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MycoBank Taxonomy: Fungi, Dikarya, Basidiomycota, Agaricomycotina, Agaricomycetes, Agaricales, Tricholomataceae, Tricholoma Oh, matsutake: The excitement before finding them. –Yamaguchi Sodo (1642–1716) Matsutake… the name alone evokes something special and nostalgic: a crisp walk in the woods, a seasonal treat, a festive time. In Japan, where matsutake enjoys unrivaled popularity (matsutake means “pine mushroom” in Japanese), almost 1000 tons are consumed annually, and a single mushroom can cost up to $100. To feed this demand, mushrooms are imported from around the world, especially from China, the Pacific Northwest, and Mexico (Ota et al., 2012; Murata et al., 2023). Matsutake provides nontimber value to forests that can exceed the lumber value of the trees, promoting forest conservation and rural wellbeing (Chapela & Garbelotto, 2004). So what makes matsutake so good? Matsutake’s allure emanates from its aroma. It’s something unique and almost indescribable — the smell of matsutake — “once smelled, never forgotten”, “a taste of autumn from the old days” (Gulden et al., 2014; Yamanaka et al., 2020). I’d wager that one of the greatest pleasures in life is finding fresh matsutake emerging as mushrump masses in a duffed pine forest and getting on all fours to sniff them like a pig. Without smelling it for yourself, the best one can do is approximate it by comparing it to other, more familiar odors. Reminiscent of a warm spice like nutmeg, clove, or cinnamon, matsutake is most like the latter but categorically different and definitively funkier. David Arora famously described the smell as “a provocative compromise between ‘Red Hots’ and dirty socks” (Arora, 1986, p. 191). In the food science world, matsutake’s aroma is characterized as spicy with floral and citrus undertones, an alchemy created by the odorants 1-octen-3-one (mushroom quality), linalool (floral-citrus), hexanal (green), and ethyl (E)-3-phenylprop-2-enoate (cinnamon) (Murray et al., 2020). Like truffles, matsutake is all about the volatiles. Matsutake does not refer to a single species but rather a complex of species in the taxonomic group Tricholoma section Matsutake, all highly valued in Japanese markets for their enticing aroma. While morphologically similar, these species have varying degrees of geographic overlap and endemicity across the Northern Hemisphere, roughly matching the distribution of coniferous genera (Chapela & Garbelotto, 2004; Trudell et al., 2017; Aoki et al., 2022; Murata et al., 2023; Ding et al., 2023). Tricholoma matsutake, the eponymous taxon, is found in Asia and Europe and features a reddish brown cap. Tricholoma magnivelare, the subject of this article, is closely related to T. matsutake but grows exclusively in Eastern North America and has a lighter tannish cap. In the Western United States, one finds T. murrillianum; in Mexico, T. mesoamericanum; and in Southern Europe, T. anatolicum. When shopping for matsutake of unknown origin, you could be eating any one or even an assortment of these five different species. Tricholoma section Matsutake also contains T. dulciolens, T. ilkkae, and other less studied odoriferous oddballs that apparently have a more perfumed or fruity aroma (Intini et al., 2003; Heilmann-Clausen et al., 2017; Zhao et al., 2022). Rounding out the section are T. bakamatsutake, T. caligatum, T. fulvocastaneum, and similar undescribed species, all associated with angiosperms. Colloquially called “false matsutake” or “fool’s matsutake”, these species look like matsutake and are edible, but they sometimes taste bitter and lack the compelling matsutake aroma (Chapela & Garbelotto, 2004; Bergo, 2018; Ding et al., 2023). Oh, the disappointment of the foolish matsutake forager! Much more than an esoteric endeavor of biological classification, matsutake taxonomy is relevant to the chef and forager, too. From my experience eating Tricholoma magnivelare, I (and others) have noticed distinct smells that I have not heard associated with other matsutake species — specifically, something of a latex paint or freshly printed office paper smell. These notes become more pronounced the longer the mushroom has been out of the ground. Another notable difference is that T. magnivelaria primordia — young mushrooms whose caps have not yet expanded enough to break their partial veil — are nearly odorless. Instead, the mature, well expanded caps have the most potent and delicious smell. This contrasts with T. matsutake, whose primordia are premium grade and fetch the highest sums in Japanese markets due to their more pungent aroma. This observation has puzzled me and is something I’d like to investigate further. It’s exciting to think about sampling all the different matsutake species in one sitting to explore their nuances in taste and aroma! Like most of the highly prized/priced mushrooms on the market, matsutake fungi are ectomycorrhizal, an ecological barrier to a robust supply. Attempts at cultivating matsutake have been met with limited success, and they must instead be foraged from the wild. Tricholoma magnivelare is partial to pines in low-nutrient, well draining soils, like those found in pine barren ecosystems. To find matsutake, you should look in areas with especially stunted trees and limited to no undergrowth. The reindeer lichen Cladonia rangiferina and the bleeding tooth mushrooms Hydnellum peckii are both good indicator