V
OMPHALINISSN 1925-1858
Vol. V, No 7 Newsletter of Jul. 30, 2014 OMPHALINA 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. V, No 7 OMPHALIN ISSN 1925-1858 Jul. 30, 2014
CONTENT
Editorial ...... 2
Foray Matters ...... 3
The granitic highlands of Gros Morne Michael Burzynski ...... 3
Arrhenia obscurata
Voitk, Voitk, Mann, Lebeuf...... 5
Arrhenia griseopallida Andrus & Maria Voitk ...... 22
Arrhenia peltigerina Andrus Voitk ...... 23
Arrhenia epichysium Andrus Voitk ...... 24
Partners ...... inside back cover
Notice of Foray 2014 ...... back cover
This issue and all previous issues available for download from the Foray Newfoundland & Labrador website
OMPHALINA If your club’s logo is an Arrhenia (a former We tack on short descriptions of three other little Omphalina), and its newsletter is called OMPHALINA, brown arrhenias that we believe can be identifi ed you should have some knowledge of the omphalinoid with reasonable certainty—at least some of the time. group of mushrooms. It is an understudied group, Apart from names, the lifestyles of this group are far fi lled with as much confusion as some of the from known, and worthy of serious enquiry. other groups we have met recently. This issue will introduce you to four small, brown arrhenias with cap and central stem. See you at the Foray! The lead article (as usual, after the Foray material), with a welcome cool cover picture during the hot andrus summer, celebrates the arrival of Arrhenia obscurata fi lling into the exalted pensioner status already enjoyed PS for some years by the fi rst 75% of the authors. The fi rst. species was fi rst described 65 years ago and this year Speaking of Foray, if you are thinking of swells the ranks of ever growing gray power. coming, please register now. Space is up and registrations close August 22 or when Imagine that as the author of a mushroom book the limit is reached, whichever comes you came across an A. obscurata. The species is so uncommonly encountered that this may be the only Do not leave it to the last minute! time you will see it. You take your picture and put it in your book. That one picture will guide every ERRATUM reader to the appearance of this species. Now peruse the pictures in this issue to see the amazingly many Thanks to the two readers, who corrected an error in the last editorial. We wrote: faces of A. obscurata. A reader of your book may come across ten members of this species, not one …we have known and collected T. terreum under the resembling your picture. The poor soul will struggle name T. virgatum in the past. The two have been shown and struggle, and be frustrated with her inability to to be genetically synonymous and T. terreum is the older identify her fi nds. One thing she will know for sure: name. none is an Arrhenia obscurata, because none looks Actually, the species that has been synonymized like your picture! with Tricholoma terreum is T. myomyces (as pointed out correctly in the article describing the latter). And Ah, but that is why you also add the invaluable we have not really “known” that species, having microscopic morphology to your description. We recorded it only once, in 2003. T. virgatum is a much all know that the microscope is the tool specifi cally commoner species in our province than T. terreum devised for mushroom identifi cation. Read on, and under either of its names. To balance off this error, see how much spore size and some of the other good here is a nice note from one of authors of the recent but small things helped your reader. Nordic Tricholoma book:
If you study a species closely, you will see that most What an amazing coverage of Tricholoma in just one have a breadth of morphologic appearance. One of newsletter! the problems of texts is to illustrate ALL members of Jacob Heilmann-Clausen the species with but one picture and one description. Ed comment: The secret is to get the right contributors. 2 OMPHALINA FForayoray matters…matters… 1. REGISTRATION FORM, INFORMATION, 2. MYCOBLITZ For those able to make it, join HOW TO GET THERE, etc. Please see us at Sir Richard Squires Memorial Provincial OMPHALINA V, No 4 (April issue), as well as other ark at 11:00 AM, Fri. Sep. 12, 2014. See April material stored on our website OMPHALINA for details.
