Omphalina Vol 5 # 5

OMPHALIN

V

ISSN 1925-1858

Newsletter of

Vol. V, No 5 May 26, 2014

OMPHALINA

O MPHALINA, newsletter of Foray Newfoundland & Labrador, has no fixed 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.

is an amateur, volunteer-run, community, not-for-profit organization with a mission to organize enjoyable and informative amateur mushroom forays in Newfoundland and Labrador and disseminate the knowledge gained. Webpage:

www.nlmushrooms.ca

ADDRESS Foray Newfoundland & Labrador 21 Pond Rd. Rocky Harbour NL A0K 4N0 CANADA E-mail: info AT nlmushrooms DOT ca BOARD OF DIRECTORS

CONSULTANTS

Michael Burzynski PRESIDENT

Geoff Thurlow TREASURER

Faye Murrin SECRETARY

MYCOLOGICAL Dave Malloch NB MUSEUM

Andrus Voitk Past PRESIDENT

Jim Cornish Jamie Graham Tina Leonard Anne Marceau Michele Piercey-Normore Maria Voitk Marian Wissink

OMPHALINA

AUDITOR Gordon Janes BONNELL COLE JANES

LEGAL COUNSEL Andrew May BROTHERS & BURDEN

Issues of OMPHALINA are archived in: Library and Archives Canada’s Electronic Collection <http://epe. lac-bac.gc.ca/100/201/300/omphalina/index.html>, and Centre for Newfoundland Studies, Queen Elizabeth II Library (printed copy also archived) <http://collections.mun.ca/cdm4/ description.php?phpReturn=typeListing.php&id=162>.

The content is neither discussed nor approved by the Board of Directors. Therefore, opinions expressed do not represent the views of the Board, the Corporation, the partners, the sponsors, or the members. Opinions are solely those of the authors and uncredited opinions solely those of the Editor. Please address comments, complaints, contributions to the self-appointed Editor, Andrus Voitk: seened AT gmail DOT com, … who eagerly invites contributions to OMPHALINA, dealing with any aspect even remotely related to mushrooms. Authors are guaranteed instant fame—fortune to follow. Authors retain copyright to all published material, and submission indicates permission to publish, subject to the usual editorial decisions. Issues are freely available to the public on the FNL website. Because content is protected by authors’ copyright, editors of other publications wishing to use any material, should ask first. No picture, no paper. Material should be original and should deal with the mycota of Newfoundland and Labrador. Cumulative index and detailed Information for Authors available on our website.

COVER Gymnopus eneficola, a new species of Gymnopus, on the Pasadena Ski and Nature Club ski trails, Sept. 26, 2009. Read the lead article—our first formal description of a nomenclatural novelty—to learn about this new species. “Eneficola” signifies “denizen of en-ef = NF = Newfoundland”. Then find out about other species in the genus in this province.

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OMPHALIN

ISSN 1925-1858

Vol. V, No 5 May 26, 2014

CONTENT Editor’s comments ................................. 2 Foray matters ......................................... 3 Basic Barrens of Gros Morne Michael Burzynski ....................................

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Gymnopus eneficola spec. nov. Petersen, Hughes, Voitk ......................

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Gymnopus dryophilus clades Hughes, Aldrovandi, Petersen, Voitk ....

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The Bishop’s sketchbook ........................ 16 Dryophiles and dryophobes Voitk, Hughes, Aldrovandi ......................

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Gymnopus species in NL Petersen, Voitk .................................... 21

Mail basket ............................................ 28 Partners ......................... inside back cover Foray notice ........................... back cover

This issue and all previous issues available for download from the Foray Newfoundland & Labrador website <nlmushrooms.ca>.

OMPHALINA

Welcome W Wel We elc lcom lcom omee to to oour ur Gymnopus Gymn Gy mnop oppus issue! iiss ssue ss ue!! ue A monograph to the genus in our province, based on a review of 57 collections by Karen Hughes and Ron Petersen—not an entirely negligible task. In the course of this review we learned two interesting things. First, we found a new species, hitherto unknown to science. Second, we discovered that phylogenetically our mushrooms that look like Gymnopus dryophilus divide into two clades. One matches the European concept of G. dryophilus, while the other falls in with the European G. alpinus. Very conveniently, once we were able to divide our finds into two groups, we saw retrospectively that they grew in entirely different habitats, making future field identification possible, despite their similarity (most of the time). While the split into two clades is irrefragable, their identity is less clear. As you will see throughout this study, DNA matching can get a very good fit with a named species, but the accuracy of that identification depends on the original morphologic matching with the species description. This is difficult because many descriptions are very cursory and in many cases type specimens do not exist. As a result, the difficulty is to decide which of many matched species epithets is correct for the mushroom in your hand. If the match is with a single name only, the difficulty is to know the accuracy of the original morphologic identification. In other words, you have the same species, but is it really the species named? (Advance warning: you will see more examples of this in our next issue.) The lead article (as usual, following foray information) is the first formal report of a new species on the pages of OMPHALINA. This is a very exact process, where the format must follow the International Code of Nomenclature for algae, fungi, and plants in every detail, lest the name be declared invalid.

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Give Gi Given venn th ve that hatt OMPHALINA co considers cons nsid ns ider id erss it er its pr its pprime ime au ime im audi audience dien di ence en ce the amateur mycophiles Newfoundland th t hil off N f dl d and d Labrador, in normal times it is very much a cap-andstem type of journal. Using good English words can make even the most complex ideas accessible to the lay enthusiast, ideas which may become obscured by technical language. For us there is no advantage to pileus and stipe over cap and stem—dare we say, an affectation? However, when it comes to reporting a new species, the intended audience—in addition to mycophiles in our province—is the world, and, specifically, its mycologists. Therefore, the language is much more technical than is normal for OMPHALINA. If it wants to be a vehicle for such reports, this is the price. In our opinion the price is small. We believe that the mycophiles of Newfoundland and Labrador would much rather hear of new species in their province on the pages of their journal, even if it means reading some technical language. Part of the you-read-it-here-first policy. Read the article, and tell us your opinion! Often we complain about the skimpy descriptions for new species; we hope that this description contains sufficient information for future generations to know the characters of our species. See you at the Foray! andrus Please note correct Foray dates:

September 12-14, 2014

Foray matters… 1. 1 INFORMATION, INFORMATION REGISTRATION FORM FORM, HOW TO GET THERE, etc. Please see OMPHALINA V, No 4 (April issue), as well as material stored on our website <nlmushrooms.ca>. Past issues of OMPHALINA may be downloaded from our website.

att Si Sir Ri Richard h dS Squires i M Memorial i lP Provincial i i l ark k at 11:00 AM, Fri. Sep. 12, 2014. See the April OMPHALINA for details. 3. SHOW & SELL ARTS & CRAFTS TABLE Artists and craftspeople: Do not forget this. See last issue for more details. Please register with Glynn Bishop <fozmos AT gmail DOT com>, who is in charge of setting up the table.

2. MYCOBLITZ For those able to make it, join us

THE BASIC BARRENS OF GROS MORNE Michael Burzynski This is the first of four short articles dealing with the ecologic habitats of Gros Morne National Park, each to appear in a consecutive issue of OMPHALINA. This page shows a limestone tallus slope. The next page shows

limestone scree in the upper picture, and a typical peridiotite moonscape in the lower picture—dwarf juniper being one of the few things to grow there. The granitic highland barrens illustrated by the mountaintop on the horizon, will be discussed in the next issue. Despite their seeming lack of plant and animal life, barrens are some of the most beautiful and biologically important areas in Gros Morne National Park, .

