

0 

33 

9 

0 

0 

42 

6 

0 

24 

0 

14 

0 

0 

100 

0 

0 

60 

64 

0 

0 

33 

12 

33 

18 

100 

41 

0 

86 

52 

0 

AM only 

AM only 

AM only 

50% AM, 50% endo-M (also AM?) 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

fungal association (G), AM-like 

fungal association 

fungal association (A, B) 

fungal association (A, B) 

Cephaloziaceae 

Cephaloziellaceae 

Scapaniaceae 

Lepidoziaceae 

Geocalycaceae 

Arnelliaceae 

Plagiochilaceae 

Herbertaceae 

Pseudolepicoleaceae 

Lejeuneaceae 

Jubulaceae 

Porellaceae 

Radulaceae 

Codoniaceae 

Pelliaceae 

2 

Aneuraceae 

Metzgeriaceae 

Ricciaceae 

Conocephalaceae 

Aytoniaceae 

Marchantiaceae 

Lunulariaceae 

Blasiaceae 

Haplomitriaceae 

Characeae 

14 

6 

8 

9 

10 

1 

5 

1 

1 

11 

6 

5 

3 

2 

3 

5 

6 

1 

1 

1 

2 

1 

1 

2 

71 

67 

13 

30 

100 

20 

0 

0 

0 

0 

0 

0 

100 

100 

40 

0 

0 

100 

100 

100 

100 

0 

100 

14 

0 

13 

0 

10 

0 

0 

0 

100 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

14 

33 

75 

33 

60 

0 

80 

100 

0 

100 

100 

100 

100 

0 

0 

60 

100 

100 

0 

0 

0 

0 

100 

0 

fungal association (A) 





fungal association (B) 



fungal association (G) 


ORM-like (mycoheterotrophy via ECM) 




fungal association (G) 


bryophytes pteridophytes gymnosperms 

67 

M type 

355

356 

Fig. 1 (continued) 