THE GRANITIC HIGHLANDS OF GROS MORNE Michael Burzynski
Almost half of the park is on the highlands of thaw cycles. Rivers of ice fl owed down from the the Long Range Mountains. Low-lying areas are highland icecaps, carving existing river valleys covered in wetlands with patches of old-growth into the deep glacial gorges such as Western spruce-fi r forest. The most exposed parts of Brook, Bakers Brook, and Ten Mile Pond. The the highlands are bare outcrops of granite and highlands were the last area to be exposed as wide fi elds of ice-shattered granite and quartzite the last glaciers melted. boulders (felsenmeer). As the ice waned, seeds and spores blew in Over the two-million years of the Ice Age, from the mainland and started to re-vegetate the repeated glaciations removed all of the soil from bare rock of the Island. As conditions warmed the highlands. Exposed rock was polished by and more species arrived, the early arctic-alpine moving ice, or shattered by repeated freeze- species were pushed off the lowlands into
Late snowbed on Big Level, July 10, 2009 20 OOMPHALINAMPHALINA 3 restricted corners of harsh habitat—habitat where they could survive, but the more temperate, and faster growing, trees and shrubs could not. To this day, plants typical of the high arctic live on the exposed hilltops of Gros Morne, and in the late-melting snowbeds farther inland. The late snowbeds are a particularly unusual habitat. They form on north-facing slopes where snow builds up in winter but melts slowly in summer. Caribou favour these snowbeds when they are raising their young, since the cool air keeps large parasitic botfl ies and warble fl ies from parasitizing LLichen-coveredichen-covered ffelsenmeerelsenmeer oonn BBigig LLeveevel and weakening the calves. As the snowbeds gradually melt back during the summer, they provide constant moisture enriched with caribou droppings to specialized arctic plants that are adapted to growing, fl owering, and setting seed in only a few weeks. Most of these plants can survive nowhere else in the park. Winds are strong on the highlands; low clouds create fog-like conditions, and the temperature can be much lower than on the lowlands. If you venture up to look for plants and fungi, be prepared with a windbreaker or raincoat, water, sunscreen, and a fl eece. You will be rewarded with tremendous views of the park, and a visit to an ancient refuge of arctic plants and Rock ptarmigan on Gros Morne Mountain animals only a few hours hike from sea level.
Summit of Killdevil Hill, 2010
4 OMPHALINA 21 Arrhenia obscurata —the Phœnix of Mt. Ignoble
Andrus Voitk, Maria Voitk, Henry Mann, Renée Lebeuf
Mt. Ignoble (48°59’57”N, 57°45’04”W) is a barren mountaintop 277 m ASL near our home that we have visited it at least 6-10 times a year for over a decade. From the bare rock on top is a view onto the similarly bare rock of Easter Peaks (highest point on the horizon). Geological description by Geoff Thurlow: High grade metamorphic rock, probably originally a quartzofeldspathic conglomer- ate, compositionally like granite, with mostly quartz, feldspar and some mica and the like, arranged in a planar fabric (foliation) from strong metamorphism (heat and pressure) deep in the crust. Water seeps in along the foliation planes and freeze-thaw cycles break apart the rock, making a “soil” primarily consisting of ground up granite granules in feldspar-mica fl akes.
In 2011 we found some very small brown Weekly visits were made to the site from the omphalinoid mushrooms in this “soil”. Did the beginning of March (snow at that altitude lasts well mushrooms just move there recently? Unlikely. Once into May, but the barren mountaintop is blown bare noticed, we have been able to locate some on each by March) to the fi rst week of December, 2013 (when subsequent visit in season, suggesting that they have snow returned). Two weeks were not surveyed, when always been there, but it just took us over 10 years to we were away. Note was taken of the number of notice them. Why? Well, the mushrooms are obscure: mushrooms, their size, growth, habitat, duration, other dark and tiny, blending inconspicuously into their macroorganisms, weather and any other notable surroundings—surroundings that are unlikely places to event. At least one fruiting body was photographed look for mushrooms in the fi rst place. We set out to and at least one collected (if there) at each visit and a study them more closely. microscopic examination done.
OMPHALINA 5 The cap measured 3-14 mm. in diameter. It was varied from bowl-shaped to fl at, but almost always approximately the same as stem length. Its shape retained an inturned margin.
The margin was even, but became crenulate (sinuate radially ribbed with raised regions above the gills and or wavy) with time. At the same time the cap became lower areas between.
There navel varied from several mm in depth to a The cap was translucently striate (the darker radial shallow central depression, to bowl shape with no stripes over the gills show through) when moist. depression.
6 OMPHALINA Hygrophanous and translucent, the cap became fl ecked with brown pigment. opaque and lighter in colour when it dried, often
Cap texture was initially coarsely granular, becoming fi nely bumpy texture. The projections seemed to be minutely pruinose, and fi nally merely of a pebbly or end hyphae, not cystidia.
