Truffles in Context: A Mycological Overview By Matt Trappe, Oct. 2006

1. Truffles in Context: A Mycological Overview By Matt Trappe, Oct. 2006

2. Truffles are here!! Truffles in Context: A Mycological Overview By Matt Trappe, Oct. 2006

3. or, “Where on Earth do Truffles fit in?”

4. or, “Where on Earth do Truffles fit in?”

5. Taxonomy: Slicing the Pie… • Many ways to divide and organize people • Diet • Morphology (height, weight) • - Family group, ethnicity

6. Taxonomy: Slicing the Pie… • Many ways to divide and organize people • Diet • Morphology (height, weight) • - Family group, ethnicity • Many ways to divide and organize fungi • Diet (trophic status) • Morphology (shape, growth habit) • Family group, genetic taxonomy

7. Taxonomy: Slicing the Pie… • Many ways to divide and organize people • Diet • Morphology (height, weight) • - Family group, ethnicity • Many ways to divide and organize fungi • Diet (trophic status) • Morphology (shape, growth habit) • Family group, genetic taxonomy

8. Taxonomy: Slicing the Pie… • Many ways to divide and organize people • Diet • Morphology (height, weight) • - Family group, ethnicity • Many ways to divide and organize fungi • Diet (trophic status) • Morphology (shape, growth habit) • Family group, genetic taxonomy

9. Organizing Fungi by their Diet (trophic status) • Mycorrhizal Fungi • Form mutually beneficial associations with plants • “Myco” = Fungus, “Rhiza” = Root • Saprobic Fungi • Digest dead material; wood and litter decomposers • Pathogenic Fungi • - Attack plants (& animals!) to feed off them

10. Mycorrhizal Fungi • Form mutually beneficial associations with plants • Ectomycorrhizae • Endomycorrhizae

11. Ectomycorrhizal Fungi “Ecto” = outside Ectomycorrhizal fungi wrap outside of root with a “mantle” from which foraging hyphae emanate.

12. Ectomycorrhizal Fungi This is a long-term structure, growing at the tips of the roots and extending with them through the soil.

13. Ectomycorrhizal Fungi The fungi also grow between the root cells but do not penetrate them. This is called a “Hartig Net” and is where nutrient exchange occurs. Hartig net Root tip cross section Mantle M.Brundrett photo

14. Ectomycorrhizal Fungi Summary Almost all truffles are the fruiting bodies of ectomycorrhizal fungi. Many mushrooms(half? less in tropics!) are the fruiting bodies of ectomycorrhizal fungi. Ectomycorrhiza is formed by members of Ascomycetes, Basidiomycetes, and Zygomycetes (we’ll get to them later) There are about 12,000 species of ectomycorrhizal fungi that associate with about 2000 species of host plants.

15. Ectomycorrhizal Fungi Hosts Some major plants that are hosts to ectomycorrhizal fungi in the Pacific Northwest: Douglas-fir Pines Firs Oaks Alders Birches Poplars Hazel Eucalyptus Beech Spruce

16. Endomycorrhizal Fungi, a.k.a. “VAM Fungi” “Endo” = inside Endomycorrhizal fungi do not form a mantle, but rather penetrate root cells and form temporary structures inside the root cells called “vesicles” and “arbuscules”. Thus, the term “VAM” is an abbreviation for “Vesicular-Arbuscular Mycorrhizae”

17. Endomycorrhizal Fungi, a.k.a. “VAM Fungi” Vesicles are balloon-like structures thought to be for energy storage. Arbuscules are tree-shaped (“arbor”) structures where nutrient exchange occurs. Vesicles Arbuscule

18. Endomycorrhizal Fungi, a.k.a. “VAM Fungi” Foraging hyphae extend through the host root cell wall into the soil. Vesicles and arbuscules have a relatively short life; individual structures can form and senesce within days.