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Soils are called basic (or alkaline) if they result from the breakdown of limestone, basalt, peridotite, or other rocks that have a high content of calcium or magnesium—giving them a pH greater than 7 (pure water has a pH around 7). The basic soils derived from the Green Gardens basalt are extremely rich, because of the other nutrients in the rock. However, soils associated with limestone and peridotite often support very little vegetation, earning them the name barrens. The park’s barrens may look a bit like deserts, but that is not due to a lack of water. Peridotite is a dark green ultramafic rock, originally from far below the ocean floor. It forms the Tablelands plateau in the south of the park. This rock is high in magnesium, iron, chromium, and nickel, and is chemically unstable in the presence of air and water. Groundwater has altered the rock into serpentinite, and the iron has oxidized (rusted), turning the outside of the rocks orange-brown. The high metal content, alkalinity, and lack of regular soil nutrients (nitrogen, potassium, and phosphorus) make it very difficult for most plants to grow. Without plants to feed on, it is similarly very difficult for most fungi to survive there. However, the difficult conditions also mean that what fungi grow there might be very unusual, so it is always worth looking! Even lichens have a difficult time in the Tablelands because of the crumbly rock surfaces and metals, and only a handful are found there—most rocks are completely bare, even after over 10,000 years of growing time since the last glaciation. The exception is seen on erratics, rocks from elsewhere dropped by melting glaciers. Those are covered in bright lichens, showing how important substrate can be to growth. Limestone bedrock forms a band in the southern and eastern parts of the park. For the most part, the limestone has been covered with a layer of soil, and lies beneath forest and wetland. However, in places it pops out of the soil as small outcrops, and along valleys, bays, and the outer coast there are steep cliffs with crumbling tallus (or scree) slopes, This is another extremely important habitat for rare specialized plants, often with arctic and alpine affinities. Because of the steepness of the cliffs, fieldwork is extremely difficult, and there is probably a lot left to learn about the plants, fungi, and lichens that grow there. Because of the lack of vegetation and exposed locations of the barrens, climate also has a strong effect on the organisms that try to grow there. When you visit, be prepared for high winds and cooler conditions than elsewhere in the park. You will be rewarded by the raw beauty of the scenery, and the exotic nature of the life that you see.

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Gymnopus eneficola—species nova from Newfoundland Ron Petersen,1 Karen Hughes,1 Andrus Voitk2

Introduction In the course of a survey of dried collections of Gymnopus collected over some years in Newfoundland by AV and Foray Newfoundland & Labrador, a taxon was discovered which did not conform to other known eastern North American or European taxa of the genus.1,2 Occasional reports of taxa now placed in Gymnopus (formerly included in an expansive concept of Collybia) include those concerning Nova Scotia,3 summarized by Gourley,4 and Michigan,5,6,7,8 none of which, nor those of the even older publication by Coker and Beardslee9 circumscribed the proposed species from Newfoundland. In addition to morphological distinctions, DNA analysis also indicated that nuclear ribosomal LSU and ITS sequences from the new taxon did not match those deposited in GenBank <http://www. ncbi.nlm.nih.gov/nucleotide> or sequences in the Gymnopus files at TENN.10 The new taxon is proposed, illustrated and described below. Materials and methods

Methods for DNA extraction, PCR of the nuclear ribosomal LSU and ITS region and sequencing were Collection of fresh specimens employed typical field carried out as described previously.13 Sequences were techniques. In several cases, photographs were made aligned in GCG14 and a blast search was used to in the field, and basidiomata were dried and preserved query GenBank. GenBank accession numbers for G. in the fungarium of AV. In situ observation of eneficola ITS sequences are KJ128262-KJ128268; LSU macroscopic characters by AV have been augmented sequences are KJ189586-189590, and all are listed by deductions from dried material, but chiefly from individually under additional specimens examined. several photographs, some of which are reproduced A preliminary analysis of our entire (TENN) LSU here. Microscopic features were observed exclusively Gymnopus library showed that G. eneficola was most from dried basidiomata. Microscopy and “barcoding” closely related to G. menehune, G. confluens and G. (i.e. production of ITS sequences) took place biformis. The closest outgroup was Gymnopus ramealis. using small bits of dried material. Microscopy was LSU sequences from collections of these species were accomplished using an Olympus BX60 microscope used for parsimony analysis to demonstrate placement under Phase Contrast Microscopy (PhC), using of G. eneficola (Figure 10). 3% KOH with no stains. Abbreviations: TENN = Results herbarium, University of Tennessee; Q = spore length divided by spore width; Qm = mean Q value for all Gymnopus eneficola R.H. Petersen, species nova spores measured; Lm = mean length of all spores MycoBank no. 807536 measured. Colours within quotation marks from 11 Ridgway, and those in parentheses from Kornerup Holotype: Canada, Newfoundland, Pasadena Ski and and Wanscher.12 Nature Club ski trails, coll. A. Voitk, 49° 0’ 6” N, 57° 35’ 1

Ecology & Evolutionary Biology, University of Tennessee, Knoxville, TN 37996-1100, USA. repete@utk.edu 2

Box 2312, RR #1, Corner Brook, NL, A2H 2N2 CANADA

36” W, 41 m ASL, 26.IX.2009, coll no. 09.09.26av13 (TENN 69123) KJ128264, KJ189586.

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1 cm Figure 1. Gymnopus eneficola, basidiomata. Type collection from The Pasadena Ski and Nature Park ski trails, Pasadena, NL, 09.09.26av13 (TENN 69128). Note: the camera angle makes stipes look shorter than they are.

1 cm Figure 2. Gymnopus eneficola, basidiomata. 06.08.19av01 (TENN 69126), Humber Village, Newfoundland and Labrador. Note: camera angle makes stipes of lying basiomata seem shorter than in real life.

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Etymology: En = “N;” ef = “F” (NF = abbreviation for Newfoundland); + -icola = dwelling in or preference for. English diagnosis: Basidiomata collybioid or marasmioid. Pileus white when young, remaining so or mellowing to cream color; lamellae subdistant, adnate with slight tooth, white when young, mellowing to off-white by maturity; stipe terete when young becoming compressed or grooved by maturity, lightly vestured above, more strongly so downward, disappearing into a felty mycelial mass binding substrate. Pileipellis a modified dryophila structure with erect pileal hairs. Cheilocystidia 30-40 × 8-13 μm, variously contorted, lobed and roughly coralloid. Caulocystidia abundant, hyphal, downward long and branced. Basidiospores 7.5-9 × 3.5-5 μm (Qm = 2.32; Lm = 8.30 μm), ellipsoid, slightly flattened adaxially, thin-walled, hyaline, smooth, inamyloid.

1 cm

Description MACROSCOPIC Basidiomata collybioid (Fig. 4), marasmioid, Figure 3. Gymnopus eneficola basidiomata. 05.10.12av05 (TENN 69121), Humber Village, Newfoundland and Labrador.

1 cm 1 cm 1 cm 1 cm

Figure 4. Gymnopus eneficola, basidiomata. MR3-016 (TENN 69120), 2011 Faculty Foray, Main River, Deep Section. Photo: Roger Smith.

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Figure 5. Known distribution of Gymnopus eneficola. Limitation to the west coast region of the Island likely reflects the location of AV in that area. However, it is noteworthy that annual forays in other areas have not collected it.

b

a

20 μm

c

Figure 6. Gymnopus eneficola: Cells of pileipellis. a) Cells of pileipellis surface [09.09.26av13 (TENN 69128)]. b) Repent terminal cells of pileus surface [09.09.26av13 (TENN 69128)]. c) Erect slender hyphal tips (“pileal hairs”) [06.09.02av01 (TENN 69127)].

