V
OMPHALINISSN 1925-1858
Vol. VI, No I Newsletter of Feb. 2, 2015
OMPHALINA 3 OMPHALINA, newsletter of Foray Newfoundland & Labrador, has no fi xed schedule of publication, and no promise to appear again. Its primary purpose is to serve as a conduit of information to registrants of the upcoming foray and secondarily as a communications tool with members.
Issues of OMPHALINA are archived in: is an amateur, volunteer-run, community, Library and Archives Canada’s Electronic Collection
OMPHALINA V Vol. VI, No 1 OMPHALIN ISSN 1925-1858 Feb. 2, 2015
CONTENT
Editorial ...... 2 Global distribution of fungi Leho Tedersoo ...... 3 Sarcoma globosum in NL Sveshnikov, Mann, Tedersoo, Voitk ..... 7 Cortinarius boulderensis Joe Ammirati ...... 11 Lactarius aurivolla Kuulo Kalamees ...... 12 Clavaria acuta Andrus Voitk ...... 13 Russula griseacea Andrus Voitk ...... 14 Morchella conica or Mel-19? Andrus Voitk ...... 15 Tuber of NL Andrus Voitk ...... 17 Bishop’s sketchbook ...... 19 Mail basket ...... 24 Partners ...... inside back cover Notice of Foray 2015 ...... back cover
This issue and all previous issues available for download from the Foray Newfoundland & Labrador websiteO
Happy 2015! How could we resist this backpedaling volte face? Better make the most of it, because this may be our andrus only opportunity to participate in a study like this. Living on an island, obviously we cannot go on
2 OMPHALINA GGloballobal diversitydiversity andand bbiogeographyiogeography ofof fungifungi
Leho Tedersoo
LOW HIGH
Fungal diversity is estimated at 1.5-5 million species, the largest group of eukaryotes after insects. Despite their enormous diversity and pivotal role in ecosystem function, little is known about the distribution of diversity and biogeography of functional fungal groups on a global scale. This investigation aims to 1) defi ne the relative roles of climate, soil, plant diversity, latitude and biome variables on global fungal diversity, and 2) examine biome and continental relationships based on major fungal functional and taxonomic groups.
Methods diversity of saprotrophs and animal parasites. 40 soil samples from each of 365 sites across the Soil world (see next page) were analyzed for fungal Calcium concentration or pH was the most important DNA. Single species sequences (44.6%) were ex- soil predictor of fungal diversity. ECM fungal diver- cluded from the analysis. sity correlated best with soil pH (greatest in slightly Results acidic to neutral soils), and the diversity of basidi- Overall omycetes, especially agarics, correlated best with increased soil calcium concentration. Saprotrophs, Over 80,000 “species” of fungi were identifi ed; the especially white rot decomposers, were more diverse study group was reduced to 44,563 after removing in moderately to strongly acidic soils. Diversity of singletons. All major fungal phyla and classes, ex- plant pathogens declined with the soil Carbon to cept Microsporidia were represented. The common- Nitrogen ratio. Phosporus concentration or its ratio est soil phyla were ascomycetes (48.7%) and basidi- to Carbon was also a predictor for some groups, e.g. omycetes (41.8%). Nearly 6% could not be assigned geoglossums and mucromycetes. to any known class of fungi. Distribution by lifestyle was saprotrophs, 43.8%, ectomycorrhizal (ECM) Plant life mutualists, 23.3%, plant pathogens, 4.0%, other Plant and fungal diversity were positively correlated, <1%), and uncertain 27.7%. Only 9.8% matched but the plant to fungus diversity ratio decreased fully identifi ed specimens with a scientifi c binomial. exponentially toward the poles (Figure 1). Major Climate functional and taxonomic groups showed dramatic differences in latitudinal diversity distribution. ECM Mean annual precipitation was a powerful predic- fungal diversity was directly proportional to the tor of fungal diversity, correlating strongly with the number and diversity of ECM plants. OMPHALINA 3 Photos of other teams and sites around the world. Clockwise from upper left: Guyana, photo courtesy Terry Henkel; Panama, photo courtesy Meike Piepenbring; Benin, West Africa, photo courtesy of Nourou Yorou; same as previous; New Zealand, photo courtesy Gwen Grelet; Panama, photo courtesy Meike Piepenbring; Malaysia, photo courtesy Su See Lee; middle insert principal investigator Leho Tedersoo in Benin, photo courtesy Nourou Yorou. For Newfoundland and Labrador team in action, see next article. 4 OMPHALINA Latitude Fungal diversity: • Increased/peaked toward the equator: - Ascomycota, in particular Archaeorhizo- mycetes, Dothideomycetes, Eurotiomycetes, Orbiliomycetes, and Sordariomycetes - Saprobes, soil fungi, plant pathogens, animal parasites • Increased/peaked toward the poles: - Lecanoromycetes, Leotiomycetes, Microbot- ryomycetes, Mortierellomycotina, and Mu- coromycotina Figure 1 • Peaked in the middle temperate zones: - Agaricomycetes, Pezizomycetes, and Tremel- lomycetes - ECM fungi Multivariate analysis • Greater in the Northern Hemisphere: Combined climate, soil and plant variables were the - Agaricomycetes strongest predictors for all fungal, ECM, and plant pathogen community compositions, but had little • Greater in the Southern Hemisphere: effect on saprobe distribution, which was mostly - Microbotryomycetes, Tremellomycetes, Wal- related to latitude. Of specifi c variables, potential lemiomycetes evapotranspiration (PET) and soil pH exerted a Biomes signifi cant effect on total fungal community compo- • Taxonomic distribution sition. Specifi cally, PET contributed signifi cantly to - The Ascomycota:Basidiomycota ratio was the distribution of saprotrophs, plant pathogens, and highest in grasslands (1.86), shrublands (1.86), yeasts. Distance from the equator and soil pH were and tropical dry forests (1.64), and