2 McIlvainea

A Mycological Florilegium: A Selection of Articles Published in 2007 and 2008 Else C. Vellinga*

MYCOLOGY IS is a booming and blooming fi eld Lots to Do about Fly Agarics of science, and mycologists are prolifi c writers. Geml, J., R. E. Tulloss, G. A. Laursen, N. A. Sa- The result is that many fungal papers are published zanova, and D. L. Taylor, 2008. Evidence for every year, and they range widely; recently there are strong inter- and intracontinental phylogeo- biochemical studies of how a gene works in the de- graphic structure in Amanita muscaria, a wind- velopment of a small dung denizen, inventories of dispersed ectomycorrhizal basidiomycete. Mo- the mycofl ora of a hemlock forest in Ontario, the lecular Phylogenetics and Evolution 48: 694–701. description of new species and a comprehensive phylogenetic study of the entire kingdom Fungi. Geml, J., G. A. Laursen, K. O’Neill, H. C. Nus- An eclectic and highly personal choice of baum, and D.L. Taylor, 2006. Beringian ori- topics is presented here to bring you some of the gins and cryptic speciation events in the fl y most recent and noteworthy literature about your agaric (Amanita muscaria). Molecular Ecology 15: favorite organisms. The selection covers all types 225–39. of research, comes from all over the globe, and Oda, T., C. Tanaka, and M. Tsuda, 2004. Molecular in several cases involves professional and amateur phylogeny and biogeography of the widely mycologists working together as a team. distributed Amanita species, A. muscaria and What stands out for me, after compiling A. pantherina. Mycological Research 108: 885–96. this overview, is that with molecular data adding a whole new dimension to what we knew about The fl y agaric has recently been the subject of in- mushroom morphology and classifi cation, the tensive study, starting with the Japanese research- world of mushrooms has become more fascinat- ers Oda, Tanaka, and Tsuda who concluded that ing, more complex and more challenging than we there are two distinct taxa under the name Amanita could have dreamt. The number of questions to be muscaria, one in America, and one in Eurasia, as answered just keeps growing. well as the high-latitude and high-altitude species Also striking is how many of these studies rely Amanita regalis. The Alaska based group of Lee on fi eld observations, and well-described collec- Taylor and Gary Laursen did a follow-up study tions, showing that fi eld mycology is still the basis and added Laursen’s Alaska collections to those of for all other types of research. Amateur mycolo- the Japanese. Lo and behold! All three taxa occur gists can really help. in Alaska; could it be that this is the original birth If this smorgasbord makes you hungry for place for all fl y agarics? But this was not the last more mycological information, but you do not word; a recently published third study includes live next door to a good mycological library, be collections from far eastern Russia, just across the aware that most journals post abstracts of their Bering Strait, and also from many other parts of articles online— just use the name of the journal the U.S.A. With more samples from more places, in a search engine to start exploring. Some journals the picture becomes clearer and more confused now allow free online access to the whole article, at the same time. One group is found to be wide- either immediately or after a modest delay; this is spread throughout the northern hemisphere from indicated below when it applies to one of the ar- Europe eastwards to Alaska (though sampling ticles on my list. from northern Canada and Greenland is spotty). Another big group is represented in various parts * Department of Plant & Microbial Biology, University of North America. Then, there are several really of California at Berkeley different (different in this case means different in

2 McIlvainea 18 (1) 2009 Volume 18, Number 1, 2009 3 genes), smaller groups in the southern states, in the through the New York Botanical Garden) de- Pacifi c Northwest, and especially on the Channel scribes many species, but does not place them in a Islands off the coast of southern California. Santa phylogenetic context, nor makes it easy to identify Cruz Island, the biggest of these islands, is rugged, them. hardly inhabited, and harbors at least four different Some progress has recently been made by sev- types, only one of which falls within the big North eral research groups tackling the genus in a wide American group. Such variation on such a small context, examining only a few sections, or focusing scale is remarkable, and