In colour it was a dark mushroom, almost black photo. The mushroom colour is darker in real life than brown, but varied to a mixture of light browns and shown on the photos. Narrow spot metering, fi xed tan. Radial lines over the gills were darkest, with lighter on the mushroom, causes the camera to compensate areas between. Colour lightened as it dried. Note for the darkness to bring out a wider range of colours presence of some red tones, especially the far right than seen with the naked eye.
OMPHALINA 7 The gills were distant, arcuate and moderately to than the cap or stem, with an edge that was usually deeply decurrent. Occasionally they forked near the somewhat darker. There was rarely more than one cap edge. Cross-veining was very rare, seen minimally level of lamellulae. in old specimens only. Gills were lighter in colour
The stem was 4-15 mm x 0.6-2 mm, minutely the cap. pruinose, becoming smooth with age, with a few Average stem length was equal to or only slightly whitish wisps of mycelia at its base. Old stems showed longer than cap diameter for most mature longitudinal fi brous striations. It was dark in colour, like mushrooms.
8 OMPHALINA The basidia (club-like reproductive cells on the gill attempts to focus at several depths, it is easy to surface), illustrated here from two fruiting bodies, overestimate the proportion of bisporigerous basidia. were about 30 x 6 μm and had a basal clamp Several basidia are shown, some with spores in various connection. In fact, clamp connections were prominent stages of development. Two views show full-sized throughout all tissues. Most basidia had four sterigmata spores for comparison. There were no cystidia (club- (prong-like projections or prongs, on which spores like larger and often sharp projections from tissues) are formed), with 20-25% bisporigerous (2-pronged). on the gill faces, edges, cap or stem. Because sometimes all prongs cannot be seen despite
The hyphae (long cells) of the pileipellis (pilei=of the granular surface. The lower pictures show some the pileus or cap, pellis=skin) run parallel to the cap hyphae with crosswise stripes—the incrusted brown surface, forming a cutis (skin). The upper pictures show pigment. Also note the clamp connections (bumps like interlaced, winding hyphae, roughly parallel to the swollen knee caps at cell junctions), abundant in all surface, with some ends extruding, probably creating tissues.
Spores of two mushrooms (upper: ammonia, Congo widely. Size for 24 mushrooms: 6.5-10 x 3.5-6 μm, with red, Meltzer’s; lower: Congo red & Meltzer’s). Spores an average Q (length divided by width, an indicator were hyaline with a prominent lateral apiculus, and of shape) 1.7 (range, 1.4-2.0). Spores from a fresh mature spores contained granules and vacuoles or sporeprint tended to be more uniform in shape, more droplets. Size range for individual mushrooms was elliptical than pip shaped, and tended to be in the narrow, but the range between mushrooms varied upper half of the expressed range. OMPHALINA 9 A. griseopallida A. obscurata A. umbratilis A. velutipes A. parvivelutina x A. obatra A. rigidipes
A. baeospora
A. rustica
The left image shows parallelograms for reported species is that over 50% of spores are over 10 spore size range of nine macroscopically similar little μm long. Not so on Mt. Ignoble—provided our brown arrhenias.1 The spore size range of A. obscurata identifi cation is correct. (yellow panel) is the most central and largest of 2. Only one mushroom had spores that covered the depicted species, overlapping all the others (4 most of the reported range for the species. totally, 2 over 70% and two 20-35%). This means 3. The range for most mushrooms was very narrow. that any specimen of any of these species could The ranges for several single collections differed have spores in the size range of A. obscurata, and the so much from each other that they could be vast majority will. Thus, spore size outside this range suspected to be different species. may exclude A. obscurata, but a match does not help 4. The range for the majority of individual collections distinguish it from several others. A few have other fi t entirely within reported ranges for other microscopic features that help identify them, but the species, and if a single collection, could be mistaken majority cannot be distinguished from each other for any of these. microscopically. 5. About 15 collections were required before it Let us say, for example, that you found a fruiting body became apparent that all clustered within the of Arrhenia parvivelutina, with spores closely scattered wide range of A. obscurata, and did not represent a around the middle of its reported range (the white variety of different species. X). That spore size is within the range of A. obscurata, With this much variation in spore size, is it possible A. umbratilis, A. velutipes and A. rigidipes. A little longer that our collections are, indeed, of several species? spores would bring in A . griseopallida and a little Theoretically, yes, but our combined many hundred thinner, A. obatra. Good luck! years of observing life on this earth argue strongly that The right image shows the spore size ranges for 23 of it is unlikely to fi nd another, let alone several, species our individual collections of the putative A. obscurata, elbowing each other for a chance to grow on and out with the same yellow panel of reported ranges in of bare rock during frost and snow. However, we are the background. A minimum of 10-30 spores were not too old to learn, so please do not hesitate to let measured each time. Note that: us know if you think we err. 1. Only 2 mushrooms had spores of or minimally Note: the parallelogram is a quick way to represent the above10 μm. This is signifi cant, in that many spore size range, but is not as accurate as a mathematically calculated or computer-generated scatter diagram, based authorities state that one way to recognize this on a large number of measurements.