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especially sect. Globulares; (i.e. adult pileipellis smooth, with pileus diameter/stipe length 1:1-1:1.5; Figs. 1, 2), when young sometimes mycenoid (pileus diameter/ stipe length 1:4-1:2) or hygrophoroid (Fig. 3), solitary, gregarious or cespitose in small groups of 2-3 (Fig. 1), with obviously vestured stipe (Fig. 3) and stipe base disappearing into felty-arachnoid hyphal mass involving minute bits of substrate (Fig. 2, right; 4). Pileus (Figs. 1-3) 11-24 mm broad, thin, convex when young (Fig. 1) and then with suggestion of an umbo, expanding and flattening through development, finally nearly applanate with thin, downturned but not inrolled margin (Figs. 1, right; 2, left), sometimes with a suggestion of central depression, smooth under lens, matte or occasionally plushy as a peach (60×), appearing polished in small areas especially in age, not striate, not hygrophanous, white when fresh (Fig. 1, center; 3), sometimes with very slight blush of “pinkish buff ” (6A3) over disc (Fig. 1, left), slowly mellowing to cream-colored in age (Figs. 2, 4) from margin inward. Lamellae subdistant, adnate with slight tooth, ventricose, seceding in drying, sinuate, thickish, white when young (Figs. 1, 3), discoloring slightly over time through “pale cinnamon pink” (5A2) or “tilleul buff ” (7B2) to “cinnamon buff ” (6B4) or “cinnamon” (6B5) in age and drying; lamellar edge entire or slightly eroded (Figs. 1, 2) or undulate but not serrate; lamellulae in three ranks. Stipe (Fig. 2) 30-55 × 1.5-5 mm broad, initially terete (Figs. 1, 3), often becoming grooved or compressed (Fig. 2, right; 4, right), equal through upper portion, slightly expanded in lowest portion and there covered with loosely felty to sublannose mycelium (Figs. 2, 3, 4) and disappearing into and binding superficial substrate (Fig. 2), white when fresh becoming pallid creamy gray on drying, perhaps tan where chafed; vesture of upper stipe appearing frosted or minutely granular (Figs. 2, 3), downward vesture becoming thinly felty, then substrigose and finally at base densely arachnoid to lannose, especially when young (Figs. 3, 4), vesture easily crushed or removed where chafed (Fig. 2, right; 4, left); 3% KOH applied to lower stipe surface (dry) with off-white sublannose vesture = dull orangebrown (no evidence of green or citrine). HABITAT, PHENOLOGY AND DISTRIBUTION “Mixed woods,” (Betula, Abies balsamea, Picea, Larix; sometimes dominated by Betula, and no record without Betula), fruiting mid-August through early Novermber. At the moment, only known from a limited area of the Island of Newfoundland in the Canadian province of Newfoundland and Labrador (Fig. 5).

MICROSCOPIC

a

Pileipellis composed of generally repent, radially oriented hyphae 4-10 μm diam, often broadly “free-form” (i.e. individually resembling cells of a “dryophila structure;” Fig. 6a) firm- to thick-walled (wall -0.7 μm thick), not or hardly ornamented (ornamentation very vague bands under PhC but not raised or producing profile calluses, not visible in BF), frequently branched, conspicuously clamped, hyaline (PhC); terminal cells of pileipellis hyphae (Fig. 6b) 35-80 × 4-7 μm, single or in processes of 2-4 cells with clamp connections at septa, usually simple, occasionally tibiiform or substrangulate near terminus, firm-walled (wall -0.5 μm thick), hyaline (PhC). Occasional slender branches from pileipellis hyphae erect (“pileal hairs,” Fig. 6c), -225 × 2-3(-4.5) μm, hyphal, unbranched, not necessarily arising at a clamp connection, usually internally clamped, firm-walled, hyaline. Hyphae of pileus trama Figure 7. Line drawings of G. eneficola hymenial structures. 4-8 μm diam, firm-walled, frequently branched, a) Basidioles [09.09.26av13 (TENN 69128)]. frequently clamped, easily disarticulated, never b) Immature basidia [09.09.26av13 (TENN 69128)]. sculptured. Lamellar trama generally longitudinal; c) Mature basidia (note 4- and 2-sterigmate individuals) tramal hyphae 4-7.5 μm diam, thin-walled, usually [09.09.26av13 (TENN 69128)]. constricted at septa, clamped. Subhymenium a d) Cheilocystidia [09.09.26av13 (TENN 69128)]. e) Cheilocystidia [07.11.07av01 (TENN 69125)]. tight tissue of tortuous/branched hyphae 2-2.5 μm diam, bearing basidia on subsymbodial branches. Basidioles (Fig. 7a) 29-31 × 5-7 μm, digitate when young, becoming clavate/fusiform, clamped, thin-walled; contents homogeneous. Basidia (Fig. 7b, c) 23-30 × (7-)9-10 μm, broadly to bulbo-clavate, 2-4-sterigmate (sterigmata – 11 μm long when 2, -5 μm long when 4), clamped; contents homogeneous. Pleurocystidia not observed. Cheilocystidia (Fig. 7d,e) very locally common, overall occasional, (25-)30-40(-60) × 8-13(-17) μm, usually arising with basidia, hyaline, occasionally rising in subhymenium or outer trama, free-form with irregular lobes and curved branches, clamped. Stipe surface hyphae 4-11 μm diam, adherent, thick-walled (wall -1.5 μm thick), refringent (PhC), seldom but prominently clamped, hyaline. Caulocystidia from stipe apex (Fig. 8a) of two types: 1) a solid turf of short (-70 × 3-4 μm) hyphal tips arising as side branches and termini of stipe surface hyphae, firm- to thickwalled (wall -0.7 μm thick), internally clamped; Figure 8. Line drawings of Gymnopus eneficola caulocystidia. and 2) a tangle of common longer (-175 μm a) Short caulocystidia with longer individuals from upper long × 3-4 μm diam) hyphae, similar to shorter stipe [09.09.26av13 (TENN 69128)]. caulocystidia, branched 1-3 times at conspicuous b) caulocystidia from lower portion of stipe [06.09.02av01 clamp connections, hyaline, internally clamped (TENN 69127)].

b

c

20 μm

d

e

a

20 μm

b

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5 μm

Figure 9. Line drawings of G. eneficola basidiospores [06.09.02av01 (TENN 69127)].

so terminal cells of long caulocystidia 17-27 μm long. Caulocystidia from stipe midsection (Fig. 8b) -240 × 3-4 μm, often branched near origin, hyphal, firm- to thick-walled (wall -0.7 μm thick), seldom clamped and almost never near cystidial apex; when branched, one branch consistently short and less developed; turf of shorter caulocystidia characteristic of upper stipe present only as scattered individuals. Basal mycelium inward organized into tissues of adherent hyphae 3-4 μm diam, firm- but not thick-walled, shearing into plates indicating adherent surfaces; outward hyphae disorganized, 3-5 μm diam, similar to caulocystidial hyphae, conspicuously clamped, firm-walled, hyaline, but macroscopically slightly discolored toward pinkish cinnamon, involving small bits of debris including welldecayed leaves, tiny twigs or petioles and occasionally flattened needles, probably of Abies. Basidiospores (Fig. 9) (7-)7.5-9 × (3-)3.5-5 μm (Q = 2.00-2.73; Qm = 2.32; Lm = 8.30 μm), ellipsoid, somewhat flattened adaxially, thin-walled, hyaline, inamyloid; contents apparently homogeneous. PHYLOGENY LSU sequences place G. eneficola well within the Gymnopus clade (clade Omphalotaceae, not Marasmiaceae in the Agaricales). Neither LSU nor ITS sequences from collections of G. eneficola match any GenBank sequences more closely than 92%; the closest ITS blast match was to GenBank accession GU234141, Gymnopus alkalivirens from the Svalbard. An abbreviated LSU phylogram is shown in Figure 10. Discussion From notes and photos accompanying specimens, habitat might be inferred as decaying deciduous leaves and small twigs, favoring Betula. Most specimens were collected as G. confluens, also fruiting in such habitat, with which they are similar in vestured stipe, which is usually compressed or grooved, and presence of a basal mycelial mass which binds minute bits