makes one wonder how on the dung-dwellers. speciation takes place, and whether spore dispersal The fi rst study tackles the whole family Psathy- is very short-range. Would more sampling unearth rellaceae, including the ink caps Coprinellus, Parasola, more variation, especially in northern Canada, in and Coprinopsis. By the time these three genera were Mexico and its western islands, Central America, recognized as separate from Coprinus in the strict and in the southern states? sense (with shaggy mane, C. comatus, and C. sterqui- The genetic variation and the biogeographical linus, its sole surviving members, expelled to the structure within “Amanita muscaria” have been pre- Agaricaceae), it was clear that their relationships with sented but not yet the morphological descriptions Psathyrella had to be re-examined. It now appears, and conclusions. that the family Psathyrellaceae as a whole is mono- Inevitably, the studies raise questions about phyletic (meaning that all its members share a other well known and easily recognized species. most recent common ancestor and all descendents Is there any such thing as a cosmopolitan species? of this ancestor are members of the family), but And do we have more than one Boletus edulis and that the genus concepts have to be reconsidered. Cantharellus cibarius in North America? Thirteen different groups stand out; some consist If you are interested in further aspects of the only of species from one morphologically defi ned fl y agaric and the roles it has played in cultures and genus, but others are a mixed bag. For instance, folk tales, check out Dugan’s article in North Ameri- Psathyrella conopilus forms a group together with can Fungi 3 (see below). Parasola species (ink caps with pleated caps, such as P. plicatilis) while some Psathyrella species, like Ps. Psathyrella and the Demise of a Genus marcescibilis, fall smack in the middle of Coprinopsis (the genus for C. atramentarius, C. lagopus, C. radiatus, Padamsee, M., P. B. Matheny, B. T. M. Dentinger, C. friesii, C. narcoticus and many others). Though this and D. J. McLaughlin, 2008. The mushroom paper clearly indicates that the old genus Psathyrella family Psathyrellaceae: Evidence for large-scale cannot be maintained in its present form, new gen- polyphyly of the genus Psathyrella. Molecular era or groupings are not proposed. Phylogenetics and Evolution 46: 415–29. The Swedish paper is a rare example of a Larsson, E., and L. Örstadius, 2008. Fourteen combined approach, with descriptions of all the coprophilous species of Psathyrella identifi ed fourteen species encountered, including two new in the Nordic countries using morphology ones, and placement of all species in a phylogenet- and nuclear rDNA sequence data. Mycological ic context using molecular data. The ecology is also Research 112: 1165–85. treated, and a key to the European dung-inhabiting species is given. Vašutová, M., V. Antonín, and A. Urban, 2008. Phy- The third paper looks in detail at two sections, lo genetic studies in Psathyrella focusing on sec- section Spadiceae in which the species have rela- tions Pennatae and Spadiceae—new evidence for tively small spores, and thick-walled cystidia, and the paraphyly of the genus. Mycological Research section Pennatae, also with small spores, but with 112: 1153–64. fl ask-shaped to wedge-shaped, often thick-walled Much less visually appealing than the fl y-agarics pleurocystidia. A nice thing about this paper is that are members of the genus Psathyrella. Many grey, the morphological characters are indicated in color brown, purple, cream species with pale to dark on the phylogenetic tree. brown spores cry out to be classifi ed. The clas- The general conclusions are the same as in sic monograph by Alexander H. Smith (available the fi rst paper: Psathyrella is a mess and needs to be 4 McIlvainea

sensibly split up, with some reshuffl ing of species. Agaricus—Names and Occurrences We are waiting for a courageous person to do the Geml, J., G. A. Laursen, and D. L. Taylor, 2008. carving. Molecular diversity assessment of arctic and boreal Agaricus taxa. Mycologia 100: 577–89. Polypores in the Age of Molecular Phylogenetics Kerrigan, R. W., (“2007”) 2008. Lectotypifi cation Lindner, D. L, and M. T. Banik, 2008. Molecular of Agaricus brunnescens. Mycologia 99: 906–15. phylogeny of Laetiporus and other brown rot polypore genera in North America. Mycologia It is very easy to recognize a member of the genus 100: 417–30. Agaricus as such, but species recognition is hard. The species are not very rich in morphological Sotone, K., H. Tsutome, Y. Ota, C. To-anun, B. characters