b 

core eudicots 

Gunneraceae 

Buxaceae 

Proteaceae 

1 

Ranunculaceae 

Berberidaceae 

Menispermaceae 

Papaveraceae 

Piperaceae 

Aristolochiaceae 

Lauraceae 

Annonaceae 

Magnoliaceae 

Myristicaceae 

Costaceae 

Zingiberaceae 

Cannaceae 

Heliconiaceae 

Musaceae 

Commelinaceae 

Poaceae 1 

Restionaceae 

1 

Cyperaceae 

Juncaceae 

Bromeliaceae 

Typhaceae 

Sparganiaceae 

Arecaceae 

Ruscaceae 

Asparagaceae 

Agavaceae 

Hyacintheaceae 

0 

Themidaceae 

Amaryllidaceae 

Alliaceae 

0 

20 

8 

Asphodelaceae 

0 

Hemerocallidaceae 

0 

Iridaceae 

Hypoxidaceae 

13 

0 

Asteliaceae 

0 

1 2 

Orchidaceae 

2 

Liliaceae 

Smilacaceae 

Melanthiaceae 

20 

0 

33 

Colchicaceae 

0 

Alstroemeriaceae 

0 

Corsiaceae 

2 

0 

Cyclanthaceae 

0 

Pandanaceae 

0 

AM only 

Triuridaceae 

2 

Dioscoreaceae 

2 

Burmanniaceae 

2 

2 

50 

100 

0 

0 

0 

50 

0 

mycoheterotrophy via AM 

AM only 


Nartheciaceae 

1 0 100 0 AM only 

2 

Petrosaviaceae 2 100 0 0 mycoheterotrophy 

AM only 

AM only 

Potamogetonaceae 

Juncaginaceae 

Limnocharitaceae 

Alismataceae 

Butomaceae 

Hydrocharitaceae 

Araceae 

Tofieldiaceae 

Acoraceae 

Nymphaeaceae 

monocots magnoliids basal eudicots 

Total 

1 

1 

6 

39 

5 

1 

6 

1 

13 

6 

3 

5 

2 

5 

1 

1 

1 

3 

267 

2 

253 

27 

4 

3 

5 

9 

6 

2 

9 

5 

1 

5 

12 

3 

1 

8 

1 

1 

83 

5 

4 

3 

1 

1 

1 

1 

3 

2 

7 

2 

1 

7 

1 

4 

18 

1 

1 

5 

100 

100 

67 

54 

80 

100 

83 

0 

100 

83 

67 

100 

100 

100 

0 

100 

100 

67 

71 

100 

35 

37 

75 

33 

40 

89 

17 

100 

100 

100 

100 

80 

92 

100 

100 

75 

100 

100 

98 

80 

100 

67 

100 

100 

100 

0 

33 

100 

non-angiosperm land plants 

M% FM% NM% 

0 

0 

0 

57 

0 

25 

78 

100 

100 

20 

0 

0 

0 

33 

0 

0 

0 

100 

0 

0 

0 

0 

0 

0 

0 

0 

0 

33 

20 

0 

19 

37 

0 

67 

20 

0 

83 

0 

0 

14 

50 

0 

29 

0 

0 

0 

0 

0 

0 

0 

0 

33 

13 

20 

0 

4 25 25 50 

17 

0 

0 

17 

33 

0 

0 

0 

100 

0 

0 

0 

9 

0 

46 

26 

25 

0 

40 

11 

0 

0 

0 

0 

0 

0 

0 

0 

0 

13 

0 

0 

0 

0 

0 

0 

0 

0 

0 

100 

67 

0 

M type 

AM only 

AM only 

AM only 

97% AM, 3% AM+ECM 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

99.6% AM, 0.4% AM+ECM 

AM only 

99.3% AM, 0.7% AM+ECM 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

82% ORM, 4% AM, 8% ORM+ECM, 6% mycoheterotrophy via ECM 

AM only 

AM only 

AM only 

AM only 

AM only 


86 

50 

100 

14 AM only 

100 

75 AM only 

22 AM only 

0 AM only 

0 AM only 

80 AM only

357 


c 

rosids 

Valerianaceae 

Dipsacaceae 

1 

Caprifoliaceae 

Adoxaceae 

1 

Asteraceae 

1 

Goodeniaceae 