lowest in the strongest predictors of ECM fungal community temperate deciduous forests (0.88). composition. Mean annual temperature and distance - The diversity of moulds peaked in the tundra from the equator correlated best with variations in and was lowest in tropical dry forests. animal parasite and mycoparasite communities. • Functional distribution Cross-biome and cross-continental relationships of - ECM fungi made up 34.1% of all taxa in functional and taxonomic groups northern temperate deciduous forests, but only The mean latitudinal range of fungi increased strong- 11.9% in grasslands and shrublands. ly toward the poles, linearly for ECM fungi and plant - The distribution of ECM fungi was similar pathogens, and unimodally with a temperate peak for southern temperate forests, tropical mon- for saprotrophic fungi. Diversity of saprotrophic and tane forests, grasslands, shrublands, lowlands ECM fungi decreased with increasing latitude. Major and Mediterranean biomes. taxonomic and functional groups of fungi differed - The distribution of plant pathogens and yeasts markedly in the frequency of occurrence across sites. was similar in tropical montane forests and Saprotrophs and plant pathogens had a broader distri- tropical lowland biomes. bution range than ECM and AM root symbionts. - Plant pathogens were more diverse in tropical Moulds and parasites were most widely distributed, forests. with the greatest range for animal parasites. • Biome distribution Figure 2 shows the clustering of geographic zones, - Boreal and temperate forests shared the most and biomes, as determined by shared fungal groups. fungal groups and species. Europe, West Asia, East Asia and North America clustered tightly, forming a sister group to Southern OMPHALINA 5 patterns suggest modifi cation by more recent fac- tors. Favorable climatic or soil conditions may have stimulated radiation (adaptive speciation) of these taxonomic groups in some biogeographic regions. Most fungal classes responded strongly to environ- mental predictors; even more striking effects can be expected at the order to genus levels, due to greater niche conservatism (retention of ancestral traits) in younger groups. These biogeographic analyses complement commu- nity-level analyses suggesting that both climate and biogeographic history shape fungal macroecological patterns. Co-migration with hosts over Pleistocene land bridges (Beringia, Wallacea, Panama, etc.) and long-distance dispersal by spores play important roles in shaping fungal distribution. Climate and associations with host plants probably contribute to Figure 2 differences in distribution range and biogeographic relationships among fungal functional groups. Hemisphere regions. Despite the large geographi- Our analyses revealed a strong infl uence of host cal distance and lack of terrestrial contact, tropical density on ECM fungal diversity, suggesting that the biogeographic regions clustered together. type of root symbiosis drives the functional composi- Biogeographic clustering of regions deviated mark- tion of soil fungi. ECM associations have evolved edly for certain functional groups of fungi. Based on relatively recently in multiple plant and fungal ECM fungi, the Southern hemisphere and tropical taxonomic groups. The peak of their taxonomic and regions clustered together. Based on yeasts, New phylogenetic richness in northern temperate biomes Zealand and Southern South America clustered with coincides with the geographical distribution and the Northern Hemisphere regions, and Australia clus- dominance of the Pinaceae, the oldest extant ECM tered with Central Africa and Madagascar. host. AM fungi exhibit even more promiscuous Discussion associations with tree and understory species and, therefore, their diversity is expected to be causally This is the fi rst study of the distribution of fungal unrelated to plant diversity. Finally, the paucity of taxonomic and functional groups on a global scale. matches to available database sequences indicates Analysis of over 1,200 soil samples distributed glob- that a vast majority of soil-inhabiting fungi have ally is without precedent, revealing an unprecedented remained undetected. fungal diversity and several striking biogeographic and macroecological patterns. Our results are strong- Conclusions ly at odds with current fungal diversity estimates, Except for ectomycorrhizal symbionts, richness of based on a constant plant to fungus ratio. The un- fungal functional groups is unrelated to plant diver- precedented size of our dataset notwithstanding, it sity; plants respond similarly to climate resulting in cannot be used for reliable richness estimates. These a similar diversity pattern. The plant:fungus ratio rely on the frequency of the rarest species, recovered declines exponentially towards the poles, question- but a single time, which make up almost one-half of ing the validity of current fungal diversity estimates. species lists, but which were rejected in our study to Climatic factors, followed by soil and latitude, con- exclude sequencing errors. stitute the best predictors of global fungal diversity. Fungal taxonomic groups originated >150 million ECM fungi differ from other groups by exhibiting years ago during the existence of the Pangaea super- a reversed latitudinal diversity gradient and strong continent, when they had suffi cient time to disperse dependence on the density of their host plants. over all the land. Current differences in distribution
6 OMPHALINA Dmitry Sveshnikov, Henry Mann, Leho Tedersoo, Andrus Voitk
ddoesoes ssarcosomaarcosoma gglobosumlobosum growgrow inin NL?NL?
This report makes a brief foray into soil samples from western Newfoundland in a global study,1 to learn what they may divulge of local mycota.