and, on top of that, their appearance Salleh, and M. Kakishima, 2008. Phylogenetic changes signifi cantly during maturation. Micro- relationships of Polyporus and morphologically scopically, too, only a few characters, in particular allied genera. Mycologia 100: 603–15. spore size, are informative. This paucity of mor- We are slowly getting used to the notion that the phological characters is paralleled at the molecular genus Laetiporus, the sulphur shelf, is represented level in the low differentiation of the ITS region by more than one species in North America; it (Internal Transcribed Spacer, a part of the DNA is not all L. sulphureus,. We have learned that the that does not code for the making of proteins). deep orange species growing on conifers in the In many species of mushrooms, ITS is a good West is not the same as the white-pored pink- species recognizer, and can be used as a universal ish species on hardwoods in the Midwest, and ‘bar code’, but closely related Agaricus species often indeed that there are at least fi ve species in the differ only in a few base pairs in the ITS region. continental USA. Mating studies, molecular data, Perhaps other parts of the DNA might serve bet- and morphology showed this quite convincingly. ter for species recognition, but so far nobody has Now, the genus Laetiporus is put in a phylogenetic looked into it seriously. The present study focuses context and compared with genera like Wolfi poria, on the Alaskan species, which represent only a few Phaeolus, and Antrodia. The rare species L. persicinus, sections. The Arvenses and the Campestres, both known from the Gulf states, is separate from all notoriously diffi cult groups, are well represented. the other Laetiporus species. Wolfi poria is split up; Agaricus bisporus is found too but the reddening W. dilatohypha becomes a sister to Laetiporus, while species, and the toxic phenol-smelling taxa around the well-known and sought after W. cocos becomes Agaricus xanthodermus, were not found despite ex- a sister to the dyer’s polypore Phaeolus schweinitzii. tensive sampling in time and space. The authors Several new taxa are discovered, e.g. one sulphur refrain from naming their species, but suggest that shelf species growing on Koa trees in Hawaii, but their area contains several which are undescribed. none is offi cially named and described, nor are the One name in Agaricus that has received a lot non-Laetiporus species placed in new or other exist- of scrutiny is Agaricus bisporus (J. E. Lange) Imbach, ing genera. since it is used for the most commonly cultivated A similar situation appears to be true for mushroom species, the button mushroom. But Polyporus which splits up into fi ve rather non-diver- should we be calling it Agaricus brunnescens Peck? gent groups. One part of the genus (P. brumalis and Kerrigan becomes a historical sleuth to tackle P. arcularius) separates from all the rest and should this lingering problem; he looks at Peck’s letter be considered a separate genus that is close to Len- exchanges in the late 19th century and examines tinus. Some species from other genera are found Peck’s collections which are still available. Just in the middle of the old Polyporus. The authors like any modern-day detective, he uses molecular consider making separate genera for all the new forensic methods to determine what the original groups, but this is going beyond the data and is Agaricus brunnescens collection really is. It turns out not a road I am ready to take. A mixed approach, that Peck packed two different species under one combining morphology with molecular, and mat- label. That does happen. One part of the collection ing data, is necessary to clear up the confusion in (the 22 different bits and pieces were carefully split this genus. Volume 18, Number 1, 2009 5

between the two species) is chosen to represent nominated for whole genome sequencing. Other what Peck meant by the name A. brunnescens, the well-known ectomycorrhizal species, such as those rest is Agaricus subrufescens. The Agaricus brunnescens in Russula and Tricholoma, are very hard to grow in part is the same as what we have been calling A. culture, and impossible to experiment with. bisporus. Because A. brunnescens is the earlier name, To fi ll out the picture, the non-mycorrhizal a formal proposal will be put forward to keep the species of Paxillus were transferred earlier to Tapi- well-known name A. bisporus instead of renaming nella—you can now fi nd P. panuoides and the dye the button A. brunnescens. mushroom P. atrotomentosus in that genus.

