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Appendix- Common and Scientific Names

Chapter 1: Matsutake: An Ancient Tradition

History of Matsutake

Matsutake have been used and revered by the Japanese people for more than a millennium and have become more than just a seasonal delicacy. They also symbolize fertility, and by extension, good fortune and happiness. A gift of matsutake is considered special and is cherished by those who receive it. According to Ohara (1994), one of the earliest records extolling its virtues is found in a 759 A.D. poem. Later references to matsutake often were related to activities of nobles and priests. Records from the 13th to 17th centuries indicate that nobility enjoyed mushrooming events and often sent matsutake as gifts, a tradition that persists today, especially in the corporate world. During the 11th century in the Imperial Court of Kyoto, women were prohibited from saying "matsutake" openly but instead were required to speak of it with the honorific marker "0," as O-Matsu. Until the 17th and 18th centuries, matsutake consumption was strictly limited to the imperial court. As matsutake consumption became more common among the public during these centuries, vulgar (phallic symbolism) and graphic short stories came into vogue, portraying comical characters attempting to conceal their matsutake picking areas and indulging in risque talk about the mushroom. It was in this same period that the first stirrings of scientific interest were recorded: A Buddhist priest in Kyoto recorded the annual productivity of matsutake (that is, mushroom numbers) in a mountain forest (Kinkakujiyama). Later, based on the priest's diary, Professor M. Hamada of Kyoto University was able to approximate the seasonal precipitation, temperature conditions, age and ecological status of the mountain forest from 1636 to 1667 (Ohara 1994).

The Decline of Japanese Matsutake

Matsutake shiros were once widespread and common in mixed pine forests of Japan from Hokkaido in the north to Kyushu in the south. After World War II, they became increasingly scarce, in spite of efforts to enhance productivity in local forests. By 1981, productivity had declined to one-tenth of the pre-War levels, and imports of Japanese matsutake, especially from South Korea, increased greatly to meet demand (Kawai and Ogawa 1981). The quantity imported today differs from season to season but can exceed 60 percent of the total retail market (EAITC 1990). Imported matsutake, especially species other than T. matsutake, are generally perceived as lacking freshness, proper color, taste, and fragrance and therefore are considered lower in quality and value. Whether or not this perception is true, improved grading, handling, and marketing strategies, coupled with a continuing decline of Japan's matsutake production, will certainly improve the image and market value of imported mushrooms.

Since 1905, the matsutake forests of Japan have been plagued by the pine nematode or pine weevil.' The nematode is transmitted to living pines by the Japanese pine sawyer, a longhorn beetle. Invasion of vascular tissue by the nematode results in wilt and rapid death (Futai 1979, 1980a, 1980b, 1980c; Futai and Furuno 1979). Most host pines of matsutake, including the Japanese black and red pines, are very susceptible to this devastating pathogen. Since the introduction of the nematode at the start of the 20th century on the southern island of Kyushu, it has steadily spread north-eastward. The current blight is the fourth in a series of epidemics since 1905. The third epidemic lasted a decade, peaked in 1979, and caused an estimated loss of 2.4 million cubic meters of pine wood. The current epidemic began in 1990, and killed enough trees in one year to build 50,000 houses. Recent reports indicate that the disease has also spread to forests of Okinawa, Taiwan, South Korea, North Korea, and China. A combination of climatic, socio-economic, and biological factors in Japan tends to increase the magnitude of the blight. Pine mortality from the nematode often increases after prolonged drought and high temperatures, which weaken the resistance of the pines to the parasite.

A change in the way communities use local forests also has adversely affected the health of pine forests and, indirectly, matsutake productivity. For at least a millennium, routine harvesting of understory shrubs and oak trees for charcoal favored growth of shade-intolerant pines and the matsutake associated with them. In post-World War II years, traditional charcoal- and wood-burning stoves were replaced by natural gas burners, and the removal of woody shrubs for charcoal diminished. Trees and shrubs that do not form matsutake mycorrhizae have consequently formed dense, brushy understories that block sunlight needed by pine seedlings. While mature pines die from nematode attacks, new pine seedlings find themselves at a competitive disadvantage and have difficulty becoming established; hence, matsutake shiros disappear along with their pine hosts as other tree species become dominant.

