Cool and their fungal friends

Andy MacKinnon Metchosin Cool plants and their fungal friends

What is myco-heterotrophy? What plants are mycoheterotrophic? What fungi are involved? What is the of the relationship? What are plants? What is ‘mixotrophy’? What are fungi?

SomeCool local plants examples. and their fungal friends • lichens Lessons• frommycorrhizae myco-heterotrophy. • mycoheterotrophs and mixotrophs Cool plants and their fungal friends

What is myco-heterotrophy? What plants are mycoheterotrophic? What fungi are involved? What is the nature of the relationship? What are plants? What is ‘mixotrophy’? What are fungi?

SomeCool local plants examples. and their fungal friends • lichens Lessons• frommycorrhizae myco-heterotrophy. • mycoheterotrophs and mixotrophs

Cool plants and their fungal friends

What is myco-heterotrophy? What plants are mycoheterotrophic? What fungi are involved? What is the nature of the relationship? What are plants? What is ‘mixotrophy’? What are fungi?

SomeCool local plants examples. and their fungal friends • lichens Lessons• frommycorrhizae myco-heterotrophy. • mycoheterotrophs and mixotrophs

Cool plants and their fungal friends

What is myco-heterotrophy? What plants are mycoheterotrophic? What fungi are involved? What is the nature of the relationship? What are plants? What is ‘mixotrophy’? What are fungi?

SomeCool local plants examples. and their fungal friends • lichens Lessons• frommycorrhizae myco-heterotrophy. • mycoheterotrophs and mixotrophs Cool plants and their fungal friends

lichens mycorrhizae mycoheterotrophs and mixotrophs Cool plants and their fungal friends

lichens mycorrhizae mycoheterotrophs and mixotrophs

Viktoria Wagner and Toby Spribille Edible Horsehair Lichen Inedible Horsehair Lichen (Bryoria fremontii) (Bryoria tortuosa) A fluorescent microscope image shows the location of different cell types in a bryoria lichen, cut at the ends and lengthwise through the middle. Green are the yeasts, blue are the fungi, red are the algae. (Toby Spribille) Cool plants and their fungal friends

lichens mycorrhizae mycoheterotrophs and mixotrophs

Greek μύκης mykēs, “” Greek ρίζα, riza, "root"

A mycorrhiza is a symbiotic association composed of a fungus and roots of a vascular . In a mycorrhizal association, the fungus colonizes the host plant's roots, either intracellularly as in arbuscular mycorrhizal fungi (AMF or AM), or extracellularly as in ectomycorrhizal fungi. The association is generally mutualistic.

ectomycorrhizae VA (endo-) mycorrhizae

ericoid mycorrhizae salal Cool plants and their fungal friends

lichens mycorrhizae mycoheterotrophs and mixotrophs What is myco-heterotrophy? What plants are mycoheterotrophic? What fungi are involved? What is the nature of the relationship?

• WhatWhat is mycois ‘mixotrophy’?-heterotrophy? • What plants are myco-heterotrophic? • WhatSome fungi local are involved? examples. • Some local examples. • LessonsLessons from from myco -mycoheterotrophy.-heterotrophy. Mycoheterotrophy

Greek μύκης mykēs, “fungus” Greek ἕτερος heteros, “another”, “different” Greek τροφή trophe = "nutrition", "growth”

Myco-heterotrophy is a symbiotic relationship between certain kinds of plants and fungi, in which the plant gets all or part of its food from upon fungi rather than from . A myco-heterotroph is the parasitic plant partner in this relationship. Movement of carbon (black arrows) and nutrients (white arrows) in myco-heterotrophy. (from Leake and Cameron 2010) Mycoheterotrophic plants generally …

Lack Have small ‘dust’ seeds with undifferentiated embryos Have leaves that are scale-like or absent Have reduced vascularization of the stem Lack stomata Have reduced roots that lack root hairs Have elevated δ13C and δ15N values (compared to green plants) Mixotrophic plants generally …

Are green have δ13C and δ15N values between those of mycoheterotrophs and full autotrophs (regular green plants), indicating that they get some of their C and N from other plants through their fungal partner

Are related to mycoheterotrophs What is myco-heterotrophy? What plants are mycoheterotrophic? What fungi are involved? What is the nature of the relationship?

• WhatWhat is myco is ‘mixotrophy’?-heterotrophy? • What plants are myco-heterotrophic? • WhatSome fungi local are involved? examples. • Some local examples. • LessonsLessons from from myco -mycoheterotrophy.-heterotrophy. Plant families that have evolved myco- heterotrophy. (from Merckx and Freudenstein 2010) Plant families that have evolved myco-heterotrophy. Plant families that have evolved myco-heterotrophy. Plant families that have evolved myco-heterotrophy.

Epirixanthes verticillata

5 species of Epirixanthes

Voyria parasitica

25 species of Voyria, Voyriella, Cotylanthera and Sebaea Plant families that have evolved myco-heterotrophy.

Geosiris aphylla

Burmannia bicolor Plant families that have evolved myco-heterotrophy.

Striped Coralroot (Corallorhiza striata) Plant families that have evolved myco-heterotrophy.

Indian-pipe (Monotropa uniflora) What is myco-heterotrophy? What plants are mycoheterotrophic? What fungi are involved? What is the nature of the relationship?