Menyanthaceae 

Campanulaceae 

1 

1 

Apiaceae 

Pittosporaceae 

Araliaceae 

1 

Aquifoliaceae 

Solanaceae 

Convolvulaceae 

Lentibulariaceae 

Scrophulariaceae 

Acanthaceae 

Lamiaceae 

Verbenaceae 

Bignoniaceae 

Plantaginaceae 

Pedaliaceae 

Gesneriaceae 

1 

Oleaceae 

Apocynaceae 

Loganiaceae 

2 

Gentianaceae 

1 

Rubiaceae 

Boraginaceae 

Icacinaceae 

1 2 

Ericaceae 

Clethraceae 

Actinidiaceae 

Sapotaceae 

Lecythidaceae 

Diapensiaceae 

Styracaceae 

Myrsinaceae 

Primulaceae 

Theophrastaceae 

Ebenaceae 

Theaceae 

Fouquieriaceae 

Polemoniaceae 

Tetrameristaceae 

Balsaminaceae 

Cornaceae 

Loasaceae 

Santalaceae 

Loranthaceae 

Olacaceae 

Portulacaceae 

Cactaceae 

Nyctaginaceae 

1 

Phytolaccaceae 

Aizoaceae 

Amaranthaceae 

1 

Caryophyllaceae 

Polygonaceae 

1 

Plumbaginaceae 

Tamaricaceae 

Frankeniaceae 

Droseraceae 

Dilleniaceae 

caryophyllids asterids 

Total 

25 

83 

69 

0 

84 

25 

17 

31 

0 

13 

50 

0 

33 

100 

3 

2 

100 0 0 

0 0 100 

14 5 

24 11 

0 0 

0 33 

0 0 

10 5 

0 8 

0 100 

26 34 

0 0 

16 6 

0 0 

0 11 

0 0 

0 0 

0 0 

0 0 

0 17 

0 0 

19 6 

9 5 

22 19 

0 0 

0 0 

0 0 

0 0 

0 0 

0 0 

0 0 

0 0 

0 0 

44 0 

0 0 

0 0 

0 0 

0 0 

0 0 

0 

60 

0 

0 

100 0 

0 100 

0 75 

4 

6 

13 

1 

233 

10 

21 

46 

1 

9 

3 

20 

12 

2 

58 

6 

69 

15 

9 

10 

1 

2 

13 

35 

2 

16 

55 

36 

1 

78 

1 

1 

8 

3 

1 

1 

5 

18 

1 

1 

1 

1 

1 

1 

5 

1 

2 

4 

1 

3 

8 

31 

8 

1 

4 

42 

54 

38 

3 

2 

2 

3 

1 

M% FM% NM% 

81 

65 

100 

67 

100 

85 

92 

0 

40 

100 

78 

100 

89 

100 

100 

100 

100 

83 

100 

75 

85 

58 

100 

100 

100 

100 

100 

100 

100 

100 

100 

56 

100 

100 

100 

100 

100 

100 

40 

0 

0 

25 

0 

100 

13 

90 

38 

100 

25 

36 

15 

29 

33 

100 

50 

0 

100 

0 

0 

13 

0 

13 

0 

50 

19 

26 

24 

67 

0 

0 

67 

0 

M type 

AM only 

AM only 

92% AM, 8% AM+ECM 

97% AM, 3% AM+ECM 

70% AM, 30% AM+ECM 

95% AM, 5% AM+ECM 

98% AM, 2% ECM 

AM only 

AM only 

67% AM, 33% AM+ECM 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

92% AM, 8% AM+ECM 

AM only 

AM only 

53% AM, 47% mycoheterotrophy via AM 

98% AM, 2% AM+ECM 

AM only 

AM only 

74% ERM, 8% AM+ERM, 5% EEM, 4% MTM, 9% others* 

AM only 

AM only 

AM only 

AM only 

ERM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

100 

0 

75 

10 

50 

0 AM only 

25 AM only 

45 AM only 

59 

47 

0 

0 

50 AM only 

AM only 

AM only 

AM only 

50% AM, 50% ECM 

95% AM, 5% AM+ECM 

80% AM, 10% ECM, 10% AM+ECM 

AM only 

AM only 

33 AM only 

0 AM only 

* Ericaceae: MTM is mycoheterotrophic. 9% others include 1.3% AM, 1.3% endoM, 1.3% ABM, 1.3% ABM+ECM, 