Essentially, the red and black pines introduced from the Korean Peninsula flourished in the past under conditions no longer existing in Japan. Although the Japanese have developed many silvicultural strategies to manage matsutake forests (discussed below), matsutake production has disappeared from the vast stretches of mountain pine forests that die each year. Indeed, Japan's temples and public parks may eventually become the last refuge for matsutake pine forests, as suggested by a respected Kyoto gardener, Mr. Shiro Nakane (c. 1992):

To me, this already seems to be the case. When I was a kid, I'd walk on
the paths right up there and gather matsutake from the many, many red
pine trees. Now I sometimes go up to the mountain with my two boys and
my dog, but the forest there has changed. The trees I remember are gone.
Still, different trees are appearing, and maybe this is nature's way.

Other Species of Matsutake
Two species of matsutake are recognized in Japan: Tricholoma matsutake and T. bakamatsutake. The first is the principle Asian species and grows widely in Japanese red pine (known as "aka-matsu") forests throughout Japan, South Korea, North Korea, northeastern China, and Primorsk Kray, Russia. In Japan, T. matsutake also associates with Japanese black pine, Japanese stone pine, Japanese white pine, southern Japanese hemlock, northern Japanese hemlock, and Sakhalin spruce. On the Korean Peninsula and in Manchuria, it associates with Korean pine; in Sakhalin with Maries fir; and in Taiwan and along the east coast of China, with Taiwan red pine (Hamada 1966, Ogawa 1982, Ohara 1981). It occurs in at least eight provinces of China under various pines (often mixed with oaks), but most often under Japanese red pine in the northeastern province of Jilin (Wang c. 1982). Tricholoma bakamatsutake or "foolish pine mushroom," is less common and differs from T. matsutake by its association with broadleaf trees, particularly Mongolian oak, serrated-leafed oak, edible Asian tanoak, and chinkapin species (Ogawa and Ohara 1978). It, too, ranges from Hokkaido to Kyushu in Japan, and is found in Taiwan, South Korea, North Korea and China. It has the same general morphology and strong matsutake odor and taste of T. matsutake, but it differs from the latter in being slightly smaller and more fragile, and in having cheilocystidia on the gills (Hongo 1974). Thus, the two can be easily confused morphologically and indeed may represent only ecotypes or variants of one "plastic" species.

The principal North American counterpart of the Asian matsutake species is Tricholoma magnivelare. It is similar in texture, taste, and odor to Asian species and has long been a favorite of Japanese-Americans and certain tribes of Native Americans. Mushroom guides often refer to the American matsutake as either T. ponderosum (Peck) Singer or Armillaria ponderosa (Peck) Sacc. The epithet "ponderosa," meaning large and heavy, is descriptive of the species. The earliest name applied to the American matsutake was Agaricus magnivelare. "Magnivelare" also describes the species well; it refers to the large well-developed veil, which breaks or tears, leaving a membranous ring on the stalk. In his 1984 review of the literature, Redhead determined that the nomenclaturally correct name is T. magnivelare (Peck) Redhead. The species is morphologically distinct from the Japanese species and is described by Smith and Weber (1980:127) as follows: "This large white mushroom gradually develops cinnamon stains as it ages, and in age is quite discolored. The cap may be up to 35 centimeters broad and covered with cinnamon colored patches of tissue. The gills are white and slowly stain vinaceous cinnamon." Arora (1986:191) ventures the most intriguing description of its unique spicy odor as a "cross between red-hot [spicy cinnamon candy] and dirty socks.