10 OMPHALINA This view is from Easter Peaks, along the same axis as Kalmia and stunted Picea among Cladonia rangiferina on the title banner, but the opposite direction. And a and other reindeer lichens made up the part of the bit later in the year. The barren Mt. Ignoble is indicated mountaintop covered with thin soil, and on the slopes by the arrow, with Marble Mountain ski runs and the leading to the top there was a mixed boreal forest of Humber River behind it. Abies, Betula and Picea. People, dogs, snowshoe hare, The general habitat of this mushroom was impressive: squirrel, voles, various birds and moose and caribou bare rock! Not areas of turf nearby, or gouged out visited the area. The fi rst basidiolichen (Lichenomphalia depressions with soil and trees or other vascular umbellifera) was seen May 27, the fi rst saprobe (an plants, but bare rock. As the photo shows, at unidentifi ed Mycena) July 31, and the fi rst mycorrhizal 277 m ASL, Mt Ignoble is not above the normal mushroom (Inocybe lacera) Aug 5. On one occasion timberline. However, its top has suffered some we collected Microglossum atropurpureum in one of stochastic event removing the trees, and erosion has the cracks on bare rock. Examination of this crack removed the soil in places, to lay part of the rock showed it to have been fi lled with caribou droppings. bare. The bare rock was covered with a plethora of Easter Peaks is 2 Km due NE from Mt. Ignoble and lichenized ascomycetes. In more fertile cracks, moss 100 m higher in altitude. It has similar topography, grew, followed by heath plants such as Empetrum with bare areas of similar rock to that of Mt. Ignoble. hermaphroditum, E. eamsei and Vaccinia spp. Associated To test our suspicion that this mushroom actively with these were Lichenomphalia umbellifera and seeks out such habitat, we explored Easter Peaks. The Clavaria argillacea. Among them small conifers and same mushroom grew there, in the exactly the same birch gained a toehold, at least for a while. Around microhabitat. these we have collected Inocybe lacera (with birch) The forest between the two peaks was clearcut a few and small Lactarius hibbardae (with conifer), and years ago by Corner Brook Pulp and Paper Company. occasional unidentifi ed Mycena spp. Heath with
OMPHALINA 11 The most striking thing about this mushroom was yet accumulated. These places had no evident organic not what it looks like, but the substrate on which it nutrients, but water could be assumed to be present chose to fruit. In this choice it was very consistent. in the thin space between opposing foliation plate Its favourite place was between cracks of foliation surfaces due to capillary action. Here the mushroom plates that have undergone minimal separation, a stood out as the only large macroorganism. place where “soil” from crumbling of granite had not
An alternate habitat was pioneer soil fi lling small not contain enough organic matter to sustain a large crevasses in the rock face. The soil consisted of visible saprobe. In these furrows a small number of lichens grains of granite and quartz, mixed with fi ner feldspar- could be seen, and very small beginnings of moss. A mica silt. This gravel-sand contained some organic botrydinal mat was always found. Mushrooms were matter. Because our mushroom was the largest not found in older furrows overgrown with low moss, multicellular organism there, the soil probably did or those with established vascular plants. 12 OMPHALINA Botrydinal mat covering the “soil” around the Strands to the right in upper picture look like a leafy mushrooms. Not always so prominent, but with liverwort, and in the lower some moss is evident. See magnifi cation algal globules could always be found. next page for magnifi ed views of these. OMPHALINA 13 Although there were no large vascular plants microscopically small buds and branches were not inhabiting the furrows where these mushrooms grew, unusual. Small moss buds and a few sprigs were not there were other organisms containing chloroplasts unusual, if very carefully sought. However, of interest is (small organelles with chlorophyll, allowing the that arrhenias were not found in furrows where there organism to use the sun’s energy to convert was more mature moss, or moss that had covered the
atmospheric CO2 into sugars—photosynthesis). The soil. Algal globules were always found on soil around photo shows enlarged details from the photos on the mushrooms, although in certain sites they were sparse previous page. The organisms, left to right, are a leafy and not obvious. Note on this picture the network of liverwort, beginning small mosses, and algae in globules strands going to and over the algal globules. We may (Botrydina). guess, but have not confi rmed them to be mycelial or Liverworts of this size were not commonly seen, but to be directly connected to the algal globules.