1.3% ABM+ERM, 1.3% Mono M+ERM, 1.3% ERM+ABM+ECM+EEM

358 


d 

Total 

M% FM% NM% 

M type 

1 

Sapindaceae 

Burseraceae 

Anacardiaceae 

Simaroubaceae 

Meliaceae 

Rutaceae 

1 

Malvaceae 

Dipterocarpaceae 

Cistaceae 

1 

Bixaceae 

Thymelaeaceae 

Brassicaceae 

Capparidaceae 

Resedaceae 

Bataceae 

Caricaceae 

Moringaceae 

Tropaeolaceae 

Casuarinaceae 

1 

Betulaceae 

1 

Juglandaceae 

1 

Myricaceae 

Fagaceae 

1 

Tetramelaceae 

Begoniaceae 

Cucurbitaceae 

Urticaceae 

Moraceae 

Cannabaceae 

Ulmaceae 

1 

Elaeagnaceae 

1 

Rhamnaceae 

Rosaceae 

1 

Fabaceae 

1 

1 2 

Polygalaceae 

Elaeocarpaceae 

Cunoniaceae 

1 

Oxalidaceae 

Chrysobalanaceae 

Ochnaceae 

Elatinaceae 

Hypericaceae 

Clusiaceae 

Turneraceae 

1 

Salicaceae 

Violaceae 

Euphorbiaceae 

1 

Erythroxylaceae 

Linaceae 

Rhizophoraceae 

Pandaceae 

Humiriaceae 

Celastraceae 

Parnassiaceae 

Zygophyllaceae 

Krameriaceae 

Melastomataceae 

Myrtaceae 

1 

Lythraceae 

Onagraceae 

Combretaceae 

Geraniaceae 

Vitaceae 

Saxifragaceae 

1 

Grossulariaceae 

Haloragaceae 

Crassulaceae 

Hamamelidaceae 

1 

rosids 

1 

19 

4 

16 

2 

21 

12 

74 

75 

100 

100 

90 

100 

80 

100 

100 

83 

15 

50 

0 

0 

100 

100 

100 

100 

93 

67 

40 

86 

100 

100 

100 

60 

93 

0 

83 

100 

88 

84 

94 

87 

100 

100 

86 

100 

100 

100 

38 

78 

100 

93 

60 

87 

0 

50 

50 

100 

100 

100 

0 

78 

100 

80 

98 

67 

74 

100 

62 

100 

11 

80 

26 

25 

0 

0 

5 

0 

0 

0 

0 

0 

5 

0 

42 95 5 0 

5 

12 

1 

6 

55 

2 

2 

1 

1 

1 

1 

4 

15 

3 

5 

14 

1 

3 

8 

10 

14 

1 

6 

4 

17 

80 

315 

6 

3 

1 

7 

1 

1 

1 

8 

9 

1 

41 

15 

69 

1 

6 

2 

1 

1 

5 

1 

9 

1 

5 

52 

6 

23 

2 

13 

5 

9 

5 

2 

16 

1 

50 

31 

100 

0 

0 

0 

17 

33 

0 

100 

0 

0 

0 

0 

0 

7 

33 

60 

7 

0 

0 

0 

20 

0 

100 

0 

0 

12 

15 

3 

13 

0 

0 

14 

0 

0 

0 

13 

11 

0 

7 

40 

9 

0 

33 

0 

0 

0 

0 

100 

0 

0 

0 

0 

17 

22 

0 

31 

0 

44 

20 

0 

19 

0 

20 

0 

0 

0 

53 

50 

0 

100 

0 

0 

0 

0 

0 

0 

0 

7 

0 

0 

0 

20 

7 

0 

17 

0 

0 

1 

3 

0 

0 

0 

0 

0 

0 

0 

50 

11 

0 

0 

0 

4 

100 

17 

50 

0 

0 

0 

0 

22 

0 

20 

2 

17 

4 

0 

84% AM, 16% AM+ECM 

AM only 

AM only 

AM only 

AM only 

AM only 

93% AM, 5% ECM, 2% AM+ECM 

75% AM, 25% ECM 

8% AM, 42% ECM, 8% AM+ECM, 17% ECM+EEM, 25% AM+ECM+EEM 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

25% AM, 50% ECM, 25% AM+ECM 

7% AM, 67% ECM, 7% AM+ECM, 13% ECM+EEM, 7% AM+ECM+EEM 

33% AM, 33% ECM, 33% AM+ECM 

AM only 

8% AM, 92% ECM 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

80% AM, 20% AM+ECM 

AM only 

94% AM, 6% AM+ECM 

84% AM, 3% ECM, 13% AM+ECM 

86% AM, 7% ECM, 7% AM+ECM, 0.3% AM+ECM+EEM 

67% AM, 17% AM+ECM, 17% mycoheterotrophy via AM 

AM only 

AM+ECM 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

15% AM, 49% ECM, 34% AM+ECM, 2% ECM+EEM 

AM only 

92% AM, 5% ECM, 3% AM+ECM 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

AM only 

75% AM, 25% AM+ECM 

53% AM, 29% ECM, 17% AM+ECM 

AM only 

AM only 

AM only 

8 AM only 

0 AM only 

44 AM only 

0 80% AM, 20% AM+ECM 

50 AM only 

88% AM, 12% endo-M (also AM?) 

50 

0 AM only

359 

The very different evolutionary pattern of ECM from 

that of AM might suggest a different underlying genetic 

mechanism. Also, the structure is quite different between 

the two types of mycorrhizas. In ECM, the fungal hyphae 

do not penetrate the cell wall, and instead form a Hartig net 

(an intercellular meshwork in the root epidermis and 

cortex) and a sheath around the root derived from this net. 

On the other hand, the fungal hyphae in AM penetrate the 

cell wall and often form arbuscules. At present, little is 

known about the molecular genetic basis of ECM development. 

All plant genes involved in mycorrhizal symbiosis 

identified so far control AM development (Stracke et al. 

2002; Liu et al. 2003; Ane et al. 2004; Demchenko et al. 

2004; Levy et al. 2004). One of these genes has a homolog 

in the moss Physcomitrella patens and several other early 

land plants (Wang and Qiu, unpublished data). Hence, it 

can be inferred that there might be a general genetic 

program in plants that mediates plant–fungus interaction, a 

small portion of which might even be involved in plant– 

fungal pathogen interaction as well (Harrison 1999). This 

program was most likely established at the beginning of 

land plant evolution, if not earlier, and mostly targets 

glomeromycetous fungi. It has been inherited in land plants 

since its inception and is responsible for the prevalence of 

AM seen today. When ECM evolved in a particular plant 

lineage, the homobasidiomycetous fungi might have 

adopted much of this general genetic program, but also 

modified some aspects of it to suit a slightly different 

plant–fungus partnership. One intriguing question is how 

different plant lineages, particularly those as diverse as 

ferns, gymnosperms, and angiosperms, adapted the general 

genetic program for ECM development, as the structures of 

mycorrhizas are apparently very similar between these 

diverse lineages. Perhaps different genes are involved in 

each case, thus, the host specificity developed with 

different fungal symbionts in each case of ECM (as 

opposed to AM). Indeed, difference in gene expression was 

observed to be related to host specificity (Le Quere et al. 