Paxillus —More Species Th an We Th ought Spore Secrets Hedh, J., P. Samson, S. Erland, and A. Tunlid, 2008. Bruns, T. D., K. G. Peay, P. J. Boynton, L. C. Multiple gene geneaologies and species recog- Grubisha, N. A. Hynson, N. H. Nguyen, and nition in the ectomycorrhizal fungus Paxillus N.P. Rosenstock, 2009. Inoculum potential of involutus. Mycological Research 112: 965–75. Rhizopogon spores increases with time over the fi rst 4 yr of a 99-yr spore burial experiment. Paxillus involutus is often one of the fi rst mushrooms New Phytologist 181: 463–70. one learns, with its slightly ribbed cap margin, and gills that peel off the underside of the cap as eas- Nara, K., 2009. Spores of ectomycorrhizal fungi: ily as the tubes come off a bolete. It is distinctive ecological strategies for germination and dor- and ubiquitous. European research had established mancy. New Phytologist 181: 245–48. the presence of more than one species, that were Ishida, T. A., K. Nara, M. Tanaka, A. Kinoshita, not mating with each other, but not all had been and T. Hogetsu, 2008. Germination and infec- named. Now, data from fi ve different gene regions tivity of ectomycorrhizal fungal spores in re- in 50 collections show that four different Euro- lation to their ecological traits during primary pean taxa can be distinguished: P. involutus sensu succession. New Phytologist 180: 491–900. stricto, P. validus, P. obscurosporus, and one unnamed species. The alder associated Paxillus rubicundulus Kauserud H., J. E. Colman, and L. Ryvarden, 2008. is totally separate from the P. involutus group. It Relationship between basidiospore size, shape still belongs in Paxillus though its closest relative, and life history characteristics: A comparison Paragyrodon sphaerosporus, has pores instead of gills. of polypores. Fungal Ecology 1: 19–23. The North American species Paxillus vernalis, rep- One of the big questions in mushroom ecology is resented with one sample in this study, is found to what role the spores play in dispersal of individu- be close to P. validus and P. obscurosporus. That is also als and in distribution of species. Mushrooms are my own conclusion, based on work on the Califor- spore factories, pumping out millions of spores, nian species. Here in the west of the USA we fi nd but how many actually form a new individual, how P. vernalis in natural habitats associating with oaks, far away from the “mother” mushroom do they and aspens, the unnamed species with introduced get and how long do they stay viable? These are all birch, and a third, chunky, species, whose identity open questions. is not yet known, growing with introduced Cedar After a disastrous wildfi re on the California (Cedrus deodara), and with our native coastal pines. coast in 1995, Bishop pine seedlings sprouted But the real P. involutus has been found in the mid- in many of the burned areas. They could do so west and the east. because of the presence in the soil of Rhizopogon For this group at least, the single universally (false truffl e) spores that survived the heat of the used mushroom bar-coding ITS region is as good fi re. During the succession and the aging of the at distinguishing species as fi ve gene regions com- forest, the composition of the mycofl ora changes, bined. different Rhizopogon species, and many other spe- Paxillus involutus grows easily and fast, making cies occupy and compete for the tree roots. The it the perfect lab rat for ectomycorrhizal research, early colonizers are the species whose spores stay and causing it to be one of the few mushrooms viable in the soil for extended periods. To assess 6 McIlvainea how long, very long-term studies are under way A different kind of study looked at the size and the fi rst results, obtained four years after the and shape of spores of 303 species of European experiment began, are now reported. It is very polypores, trying to fi nd a correlation between refreshing to see a research plan with a focus that size and shape of the spores and the character- extends so far beyond the here and now that it will istics of the fruitbodies—their size and life style occupy several generations of researchers. (saprotrophic or parasitic), the kind of rot they A grassy area where Rhizopogons do not oc- produce and whether they grow on the wood of cur and pine trees are not found, was selected to conifers or hardwoods. The conclusions were that bury pots containing soil mixed with Rhizopogon the bigger the fruitbodies, the bigger the spores, spores. The pots cannot be penetrated by tree and the bigger the spores the more globose they roots and one by one, over a period of 99 years, are. Parasites tend to have bigger spores and there they will be dug up, and the soil they contain used is a tendency for white rot species and parasites to to grow pine trees. After the treelets have been in have more rounded spores than brown rotters and the soil for several months, they will be dug up and saprotrophs. Now we only need to fi nd out what, the viability of the