Methods an identifi ed species may have been shows their distribution in the two questionable, the original study referred woods, divided into saprobes and Two habitats were surveyed with to the genetically distinct entities as mycorrhizals. The total number 40 samples from each: a deciduous Operational Taxonomic Units (OTUs). of species from each woods woods (primarily Betula—Figure 1) 2. The relative number of recordings of was almost the same, but the leading to Barry’s Lookout behind ribosomal DNA is used as an indicator communities were different, with Humber Village, and two coniferous of frequency or commonness. Strictly, only 7% of the total species found woods, one near Pasadena this number represents the relative in both sites. (primarily Picea) and the other near amount of an organism’s rDNA in the the campus of Sir Wilfred Grenfell soil sample when compared to that of Table 3 lists the sequences College in Corner Brook (primarily others, which may not be the same as recovered 10 or more times that Abies). Soil was collected and habitat the relative frequency of the organism matched named species. observations made according in the soil. to a standardized protocol and Results Phylum Nr spp. DNA and related analyses done as NL samples yielded 665 genetically reported in the global study.1 Basidiomycota 357 distinct fungal “species,” from which Terminology 129 sequences matched species Ascomycota 193 For ease of interpretation, this report named in publicly available genetic Zygomycota 85 makes two assumptions, which may not databanks. The full list is available Chytridiomycota 12 be true, using terms or concepts more on request to the editor. loosely than strict scientifi c observation Roziellida 10 would support. 1.“Species” is used Table 1 lists the phyla from Table 1. Fungal phyla from NL soil to refer to a genetically distinct entity. kingdom Fungi in order of samples in order of diversity. The In the study, because all distinct diversity. Basidiomycetes and ascomycete:basidiomycete ratio is DNA units were not identifi ed to ascomycetes accounted for 84% low. Note that Roziellida is here con- species, and those that did match of the species found. Table 2 sidered in kingdom Fungi. OMPHALINA 7 Discussion Ascomycetes Basidiomycetes The original study was ECM Sap- Undeter- Total ECM Sap- Undeter- Total designed to examine robe mined asco robe mined basidio global diversity and Deciduous 16 63 26 106 114 51 13 178 distribution of fungi. woods Applying these data to Conifer- 16 61 30 107 113 78 15 206 the study of regional ous woods fungi at the species level Table 2. Ascomycetes and basidiomycetes divided into ectomycorrhizals (ECM), saprobes may not be reliable. and undetermined, for both deciduous and coniferous woods. One lichenized ascomycete For example, blasts was classifi ed as a mycorrhizal, and parasites were classifi ed as saprobes (a photobiont is (techniques of matching a photobiont, regardless of size, and organic matter is organic matter, regardless whether DNA to that available living or dead). on record) depend on various algorithms, each with its own strengths and Species T D C weaknesses in any given situation. An algorithm suited Cortinarius obtusus 402 0 402 for the purpose of the original study may select the most “convenient” sequence segment for matching, Melanophyllum haematospermum 117 117 0 not always the closest. To be in the same clade as Humidicutis cf. marginata 70 70 0 Species X is different from being Species X. Even if the Trichosporon porosum 43 18 25 DNA really does match fully, whether it is the species Cortinarius duracinus var. 25 0 25 identifi ed depends on how accurately the deposited raphanicus material was identifi ed. This is a problem that we have Tomentella subclavigera 24 24 0 encountered in many of our other studies, and much more a problem when dealing with DNA from soil Cryptococcus podzolicus 24 24 0 samples, where there are no fruiting bodies to fall Cortinarius boulderensis 22 0 22 back on. Less than 20% of unique DNA samples from Russula griseascens 22 0 22 NL matched material in public genetic depositories Cryptococcus terricola 21 20 1 identifi ed to species with a binomial name. Therefore, even if the matching is impeccably reliable, this species Byssoporia terrestris 20 0 20 list sampling but 20% of recovered mycota may not be Cortinarius acutus 19 0 19 representative of the mycota of our region. Clavaria acuta 19 19 0 This is not a provincial study: the samples were taken Cortinarius mucifl uus 18 0 18 from two forests (coniferous and deciduous) in a Lecythophora cf. mutabilis 12 12 0 small area of Western Newfoundland, in one of the Hygrocybe laeta 12 0 12 nine ecoregions of the Island. Even with 80 samples Piloderma sphaerosporum 11 11 0 from this small region, not all regional fungal taxa are recovered. People familiar with the territory Lactarius tabidus 11 11 0 will recognize several species that have not been Lactarius aff. lignyotus 11 0 11 recovered, some of them relatively common (e.g. the Cenococcum geophilum 10 1 9 total absence of Cortinarius armillatus, the commonest Table 3. Species matched to deposited sample with observed macrofungus in the sampled birchwoods). identifi ed binomial, recorded 10 or more times (T= total, Given these caveats, the broader fi ndings described D=deciduous woods, C=coniferous woods). Specifi c ex- under Results, may be expected to be more reliable amples of interest will be discussed in the text. than specifi c ones, as the former are more in line with the aims of the original study. the primary trees involved. For example, some balsam The marked difference in the funga of broadleaved fi r and spruce can always be found in our birchwoods, and coniferous forests is interesting. We seem more yet the majority of organisms from coniferous woods ready to accept specifi city in mycorrhizal relationships, were not recovered from soil in the birchwoods. This but clearly the preferences of decomposers are every bespeaks of systems with complex interdependence bit as specifi c, even when they are not decomposing of their parts. The obverse is that extinction of a