2004). Future work will likely reveal more details to help 

us understand how many independent conversions of ECM 

from AM had occurred. 

The ericaceous mycorrhizas and their close 

relationship to ectomycorrhiza 

There are six types of mycorrhizas in Ericaceae. With the 

exception of AM, five of them are partnerships between 

ascomycetous or basidiomycetous fungi and plants: ECM, 

ectendomycorrhiza (EEM), arbutoid mycorrhiza (ABM), 

monotropoid mycorrhiza (MTM), and ericoid mycorrhiza 

(ERM). Among these five types, ECM occurs in many 

plant lineages outside of Ericaceae (see above), and EEM is 

found in Pinaceae, Araucariaceae, Ericaceae, Salicaceae, 

Fabaceae, Betulaceae, and Cistaceae (Fig. 1). ABM, MTM, 

and ERM can be deemed to be genuinely ericaceous 

mycorrhizas, as they occur almost exclusively in this 

family. The only exception is ERM, which has also been 

found in Diapensiaceae, a close relative of Ericaceae. The 

ericaceous mycorrhizas share many characteristics with 

ECM, but exhibit a high degree of intracellular penetration 

(Smith and Read 1997). Brundrett (2002) suggested that 

ABM, MTM, and ERM were derived from ECM. Our 

mapping of ECM and these other types of mycorrhizas 

onto the land plant phylogeny and examination of their 

distribution within Ericaceae supports this interpretation. 

Further, we add that EEM probably represents a transitional 

stage between ECM and the three types of ericaceous 

mycorrhizas, ABM, MTM, and ERM, as ECM and EEM 

are found only in the basal lineages of Ericaceae, 

Monotropoideae, and Arbutoideae (Kron et al. 2002). 

Previous studies have identified the fungal symbionts of 

the ericaceous mycorrhizas. One interesting phenomenon 

is that the fungal strains isolated from these three types of 

mycorrhizas can also form ECM with other plant species 

(reviewed by Smith and Read 1997). This observation 

strongly suggests a close relationship between ECM and 

the ericaceous mycorrhizas. An interesting question that 

arises is why the same fungi cannot penetrate into root cells 

of ectomycorrhizal hosts yet are able to do so with 

ericaceous plants. One answer could be that host plants 

play an important role in controlling development of 

different types of mycorrhizas. 

Orchid mycorrhiza 

Orchid mycorrhiza (ORM) is generally thought to be 

restricted to the Orchidaceae, which involve mostly 

basidiomycetous fungi (Leake 1994). Occurrence of this 

type of mycorrhiza in other plants such as the subterranean 

nonphotosynthetic liverwort Cryptothallus mirabilis 

(Ligrone et al. 1993; Bidartondo et al. 2003) andthe 

monocot Thismia sp. (Brundrett 2002 see http://www. 

ffp.csiro.au/research/mycorrhiza/) probably represents 

functional and structural convergence, as the fungal 

partners are of different types than those of the true ORM 

in Orchidaceae (Leake 1994). In this perspective, ORM, 

like the ericaceous mycorrhizas discussed above, can be 

considered to occur exclusively in one plant family, 

Orchidaceae, and probably represents a highly specialized 

type of ectomycorrhiza. One fact supporting this 

idea is that some species of lower Epidendroideae, an 

early-diverging lineage within Orchidaceae (Cameron et 

al. 1999), still have ECM (Bidartondo et al. 2004; 

Selosse et al. 2004). 

Did evolution of ectomycorrhiza and its derived types 

of mycorrhizas spur diversification of plant lineages 

that harbor these symbiotic systems? 