spores will be measured by the if anything, these trends mean in nature. amount of mycorrhization of their roots. After the fi rst four years in the soil, spores of all four Rhi- Observations and Predictions zopogon species tested in the experiment did well. about the When and Where In fact, they actually did better than when spores Gange, A. C., E. G. Gange, T. H. Sparks, and L. were taken from fresh fruitbodies before the burial Boddy, 2007. Rapid and recent changes in experiment. Compare that to tropical Marasmius fungal fruiting patterns. Science 316: 71. species, whose spores drop and lose their ability to germinate within 24 hours. Kauserud, H., L. C. Stige, J. O. Vik, R. H. Økland, A different approach was taken by a Japanese K. Høiland, and N. C. Stenseth, 2008. Mush- group studying the shrub willows and their fungi room fruiting and climate change. Proceedings on the dry and sparsely vegetated slopes of Mount of the National Academy of Sciences of the USA Fuji. The fi rst species to arrive on the roots of 105: 3811–14. newly settled plants are Inocybe lacera, Laccaria lac- Krebs, C. J., P. Carrier, S. Boutin, R. Boonstra, and cata and L. amethystina, followed by L. murina and E. Hofer, 2008. Mushroom crops in relation Scleroderma bovista. In the last stages of develop- to weather in the southwestern Yukon. Botany ment there is a more varied mycofl ora, with some 86: 1497–1502. Russula, Hebeloma, and Cortinarius species. The researchers made spore prints of all the mush- Wollan, A. K., V. Bakkestuen, H. Kauserud, G. rooms, and sandwiched roots of willow seedlings Gulden, and R. Halvorsen, 2008. Modeling between bits of the spore prints. After a while they and predicting fungal distribution patterns examined how the spores were doing, and whether using herbarium data. Journal of Biogeography the mushroom mycelium could establish mycor- 35: 2298–2310. rhiza with the plants. Spores of ectomycorrhizal The ongoing changes in climate, especially in fungi need signals from the plant roots in order to temperature, have wide ranging repercussions in germinate at all. The fi rst colonists did reasonably the natural world. Plants are fl owering earlier in well, in so far as both germination and coloniza- spring, leaves are turning yellow and red later than tion took place, but the later-stage fungi were more before, and now two studies show that mushroom reluctant and only one Hebeloma species was eager fruiting patterns have changed dramatically in the to germinate—it was one of the late succession last 50–60 years. The fi rst study analyzed the fi nds species. The spore prints were retained, and the recorded by a group of mushroom hunters during experiment was repeated after one year, but ger- the period 1950–2005, in the course of walks in mination then was very low and in most cases just and around the English city of Salisbury. The sec- plain absent. There was enormous variation in the ond study examined 34,500 herbarium records for number of germinating spores per species. collections made from 1940 onwards in Norway. Volume 18, Number 1, 2009 7

The English mushroom season has expanded in fi ngerprints. Since the original discovery in 2003 both directions, with more early and more later from the Colorado Rocky Mountains, and thanks fruiting, whereas in Norway, the entire season has to Terry Porter, who pieced together the data, shifted since 1980, with species fruiting 13 days lat- their occurrence has been established in other er than in the 1950s. As Norway is a country with parts of North America, a tropical forest of Costa huge variation in climate and vegetation, there are Rica, dry forests of Australia and various places in considerable regional differences. Europe. The new fungi are currently known under A second Norwegian study again used her- a provisional and very prosaic name—Soil Clone barium collections to model and predict fungal Group I—and they form a new branch of the phy- distribution. Temperature was found to be a strong lum Ascomycetes, in between the Taphrinomycotina determinant for the presence of a certain species (fi ssion yeasts, leaf curl disease) and the budding fruiting at a certain place. This means, of course, yeasts of the Saccharomycotina (bakers’ yeast), with that changes in temperature will change the com- Pezizomycotina (all other ascomycetes including position of the mushroom fl ora, which will fi rst morels, truffl es, and various moulds and mildews) show in the fruiting patterns. constituting a fourth branch. Studies such as these are possible in Europe since data has been painstakingly and presciently Inocybe—What Are the Unknowns of collected over long periods. Both Norwegian stud- Which We Have Only a Genetic Footprint? ies show the special relevance of herbarium data Ryberg, M., R. H. Nilsson, E. Kristiansson, M. Tö- in documenting change, and in predicting distribu- pel, S. Jacobsson, and E. Larsson, 2008. Min- tion patterns—setting a good example for other ing metadata from unidentifi ed ITS