19. Endomycorrhizal Fungi, a.k.a. “VAM Fungi” All endomycorrhizal fungi seem to form arbuscules but not all of them form vesicles, so many folks are now calling the group “AM Fungi”. Endomycorrhiza = VAM fungi = AM fungi

20. Endomycorrhizal Fungi Summary No endomycorrhizal fungi form mushrooms. Only one genus of endomycorrhizal fungi forms truffles: Glomus. Most endomycorrhizal fungi form individual spores in the soil that are microscopic. Endomycorrhiza is formed by members of both Ascomycetes and Glomeromycetes (we’ll get to them later) There are only about 150 species of endomycorrhizal fungi that associate with about 300,000 species of host plants!

21. Endomycorrhizal Fungi Hosts Some major plants that are hosts to endomycorrhizal fungi in the Pacific Northwest: Maples Cedars Redwoods Sequoias Poison oak Most grasses Most herbs Most garden plants Most fruit trees Most annuals …and most tropical plants

22. Saprobic Fungi “Sapro” = rotten There are very few animals that can digest wood (termites, earthworms, centipedes?). Fungi do the “heavy lifting”, accounting for over 90% of wood decomposition. Without fungi, fallen trees would stack up without ever rotting. what a mess…

23. Saprobic Fungi Also called Saprophytic or Saprotrophic. Some specialize in digesting cellulose (brown “cube” rot). Some specialize in digesting lignin (white rot). Cellulose Lignin

24. Saprobic Fungi Also called Saprophytic or Saprotrophic. Some specialize in digesting cellulose (brown “cube” rot). Some specialize in digesting lignin (white rot). Brown Rot White Rot

25. Saprobic Fungi Only one genus of saprobic fungi is known to form truffles in North America: Trappea. It is related to stinkhorns. Trappea darkeri

26. Saprobic Fungi Some saprobic fungi form sequestrate (but not hypogeous) fruiting bodies, such as Nivatogastreum. It is related to the mushroom genus Pholiota. Nivatogastreum nubigenum

27. Saprobic Fungi Summary Many mushrooms (half?) and all conks are formed by saprobic fungi. There are probably thousands of species of saprobic fungi, many (most?) are microscopic. Pluerotus ostreatus Ganoderma tsugae

28. How can I tell a saprobic mushroom from a mycorrhizal one?

29. How can I tell a saprobic mushroom from a mycorrhizal one? You can’t.

30. How can I tell a saprobic mushroom from a mycorrhizal one? You can’t. Many saprobic fungi grow on the forest floor. Stropharia riparia

31. How can I tell a saprobic mushroom from a mycorrhizal one? You can’t. Many saprobic fungi grow on the forest floor. Many mycorrhizal fungi grow on decaying wood. Boletus mirabilis Stropharia riparia

32. Usually, all species within a genus are either mycorrhizal or saprobic. If you learn the genera, you can infer whether your specimen is mycorrhizal or saprobic. Morchella sp? Amanita muscaria? Lepiota clypeolaria?

33. Saprobic Genera Mycorrhizal Genera Agaricus Amanita Agrocybe Boletus Clavaria Cantharellus Clitocybe Cortinarius Collybia Gomphus Coprinus Hebeloma Galerina Hydnum Hypholoma Hygrophorus Lentinus Inocybe Mycena Laccaria Pholiota Lactarius Pleurotus Ramaria Polypores Russula Psilocybe Suillus Stropharia Tricholoma Volvariella Almost all Truffles… * We still don’t know the trophic status of many genera! • Field surveying

34. Saprobic Genera Mycorrhizal Genera Agaricus Amanita Agrocybe Boletus Clavaria Cantharellus Clitocybe Cortinarius Collybia Gomphus Coprinus Hebeloma Galerina Hydnum Hypholoma Hygrophorus Lentinus Inocybe Mycena Laccaria Pholiota Lactarius Pleurotus Ramaria Polypores Russula Psilocybe Suillus Stropharia Tricholoma Volvariella Almost all Truffles… * We still don’t know the trophic status of many genera! • Field surveying

35. Saprobic Genera Mycorrhizal Genera Agaricus Amanita Agrocybe Boletus Clavaria Cantharellus Clitocybe Cortinarius Collybia Gomphus Coprinus Hebeloma Galerina Hydnum Hypholoma Hygrophorus Lentinus Inocybe Mycena Laccaria Pholiota Lactarius Pleurotus Ramaria Polypores Russula Psilocybe Suillus Stropharia Tricholoma Volvariella Almost all Truffles… * We still don’t know the trophic status of many genera!