If AM was instrumental in the origin and subsequent 

diversification of land plants in a newly exploited niche, 

did evolution of ECM and its derived types of mycorrhizas, 

which involve Ascomycetes and Basidiomycetes, contribute 

to diversification of plant lineages that forged these 

new partnerships? Our review provides a rather positive

360 

answer to this question. In plant families where ECM and 

its derivatives are the main types of mycorrhizas or account 

for a significant percentage (>20%), e.g., Pinaceae, Orchidaceae, 

Nyctaginaceae, Polygonaceae, Ericaceae, Aquifoliaceae, 

Salicaceae, Fagaceae, Betulaceae, Cistaceae, 

Dipterocarpaceae, Myrtaceae, and Melastomataceae 

(Mabberley 1987), the species number is often high to 

very high in comparison to their close relatives that have 

AM (Fig. 1). In the case of woody plants, the members of 

these families are often dominant species in their 

communities (Malloch et al. 1980). Although there are 

many factors that may affect speciation rate, we believe 

that the correlation between high occurrence rate of ECM 

and its derived types of mycorrhizas and the species 

richness of plant clades that harbor these symbiotic 

systems are not coincidental. This observation becomes 

particularly striking when compared to mycoheterotrophy, 

another type of plant–fungus symbiosis, which will 

be discussed below. Because ascomycetous and basidiomycetous 

mycorrhizal fungi have a much higher host 

specificity than glomeromycetous fungi (Horton and 

Bruns 1998; Newton and Haigh 1998; Bidartondo and 

Bruns 2002), diversification of the plant hosts had likely 

in turn driven the speciation rate of the fungal symbionts. 

This coevolutionary arms race probably explains the large 

number of Basidiomycetes that are ectomycorrhizal today 

(Molina et al. 1992). In comparison, AM has obviously 

contributed very little to the diversification of glomeromycetous 

fungi, and the relatively low host specificity of 

these fungi may be to blame. 

Mycoheterotrophy 

Mycoheterotrophy represents a shift of balanced mutualistic 

symbiosis between plants and fungi toward an 

exploitative use of mycorrhizal fungi by plants that are 

no longer fully photosynthetic (Leake 1994). It is clearly a 

derived condition in mycorrhizal evolution. In our survey, 

we found that mycoheterotrophy had evolved many times 

independently in land plants, in the following families: 

Aneuraceae, Podocarpaceae, Petrosaviaceae, Burmanniaceae, 

Triuridiaceae, Corsiaceae, Orchidaceae, Ericaceae 

(Monotropoideae), Gentianaceae, and Polygalaceae (Fig. 1). 

The mycoheterotrophy in the moss genus Buxbaumia needs 

more thorough investigation (Leake 1994). In addition, 

gametophytes of Lycopodium, Psilotum, and Botrychium are 

also mycoheterotrophic (reviewed by Brundrett 2002). However, 

it has been suggested that mycoheterotrophy in these 

latter cases and most species of Orchidaceae may be different 

from that in other plant groups mentioned above (Bidartondo 

2005) because these plants only engage in mycoheterotrophy 

for a part of their life cycles during establishment, and they are 

photosynthetic for the rest of their life cycles and enter 

mutualistic symbiosis with mycorrhizal fungi. 

The independent evolution of mycoheterotrophy in 

many unrelated lineages of land plants is further underscored 

by the fact that it evolved repeatedly from AM and 

ECM and ECM-derived types of mycorrhizas (Fig. 1). 

Hence, evolution of mycoheterotrophy reinforces the 

evolutionary parallelism commonly seen in mycorrhizal 

evolution, which has dominated evolution of ECM as 

discussed above and loss of mycorrhiza to be discussed 

below. Although one may think that an exploitative 

relationship such as mycoheterotrophy can be evolutionarily 

unstable, which seems to be supported by the general 

paucity of species in most clades of mycoheterotrophic 

plants (Leake 1994), this unique type of interaction among 

more than two species of very different modes of nutrition 

uptake (autotrophy and heterotrophy) might have played a 

special role to facilitate some major transitions during land 

plant evolution. In the case of lycophytes and ferns, 

mycoheterotrophy probably helped to ease the transition 

from a gametophyte generation-dominant life cycle in 

bryophytes to a sporophyte generation-dominant life cycle 

in vascular plants. For most Orchidaceae, mycoheterotrophy 

has perhaps alleviated the problem of extremely small 

size of the seeds, which can be advantageous for dispersal 

but at the same time, limits nutrition packaging for the next 

generation, and thus has probably contributed significantly 

to the success of the family. These two examples once 

again highlight the important roles played by fungi not only 

in the origin, but also in many subsequent radiations of land 

plant evolution. 