A rarer and consequently less well-known species of North American matsutake is T.caligatum, or "booted" matsutake. It occurs in North America, Europe, and northern Africa. The species is generally smaller and more slender than T. magnivefare, has a broad cap with dark vinaceous to cinnamon-brown scales or fibrillose (hair like) patches, and has the same colors on the underside of the ring and the stalk below the ring or "boot." The stalk above the ring is white and the odor of the mushroom is quite variable. As Smith and Weber (1980:127) write: "One variety has an odor like that of A. ponderosa, another has an odor somewhat like that of bitter almonds, and a third no odor at all. The Japanese matsutake is one of the varieties of this collective species." Likely the variability in odor is geographic; specimens found in western North American smell more like the Japanese matsutake than those from eastern North American (Arora 1986, Smith 1979). Tricholoma caligatum may be found in pure coniferous stands, mixed coniferous and hardwood stands, and pure hardwood stands. In general, T. caligatum seems to be closely related in size, odor, and habitat to T. matsutake, T. bakamatsutake, and T. fulvocastaneum of Japan.

Kytovuori (1989) discusses the T. caligatum group in Europe: T. caligatum and T. nauseosum are species with overlapping ranges in Europe and northern Africa. After examining many specimens, he asserts that T. nauseosum of Europe is synonymous with T. matsutake of Asia, and because T. nauseosum was named first, it is therefore the valid name. Specimens originally described as T. nauseosum and T. matsutake came from opposite sides of the Eurasian continent; hence, this claim likely will require further verification (perhaps genetic comparisons) before it is widely accepted. It would, however, be ironic if one of the most highly prized mushrooms in the world ends up with the specific epithet "nauseosum" due to the priority rule of the International Code of Botanical Nomenclature (Greuter 1994), which states that the first name validly published for a species is the correct name. Kytovuori (1989) also discusses the affinities between T. nauseosum and T. magnivelare. He further asserts that the T. caligatum of Europe is distinct from both T. nauseosum and the T. caligatum of North America.

The global taxonomy of wide-ranging species like matsutake can be confusing even for experts. To further complicate matters, commercial collectors and retailers often make odor, taste, and value distinctions among specimens of the same matsutake species collected from different areas. Species distinctions often are based on descriptions of physical characteristics, and many of these have a range of gradations. With the advent of easy global communication and techniques for genetic analysis, taxonomists have powerful tools for distinguishing between actual species and local ecotypic variation within a species. Readers are referred to Wang and others (in press) for further discussion of the global distribution of closely related matsutake species.

Look - Alike

Tricholoma zeileri is another North American species related to T. magnivelare. It is not considered a matsutake and has no commercial value, but it is often abundant and is commonly found near shiros of T. magnivelare', hence, it may be considered an indicator species (Hosford and Ohara 1990). This species resembles T. magnivelare in shape, texture, and occurrence but usually is smaller and different in color, odor, and taste. The cap is viscid at first but soon dries and cracks into small scales. The cap ranges from yellow to orange and often develops distinct olive tones in spots. The stem below the well-developed ring has colors similar to the cap. The odor and taste of T. zeileri is similar to freshly ground meal (farinaceous), with an added pungent metallic taste. Tricholoma zeileri fruits prolifically at about the same time as other matsutake in the coniferous forests of the Pacific Northwest and the Great Lakes area. It is closely related or identical to T. focale of Europe and T. robustum of Japan.

The "imperial mushroom" (Catathelasma imperiale) is a species sometimes mistaken for the American matsutake by novice collectors. Although typically larger and lacking the distinctive odor of the American matsutake, the imperial mushroom otherwise has a similar appearance. It is edible and collected commercially.

Deadly Amanita mushrooms are distinctive from matsutake, but they can fruit within meters of each other (Pilz, personal observation), and poisonings from mistaking Amanita smithiana for matsutake have occurred (Benjamin 1995, Leatham and others 1997, Tullos and Lindgren 1992). Amanita mushrooms have a brittle texture, whereas matsutake have a fibrous texture (they can be peeled like string cheese). The matsutake odor is also distinctive, when one is familiar with it. The usual caveats apply: always make sure of your identification before eating any mushroom, and then start with small quantities.