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of litter and probable substrate. The extent of the basal mycelium is unknown at present (whether as widespread sheets as those of typical of G. confluens), but lamellar spacing (as a function of number), height and thickness and spore dimensions all differ substantially. Basidiomata of G. eneficola might be mistaken for Collybia tergina (Fr.) Lundell if Halling1 is used, because the stature of basidiomata and spore dimensions are similar. However, under C. tergina, Lundell and Nannfeldt15 described an organism with smaller spores [5.5-7(-8) × 2.5-3.5 μm], “stem … especially lax and almost flabby,” stipe “cylindrical and smooth, except for the base, which in some specimens is slightly villose,” and “in almost perfect agreement with Fries’s taxon” as illustrated in Fries.When Fries’s16 illustration and description are examined, his Agaricus (Marasmius) terginus exhibits a tan, striate pileus (in one individual quite pale but with tan disc) and tan stipe with no evidence of vesture. The sum of all these characters indicates an organism quite different from G. eneficola. When the key to European Gymnopus species by Antonin & Noordeloos2 is followed, a pivotal couplet concerns strong odor of basidiomata, a feature not observed for the new species. Choices which follow the strong odor lead to mismatched character fields, but if the choice of little or no odor is followed, the key permits several more choices until finally G. oreadoides is reached. Because basidiomata of G. eneficola resemble those of the Globulares section of Marasmius, G. oreadoides is a tempting choice. Numerous discrepancies are found, however, when characters of G. eneficola are compared to those of G. oreadoides, as follows: 1. the pileus of G. oreadoides is described as “probably hygrophanous” (the accompanying photo clearly shows this); 2. pileus “pale cream color when young” (the accompanying photo shows strongly convex, neutral brown pileus of young basidiomata and distinctly ochraceous disc when mature); 3. stipe “fistulose, elastic, twisted… turning orange-ochraceous, finely pubescent, glabrescent; 4. spores “5.5-7.5(8.5) × 3.0-4.0 μm … Q = 1.9-2.1;” 5. caulocystidia… “20-50 × 3.5-7.5 μm, forming a dense layer all over the stipe surface.” The line drawing accompanying the description illustrates these characters, with only short caulocystidia shown, together with encrusted pielpellis hyphae. Finally, “Gymnopus strictipes (Peck) Halling” is reported as similar to G. oreadoides, but Peck’s species is a member of Marasmius sect. Globulares.

NL

KJ189586 G. ENEFICOLA 090926AV13 NL TYPE

100

KJ189587 G. ENEFICOLA 051012AV05 NL KJ189588 G. ENEFICOLA 060902AV01 NL KJ189589 G. ENEFICOLA 090926AV12 NL KJ189590 G. ENEFICOLA 100921AV04 NL

Figure 10. Parsimony analysis of nuclear ribosomal LSU sequences for Gymnopus collections related to G. eneficola. The figure represents one of 1000 most parsimonious trees of length 74. Origin of collections indicated by country name in full or abbreviation for state/province if from the USA or Canada, appearing after the species name. Bootstrap support greater than 50% is given to the left of the supported node.

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Superficially, basidiomata are reminiscent of members of Marasmius sect. Globulares (i.e. M. cystidiosus, M. nigrodiscus, M. oreades) in general stature, lamellar morphology and attachment and pileus shape. Pileipellis construction of Gymnopus eneficola is not hymeniform, however, and DNA sequences place the Newfoundland taxon well within the Gymnopus clade. Moreover, basidiomata of Gymnopus eneficola are macromorpholigally more diminutive, have a greater stipe:pileus ratio and exhibit a vestured stipe at all ages.

coll. A. Voitk, no. 06.08.19av01 (TENN 69126); Pasadena Ski Trails, 49° 0’ 6” N, 57° 35’ 36” W, 26.IX.2009, coll. A. Voitk, no. 09.09.26av12 (TENN 69128) KJ128268, KJ189589; Pasadena Stream Trail, 49o 00’ 39” N, 57o 36’ 38” W, 21.IX.2010, coll. A. Voitk, no. 10.09.21av04 (TENN 69122) KJ128265, KJ189590; Gros Morne Nat. Park, Stuckless Pond Trail, 49o 25’ 55” N, 57o 43’ 38” W, 1.XI.2005, coll. A. Voitk, no. 05.11.01av08 (TENN 69124); same location, 2.IX.2006, coll. A. Voitk, no. 06.09.02av01 (TENN 69127) KJ128267, KJ189588; Lower Main River (deep section), Trail 1, 49o 46’ 40” N, 57o 53’ 03” W, 6.IX.2011, coll. Foray Newfoundland and Labrador, no. MR3-016 (TENN 69120) KJ128262.

At first, ventricose-rostrate immature basidia were thought to be cheilocystidia, being present consistently and abundantly, but they are present on lamellar faces (i.e. among basidia covering the lamellar surfaces) and seem to disappear near the lamellar edge. Once true cheilocystidia were observed, they could be compared to other hymenial structures for shape and size (see Fig. 7). Future caution is recommended. Caulocystidia (Fig. 8) are somewhat unique in distribution, dimensions and structure. The upper stipe appears only “frosted” without magnification, but magnification (750×) reveals a solid turf of short caulocystidia with some long indivduals present. Downward on the stipe, the turf remains but long caulocystidia become more numerous to give a somewhat more hirsute appearance to the naked eye. Both types of caulocystidia are consistently internally clamped so that the terminal cell is considerably shorter than expected. Clamp connections, especially those of basal mycelium, vary from papillate to medallion shapes to individuals with the “hook cell” elongated over the parent hyphae.

Thanks are offered to Mr. Matt Aldrovandi who performed DNA extractions and PCR. Support, in part, came from grant DEB-1144974 of the (US) National Science Foundation.

At the time of this writing (Jan. 2014), GenBank does not report any nucleotide sequences under the name Gymnopus oreadoides. Just as important, ITS sequences from collections of G. eneficola do not match any GenBank sequences more closely than 92%. However, with a blast match of 92% percent to a specimen from Svalbard, collections of G. eneficola from Newfoundland would be considered a distinct species by any criteria. LSU and ITS sequences of G. eneficola have been deposited in GenBank and are listed under additional specimens examined.

9.

Additional specimens examined (all Canada, all Newfoundland, all collected as G. confluens): Humber Village, 13 Balsam, 48o 59’ 21” N, 57o 45’ 18” W, 12.X.2005, coll. A. Voitk, no. 05.10.12av05 (TENN 69121) KJ128263, KJ189567; same location, 7.XI.2007, coll. A. Voitk, no. 07.11.07av01 (TENN 69125) KJ128266; Humber Village, Weldon’s Road, 48o 59’ 43” N, 57o 44’ 15” W, 19.VIII.2006,

15.

12 OMPHALINA

Acknowledgments

References 1.

2. 3.

4. 5.

6. 7. 8.

10.

11. 12. 13.

14.

16.