Many independent losses of mycorrhiza in land plants 

Many families of land plants have not been found to form 

mycorrhizas with fungi in their natural habitats. In 

liverworts, no fungal association has been observed in 

Blasiaceae, Ricciaceae, Metzgeriaceae, Radulaceae, Porellaceae, 

Jubuliaceae, Lejeuneaceae, or Herbertaceae. In 

pteridophytes, species in Isoetaceae and Azollaceae are 

nonmycorrhizal. Similarly in angiosperms, species in 

Nymphaeaceae, Butomaceae, Limnocharitaceae, Cyclanthaceae, 

Cannaceae, Loranthaceae, Loasaceae, Lentibulariaceae, 

Menyanthaceae, Adoxaceae, Eryothroxylaceae, and 

Bataceae are nonmycorrhizal. In addition, nonmycorrhizal 

species occur together with mycorrhizal species in many 

families, and approximately half of the ten or more species 

examined in the following families are nonmycorrhizal: 

Geocalycaceae, Jungermanniaceae, Lycopodiaceae, Hymenophyllaceae, 

Gleicheniaceae, Pteridaceae, Adiantaceae, 

Aspleniaceae, Thelypteridaceae, Dryopteridaceae, Polypodiaceae, 

Cyperaceae, Polygonaceae, Amaranthaceae, Caryophyllaceae, 

Crassulaceae, and Brassicaceae (Table 1, 

Fig. 1). The fact that these families are deeply embedded 

among mycorrhizal families and are not closely related to 

each other provides convincing evidence that these plants 

lost their ability to form mycorrhiza independently. 

There are two interesting features for phylogenetic 

distribution of nonmycorrhizal plants. One is that leafy 

liverworts and pteridophytes in general tend to have a high 

concentration of nonmycorrhizal species. Of course, 

mosses, as an entire clade, lack mycorrhiza as well (Read 

et al. 2000) except Takakia (Boullard 1988). Gymnosperms, 

on the other hand, are all mycorrhizal and almost

361 

all obligately mycorrhizal. Angiosperms generally are also 

mycorrhizal, and have only a few groups that have a high 

percentage of nonmycorrhizal species, alismatids (including 

Araceae according to Stevens’ (2004) new classification 

system, see Fig. 1b), Cyperaceae, caryophyllids, and 

Brassicaceae. This feature may be related to the fact that 

early land plants had not fully adapted to the symbiotic 

system with the fungi. Indeed, these plants generally have 

the Paris-type AM, which lacks well-developed systems of 

intercellular hyphae and do not always form arbuscules 

(Read et al. 2000). It will be interesting to see if future 

studies can demonstrate that a simpler genetic system is 

behind this rather unstable symbiotic relationship in basal 

land plants. The other feature is that almost all nonmycorrhizal 

plants are derived from ancestors that engaged in 

AM symbiosis. In the several families where ECM and 

ECM-derived mycorrhizas dominate or account for a large 

percentage, e.g., Pinaceae, Orchidaceae, Ericaceae, 

Myrtaceae, Salicaceae, Fagales, and Cistaceae, very few 

species are nonmycorrhizal. This feature is perhaps related 

to the highly specialized nature of the ECM symbiosis. 

As to the mechanisms responsible for mycorrhizal loss, 

Trappe (1987) noticed several characters that tend to be 

shared by nonmycorrhizal species, and his observations 

still hold true here. First, many nonmycorrhizal vascular 

plants grow in aquatic or wetland habitat. In these 

environments, both nutrients and water supplies are not 

as limited as in the typical terrestrial environment. Hence, 

the plants can develop independence from the fungal 

symbionts and reduce the carbon cost that would normally 

be provided to the fungal partners in exchange for nutrients 

and water. The loss of mycorrhiza in alismatids and other 

aquatic plants likely occurred through this mechanism. 

Second, species that grow in nutrient-rich environments 

tend to be nonmycorrhizal. The ruderals in caryophyllids, 

Brassicaceae, and Crassulaceae, and the spring ephemerals 

in Ranunculaceae, Papaveraceae, and Saxifragaceae belong 

to this category. Third, the plants that have a long, 

fine, and highly branched root system with well-developed 

root hairs also have a high tendency to become 

nonmycorrhizal. Cyperaceae and the ruderals perhaps 

developed independence from fungal symbionts through 

this route. It has been suggested that root hairs and 

mycorrhizal fungi were two alternative mechanisms for 

plant nutrient uptake (Baylis 1970; Koide 1991). If a plant 

can absorb sufficient nutrients through its own root hairs, 

any genetic changes preventing the formation of mycorrhiza 

with fungi would be favorably selected. Besides these 

three observations made by Trappe (1987), we also noticed 

that many liverworts that lack fungal association are leaf 

and bark epiphytes, e.g., Radulaceae, Jubulaceae, Porellaceae, 

and Lejeuneaceae. A possible explanation could be 

that fungal development is thwarted by defense mechanisms 

of the plant on which these epiphytes live. 