sequences places to follow! in Genbank: a case study in Inocybe (Basidiomy- The Canadian study examined above ground cota). BioMedCentral Evolutionary Biology 8: 50 (14 mushroom production in the Yukon over a period pp) (open access at www.biomedcentral.com). of 15 years and tried to fi nd a correlation with weather patterns during the summer months. It It is now routine to inventory mycorrhizal fungal turned out that there was a high correlation be- communities below-ground, by comparing se- tween the amount of rainfall in June of the pres- quence data derived from mycorrhizal root tips or, ent year, but even more so between the rainfall in increasingly, soil samples, with the corresponding May the preceding year, and the total weight of the data from fruitbodies. But Genbank, the public mushrooms. Whether there are any correlates with database where these sequences are deposited, the depth and length of the snow cover is still to has a dearth of sequences derived from correctly be explored. identifi ed fruitbodies. Consequently, many new sequences were entered under imprecise names Unseen, but New and Exciting such as Agaricales sp. and Inocybe sp. Careful com- parison of these unmatched sequences with data Porter, T. M., C. W. Schadt, L. Rizvi, A. P. Martin, based on identifi ed fruitbodies of the genus Inocybe S. K. Schmidt, L. Scott-Denton, R. Vilgalys, took place. Now a species name can be attached to and J. M. Moncalvo, 2008. Widespread occur- about one third of the unknown sequences, and rence and phylogenetic placement of a soil the name of a species group to many of the oth- clone group adds a prominent new branch to ers. Inocybe species have turned up in studies made the fungal tree of life. Molecular Phylogenetics and all over the globe, including the tropics and boreal Evolution 46: 635–44. and arctic regions. Several species are widespread, Schadt, C. W., A. P. Martin, D. A. Lipson, and S. K. and are now known from the entire temperate Schmidt, 2003. Seasonal dynamics of previ- zone. This paper is the fi rst to use this approach to ously unknown fungal lineages in tundra soils. identify the unnamed sequences in Genbank and Science 301: 1359–61. its success shows clearly the potential for naming and placing the sequences in a genus context. The Nobody has yet seen this newly discovered group constraint is that this is only possible when suf- of soil fungi, which are known only by their DNA fi cient sequences have already been derived from 8 McIlvainea named fruitbodies. Building up such a reference over 400 million years. Speculations abound but we database in Genbank has enormous value for envi- will probably never know for sure what this fossil ronmental sampling and providing these names is really is. Artistic renditions of the organism and its a project to which all mycologists and mushroom environment have been featured widely, e.g. on the lovers can contribute. back of the April 2008 issue of Field Mycology. Just as speculative are the claims that fungal Fossils remnants in amber from the Cretaceous period, some 100 million years ago, represent nematode- Boyce, C. K., C. L. Hotton, M. L. Fogel, G. D. trapping fungi. Some hyphal remnants, some Cody, R. M. Hazen, A. H. Knoll, and F. M. spores and a lonely nematode were the grounds for Hueber, 2007. Devonian landscape heteroge- this claim, on which a new genus and species were neity recorded by a giant fungus. Geology 35: described. The authors speculate that the species 399–402. was present in a fi lamentous stage and in a yeast- Schmidt, A. R., H. Dörfelt, and V. Perrichot, 2007. like stage. Evidence, however, is scant and we will Carnivorous fungi from Cretaceous amber. probably never know whether these were indeed Science 318: 1743. nematode trappers. Schmidt, A. R., H. Dörfelt, and V. Perrichot, 2008. Megacollybia Palaeoanellus dimorphus gen. et sp. nov. (Deu- teromycotina): a Cretaceous predatory fungus. Hughes, K. W., R. H. Petersen, J. L. Mata, N. H. American Journal of Botany 95: 1328-1334. Psurtseva, A. E. Kovalenko, O. V. Morozova, E. B. Lickey, J. Cifuentes Blanco, D. P. Lewis, Today’s fungi are very intriguing and in many cases E. Nagasawa, R. E. Halling, S. Takahashi, M. we have no clue how they live. Structures preserved C. Aime, T. Bau, and T. Henkel, 2007. Megacol- from the past pose an even greater challenge lybia (Agaricales). Report of the Tottori Mycological for interpretation and understanding. One such Institute 45: 1–57. enigma is Prototaxites, which lived 400 million years ago; their fossils are known from many different A combined molecular-phylogenetic and morpho- places and are preserved as pillars up to 9 m high. logical study covering many parts of the Northern This structure has been called a tree, a red alga, a Hemisphere provides an in-depth study of Megacol- lichen or just a fungus. Most recently, researchers lybia. This genus was always considered to contain who cross-sectioned the structure have compared one species, M. platyphylla. The differences