8 OMPHALINA Photo: Maria Voitk Sampling of Newfoundland birchwoods. Primarily Betula Here we see 75% of the authors (L to R: HM, AV, DS), payrifera, with 20% B. alleghanensis, and 10% Acer, 67% of them doing 99% of the sampling associated work. Abies, Picea, plus a deciduous arbuscular understory. species from its econiche may have Clearly, there can be quite a wide Northwest. However, the high far wider repercussions than the dichotomy between the proportion number of occurrences in our soil loss of a single species. of fruiting bodies and environmental is diffi cult to ignore. How reliable DNA samples. Of the many other are the matches? Well, on blast they While the original study was not interesting fi nds in the data, three make a 99% match with DNA from designed to identify or study deserve special mention. All were a specimen from British Columbia,2 individual species, several such reasonably commonly encountered interesting observations were seen and the DNA of that specimen macrofungi, in soil samples and 3 in the data for our region. For matches that for the holotype. should, therefore, be familiar, yet all example, the three commonest Thus, despite the potential bear species names we have not birchwoods macrofungal species inaccuracies of the methodology encountered hitherto. were Humidicutis cf. marginata, for identifying individual species, Clavaria acuta and Lactarius tabidus. Russula griseascens (99% match the likelihood is quite high that our The fi rst and last have been with 2 collections determined by soil Isolate is indeed Cortinarius documented at our forays, but there Jukka Vauras) was encountered boulderensis. is no record of Clavaria acuta. AV’s 22 times. This is a member of the A somewhat similar situation occurs personal collections had but one Russula emetica complex, and since with Lactarius auriolla, found fi ve example of this “common” species. R. emetica was not recovered, it is times. It was described in 1984 conceivable that in our coniferous In the coniferous forest, Cortinarius by Kytövouri, who described a woods R. griseascens may be the obtusus was by far the commonest series of similar, hairy-capped, commonest species of the complex. 4 recovered species: identifi ed slimy, yellow Lactarius species. The 402 times in a database where Cortinarius boulderensis was DNA of our soil fungus matches the second-commonest species identifi ed 22 times (third- 100% with DNA found in the soil (Cortinarius duracinus var. raphanicus) commonest macrofungus) in the in Alaska, which, in turn, is a 99% was identifi ed only 25 times, and soil samples from the coniferous match for DNA extracted from most species only once. While C. forest, yet the species has been a specimen near the type locality, obtusus is seen occasionally in these unknown here. Hitherto it has been its identifi cation confi rmed by its woods, the fruiting body is not thought to be a North American author.5 This makes the likelihood encountered nearly as frequently. species limited to the Pacifi c very high that this European
OMPHALINA 9 information such studies can provide. Because of potential problems with reliability due to methodology, matches need to be examined and referred to type or other reliable material, as illustrated here, before conclusions can be formed. The power of environmental sampling and DNA analysis is readily evident: it took FNL six forays of 45-65 participants before we had accumulated 665 species. This approach, combined to the power of statistical analysis can unleash massive amounts of information; the global study from which this report is culled has barely scratched the surface of the potential of this technology. Oh, and do we have Sarcosoma globosum in NL? According to this study, no. Is that certain? No. After all, soil samples were only studied from a very limited area of the province, and not from areas most likely to harbour the species. However, the entire global database contained very few samples identifi ed as Sarcosoma, and none were identifi ed to species, so possibly the technique was not ideal to answer this question in any case. These considerations aside, it is our best guess that Sarcosoma globosum does not exist in this province. But soil for sampling does.
References 1. Tedersoo L, Bahram M, Põlme S, Kõljalg U, Yorou NS, Photo: Henry Mann Wijesundera R, Ruiz LV, Quang Thu P, Dang T, Suija A, Smith ME, Sharp C, Saluveer E, Saitta A, Rosas M, Ratkowsky D, Sampling of Newfoundland softwoods. Primarily a pure Pritsch K, Põldmaa K, Piepenbring M, Phosri C, Peterson M, Picea glauca stand, although a small Abies balsamea Parts K, Pärtel K, Vasco-Palacios AM, Otsing E, Nouhra E, can be seen in the foreground, on the left. One co-author Njouonkou AL, Nilsson RH, Mayor J, May TW, Majuakim L, examines the soil for gold nuggets (to be removed before Lee SS, Larsson KH, Kohout P, Hosaka K, Hiiesalu I, Henkel sending for DNA), and the other fi ne-tunes the digger’s TW, Harend H, Guo L-D, Greslebin A, Grelet G, Geml J, carburetor. Gates G, Dunstan W, Dunk C, Drenkhan R, Dearnaley J, de Kesel A, Chen X, Buegger F, Brearley FQ, Bonito G, Anslan S, Abell S, Abarenkov K: Global diversity and biogeography of species also exists in Newfoundland and Labrador soil fungi. Science, Vol. 346, DOI: 10.1126/science.1256688. (as well as Alaska), and has been overlooked to date. 2014. These results suggest that we need to work up the 2. Harrower E, Ammirati JF, Cappucino AA, Ceska O, large genera, Cortinarius, Lactarius, and Russula, in our Kranabetter JM, Kroeger P, Lim SR, Taylor T, Berbee ML: Cortinarius species diversity in British Columbia and province. molecular phylogenetic comparison with European specimen Among several other interesting observations was sequences. Botany 89:799-810. 2011. the fi nding of two real truffl es, unnamed species of 3. Niskanen T, Liimatainen K, Kytövouri I: Taxonoimy ecology and distribution of Cortinarius rubroviolipes Tuber, and our cosmopolitan morel, Mel-19, in the and C. hinnuleoarmillatus (Basidiomycota, Agaricales) in process of being formally reported. Because the Fennoscandia. Karstenia 46:1-12. 2006. species mentioned here may represent undetected 4. Kytövouri I: Lactarius subsection Scrobiculati in NW Europe. but relatively common species in our woods, or are Karstenia 24:41-72. 1984. otherwise of interest, they are described in more 5. Verbeken A, Stubbe D, van de Putte K, Eberhardt detail elsewhere in this issue. U, Nuytinck J: Tales of the unexpected: angiocarpous representatives of the Russulaceae in tropical southeast Asia. This mere glimpse should give a hint at the sort of Persoonia, 32:13-24. 2014.