36. Pathogenic Fungi “Pathos” = disease Unlike saprobic fungi, pathogenic fungi just can’t wait until something dies on its own. They attack living organisms, sometimes killing the host and sometimes just causing sickness. Sudden Oak Death SOD bark lesion

37. Pathogenic Fungi Fungal Plant Pathogens Armillaria mellea Phaeolus schweinitzii Phellinus weirii White Pine Blister Rust Swiss Needle Cast Chestnut Blight Dutch Elm Disease Sudden Oak Death* Potato Blight* Various Rusts and Smuts (more than 6000 species!) Armillaria mellea Phaeolus schweinitzii

38. Pathogenic Fungi Fungal Human Pathogens (a digression…) Tinea (athlete’s foot, toenail fungus) Candidia (yeasts) Aspergillus (molds) Coccidioides (valley fever) Pneumocystis (pneumonia) Schizophyllum commune Schizophyllum commune See www.doctorfungus.com for gruesome pictures!

39. Pathogenic Fungi Fungal Human Pathogens (a digression…) Tinea (athlete’s foot, toenail fungus) Candidia (yeasts) Aspergillus (molds) Coccidioides (valley fever) Pneumocystis (pneumonia) Schizophyllum commune Schizophyllum commune See www.doctorfungus.com for gruesome pictures! Luckily we have Penicillium!

40. Fungal Plant Pathogens Summary No pathogenic fungi form truffles. Armillaria spp. form mushrooms. Some (Phellinus, Phaeolus) form conks. Most are in habitats other than forests. Most are inconspicuous. They are ubiquitous. Maple leaf spot Rhytisma punctatum Rose blight Phragmidium sp.

41. Summary: Fungal Trophism and Morphology TrufflesMushroomsConks Mycorrhizae Ecto- Many Many None Endo- Few None None Saprobes One Many Many Pathogens None Few Few

43. The Big Picture - Kingdoms

44. The Big Picture - Kingdoms

45. The Big Picture - Kingdoms Animals – multiple cells with nuclei - internal nutrient digestion Fungi – multiple cells with multiple nuclei - external nutrient absorption Plants - multiple cells with nuclei - photosynthesis Protozoans - single cell with nucleus - photosynthesis (algaes) Bacteria – single cell without nucleus - external nutrient absorption

46. The Big Picture - Kingdoms Loss of external enzymes Gain of internal digestion (and a few other things) Animals – multiple cells with nuclei - internal nutrient digestion Fungi – multiple cells with multiple nuclei - external nutrient absorption Loss of chlorophyll Gain of enzymes Plants - multiple cells with nuclei - photosynthesis Gain of multicellular organization Protozoans - single cell with nucleus - photosynthesis (algaes) Gain of nucleus & chlorophyll Bacteria – single cell without nucleus - external nutrient absorption Primordial Bacteria

47. The Big Picture - Kingdoms Animals – multiple cells with nuclei - internal nutrient digestion Fungi – multiple cells with multiple nuclei - external nutrient absorption Plants - multiple cells with nuclei - photosynthesis Protozoans - single cell with nucleus - photosynthesis (algaes) Bacteria – single cell without nucleus - external nutrient absorption

48. The Big Picture - Kingdoms Animals – multiple cells with nuclei - internal nutrient digestion - no truffles Fungi – multiple cells with multiple nuclei - external nutrient absorption - truffles! Plants - multiple cells with nuclei - photosynthesis - no truffles Protozoans - single cell with nucleus - photosynthesis (algaes) - no truffles Bacteria – single cell without nucleus - external nutrient absorption - no truffles

49. The Big Picture - Kingdoms Fungi – multiple cells with multiple nuclei - external nutrient absorption - truffles!

50. Phyla in the Fungal Kingdom Basidiomycetes – mycorrhizae, saprobes - spores from basidia Ascomycetes – mycorrhizae, saprobes - spores from asci Fungi Glomeromycetes – mycorrhizae - spores formed from hyphae Zygomycetes – mycorrhizae, molds - spores formed on suspensors Chytrids – microscopic soil, puddle, & gut fungi - spores have flagella