Areas of further study 

This study and those by Trappe (1987) and Harley and 

Harley (1987) represent the three most extensive surveys of 

mycorrhizal occurrence in land plants, but the total number 

of species included in the three studies is only slightly over 

10,000 (assuming that there is little overlap among the 

three surveys), which is about 3% of all listed land plant 

species that live on our planet. As can be seen from Table 1 

and Fig. 1, for most plant families we have knowledge of 

the mycorrhizal status for only one or a few species. Hence, 

the first area that deserves more attention in the future is the 

investigation of mycorrhizal status of more species. From 

this study and those by Trappe (1987) and Harley and 

Harley (1987), as well as several reviews published earlier 

on this topic (Pirozynski and Malloch 1975; Malloch et al. 

1980; Selosse and Le Tacon 1998; Read et al. 2000; 

Brundrett 2002), it is clear that mycorrhizas were instrumental 

in the origin and subsequent diversification of land 

plants and are continuing to play a vital role in maintaining 

floristic diversity and ecosystem function on earth. Thus, 

mycorrhizal symbiosis should be viewed as an integral part 

of any floristic, ecological, or evolutionary studies in the 

future. 

Besides the general increase of mycorrhizal research, a 

few specific areas should be pursued. First, more basal land 

plants should be investigated, as they occupy an especially 

important position in our understanding of the origin of 

mycorrhizal symbiosis in land plants. Despite several 

studies on basal land plants (Pocock and Duckett 1985; 

Duckett et al. 1991; Read et al. 2000), our knowledge on 

phylogenetically critically positioned taxa such as Treubiaceae, 

many simple thalloid liverworts, mosses, and some 

hornworts, remains fragmentary. In this regard, mosses 

deserve special attention, as they represent the only major 

clade of land plants that is nonmycorrhizal (Read et al. 

2000), a situation which by itself is very puzzling. Several 

basal moss lineages such as Andreaeaceae, Tetraphidaceae, 

Polytrichaceae, Buxbaumiaceae, and Diphysciaceae all 

grow on nutrient-poor, sandy soil or rocks. It will be 

interesting to know how they manage to grow on these 

poor substrates if they are truly nonmycorrhizal. Takakia,a 

sister group to all other mosses, is reported to be 

mycorrhizal (Boullard 1988) and grows on thin soil and 

rocks (Gao 2000). Second, several early-diverging lineages 

of angiosperms, Amborella, Austrobaileyales, Chloranthaceae, 

and magnoliids should be investigated; the information 

on their mycorrhizal status is currently lacking. Even 

when Trappe’s (1987) survey is included, our knowledge 

of these plants is still quite poor. A focused study on these 

plants will fill an important gap in our understanding of 

mycorrhizal evolution in early angiosperms. Finally, as we 

learn more from studies of model organisms such as 

legumes (Stracke et al. 2002; Liu et al. 2003; Ane et al. 

2004; Demchenko et al. 2004; Levy et al. 2004), characterization 

of molecular aspects of mycorrhizas in 

nonmodel organisms will significantly expand the dimension 

and depth of our knowledge of this important 

symbiotic system. In summary, the mycorrhizal research

362 

conducted over the last several decades has brought this 

long neglected field into mainstream biology, and future 

research will undoubtedly further enhance our understanding 

of this important biological interaction and its 

impact on the evolution of both plants and fungi, as 

well as the establishment and functioning of the terrestrial 

ecosystem. 

Acknowledgements We would like to thank Rong-rong Xu for 

helping with figure preparation, Malini Jane Sridharan for critically 

reading the manuscript, and Jim Trappe and two reviewers for their 

insightful comments. This work was supported by an Early Career 

Award (DEB 0332298) and ATOL grants (DEB 0431239, DEB 

0531689) from NSF to Y-L Qiu. 

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Phylogenetic distribution and evolution of mycorrhizas in land plants

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