Ecology of the Japanese Matsutake
The roots of ecological knowledge stem from ancient tradition. During the Edo Period (1603-1867), a number of books contained descriptions (including habitat) and illustrations of matsutake (Fig. 1). The book Honcho Shokkan (1697) was the first to recognize the symbiotic relationship between matsutake and pine and to promote matsutake cultivation by transplanting soil from matsutake shiros. No data on either the success or failure of transplanting is mentioned. Early research  by Inoue in 1930, Kaneyuki in Hiroshima in 1955, Senbara in Kyoto in 1937, and Asada (1937) added greatly to our knowledge of matsutake biology and ecology and promoted further study. Senbara focused ecological research on the shiro and its management and developed techniques later refined by Professor M. Hamada at Kyoto University. His principal graduate students, M. Ogawa, H. Ohara, N. Sagara, and K. Kinugawa, contributed information on matsutake mycorrhizae, microbial interactions in the shiro, chemical treatment of the shiro, and effects of temperature on formation of the mushrooms. The research of Hamada (1950), his students and others, such as Tominaga (1963, 1973, 1978), is embodied in the following discussion of matsutake shiros and forest management.
Ohara (1994:29) defines a matsutake shiro as "...a subterranean biotic community where mycorrhizal development plays a leading part over the soil constituents, especially soil microbes." Ohara's definition emphasizes the key importance of the fungus-root association in a shiro. The annual growth of a matsutake shiro (initially located by the emergence of mushrooms) is typically followed by inserting bamboo or flag markers at the exact point of fruiting. A few seasons of such marking provide an accurate record of mushroom production and a map of the shiro's growth. The growth of the shiro is directly correlated to mycorrhizal development in the soil beneath the organic layer. Several zones are recognized in a soil profile of a shiro (Fig. 2). Zone III, the currently active mycorrhizal zone (AMZ), contains the mycorrhizae directly below the mushroom. When fruiting occurs, the soil in this zone becomes strongly desiccated and the mycorrhizae begin to deteriorate. The AMZ contains about 10 times the number of fine rootlets (of Japanese red pine in this case) compared to the zones on either side of it. Zone II contains young, physiologically active mycorrhizae that will produce mushrooms the following season. The mycorrhizae of both zones II and III appear as broomlike clusters. Structurally, the mycorrhizae are classified as ectomycorrhizae but, unlike most ectomycorrhizae, have loose, soft mantles with poorly developed Hartig nets. Zone I contains mycelium and structural roots but no mycorrhizae. In zones IV and V (representing the previous two years of fruiting), the soil is still granular and desiccated and contains decayed mycorrhizae. Finally, in zone 0, there is no mycelium, and zone VII shows complete recovery from the earlier effects of active mycorrhizae. All zones (O-VII) can be recognized in the field by variation in the color of the soil profile, as well as soil texture, relative moisture, and odor (the unique matsutake aroma). Ogawa and Hamada (1965), Ohara (1966), and Ohara and Hamada (1967) also report qualitative and quantitative differences in microbial populations for each zone, especially the AMZ. Fruiting bodies begin to develop in the AMZ when soil temperature falls below 19 C. About 20 days after initiation, the cap of the fruiting bodies opens (Kinugawa 1963). Fruiting typically ceases when shiro temperatures fall below 10 C. The species of plants and fungi present, their densities, health, age and ecological effects on shiro productivity have been carefully studied (Ogawa 1982, Ohara 1974). The results have been applied to forest management techniques intended to enhance matsutake production.

Enhancing Japanese Matsutake Production

The following sections summarize management techniques employed to enhance the production of matsutake (numbers and size of the mushrooms) in Japanese red pine forests. More complete discussions of the subject can be found in books (in Japanese) by Ogawa (1982) and Ogawa and lto (1989) and papers by Ohara (1994) and Sagara and Hamada (1965). Although this discussion pertains to research with the Japanese matsutake, many of the techniques may prove applicable to the American matsutake.