10 OMPHALINA Photo: Kare Liimatainen Cortinarius boulderensis Smith Joe Ammirati Cortinarius boulderensis Smith was described from colored fi brils or a sheath-like covering below. The an old growth conifer forest along Boulder Creek, medium sized spores are not particularly distinctive, Olympic Hot Springs, in Olympic National Park. but are helpful in separating it from unrelated species Alex Smith made the type collection of this very with similarly colored veils. distinctive Telamonia in 1941, and since then it has Alex Smith considered Cortinarius boulderensis been documented in the Pacifi c Northwest from to be related to C. armillatus and other telamonias several locations scattered along the west side of the with more or less reddish colored universal veils. Cascades and Olympic Mountains from Washington Phylogenetic studies by Kare Liimatainen and south into western Oregon and northwestern Tuula Niskanen and have shown that telamonias California, and north into western British Columbia. with reddish universal veils represent different Cortinarius boulderensis prefers moist, well evolutionary lineages and that C. boulderensis established conifer forests of Abies, Tsuga and/or and C. rubrovioleipes are sister taxa that are Pseudotsuga, sometimes also including Picea. morphologically the same and genetically similar. The coloration of fresh specimens in prime condition Both of these taxa are related to a western subalpine is remarkable. This medium sized Telamonia features species, C. pseudobovinus, which feature a greyish a slightly violaceous brown to rich brown, moist, brown to fuscous universal veil. silky pileus with slight striations and a pale to Tuula Niskanen, Kare Liimatainen and Ilkka whitish edge. The lamellae are grayish lilac at fi rst, Kytövuori published on Cortinarius boulderensis later becoming dull cinnamon, and the silky stipe in Karstenia in 2006. This prompted me to contact is violaceous above becoming brownish below, and them about this species which resulted in a very ending in a slightly bulbous base. The most striking enjoyable and rich collaboration on Cortinarius feature is the universal veil that leaves a distinct taxonmy and evolution, for which I am thankful. testaceous band on the upper stipe and similarly OMPHALINA 11 Photo: Vello Liiv Lactarius auriolla Kytövouri Kuulo Kalamees
CAP: yolk yellow, deeply funnel shaped, edge Photo: Vello Liiv inrolled for a long time, from which hang distinct whitish threads, glutenous to slimy, up to 7 cm diameter. GILLS: cream to light yellow, crowded, broadly attached to decurrent. STEM: Concolorous with the cap or lighter, no or few yellowish scrobiculations, dry, slightly longer than cap diameter, cylindric, up to 1.7 cm wide. FLESH: white, cut surface quickly turns straw coloured, fruity smell, mild taste. MILK: white, turns lemon yellow immediately on exposure to air, mild to acidic taste. SPOREPRINT: pale cream. SPORES: 7-8.5 x 6-6.5 μm, reticulate to ribbed, broadly ellipsoid. HABITAT: calciphilic, in wet spruce forest. Fruits in August to September. Considered rare to uncommon in regions where it is known to fruit. DISTRIBUTION: hitherto known from limestone in Scandinavia and Estonia; photos and description based on Estonian collections.
SIMILAR SPECIES: Lactarius scrobiculatus, L. leonis, L. tuomikoskii. Distinguished by its prominent threads (wider than hairs) hanging from the cap edge.
12 OMPHALINA its DNA recorded in the soil study. This means that it may or may not exist in NL. There were seven big (about 20 cm tall) and four small clubs in a lawn near the soil sampling site, but, of course, I had to photograph the only uncommon shape, the forked one, with the main branch broken. The common shape resembles single, pointed spindles, the pointed tip giving rise to the species Clavaria acuta Sowerby epithet, acuta. Probably the species is commoner Andrus Voitk than thought, but not often recognized because of its small size and scattered growth habit. Finding at Clavaria acuta is a small, white, club-like Clavaria least one collection in the region of the soil samples with a pointed tip, that usually fruits as a few increases the likelihood that this was a reliable scattered individuals. Its characterizing features are fi nding. the relatively small size, pure white fertile surface, a Roger Smith’s photo of C. fumosa comes from grayish-translucent stem, and a non-cespitose mode the fi elds around Killdevil Camp in Gros Morne of growth. The somewhat similar C. vermicularis National Park, site of our 2014 foray. It has also and C. fumosa differ by being larger, having a hollow been collected from Sir Richard Squires Memorial fruiting body, being without a transparent stem and Provincial Park, site of the 2014 mycoblitz. The growing in cespitose clusters. Microscopically these cespitose growth pattern and off-white, smoky colour latter two are alike, but in colour C. vermicularis are evident. Comparing the moss on the two photos is white like C. acuta and C. fumosa is off-white, also gives an idea of the size difference between the varying from gray, straw to pinkish tones. Clavaria two species, C. fumosa being at least three times vermicularis has not been recorded by FNL, nor was taller. In addition, there are microscopic differences.