Halling RE: The genus Collybia (Agaricales) in the northeastern United States and adjacent Canada. Mycol. Mem.8: 1-148. 1983. Antonin V, Noordeloos ME: A monograph of marasmioid and collybioid fungi in Europe. IHW Verlag. 478 pp. 2010. Smith AH, Weymeyer LE: Contributions to a study of the fungous flora of Nova Scotia. II. Agaricaceae and Boletaceae. Pap. Mich. Acad. Sci. 21: 163-197. “1935” [1936]. Gourley CO: An annotated index of the fungi of Nova Scotia. Proc. Nova Scotian Inst. Sci., 32(2/3), 75-293. 1982. Kauffman CH, Smith AH: Agarics collected in the vicinity of Rock River, Michigan, in 1929. Pap. Mich. Acad. Sci. 17: 153-200. “1932.” [1933] Smith AH: “1935” [1936]. Unusual agarics from Michigan. III. Pap. Mich. Acad. Sci. 21: 147-161. Smith AH: New and unusual agarics from Michigan. II. Pap. Mich. Acad. Sci. 26: 61-68. “1940” [1941]. Smith AH: New records of fleshy fungi in Michigan. Asa Gray Bull. 1(2): 185-194. 1952. Coker WC, Beardslee HC: The collybias of North Carolina. J. Elisha Mitchell Sci. Soc. 37: 83-107. 1921. Mata JL, Hughes KW, Petersen RH: An investigation of / omphalotaceae (Fungi: Euagarics) with emphasis on the genus Gymnopus. Sydowia 58: 191-289. 2007. Ridgway R: Color standards and color nomenclature. Publ. priv. Washington, DC. 1912. Kornerup A, Wanscher JH: Methuen handbook of colour. Methuen & Co., London. 1967. Hughes K W, Petersen RH, Lodge DJ, Bergemann S, Baumgartner K, Tulloss RE, Lickey E, Cifuentes J: Evolutionary consequences of putative intra- and interspecific hybridization in agaric fungi. Mycologia 105: 1577-1594. 2013. GCG: Wisconsin Package, Version 10.3. San Diego, California, Accelrys Inc. 2000. Lundell S, Nannfeldt JAF: Fungi Exsiccati Suecici, Fasc. XXIIIXXIV, no. 1109. Praesertim Upsalaensis. 1942. Fries EM: Icones selectae. Hymenomycetum nondum delinatorum. Volume 2: Pl. 174, Fig. 4. Uppsala. 1877.

Gymnopus-dryophilus-like Gymnopus-dryophiluslike mushrooms in NL Karen Hughes, Matt Aldrovandi, Ron Petersen, Andrus Voitk The beautiful Gymnopus dryophilus has a varied appearance, which may not have mattered to most mycophiles, were it not that it is reputed to be one of the commonest mushroom species in North America. It is said that rare is the foray on this continent, regardless of location or season, that does not have it on its list. Not surprisingly, many related taxa have been described, some with more, some with less convincing supporting evidence. Most of the older species descriptions lack detail and type specimens. Many seem to overlap, and many have been reinterpreted by later investigators, synonymized and then split again, and, of course, new taxa described. An accurate description of the result is: a mess! Enter the age of phylogeny. Surely DNA can solve all messes? Well, for species within the Gymnopus sect. Levipedes group (G. dryophilus and related species), not quite. The DNA barcode selected for fungi (part of the gene for ribosomal RNAs) does not separate these collections into reliable species groups, and species boundaries are unclear. Geographically, it is only in situations where species diversity is limited (such as Newfoundland) that it becomes possible to distinguish species using this barcode. Further difficulties occur if the original morphologic identifications have been problematic: the resulting clusters contain specimens bearing various species names, and often specimens bearing the same names end up in several distinct clusters!1 Deciding which is the correct name for each cluster becomes more difficult than defining the phylogenetic groups. How did the NL mushrooms that look like G. dryophilus negotiate this abyss? Over a decade, Foray Newfoundland & Labrador and AV together had accumulated Group 1 Group 2 Total Gymnopus alpinus

1

Gymnopus aquosus

1 2

2

13

4

17

1

1

2

Gymnopus loiseleurietorum

7

7

Gymnopus ocior

3

3

1

3

1

1

Gymnopus dryophilus Gymnopus earlei

Gymnopus subsulphureus

2

Gymnopus terginus Rhodocollybia butyracea TOTAL

1 18

1 19

37

Table 1. Original identifications of the collections in Groups 1 & 2. Nine taxa divided between two clades. Three taxa are found in both clades.

57 collections of mushrooms that look like Gymnopus dryophilus. These were examined morphologically and phylogenetically, with more than their share of difficulties because of mould contamination and relative paucity of material in many collections. Good DNA was obtained from 37 collections of Gymnopusdryophilus-like mushrooms. On nuclear phyleogenetic analysis they fell into two distinct clades, Group 1 and Group 2. Original identifications are listed in Table 1. As can be seen, our experience was the same as described above: two groups contained nine putative species, three of which fell into both. Morphologically, Group 1 fit with the current concept of Gymnopus dryophilus, and its DNA was 99.4% homologous with G. dryophilus, as defined by Antonin and co-workers.2 Group 2 matched a species complex in the University of Tennessee database that extends across Northern Europe and North America, with a diverse morphological expression. It grouped together with G. alpinus of Antonin,2 but also with collections with different names. Hybrids between the two groups were not detected. Figure 1 shows the phylogram for the two groups.

OMPHALINA 13

N L G R O U P 1

N L G R O U P

2 difference is considered normal, attributed to geographic and individual variation. A difference This is an unrooted tree (branches over 2% has been shown to indicate sufficient divergence of connecting the studied groups to the evolutionary DNA pathway to their other relatives and common represent a different species 97% ancestors are omitted), which 3 of the time. In this analysis, the allows clear focus on the derived difference was 0.53% between the groups. Origin of the collections European neotype of G. dryophilus is indicated after the collection (pink panel) and our Group 1. number: NL=Newfoundland This makes it highly likely that and Labrador, GRL=Greenland, all collections on the dark green TN=Tennessee, MI=Michigan, panel are the same species. The ENV= environmental sample (soil difference between the neotype or litter); h1 & h2= haplotypes of G. dryophilus and Group 2, of the same fruitbody; bootstrap values noted above the branch, left was much greater, 2.64%, clearly indicating that they are not the of the node. same species. Therefore, it should Our collections cluster in two very not be surprising that Group 2 also distinct clades, NL Group 1 (light differed from Group 1 (3.31%), green panel) and NL Group 2 (light confirming that they are two brown panels). different species. The second group aligns with European Gymnopus The DNA sequence in the selected alpinus (teal panel), along with ITS site of the DNA should be North American collections given similar for each species. A small different names (white panel). The Figure 1. Phylogeny of the Gymnopus-dryophilus-like mushrooms in NL.

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present analysis, limited to ITS sequences only, does not permit us to determine whether Group 2 is G. alpinus or another species with a nearly identical ITS sequence but it is likely that the Newfoundland Group 2 is, itself, a single species. Bootstrap values measure the probability (as per cent) that the node is “real” and not a chance observation. The values are determined by random sampling of sites in a set of sequences a set number of times, generating a series of trees. High bootstrap values (>70%) indicate that the tree shown appears at high frequency regardless of which sites were sampled. Since both Group 1 and Group 2 have bootstrap values of 100%, this indicates that alternate arrangements of taxa (trees) were not found by this process, and the tree is an extremely reliable representation of the “true” relationship of these taxa.

Group 1 and Group 2 both contained collections with varied morphology, and no characters could be identified that separated them consistently. Specifically, both dark and light coloured caps, as well as yellow gills were seen in both groups. Size varied within but not between groups. Fruiting times were similar for both (Figure 2). The only difference noted for the groups was in habitat. Group 1 was a woodland species (1 of 18 from barrens) and Group 2 was a species of open areas like heaths, bogs, coastal and mountain barrens (3 of 19 from wooded areas) (Figure 3). Chi squared analaysis of the difference in habitat was extremely significant (p<0.0001). Discussion To find nine putative species reduced to two is in keeping with observations of other genera, where the diversity of Newfoundland and Labrador mycota is much less than on the continent, presumably related to our low diversity of vascular plants. Our experience confirms that correct morphologic identification of species in this complex is difficult.