The soil from one collection was placed in a covered wearing what looks like a necklace of blue-green algae, moist glass container on a cool windowsill to and some tiny cup-forming lichens, likely Cladonia encourage growth, and examined with a dissecting species. On the right is a mat of algal globules, the microscope after 2 weeks. Upper two left photos commonest visible green organism at the time. show a leafy liverwort and young moss beginning to Photographed at 400 x magnifi cation; same scale kept sprout. Below from the left are one of three peacefully for all pictures in this composite. grazing purple hypogasturids (more can be read about them in the latest Osprey2), some early lichens, one
14 OMPHALINA Photomicrographs (original magnifi cation 400 x) (small green granules) in the cells of these plants. showed again the liverworts (four photos on the We consistently found a few long hyaline (clear and right) and moss (two on the left). Note the different uncoloured) cells, presumed to be hyphae, in the cellular structure. Note also the visible chloroplasts vicinity of liverworts, but none around moss.
The algal globules consisted of spherical attached to empty cell walls, and chloroplasts are conglomerates of algal cell, surrounded by fl at hyaline released. cells. Note the profusion of long hyaline strands These globules are similar to the botrydinal granules traveling to, from and around the globules, in intimate Oberwinkler described for some basidiolichens.3 Algae contact with them, and seemingly extending from are contained within a globular sac of hyphae, with the fl at cells that make up the walls of the globules. hyphal cells also between the algae. As algae multiply, Arrows point to what seem to be clamp connections, they become more compressed. The globules enlarge, two enlarged in the left upper panel. but seemingly not at the same speed. This is where the As opposed to the organized but free cells of moss exchange with the photobiont takes place: water and and liverwort, the algal cells seem compressed and minerals for sugars. The connecting hyphae transport chloroplasts cannot be discerned within each alga. the exchange materials in the required direction. Not However, if the preparation is compressed to crush unlike ectomycorrhizal exchange, except that the the cells (lower left photo), the hyaline strands are photobiont is noticeably smaller! OMPHALINA 15 RRoRockockc aaroundroounu d ththehe bbabaseasese ooff thtthehe stsstemtemem ooff ththee mushroommumushshrrooomo iinn thtthehe iniinsert.nses rtrt. LoLLowerowewer phpphotohototo sasameameme aass upper,upupppeer,r, withwitith yellowyeyellllowow lineslinineses andandnd greengrereenn ddotsotts drdrawnrawwn inin toto hhelpele p lollocateocacatete pputativeuutatativve mymmycelialycecelilialal tthreadhrhreaead anandnd alaalgalgag l globulesglg obo ululeses iinn upuupperpppeer phpphoto.otto.o IIff thtthesehesese araarere hhyhyphae,yphphaeae, ththishis isis a hhigh-riskigigh--ririskk wwayay ttoo llilive.vee. OnO bbarearare rorrock,occkk, thttheyheyey aarerree exposedexexpoposes d toto ssunuunn aandndn wwind,inndd,, eevervveer att rriskisisk fofforr drddryingryiyingng outoout anaandnd ththerebyheerrebeby cuccuttinguttttining ofoofff tththee ssosourceoururccee ooff fofoodoodod ttoo thetthhe fuffungus,unggusus. AnAand,nd,d , probably,prorobababblly,y thethehe sourcesouourcrce ofofof waterwataterr andanndd mineralsmineneraalsls ffororor tthehehe aalgae.lglgae. PePPerhapserhhapaps thtthishis iiss wwhwhyhy frfruitingruiuitit ngng bodiesboodieses oonn rorrockckk ttendedeendeded tot llastaasst hahhalfalff aass lolongonngg aass ththosehoso e ono ““soil”.sos il”.”