in mor- the thick-walled cells to those of polypore hyphae. phology, though mostly qualitative, in sequence Carbon from Prototaxites fossils was compared with data and in distribution seem to be suffi cient to carbon from plant fossils in the same environment distinguish six species besides M. platyphylla, with and of the same age. Differences were found that the latter not occurring in America. The genus suggest a different lifestyle and appear to corrobo- splits up nicely into a Eurasian group and a New rate that, indeed, Prototaxites was a fungus. How- World group with the new species M. rodmani, M. ever, the organism lived at a time when there were subfurfuracea, M. texensis, and the transferred Tri- only some small plants, and it would have towered cholomopsis fallax, now Megacollybia fallax, in North over them. The big question is what food source America. The article was published in a less acces- did it have? Was the bacterial crust rich enough sible journal, but fortunately the pdf is available on in minerals and proteins to sustain this huge the fi rst author’s web page. fungus, or was it just very long-lived? Could the small plants furnish all the cellulose and lignin the Laccaria Bicolor Genome fungus needed? Unfortunately, the evidence from Martin, F. + many other authors, 2008. The ge- carbon is not entirely convincing, as there is a big nome of Laccaria bicolor provides insight into variation over time, suggesting different lifestyles ectomycorrhizal symbiosis. Nature 452: 88–92. at different times. Neither is it clear that carbon content and isotope ratios remained unchanged The sequencing of the genome—that is, every gene—of Laccaria bicolor is truly a milestone in Volume 18, Number 1, 2009 9

mushroom knowledge. This is the fi rst ectomycor- The issue of Economic Botany in which these rhizal species to be completely sequenced, and, new species are described is totally devoted to eth- also the fi rst mushroom species of any kind for nomycology. The other articles treat the usage of which complete data are publicly available (the mushrooms by indigenous people in Burkina Faso, fi rst fungal species, however, was the bakers’ yeast Mexico, China, Japan, and many other countries, Saccharomyces cerevisiae). A fi nding that surprised the with attention to economic changes that are due authors is that Laccaria bicolor lacks the necessary to mushrooms. enzymes to break down lignin and cellulose. Those In the western mountains, morels are found enzymes are necessary for fungi which live sapro- in greatest abundance after forest fi res. There are trophically from wood and litter, but not for a spe- defi nitely several species but they still go nameless; cies that receives its carbohydrates and sugars from for one of them, that situation has changed, and a mycorrhizal partner. Laccaria has around 20,000 Kuo (known from his Mushroomexpert website) genes that encode for proteins—compare that to named it offi cially Morchella tomentosa, because of 20,000 to 25,000 for humans! There are also many the dark tomentose stem. Dark pigmented hairs on pieces of DNA that are not incorporated in genes, the stipe make up that tomentum. but the function of these is not known—they are often, but perhaps wrongly, called junk DNA. Real Birds’ Nest Fungi New techniques are slashing the cost and more Freymann, B. P., 2008. Physical properties of fun- and more species are in line to have their genomes gal rhizomorphs of marasmioid basidiomyce- sequenced. A wealth of new data will soon illumi- tes used as nesting material by birds. Ibis 150: nate the origin of different lifestyles and habitat 395–99. preferences. Prangle, S., and D. H. Nelson, 2006. Use of fungal Edibles Named rhizomorphs as nesting material by Glaucomys volans (Southern fl ying squirrels). Southeastern Arora, D., 2008. California porcini: three new taxa, Naturalist 5: 355–60. observations on their harvest and the tragedy of no commons. Economic Botany 62: 356–75. The Streak-backed Oriole, a bird occurring from southern Arizona to northwest Costa Rica builds Arora, D., and S. M. Dunham, 2008. A new, com- hanging nests over two feet long. The outer, hang- mercially valuable chanterelle species, Canthar- ing parts of these nests are made entirely from ellus californicus sp. nov., associated with Live long rhizomorphs of Marasmius species, whereas Oak in California, USA. Economic Botany 62: the lining of this hammock or basket consists only 376–91. of short grassy fi bers. The birds go to great lengths Kuo, M., 2008. Morchella tomentosa, a new species to gather the material for their nests, as there is a from western North America, and notes on lot of grass or other vegetation around, but fi nd- M. rufobrunnea. Mycotaxon 105: 441–46. ing Marasmius rhizomorphs requires much more time and effort. What are the advantages of such Three big conspicuous California mushrooms are a structure? First of all, the rhizomorphs take up now named: Boletus rex-veris, the reddish capped much less water than the