OMPHALINA 13 Photos: Roger Smith
Fogo, 2013Labrador, 2005 Terra Nova, 2011 Terra Nova, 2012 Russula griseascens (Bon & Gaugué) Marti Andrus Voitk
Russula emetica is a common, relatively small, from 2013, as well as three photos of R. “emetica” beautiful, red-capped Russula with a very acrid taste, with similar graying stems. What does this mean? famous for inducing vomiting in the mycophagist Scientifi cally, nothing. All material needs to be foolish enough to eat it. Although found in moist sequenced, analyzed and matched (or not) with a areas of coniferous forests, it is suspected to be reliable reference collection, before a defi nitive mycorrhizal with birch, not conifers. It forms a conclusion can be made. However, what is the species complex with several similar species. Among likely implication? Given the prevalence of putative these we have identifi ed R. aquosa and R. nana. R. griseascens in our soil samples, the absence of In 1975 Bon and Gaugué described Russula emetica R. emetica in the same samples, the genetically var. griseascens, characterized by graying of the confi rmed R. griseascens of 2013, and the presence stem with age, trauma or handling. It was elevated of similar gray stems on photos of most collections to species status by Marti in 1984. Russulales news labelled “R. emetica”, there is a very real likelihood states that in the Alps this is the commonest species that R. griseascens grows in Newfoundland and of the complex. At our foray, R. griseascens was Labrador. Our diversity here is low and usually we identifi ed from Fogo Island in 2013 because nuclear do not have several similar species occupying the studies showed the closest match to a named species same econiche. Therefore, R. griseascens may be was at 97.5% to two Finnish collections identifi ed as our commonest member of the R. emetica complex, R. griseascens by Jukka Vauras. In the soil samples possibly hitherto misidentifi ed as R. emetica, which the third commonest identifi ed species in our may not even grow here. Of course, we may also coniferous woods matched those same two Vauras have both, but our low diversity and the absence collections at 99%. At the same time, there is no of DNA from R. emetica in the soil argues for one record of DNA marching R. emetica. Likely the soil species only. fungus and the Fogo fungus are the same species, Until the question is confi rmed, the above must be and provided Jukka Vauras identifi ed the species of viewed as speculation only. As is the tenet that R. Bon and Gaugué, both are R. griseascens. emetica grows here. To me the latter speculative I reviewed all photographs of R. emetica from past position seems considerably less likely. Would we be forays. Five of nine photos showed mushrooms with more likely to be accurate, were we to list anything some graying on the stem. The composite photo, identifi ed as R. emetica as R. griseascens until more above, shows the genetically matched R. griseascens clarity is brought to the question?
14 OMPHALINA Morchella sp. Mel-19 Andrus Voitk Photo: Vello Liiv Photo: Vello
We have documented three identifi ed by the depositor. The Labrador. Finding it in Estonia is also morel species in Newfoundland DNA from our soil matched that not surprising, because the species and Labrador: M. importuna (the from a collection identifi ed as M. is known from nearby Scandinavia. mulch morel), and two undescribed conica that came from limestone Finding it on limestone barrens species, temporarily code named barrens of an Estonian island. The fi ts with our experience, because Mel-36 and Mel-19.1 In addition, photo in the title banner is of that the majority of our collections of the DNA of Morchella conica was collection, taken by Vello Liiv, whom Mel-19 also come from this habitat. reported from the soil samples we foray veterans may remember as a submitted for study; this piqued our member of our fi rst faculty in 2003. This experience illustrates the curiosity. M. conica is a European diffi culty of identifying an organism taxon, described by Persoon in If M. conica does not exist here, by matching its DNA with that 1818, reputedly not native to North there is reason to suspect that on hand in gene depositories: America.2 Current opinion holds the reference collection may be the accuracy depends on the that it is not a valid taxon at the misidentifi ed. To test this, Kerry accuracy of the identifi cation of the 3 O’Donnell ran the DNA on fi le deposited collections. One review species level. Since M. conica is not for the Estonian collection against revealed that only 33% of morel a valid species and not found on known morel DNA sequences. On collections that were deposited this continent, and since we have preliminary analysis, the Estonian with a species name in GenBank not found other morel species after collection matched Mel-19. Mel-19 were identifi ed correctly.5 In a decade and a half of collecting, it is is also known from Washington other words, if you were a betting most likely that the DNA found in State, Newfoundland and Labrador, woman, you could win 2:1, if you our soil is that of one of the three Scandinavia, The Netherlands, just bet that every named morel in species we have already identifi ed. and various parts of China4.The GenBank is misidentifi ed. DNA from soil is identifi ed as match was so good, and the other Why such seeming unreliability? a certain “species” by matching evidence fi t so well, that we predict For one thing, Mel-19, although it up with DNA on deposit in that multilocus analysis will likely known from around the world, has depositories, where investigators confi rm Mel-19 in Estonia. Similar not been described or named yet. have fi led DNA sequences of DNA was also recorded from Thus, most workers are not able to collections they have identifi ed. another collection in the same area consider it in their identifi cation. As If soil DNA matches that of a (photo, also by Vello, next page). for the described species, identifying collection, it is assumed to come This fi ts well, because we know often relies on a morphological from the same species as was Mel-19 grows in Newfoundland and species concept. This does not work OMPHALINA 15 well with the genetically discovered has culminated in a detailed study, References morel species: with a few distinctive where species have been described 1. O’Donnell K: True morels in Newfoundland exceptions, most members of again and new type collections with and Labrador. OMPHALINA 5(2): 3-6. 