Site Selection and Vegetation Management

Important criteria for site selection include age of the pine trees; composition, density, and stratification of tree species; canopy coverage; slope; and soil characteristics, including moisture and temperature. Pines 40 to 50 years old are best for matsutake production, but those over 50 exhibit declining production. Forests consisting predominantly of pines, with few other tree and shrub species, are most favorable for matsutake production. Ideal conditions also include an open canopy that allows light to penetrate to a sparsely vegetated forest floor, and warm, well-drained soils with thin litter and organic layers. Pine forests positioned on southwestern slopes and ridge tops tend to be most suitable. Unfortunately, few sites are naturally optimal for matsutake production.

Forest management treatments employed for enhancement of matsutake production differ from those used to maximize timber production. The primary objective is to alter tree species composition, stand density, and soil conditions to encourage shiro development. Ogawa (1982) has designed a schedule of silvicultural treatments that produces ideal conditions for the establishment, growth, and fruiting of matsutake shiros in a 30-year-old Japanese red pine forest. A similar set of treatments also may be applied to younger stands over a longer period. This management regime produces an open canopy and forest floor that is warm, dry, and free of excessive debris and organic material. Under these conditions, pine roots proliferate near the soil surface while many non host roots, bacteria, and saprobic and other mycorrhizal fungi decline. In an experimental forest near Kyoto, these management procedures increased the number of shiros from 12 to73 over 15 years (Ogawa 1982).

The first year includes thinning of pines and other competing tree species, removal of shrubs, herbs, dead branches, and damaged or diseased trees. The increased light penetration and air circulation reduce soil moisture and increase soil temperature.

Vegetation management during the second year includes removal of new sprouts, weeding, and tree trimming. Soils with excess accumulation of litter and organic matter are raked to expose the mineral soil. In effect, this eliminates pine roots and pests, which tend to accumulate in the litter layer, decreases soil moisture, and increases soil temperature to levels conducive to mushroom production. On the other hand, if needed, a bare forest soil may be mulched with rice straw, to increase moisture and lower temperature to levels again more conducive to the matsutake. The third year of treatment is much the same as the second, but may include another thinning of pine and pruning of non dominant trees. New sprouts, weeds, and excess litter are again removed. By autumn of the third year, matsutake shiros usually appear and marking of mushrooms begins. After the third year, management consists of annual inspection and cleanup of the forest floor, removal of dead trees, and biannual pruning of understory vegetation. Vigorous growth and gradual increase in numbers of shiros usually occurs between the fifth and tenth years of treatment. After the tenth year, production of mushrooms drops as the growth rate of maturing red pines declines.

Fruiting Enhancement

Several techniques, other than those discussed above, have been used to stimulate the formation of matsutake primordia (small "protomushrooms" with the potential to develop), increase the number of mushrooms that actually develop from the primordia, or increase the eventual size and weight of the mushroom. Most of these techniques operate by controlling humidity and temperature.

The "Hiroshima tunnel" (Plate 8) is one such method, where a tunnel like construction is placed over the physiologically active mycorrhizal zone (AMZ 11-111). The tunnel roof is formed with fine-mesh netting or screening (as used in greenhouses) stretched over wire wickets arching to a height of 30 centimeters and spanning up to 1 meter in width. The length of the tunnel differs with the length of the active shiro front. The tunnel shades and cools the soil to 20 C or less for 4 to 5 days to initiate the development of primordia. The tunnel netting also maintains soil moisture levels and may control some insects.

Tominaga (1975) and other Japanese have reported success with this method. Tominaga (1975, 1978) compared two shiros over 3 years by using variable treatments, including artificial sprinkling of water, air-conditioning, and ice, beneath the tunnel to control moisture and temperature. Some treatment combinations resulted in two to six times greater mushroom production and 12 to 20 days earlier production than without a tunnel.