grass fi bres and the fungal Sierra Nevada spring King Bolete; Boletus regineus, fi bres carry much higher weights before snapping the Queen bolete, erroneously called B. aereus, a than the grass fi bres—two huge advantages. European species; and Cantharellus californicus, the You don’t need to go to the tropics to see fun- big muddy chanterelle, growing under coastal gi in nests—lichens line humming birds’ nests, and live oaks. A big variant of the King Bolete with several birds (Bicknell’s thrush, Yellow-rumped a brown to cinnamon pore surface when mature warbler, and Blackpoll warbler to name a few) in is given the name Boletus edulis var. grandedulis. It is the boreal and subalpine zones of the northeast- amazing that some of our big conspicuous edible ern USA use Marasmius androsaceus rhizomorphs, mushroom species went unnamed or under cover another horsehair fungus, as nest material. of European names for so long! Flying squirrels use mosses, lichens, feathers, 10 McIlvainea

Spanish moss, and a huge variety of all kinds of 2008 issue carries a long article treating the British (man-made) material for their nests, depending on Xerocomus species (the velvety Boletus species) with what is available. But on top of all that, Southern many photos of the details used in the accompany- fl ying squirrels also use Masrasmius brevipes rhizo- ing synoptic key, including the cap and stipe sur- morphs as nesting material. faces, and the spore shape; descriptions of the spe- cies and photos of the less common taxa are also “North American Fungi” (formerly Pacifi c provided. To wander off on a side-track, Xerocomus Northwest Fungi): www.pnwfungi.org is not a monophyletic group, and has recently been split up into smaller genera, based on morphology; Glawe, D. A., and J. F. Ammirati, editors. 2008. A new are Xerocomellus, and Hemileccinum, whereas Festschrift in honor of Professor Jack D. Rog- Pseudoboletus (for the parasite on Scleroderma) had ers. North American Fungi 3(7): 1–267. A selec- been described earlier. tion of the articles: A series of articles on the genus Hyopoxylon Dugan, F. M. 2008. Fungi, folkways and fairy tales: appeared throughout the year. mushrooms & mildews in stories, remedies & The Farlow Herbarium librarian Lisa DeC- rituals, from Oberon to the Internet. North esare, together with George Hasapidis, wrote an American Fungi 3(7): 23-72. article on the Danish 18th-century mycologist Theodor Holmskjold, whose most infl uential work Norvell, L. L., J. F. Ammirati, and S. A. Redhead. is called Beata ruris otia fungis Danicis impensa (Happy 2008. Woody desert puffballs of the Pacifi c rural interludes in the country studying Danish Northwest. 1: Chlamydopus meyenianus. North Fungi). His illustrations are beautiful, and depict American Fungi 3(7): 127–38. the mushrooms in their natural habitats. He was Pfi ster, D. H. 2008. Early illustrations of Xylaria the fi rst to realize that Cordyceps militaris grows on species. North American Fungi 3(7): 161-166. dead insects buried in the soil; previously, insects were supposed to transform miraculously into Hemmes, D. E., and D. E. Desjardin. 2008. An- fungi. notated List of Boletes and Amanita in the Hawaiian Islands. North American Fungi 3(7): Mushrooms from Ecuador 167–76. Læssøe, T., and J. H. Petersen, 2008. Svampelivet Burdsall, H. H., Jr., and T. J. Volk. 2008. Armillaria på ækvator. Svampe 58: 1-52. [Equatorial solidipes, an older name for the fungus called fungi—mycological biodiversity in Ecuador] Armillaria ostoyae. North American Fungi 3(7): www.mycokey.com/Ecuador.html 261–67. The last item I want to draw attention to for arm- The completely online, Washington-based journal chair myco-travellers is the autumn 2008 issue of North American Fungi publishes articles on fungi the Danish journal Svampe. The main part of the (including lichens) from North America. This issue is taken up with a lusciously illustrated article journal is freely accessible and anyone can down- on the fungi of Ecuador. Wow! What beauties and load articles without cost. A special issue on the weirdos are there! Waxcaps without gills, stink- occasion of Jack D. Rogers’ 45 years of service at horns with the gooey part halfway up the stalk Washington State University brings together con- under a lacy top, a pink telescope-shaped Favolas- tributions on Xylaria and its allies, Jack’s pet fungi, chia species, small black chanterelles with spines and many other species and topics. A selection is on their cap, and so on and so forth. Pictures of listed above. the jungle with man-high buttress roots, and some scary inhabitants complement the article. Photos Field Mycology 2008 of these and many other Ecuadorian fungi (over 1200 in all) are available online at the Mycokey Field Mycology is a journal of the British Mycological website, to seduce the staunchest fungophobe. Society that focuses on fi eld studies, inventories, keys to species, and conservation issues, with a strong emphasis on the United Kingdom. The July