• WhatWhat is myco is ‘mixotrophy’?-heterotrophy? • What plants are myco-heterotrophic? • WhatSome fungi local are examples. involved? • Some local examples. • LessonsLessons from from myco -mycoheterotrophy.-heterotrophy. Main fungal groups that host myco-heterotrophic plants (after Hynson and Bruns 2010). What fungi are involved?

Generally, myco-heterotrophic Aneuraceae (liverworts), Orchidaceae and exploit ectomycorrhizal networks while myco- heterotrophic , Corsiaceae, , Thismiaceae and Triuridaceae exploit arbuscular mycorrhizal networks some myco-heterotrophic orchids are specialized on litter- and wood-decay fungi What is myco-heterotrophy? What plants are mycoheterotrophic? What fungi are involved? What is the nature of the relationship?

• WhatWhat is myco is ‘mixotrophy’?-heterotrophy? • What plants are myco-heterotrophic? • WhatSome fungi local are involved? examples. • Some local examples. • LessonsLessons from from myco -mycoheterotrophy.-heterotrophy. Probably most clubmosses and some ferns

•Gametophytes of most species of clubmosses and some ferns are non- photosynthetic, mycorrhizal and likely myco-heterotrophic

Running clubmoss (Lycopodium clavatum) •Most (perhaps all) of the 30,000 Orchids in general species of orchids require some sort of myco-heterotrophic arrangement with saprophytic or parasitic species of “Rhizoctonia” (Basidiomycetes) for germination of their ‘dust’ seeds •>100 fully myco-heterotrophic species; perhaps some mixotrophs also?

• Goodyera repens has been shown to transfer significant amounts of carbon back to its mycorrhizal fungus

Fairyslipper (Calypso bulbosa) Coralroots (Corallorhiza species)

•Four species in southwestern BC: spotted coralroot (C. maculata), western coralroot (C. mertensiana), striped coralroot (C. striata), yellow coralroot (C. trifida), all in our area (but yellow coralroot more of an interior BC species)

•All myco-heterotrophic

•Fungal associates: –spotted coralroot, 20 species in Russulaceae –western coralroot, 3 closely related Russula species –striped coralroot, Thelephora- Tomentella species

Spotted Coralroot (C. maculata) Phantom Orchid (Cephalanthera austiniae)

•Fully mycoheterotrophic •Fungal associates: 14 species in Thelephora- Tomentella

Thelephora terrestris Prince’s-Pines (Chimaphila species)

•Menzies’ pipsissewa (C. menziesii) and Prince’s Pine (C. umbellata) both in our area •Prince’s Pine is mixotrophic •Fungal associates: a variety of ectomycorrhizal species, including several Tricholomas

Prince’s Pine (C. umbellata) Wintergreens (Pyrola, Moneses and Orthilia species)

•Six species of Pyrola potentially in our area

•One-sided wintergreen (Orthilia secunda)

•Single delight (Moneses uniflora)

•White-veined wintergreen (Pyrola picta), green wintergreen (Pyrola chlorantha) and one-sided wintergreen (Orthilia secunda) are demonstrated mixotrophs

•Fungal associates: a variety of ectomycorrhizal species, including White-veined wintergreen several Tricholomas (Pyrola picta) Candystick ( virgata)

•Mycoheterotrophic •Fungal associate: pine mushroom (Tricholoma magnivelare) Gnome Plant (Hemitomes congestum)

•Mycoheterotrophic •Fungal associate: peckii and H. aurantiacum

Strawberries and cream (Hydnellum peckii) Indian-pipe (Monotropa uniflora)

•Mycoheterotrophic •Fungal associate: Russulaceae, often short- stemmed Russula (Russula brevipes) broad-leaved helleborine (Epipactis helleborine)

•Usually mixotrophic, but … •Fungal associates: Pyronemataceae, Sebacina, Tomentella, Tuber

Tuber oregonense Pinesap ()

•AKA Hypopitys monotropa •Mycoheterotrophic •Fungal associates: Tricholoma species (including T. sejunctum and T. flavovirens); European pinesap associated with European Tricholomas

Man-on-horseback (Tricholoma flavovirens) Pinedrops (Pterospora andromedea)

•Mycoheterotrophic •Fungal associate: Rhizopogon salebrosus, R. arctostaphyli

Daniel Mosquin photo Rhizopogon salebrosus What is myco-heterotrophy? What plants are mycoheterotrophic? What fungi are involved? What is the nature of the relationship?

• WhatWhat is myco is ‘mixotrophy’?-heterotrophy? • What plants are myco-heterotrophic? • WhatSome fungi local are involved? examples. • Some local examples. • LessonsLessons from from myco myco-heterotrophy.-heterotrophy. Lessons from myco-heterotrophy

These pink and white plants are not “saprophytes” It’s much too simplified to refer to some plants and fungi as having only one feeding preference The world is much more complex (and interesting) than most texts would have you believe The world is also much more complex than I’ve described tonight Rhizopogon vinicolor

KJ Beiler, K.J., D.M. Durall, S.W. Simard, S.A. Maxwell and A.M. Kretzer. 2010. New Architecture of the Wood-Wide Web: Rhizopogon spp. Genets Link Multiple Douglas- Cohorts. The New Phytologist 185(2): 543-553.

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