species that you are in the right habitat. If you do find a matsutake patch, you can expect the mushrooms to come back in the same spot year after year. The mushrooms will likely be sandy and should be cleaned thoroughly in the field. Rinsing other mushrooms is usually OK, but matsutake should never be rinsed under the tap or bathed and swished as it loses a lot of its aroma when inundated. My favorite preparation for matsutake is a rice dish called matsutake gohan, which translates to “pine mushroom rice”. This is the best way to capture matsutake’s beautiful aroma. To prepare matsutake gohan, you will need high-quality sushi rice, dashi broth, and lots of matsutake. I know — one of the core tenets of foraging is to never overindulge; anything can make you feel unwell if overconsumed. But I am such a shameless glutton when it comes to matsutake, and that has always been more than OK for me. Matsutake mushrooms are hefty and dense, so even a single mushroom can make a whole dish. After rinsing the rice, it can be cooked with the dashi broth in the usual way, with a little extra broth to account for the extra mass of the mushrooms. I add a bunch of thinly sliced matsutake pieces at the beginning and then again at the end for a light steaming, finally topped with some slivered scallions. Breaking another core tenet of foraging, I’ve always been OK eating matstuke raw or lightly steamed. Well cooked and lightly cooked matsutake together result in a rice well imbued with the volatiles and an unforgettable culinary experience. What they don’t tell you is that matsutake is an excellent culinary mushroom outside of its aroma. Like truffles, a heavily cooked matsutake loses its volatiles, but the result is still great (much better than cooked truffle). Meaty and succulent, matsutake can also be prepared like any other mushroom with excellent results. I love baking matsutake in aluminum foil with loads of garlic for juicy, flavorful mushroom bites. If you have too much matsutake (is that even possible?), you can dehydrate and powderize the mushrooms. The flavor profile of matsutake powder is wholly different from that of fresh fruiting bodies — much more robust and mushroomy, nutty even — and is a great flavor spike in any savory dish. The delicacy of matsutake has been commodified and globalized, touching rural residents and urban dwellers across much of the world. However, as Anna Tsing enlightens us in The Mushroom at the End of the World, there is something untamable in matsutake and the mushroom fever that afflicts those searching for it. Matsutake teaches us about the possibility of life in capitalist ruins and the joy and necessity of interspecies entanglements. They teach us about freedom and aliveness. SpecimenACD0587, iNaturalist #139443316; 17 October 2022; MA, USA; leg. Alden C. Dirks; GenBank OQ785316 (ITS), OQ785300 (LSU); MICH (not yet accessioned). Online ResourcesReferencesAoki W, Bergius N, Kozlan S, Fukuzawa F, Okuda H, Murata H, Ishida TA, Vaario L-M, Kobayashi H, Kalmiş E, et al. 2022. New findings on the fungal species Tricholoma matsutake from Ukraine, and revision of its taxonomy and biogeography based on multilocus phylogenetic analyses. Mycoscience 63: 197–214. Arora D. 1986. Mushrooms Demystified: A Comprehensive Guide to the Fleshy Fungi. Berkeley: Ten Speed Press. Bergo A. 2018. Tricholoma caligatum: the false matsutake. Forager | Chef. Chapela IH, Garbelotto M. 2004. Phylogeography and evolution in matsutake and close allies inferred by analyses of ITS sequences and AFLPs. Mycologia 96: 730–741. Ding XX, Xu X, Cui YY, Kost G, Wang PM, Yang ZL. 2023. A fifty-locus phylogenetic analysis provides deep insights into the phylogeny of Tricholoma (Tricholomataceae, Agaricales). Persoonia 50: 1–26. Gulden G, Trudell S, Frøslev T, Voitk A. 2014. Species of Tricholoma sect. Caligatum in Newfoundland and Labrador. Omphalina 5: 5–9. Heilmann-Clausen J, Christensen M, Frøslev TG, Kjøller R. 2017. Taxonomy of Tricholoma in northern Europe based on ITS sequence data and morphological characters. Persoonia 38: 38–57. Intini M, Dogan HH, Riva A. 2003. Tricholoma anatolicum spec. nov.: a new member of the matsutake group. Micologia e Vegetazione Mediterranea 18: 135–142. Murata H, Yamada A, Ichida H, Nakamura N, Neda H. 2023. Biodiversity of Tricholoma matsutake (syn. T. nauseosum) and its related species based on repetitive DNA and genomics. Botany 101: 138–154. Murray AF, Moore AJ, Munafo JPJr. 2020. Key odorants from the American matsutake, Tricholoma magnivelare. Journal of Agricultural and Food Chemistry 68: 9768–9775. Ota Y, Yamanaka T, Murata H, Neda H, Ohta A, Kawai M, Yamada A, Konno M, Tanaka C. 2012. Phylogenetic relationship and species delimitation of matsutake and allied species based on multilocus phylogeny and haplotype analyses. Mycologia 104: 1369–1380. Trudell SA, Xu J, Saar I, Justo A, Cifuentes J. 2017. North American matsutake: names clarified and a new species described. Mycologia 109: 379–390. Yamanaka T, Yamada A, Furukawa H. 2020. Advances in the cultivation of the highly-prized ectomycorrhizal mushroom Tricholoma matsutake. Mycoscience 61: 49–57. Zhao Z-X, Song J-G, Senanayake IC, Wu D-S, Wang G-Q, Xu B. 2022. Tricholoma tianshanense sp. nov., in Tricholoma sect. Caligata from Tianshan Mountains in China evidenced by morphological characters and phylogenetic analyses. Phytotaxa 549: 22–30.
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