2014. the Elata clade resemble each good genetic material have been 2. Kuo M, Dewsbury DR, O’Donnell K, Carter other morphologically both on “appointed” to support them.3 MC, Rehner SA, Moore JD, Moncalvo J-M, the macro- and microscopic This work should go a long way Canfi eld SA, Stephenson SL, Methven AS, Volk TJ: Taxonomic revision of true morels level, and cannot be separated to stabilize the situation for future (Morchella) in Canada and the United morphologically. Multilocus genetic investigators. Thew study was States. Mycologia 104:1159–1177. 2012. analysis is required to identify them unable to defi ne the M. elata of 3. Richard F, Sauvé M, Bellanger J-M, Clowez reliably, not available to everybody Fries, and work to elucidate this is P, Hansen K, O’Donnell K, Urban A, collecting and depositing material. still going on. It is not impossible Courtecuisse R, Moreau P-A: True morels that our Mel-19 could be defi ned as (Morchella, Pezizales) of Europe and The species concept we prefer to- that species. North America: Evolutionary relationships day rests on evolutionary genetic inferred from multilocus data and a unifi ed lineage. We have no idea of the Meanwhile, to our knowledge, all taxonomy. Mycologia, doi:10.3852/14-166. genetic sequences of the specimen morel species are desirable edibles. 2014 (defi nitive article in press. 2015). Persoon held in his hand in 1818, For the morelophagist, identifi cation 4. O’Donnell K, Rooney AP, Mills GL, Kuo when he described Morchella conica of morels to exact species may be M, Weber NS, Rehner SA: Phylogeny and historical biogeography of true morels as a new morphological taxon (or irrelevant. (Morchella) reveals an early Cretaceous the one Fries held in his hand when Unless she becomes curious, when origin and high continental endemism he described Morchella elata as a edibility may become irrelevant. and provincialism in the Holarctic. Fungal new morphological species four genetics and biology 48:252-265. 2011. years later). Those type specimens 5. Du X-H, Hansen K, Taskin H, Büyükalaca are too old to yield useful DNA Acknowledgments S, Dewsbury D, Moncalvo J-M, Douhan for the sophisticated analyses GW, Robert VARG, Crous PW, Rehner I thank Kerry O’donnell for the DNA SA, Rooney AP, Sink S, O’Donnell K: How required to determine this. Genetic matching described above, as well well do ITS rDNA sequences differentiate research over the past two decades, as review of the MS, and Vello Liiv specis of true morels (Morchella)? correlating morphologic and other for permission to use his beautiful Mycologia 104:1351-1368. 2012. characters with gene sequences, photographs.
Photo: Vello Liiv Photo: Vello Liiv Photo: Vello
16 OMPHALINA SSeekingeeking tthehe eelusivelusive NNewfoundlandewfoundland
ttruruff lele Andrus Voitk
Photo: Maria Voitk
Among interesting fungal DNA Many of the others are only known to our knowledge species of this samples recovered from our soil from genetic material recovered in genus have not been recorded for were two undescribed species environmental samplings, and the the province. Therefore, fi nding of Tuber, Tuber sp. 16 from our fruiting organism has not been seen. the DNA of two Tuber species in coniferous woods and Tuber sp. Not only are they underground, our soil was defi nitely of interest. 19 from our birchwoods. Tuber many are also very small, so they Checking with Greg Bonito, is a genus of hypogeous, or go unnoticed by most hypergeous who reviewed the genus and its underground ectomycorrhizal observers of macrofungi. Therefore, phylogeny world-wide,1 we learned ascomycetes, including the famous there are many genetic species that that fruiting bodies of these two and expensive gourmet truffl es, have not been seen to describe numbered species have not been as well as some 100 species them, and no doubt many additional seen, preventing formal descriptions. world-wide, about half in North species remain to be uncovered. The DNA of Tuber sp. 16 has been America.1 Here the famed edible recovered from environmental Our truffl e species known hitherto, ones are few, mostly limited to the samples in Québec, New York both real and false, have been Pacifi c coast, although T. lyonii s.l. State, Michigan, and California; reviewed before.2,3 Species of and T. canaliculatum are of some Tuber were not among them, and the Michigan sample came from commercial interest in the east. the roots of Epipactis helleborine.4