Figure 2. The phenology (seasonality) of the G. dryophilus complex in NL. First fruitbodies in May, peak in June, none in August. Smaller 2nd peak in September, none in October. Dark green represents Group 1 and lighter green Group 2. Both show similar pattern. Because forays are only in September, only AV collections were used here, to avoid skewing the results in favour of a September peak.

We have not identified useful foolproof morphologic characters to differentiate Groups 1 & 2, but the distinct difference in habitat, readily confirmed in retrospect on in situ photographs (Figure 3), separated them well. To our knowledge, such difference has not been documented before. As seen in Figure 1, nuclear sequence studies show the two groups to be clearly distinct. In regions with more diversity, such clear separation into clades is not found, due to many phylogenetically overlapping taxa. The evidence suggests that Group 1, the woodland species, likely is Gymnopus dryophylus. The exact identity of Group 2 is not sufficiently clear. It is likely a single species, and fits well with G. alpinus. However, more analyses using different marker sites or multigene studies, may show a better fit with a closely related species. For now, we leave the resolution of these questions to future investigations and investigators. References 1. Mata JL, Hughes KW, Petersen RH: An investigation of / omphalotaceae (Fungi: Eugarics) with emphasis on the genus Gymnopus. Sydowia 58:191-289. 2007. 2. Antonin V, Sedlåk P, Tomsovsky M: Taxonomy and phylogeny of European Gymnopus subsection Levipedes (Basidiomycota, Omphalotaceae). Persoonia 31:179-187. 2013. 3. Hughes KW, Petersen RH, Lickey EB: Using heterozygosity to estimate a percentage DNA sequence similarity for environmental species’ delimitation across basidiomycete fungi. New Phytologist, 182:795-798, 209.

Figure 3. Above, Group 1 in its typical woodland habitat, with characteristic leaf and needle litter substrate readily visible. Below Group 2 collection from a treeless habitat. No tree litter, but each photo showed a characteristic background of Sphagnum, Empetrum, Vaccinium, Loiseleuria, Dryas, or other heathland species. The mat of Salix uva-ursi (bearberry willow) in this photo is as close to a real tree as most of them got.

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The Bishop’s Sketchbook

16 O OMPHALINA MPHALINA

Dryophilic and dryophobic clade clusters Andrus Voitk, Karen Hughes, Matt Aldrovandi

Dryophilus is usually given as oak-loving. The name comes from Dryad, the Greek wood nymph. The title banner depicts a stunning scene from the Royal Ballet’s 2013 Dance of the Dryads from the ballet Don Quixote. Although particularly fond of oaks, Dryads occupy all trees and only very specialized nymphs have an especial relationship with oak. For a few, like the Hamadryads, it is obligatory; they die if their oak dies. We know that G. dryophilus does not have an obligatory relationship with oak: we have no oak in Newfoundland and Labrador, but lots of G. dryophilus. A more accurate translation of dryophilus, in keeping with both its behaviour in our woods and the original meaning of name, is tree-loving. G. dryophilus is a saprobe, a decomposer of forest litter. Saprobes excrete digestive compounds to break down organic matter so that it can be absorbed by its hyphae. The chemical structure of all organic matter is not the same, and different digestive compounds break down different organic matter. The production of such compounds is genetically determined, and, as all genetic processes, subject to evolutionary change. Let us assume that species B lives in a mixed forest, happily digesting Both conifer and deciduous debris. Either it secretes two compounds, one to digest conifers and one to digest deciduous wood, or a multipurpose compound to digest both. How much more efficient to specialize, make chemicals to digest only one? Just as some individuals are taller, others shorter, so, more than likely, some are slightly better at digesting coniferous, others deciduous wood. On leafy debris, individuals better able to digest hardwood are able to outcompete their fellows: more likely to produce fruiting bodies emitting spores with their genetic program for enhanced deciduous wood digestion. A similar story plays out on coniferous duff: individuals better able to digest softwood perpetuate

Dryads are hotly pursued by Satyrs, a relationship variously depicted in art as romantic love or brutalization. Wags observe that nobody brutalized Dryads more than Pablo Picasso in his 1908 cubistic painting, The Dryad. This may be more apt than it seems, because Picasso often used the goat symbol for himself.

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17

Oak grove

eats only Oak

Coniferous woods

Deciduous woods

O

alder, birch, oak

eats only

Coniferous

C

eats only

Deciduous

Mixed woods eats Both deciduous & coniferous

B

D

eats only Alder

A Alder bush

Adaptation and evolution. Species B, a generalist saprobe in a mixed forest, may move into deciduous and coniferous woods and evolve into specialist digesters of their respective substrate only, species D and C. Species C may travel further, into pure oak and alder woods, and there evolve into an obligatory oak or alder digester, species O and A. If some natural event, like an advancing polar ice cap, forced vegetation and its associated fungi to retreat to the original mixed woods, these derived species would exist together without hybridization, exposing the more specialized species to much greater risk of extinction from stochastic events to their obligatory nutritional substrate.

their genetic traits. These strains are sympatric (sym = same, patria = homeland, i.e. growing in the same region). Proximity allows for mixing of genetic material. So long as they can interbreed (i.e. before full speciation has evolved, when they can no longer mate to produce fertile spores), there is a continuous intermixing of their genetic material. The result will cancel any advantage in either direction. Strains mate backwards and forwards, left and right, maintaining a homogeneous population of species B, thwarting all effort at specialization. Now, let us suppose that spores of species B get carried to pure coniferous or deciduous woods. A meaningless variation in digestive enzyme composition suddenly becomes significant. If you can only digest two kinds of food, but only one is available, your survival depends on your ability to thrive on it. In the coniferous forest those individuals of species B, who are more adept at digesting coniferous wood, easily outcompete their kin, producing individuals with continuously increasing ability to digest coniferous wood. In time, through a natural selection process, a new and genetically distinct species, C, evolves, able to digest only Coniferous wood. By the same process, in

18 OMPHALINA

the deciduous woods a new species, D, evolves, able to digest only Deciduous wood. Some individuals among species D immediately grasp the benefits of specialization: the more effort they can spare from needless digestion of varied substrates, the more time and energy they have to devote to the pleasurable pursuit of procreation. The deciduous trees in their forest are alder, birch and oak. Those with a natural proclivity for alder digestion hie themselves off to a remote alderwoods, where they soon enough (say, in a thousand years) evolve as species A, a unique Alder-digesting species. Their oak loving kin do the same in a far-away oak grove, ending up with species O, an Oak-digesting fungus. Should all these species now get carried back to the original mixed woods and meet up with species B, all would live side by side: B eating Both coniferous and deciduous wood, C eating Coniferous only, D eating Deciduous only, A eating Alder only and O eating Oak only. They would be sympatric, each in its own niche, but would not hybridize, because at this stage speciation for each has gone so far that they no longer can mate with each other or their progenitor. A steady state ensues, until Janelle Smythe-Williams,