On a smaller scale, the size of individual mushrooms and their rate of maturation may be increased by covering them with damp soil or a small cup as they begin to emerge, thus maintaining higher humidity and preventing the stunting of their growth through desiccation (Lee 1989a). Plastic cups also may reduce insect damage by acting as a barrier.

In the previous section, we described how foresters open forest stands to allow sunlight to warm the forest floor, and in this section we describe techniques for cooling the mycelium and sustaining high humidity. Although these activities may appear contradictory, the vegetation management is intended to promote the establishment and growth of shiros and the formation of matsutake mycorrhizae with selected host trees, whereas the cooling and humidity control are more localized treatments to enhance fruiting.


In addition to fruiting enhancement, various inoculation or "seeding" methods have been tried over the years in attempts to establish and spread matsutake shiros to new areas. Methods include inoculating seedling roots before transplanting, transferring soil from the active mycorrhizal zone of established shiros, and spreading spores into new forest sites.

Seedling roots have been successfully inoculated in both field and laboratory. Field inoculation involves planting mycorrhiza-free seedlings in active shiros to form matsutake mycorrhizae, and subsequently (usually the next year) transplanting them into appropriate habitat lacking matsutake shiros. Laboratory inoculation involves isolating a sterile culture of matsutake from the aseptic interior of a mushroom, growing it on Hamada's medium (Hamada 1964), and placing it in contact with mycorrhiza-free seedling roots. New matsutake shiros, unfortunately, have not yet been successfully produced by planting inoculated seedlings. Soil transfers from active shiros to likely habitats also have been attempted without success at establishing new shiros. For further information on these techniques, refer to Ogawa 1982, Ogawa and lto 1989, and Lee 1989b.

The simplest inoculation method is to cut a fully opened mushroom cap from its stem and bury it near host plant roots or place it on the soil surface within a young pine stand. Litter and organic layers should be removed from the soil to improve the chance that spores will come into contact with appropriate host roots. Spore suspensions also may be created by mixing spore prints or macerated gills in water. Spore suspensions are then poured into holes in the soil near pine roots or mixed with soil placed around the roots. Japanese matsutake spores are short-lived and few apparently germinate. Repeated application as close as possible to young, non colonized host roots often is necessary. The use of spore dissemination to establish new matsutake shiros has had a long history. Those who claim success are those who cleaned the forest floor of excess organic material and sowed spores frequently over 5 years or more (Ogawa 1982).

Although attempts to establish new matsutake shiros have not yet yielded much success, plantation establishment and management for matsutake production holds the greatest promise for artificial cultivation. The high value of matsutake suggests this approach may be economically feasible. Many of the techniques for truffle (Tuber species) cultivation in plantations established with inoculated seedlings may be adapted eventually to create matsutake plantations. Progress with this approach has been demonstrated recently with chanterelle mushrooms (Danell 1994, Danell and Camacho 1997). Even if the technology to do this is developed, success will depend on using appropriate host trees, selecting adaptable matsutake strains, and establishing the plantations in appropriate habitat (especially proper soils). Artificially established matsutake plantations are unlikely to be developed on Federal lands in the United States because other species would be displaced, an effect that would conflict with management goals for preserving biological diversity and multiple use. Managing natural stands (through activities such as thinning or controlled burns) that already have matsutake populations does hold promise, however, for enhancing fruiting within the context of Federal ecosystem management goals. Plantations for American matsutake production may eventually become economically attractive for private land owners.

Pure Culture Production

Growing matsutake in pure culture and inducing it to fruit (without its mycorrhizal tree symbionts) has been a goal of researchers and commercial interests for many years. Unlike saprobic fungi that decompose organic matter, symbiotic mycorrhizal fungi have complex biochemical interactions with their plant hosts. Although most mycorrhizal fungi can be grown in pure culture, only a few have been induced to fruit without colonizing the roots of a host plant (Ohta 1994; Pantidou 1961, 1962, 1964). Both Japanese and American matsutake grow very slowly on nutrient media alone, and innovative biotechnology developments will likely be needed to induce matsutake to fruit in the absence of its tree partners.
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