OMPHALINA 17 The DNA helleborine in New York State.5 of Tuber sp. With this in mind, before the 19 has been 2014 Foray we explored the recovered earth around one patch of E. from Epipactis helleborine. The digging was dunensis in done by lichenologists Michele Austria, and Piercy-Normore, Teuvo Ahti identifi ed in and Chris Deduke. No luck. other environ- Lacking a proper search image, Above: Photo of the only known collection of Tuber sp. 19, with mental fi nding 4 mm brown pebbles permission from MycoPortal and Duke University, Rytas Vilgalys. Below: Close-up and infl orescence of Epipactis helleborine in our samples from in duff may be an unfair task birchwoods, where this introduced species has made itself quite Germany, even for non-lichenologists. at home. Not as invasive here as in parts of the mainland, it may Lithuania, serve as an indicator of some of our underground Tuber species. This article serves as a tribute Sweden, to Teuvo (Ted) Ahti on the England, New occasion of his 80th birthday England, California and New a few months before the title 4 An unidentifi ed Zealand. banner photo was taken. The collection of tiny truffl es from much younger Chris Deduke Nebraska was identifi ed by seems to have completely sequencing to be this species collapsed in the background, (illustration). toes up, but, like the Eveready The association of Tuber with bunny, Teuvo just smiles and Epipactis has been noted keeps on truffl ing. regularly. Epipactis helleborine Congratulations, Teuvo! (illustration) is a European orchid, introduced to North America, where it seems Acknowledgments to have established itself I thank Greg Bonito for reviewing quite well. A decade ago, in the MS. Newfoundland it was only References known from a site in St. John’s, 1. Bonito MG: Systematics and but now it has spread; there Ecology of Truffl es (Tuber), PhD are several pockets in the thesis. Duke University. 2009. birchwoods near our house, 2. Voitk A: The $1,000 question. Omphalina 2(8):4-6. 2011. where it thrives happily. 3. Hayward J, Horton T, Voitk A: Orchids have such small seeds, Preliminary report from the consisting of genetic material bolete underground: the false only, that to grow they need truffl es of Newfoundland and Labrador. Omphalina, 2(8):7-8. to get outside nourishment. 2011. This is supplied by a 4. Bonito GM, Gryganskyi AP, mycorrhizal fungus, and many Trappe JM, Vilgalys R: A global are quite specifi c. One of our meta-analysis of Tuber rDNA sequences: species diversity, host soil samples with Tuber DNA associations and long distance came from the birchwoods dispersal. Molecular Ecology where Epipactis has settled. 19:4994-5008. 2010. 5. Rumburg JR., Moskalenko Jess Rumburg, one of Tom M, Muska D, Horton, TR: Horton’s students, showed The invasive orchid Epipactis that Tuber sp. are the helleborine (L.) Crantz forms mycorrhizae with truffl es (Tuber commonest mycorrhizal spp.) and other Ascomycetes fungi found on Epipactis In New York state. Inoculum 59(4):54. 2008.
18 OOMPHALINAMPHALINA The Bishop’s Sketchbook
OOMPHALINAMPHALINA 19 the mail bag or why the passenger pigeons assigned to serve the lavish Corporate and Editorial offices of OMPHALINA get hernias
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Today (Nov 28, 2014) is the 10-th anniversary of the day when my wife Oluna Ceska started her macrofungal survey on Observatory Hill (Victoria, Vancouver Island, Bri sh Columbia). A er 10 years and close to 400 visits to “The Hill”, Oluna recorded 1,298 species from an area of about 75 ha (approx. 185 acres). A part of those collec ons (several hundred earlier collec ons of CorƟ narius, Inocybe, Russula and some others) have been DNA sequenced and deposited in GenBank, and some were studied and published. Observatory Hill collec ons of several species have been directly cited in the original descrip ons of some of the species listed here: h p://mushroomobserver. org/species_list/show_species_list/708. The majority of the Observatory Hill collec ons were donated to the UBC herbarium where the collec ons have been accessioned or are awai ng accessioning. Many of those collec ons have been already been included in the UBC Fungi Herbarium database h p://www.beatymuseum. ubc.ca/herbarium/index.html, and mirrored in the Consor um of the Pacifi c Northwest Herbaria h p://www. pnwherbaria.org/ “Virtual herbarium”, created in Mushroom Observer. It contains photos, microphotos, and drawings of about one third of all our Observatory Hill specimens: h p://mushroomobserver.org/species_list/ show_species_list/227. Par al results of the survey have been summarized in annual reports deposited on the GOERT web site: h p://www.goert.ca/ac vi es/2013/05/macrofungi-observatory-hill/. This project was par ally supported by the Herzberg Ins tute of Astrophysics, GOERT (Garry Oak Ecosystem Restora on Team), and private donors. Cataloguing of the collec ons has been supported by the Canadian Wildlife Service. Our special thanks go to Dr. Paul Feldman, HIA astronomer, who ini ated the project and provided all the help we needed. Adolf & Oluna Ceska, Victoria, Bri sh Columbia, Canada h p://mushroomobserver.org/observer/show_user/2873
Ed comment: Congratula ons on a phenomenal team eff ort! A suitable contrast to seeking fungi in soil samples…
20 OMPHALINA OOURUR PPARTNERARTNER OORGANIZATIONSRGANIZATIONS
People of Newfoundland and Labrador, through Department of Environment and Conservation Parks and Natural Areas Division Wildlife Division Department of Natural Resources Center for Forest Science and Innovation People of Canada, through Parks Canada Terra Nova National Park Gros Morne National Park The Gros Morne Co-operating Association Memorial University of Newfoundland St. John’s Campus Grenfell Campus Tuckamore Lodge Quidi Vidi Brewing Company Rodrigues Winery
OMPHALINA 2015 2015 2015 2015 2015 e 2015 2015 2015d ca 2015 2015de 2015 2015d decade 2015 2015on 2015 ec 2015 ssecond 2015 2015 he TThe
GROS MORNE NATIONAL PARK A UNESCO WORLD HERITAGE SITE Headquarters: Killdevil Camp, Lomond, NL September 25-27, 2015 GGUESTUEST FFACULTY*ACULTY*
Nils Hallenberg Andy Methven Andy Miller Michele Piercey-Normore Roger Smith Greg Thorn
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