cabinetmaker to Her Majesty, discovers this great source of heritage oak. She fells all alder for easy access and chops down all the oak. After the last debris is gone, there is nothing for species A or O to eat. Together with the Hamadryas, they die out, because evolution is not quick enough to respond to such stochastic events. Species D, not being a specialist, was able to survive on a birch diet until new deciduous trees moved in. Lesson: specialization is more efficient, but riskier. Which is why, at the early age of three, AV made a conscious decision to become an omnivore. Our Gymnopus-dryophilus-like mushrooms separated into two intersterile, yet morphologically very similar groups. The most obvious difference between them is that the first is a woodland species, while the second has established a niche on barrens, bogs, moors and heaths— in other words, anywhere that there are no trees. Fittingly, the woodland group was the one whose DNA matched the European (and North American) G, dryophilus.1 If Dryads are tree nymphs and a species that grows in the forest is called dryophilic, then it would be appropriate to call our second group, which grows in regions without trees, a shunner of Dryads, or dryophobic. Much as the dryophilic species has seeming host associations with several trees while remaining the same species, so the dryophobic group also seems to have several host associations. For example, an association with Loiseleuria procumbens, the beautiful mountain azalea, has led to the description of a Gymnopus species that grows with this plant: G. loiseleurietorum. There are locations where group after group of dryophobic Gymnopus fruit among Salix uva-ursi, the dwarf bearberry willow. Elsewhere clumps of similar Gymnopus fruit among Dryas (mountain aven, also named for the nymphs), and on the barren Erin Mountain behind Barachois Pond Provincial Park this same mushroom fruits profusely around every ecologically dwarfed mountain alder (Alnus viridis ssp. crispa) bush lucky enough to find a toehold. Do these associations indicate obligatory relationships, i.e. different species? Unlikely. Narrowly specialized obligatory host relationships are very risky, and in this hostile environment such species are unlikely to survive. Fungi may seek out vascular plants for other reasons than a “host” relationship. For example, it may be desirable to be near alder to get the Nitrogen it produces. In most cases the reasons for association are even more general. Barrens have far less choice of good organic debris, so whichever larger perennial vascular plant is common in an area is likely to be selected as a good food source for a saprobe. An apparent association may occur between

Gymnopus dryophilus in a deciduous woods (top), on sole rotten birch log in a primarily coniferous woods (middle), and in a coniferous woods (bottom). Did Picasso get his inspiration from a broken, clubfooted, slug-eaten and angular dryad like the top photo? Middle photo: Maria Voitk

OMPHALINA 19

totally unrelated organisms because both require the same environmental conditions to survive. The barrenlands have very limited areas with sufficient soil to keep roots or mycelium alive during the cold season. In the growing season these places also hold the most moisture for the longest time. Both plant and fungus seek them out for the same reason. Likely, associations with plants for our dryophobic group are fortuitous— opportunistic rather than obligatory. This explanation fits best with a single dryophobic species of Gymnopus. It is important to understand what our findings tell us and what they do not. We have two groups of fungi, one which falls into a clade with G. dyrophilus, and one which falls into a clade with G. alpinus. There is a difference between belonging to a group that contains G. alpinus, and being G. alpinus. In the case of G. dryophilus, we are dealing with an established species, relatively well worked out and defined. All our collections were phylogenetically close and produced a good match with the known single species. The likelihood is high that ours is that same species, minor differences being due to geographic variation. In the case of G. alpinus, the species is not that well defined, and many similar species have been described, only some of which have been defined phylogenetically. Our dryophobic mushrooms are definitely in the group around G. alpinus, but also with four other species, all equally close. Our present results do not allow us declare with certainty which of these five names is “correct”. How should we identify and record our finds? It is for this situation that the qualifier s.l. (sensu lato = in the broad sense) has been developed. The opposite is s.s. (sensu stricto = in the strict sense). “Gymnopus alpinus s.s.” means the species defined as Gymnopus alpinus, and nothing else. “Gymnopus alpinus s.l.” means either G. alpinus or a species close to it. Pending new information, it would probably be reasonable for us to identify our woodland species as G. dryophilus, and our barrenland species as G. alpinus s.l. Further investigation may reveal this name to be close, but not exact, and our species may require reassignment in the future. The choice of G. alpinus was influenced by finding that it, as described by Antonin2, fell into the same clade, matching ours with a 92% similarity., and base pair divergence under 0.3%. Our dryophobic Gymnopus alpinus s.l. with a variety of “hosts”, top down: Loiseleurium procumbens, Salix uva-ursi, Empetrum hermaphroditum, Vaccinium uliginosum. In all likelihood, none of these are true hosts and the association is opportunistic, not obligatory. This is the same species, growing near whatever plant is available at the time and place, as a source of organic debris to satisfy its saprobic lifestyle.

20 OMPHALINA

References 1. 2.

Antonin V, Sedlák P, Tomsovsky M: Taxonomy and phylogeny of European Gymnopus subsection Levipedes (Basidiomycota, Omphalotaceae). Persoonia 31:179-187. 2013. Antonin V, Noordeloos ME: A monograph of marasmioid and collybioid fungi in Europe. IHW Verlag. 478 pp. 2010.

The species of Gymnopus in Newfoundland and Labrador Ron Petersen, Andrus Voitk

The review of Gymnopus species collected over a decade from Newfoundland and Labrador revealed seven species: Gymnopus dryophilus Gymnopus alpinus s.l. Gymnopus confluens Gymnopus eneficola Gymnopus alkalivirens Gymnopus subnudus Gymnopus sp. K22 (a single collection only, not identified, and not discussed further here) This list is neither exhaustive nor exclusive. All collected material could not be studied, so that the presence of additional species cannot be excluded. Further collecting may uncover additional species, so uncommon as to be encountered less than once a decade. The species identified as Gymnopus alpinus, and to a lesser extent G. dryophilus, may be reassigned with more study, and both may contain cryptic species, to be uncovered by future analysis. A similar situation exists around the species identified as G. alkalivirens, where further analysis may be able to resolve several closely related cryptic species, so that this limestone barrens species could be reassigned to a new epithet in the complex. And for the mycodetectives, the mysterious K22 still remains. We are indebted to Karen Hughes, who contributed sequences which facilitated morphological identifications. Her work has made this descriptive article possible, but its errors should not be laid at her feet.

OMPHALINA 21

Gymnopus dryophilus Cap 20-50 mm diameter, opaque, hygrophanous, slightly greasy-waxy feel domed, becoming plane in age, wide range of colour from very light (bottom photo & title banner) to dark (upper photo—note that camera lightens the cap more than in real life), in various shades of orange-yellow tan to brown. Gills close, free or almost so, white, cream, yellow or tan.

22 O OMPHALINA MPHALINA

Stem 3-6 x 30-60 mm (slightly longer than mature flat cap diameter), even or club shaped, radicating if in rotten wood, smooth, no ring, white to cream rhizoids at base, white, cream, yellow or tan, lighter than cap. Flesh white to cream, “mushroomy” smell, mild taste. Ecology in forest on duff, primarily hardwood, but occasionally on seemingly “pure” conifer duff; at times on rotten wood. May to September.

11

Gymnopus alpinus s.l. S.l. = sensu lato = in the broad sense, i.e. may encompass other similar species.

Cap 20-50 mm diameter, opaque, hygrophanous, slightly greasy-waxy feel domed, becoming plane in age, wide range of colour from pale beige to dark, in various shades of yellow-tan to brown. Gills close, free or almost so, white, cream, yellow or

tan. Stem 3-6 x 30-60 mm (slightly longer than mature flat cap diameter), even or club shaped, smooth, no ring, white to cream rhizoids at base, white, cream, yellow or tan, but lighter than cap. Flesh white to cream, “mushroomy� smell, mild taste. Ecology outside tree zone, either in bogs, moors, heaths or barrens with some organic matter; usually with some vascular plants. May to September.

OMPHALINA 23

Gymnopus confluens

Flesh white to cream, smell & taste unremarkable.

Cap 20-50 mm diameter, opaque, hygrophanous, dry, fibrillose (loupe), bell to dome shaped, becoming plane, tan, beige, fading to cream or whitish.

Ecology on leaf litter in coniferous or mixed woods. Known elsewhere to decompose coniferous debris as well. Only collected in September, although season likely longer.

Gills very close, free or almost so, cream to tan. Stem 3-6 x 30-90 mm (equal to or longer than twice mature flat cap diameter), even or slightly widened at base, entirely coverd with dense fuzzy white hairs, no ring, white, cream, or tan.

Differences from G. dryophilus and G. ocior: hairy stem, stem-cap proportion greater, colours consistently paler, dry cap. Photos this page: Roger Smith

Gymnopus eneficola Photo below. To give an idea of relative size, this photo scaled to match that of G. confluens, beside it, left. Full description and more photos on p 4. Differences from G. confluens: smaller size, greater stem/cap ratio, paler colour, wider gill spacing.

24

OOMPHALINA MPHALINA

Gymnopus subnudus

pruinose at the top, becoming hairy toward bottom, hairs white; stem tan to pinkish buff.

Cap 10-35 mm diameter, opaque, hygrophanous, Flesh cream, smell & taste unremarkable. dry, fibrillose (loupe), domed with inrolled margin, becoming wrinkled, eventually plane, cinnamon to dark Ecology saprobic on leaf litter. Only collected in coastal barrens of Labrador Straits in August, although brown, fading to pinkish tan, margin lighter. season likely longer. Gills distant to subdistant, free or almost so, whitish to Note: Only a single collection to date, morphological pink buff, edges lighter, finely sawtoothed (loupe). and DNA identification. Description largely based on Stem 3-4 x 20-70 mm, even, often flares at top, outside sources.

OOMPHALINA 25 MPHALINA

Gymnopus alkalivriens Cap 8-28 mm diameter, opaque, hygrophanous, may have slight greasy feel, domed, becoming plane in age, very dark to mid-brown. Gills close to subdistant, free or almost so, brown with lighter edge. Stem 1-3 x 30-80 mm, even or slight basal enlargement, smooth with brown downy hairs near base, no ring, hollow, brown, dark purplish brown to nearly black brown. Flesh beige, smell & taste unremarkable. Ecology saprobic on plant litter. Only found in Dryas patches on the limestone barrens of Watt’s Point in June, although season likely longer. Plentiful in the area at the time. Note: To date only known from one small area. Morphologic identification only; did not yield DNA for confirmation. Description augmented by outside sources. Photo: Maria Voitk

OMPHALINA 26 OMPHALINA

KEYS

Both the keys and descriptions work because the number of species is small in Newfoundland and Labrador. In regions with greater mycodiversity these same keys and descriptions may be misleading; microscopic examination and other studies may be needed for accurate identification. Should our species harbour additional species, not found so far, the keys may also become less reliable. However, given that this review is based on a decade of collection, the likelihood of additional species is so low, that these keys should prove accurate for field identification the vast majority of the time. Lighter background or gray print indicates less experience, less certainty.

GYMNOPUS—SPREAD SHEET KEY alkalivirens* subnudus* dryophilus alpinus eneficola confluens CAP diameter 20-50 20-50 6-36 20-50 8-38 10-35 texture greasy greasy dry, mealy dry, mealy dry to greasy silky, wrinkled colour light to dark light to dark light beige beige to dark brown brown to browns browns to white light tan cream, often pinkish GILLS spacing close close distant very close close to medium to medium distant colour white, white, whitish beige to tan brown beige to tan cream, cream, yellow, yellow, beige beige STEM height 30-60 30-60 12-75 30-90 30-80 20-70 texture smooth smooth hairy (white) hairy (white) smooth with hairy (white) brown hairs at bottom Stem/cap diam 1.0-1.5 1.0-1.5 1.8-2.2 1.7-2.0 1.5-2.4 1.7-2.0 HABITAT forest open areas forest forest open areas? open areas? SEASON May-Sept May-Sept Sept-Nov Sept (range June (range Aug (range unknown)* unknown)* unknown)*

Gymnopus—Dichotomous key 1. a) Resembles G. dryophilus** ………....…………........................ 2 b) Does not resemble G. dryophilus ………....…........................ 3 2. a) Grows in the forest ………………….............................................. b) Grows on bogs, barrens or heaths ……...................................... 3. a) Mature fruitbody cream to white (may be beige in youth) 4 b) Mature fruitbodies some shade of tan to brown …........….. 5 4. a) Gills close, gill edges smooth ……............................................... b) Gills distant, edges finely sawtoothed (loupe)........................... 5. a) Stem smooth (may be wooly at the very bottom) ................... b) Stem definitely hairy (upper part may be smooth with age)...

G. dryophilus G. alpinus s.l.

G. confluens G. eneficola G. alkalivirens G. subnudus

*insufficient experience with the species in NL; characters from outside sources may not be reliable for the province. **variously brown, slightly greasy cap, about 35 mm diameter; near-free to free, close, white, cream or yellow gills; smooth stem same length as cap diameter or little more.

OMPHALINA 27

the mail bag or why the passenger pigeons assigned to serve the lavish Corporate and Editorial offices of

Congratulations for a fine issue (O-V-2) on morels, particularly those of Newfoundland and Labrador. I especially enjoyed Kerry O’Donnell’s overview of morels and morel study in North America. As an outsider (from that green province in O’Donnell’s Figure 6), and for readers who might collect morels in parts of Canada beyond NL, a few comments: 1) Figure 6 encapsulates a huge amount of work by Kerry O’Donnell (et al. 2011), Michael Kuo (2005, 2008, Kuo et al. 2012) and many collaborators, but might give some the impression that the work is done. I don’t think any of the authors would say that themselves, but the next two points list a couple of examples. 2) George Barron, referring to the “old name” Armillaria mellea, in contrast to the “newer name” Armillariella mellea that was in vogue in the 1980s, said “Don’t worry about too many of these new names - we will be using the old ones again soon.” (He was right, at least as far as the generic name is concerned.) At that time we had just learned to use the name Morchella elata for what we had once identified as Morchella angusticeps. Now, the correct name for the “classic black morel” of eastern North America is again Morchella angusticeps. But wait—it is not reported from Canada in any of the publications I mentioned above, and it is not reported in the map in Figure 6 of your hallowed journal. What’s up? 3) Morchella septentrionalis, its nearly look-alike sister with slightly smaller fruiting bodies and spores, is known “from roughly 44 N latitude northward” (Kuo et al. 2012), but is also not reported from Canada, nor indicated in any of the Canadian provinces north of 44 N in Figure 6. Canadian morel-hunters—we have work to do! Greg Thorn

28 OMPHALINA

OMPHALINA get hernias

Comment by Kerry O’Donnell: Hopefully the OMPHALINA issue will stimulate more research on morels in Canada. Clearly Fig 6 shows how little is known. Like you, I was surprised that Morchella angusticeps and M. septentrionalis weren’t found, but I am 100% sure that they are present in Canada. Fortunately, types exist for most of the morels in NA and Europe. More work is needed to find out what is the real M. elata. Hopefully collections near where Fries collected it will resolve this issue. Maybe you can use your newsletter to stimulate people to collect morels throughout Canada and find someone [maybe a MS student] who is interested in identifying them molecularly.

Ed comment: There you have it, folks. It took the current workers over a decade to collect representative samples, and most came from the US. If we could get all mushroom clubs, all herbaria and all Mycology departments in Canada involved, collection of representative samples could be done in one season (say, spring, 2015). A great project for some Department to take on and follow up with the sequencing the same year. And, bingo!, we’d have a much more complete picture of what morels grow in Canada. If our insular province, famed for its low diversity, yielded two new species, who knows what unknown species lurk in the Canadian woods?

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