16O OMPHALINAMPHALINA Oct 28-Nov 3 July 22-28 Dec 2-8 May 6-12 Foray week In Norway
The bars indicate the number of new fruiting bodies Light blue bars indicate frost, at least at night, during found each week. Snow persisted in the surrounding that week. The white bars indicate snowfall in the woods until mid May, but the mountaintop was blown week, even if subsequently melted. At almost 300 m bare by March. The fi rst three months with no fruiting ASL, temperatures are colder than at ground level, bodies are not shown on the graph. The fi rst Arrhenia about 30 m ASL. Week 27 had 5 consecutive days of was collected July 24. sub-zero weather with no snow cover and week 28 From the fi rst fruiting body on July 24, at least one had three. mushroom was found every week, ranging as high Note the apparent increase in fruiting with the arrival as 11 (average 4), until week 26 (cover photo). Most of cold and snow, which ceased abruptly with the grew singly, although up to 4 of varying size and onset of persistent cold. maturity could be found within the same small crack.
Photos one week apart. In temperate weather size, most mushrooms dried by the third week. If mushrooms on soil lived for over three weeks. left undisturbed, they were the same a further week Some smaller mushrooms did not survive to the later. We did not see dried mushrooms revive in wet following week, but those that had managed to weather, but frozen ones seemed to do so. Compared grow over 4 cm cap diameter usually survived. to those on “soil,” fruiting bodies growing on “bare Growth varied (presumably related to not drying) rock” rarely survived longer than one week. from 0 to 60%. Even if they had not attained full
OMPHALINA 17 Mushroom 13.11.07.av01, in one of the more species without snow. The ground was frozen and all small diverse furrows, soil depth 9 mm. The microscopic organisms growing on it. The left photo shows the pictures of soil organisms come from this furrow, as same mushroom at that time: frozen and shrivelled, do the algal granules submitted for DNA analysis. its cap shrunken to under 2 mm in diameter. This A seeming increased fruiting late in the season was followed by two days of thaw and light drizzle. suggests an ability to manage cold. Transient periods 48 hrs later the same mushroom (right photo) had of night frost, with above zero temperatures in the recovered and was of normal appearance, cap again day, began on week 23. The above mushroom was of 4 mm diameter. It produced a sporeprint, suggesting fi rst seen on week 25, when its cap diameter was that sporulation was functional. 3 mm. On week 26, snow fell for several days, then This fruiting body grew during night frosts and melted. This mushroom looked healthy and its cap snowfall, shrivelled up in response to several days of had increased to 4 mm in diameter. Week 27 had consecutive unrelenting frost, but with the appearance fi ve consecutive days of sub-zero temperature, of thaw and moisture, seemed to recover fully. Several days of frost without snow halted further fruiting in 2013. In 2012 snow fell with the fi rst frost, providing cover during cold and moisture during thaws. Three fruiting bodies are shown on the right, recovered during thaws, November 11 and 17. This is the only time we have seen the cap straighten out so that even the inrolled margin fl ares out. They sporulated well, suggesting they are able to withstand short freeze-thaw cycles. The mushroom on the left is reminiscent of a Cuphpohyllus. Maybe cryotherapy brings out family traits, because phylogenetically Arrhenia arises from a sister branch to Cuphophyllus.
18 OMPHALINA Discussion description of Arrhenia obscurata. However, Elborne These have been our observations over three years, did not mention—let alone stress—saxicoly for this brought into focus by a concentrated season of more mushroom, while describing two others specifi cally formal observation in 2013. It was a joy to pursue as denizens of pioneer soil. (Neither of these other and record the appearance and behaviour of these two fi t with the description of our species.) Arrhenia beautiful mushrooms. However, that enjoyment pales obscurata was described as growing “between low in comparison to the pleasure of the speculation these moss or on naked sandy to loamy soil from lowland to observations permitted. To compensate for over a arctic, spring and autumn.” This is presumably based on dozen pages of boring pictures, we shall discuss our Kuyper’s description, “common in The Netherlands”, speculations in three pages of solid but exciting text fruiting “between mosses on dry to moist, sandy to 4 unsullied by pictures, under the headings, Identifi cation, loamy soil” April-June and Sept.-Feb. Range, Season and Lifestyle. Even more uncharacteristic (to us) is the description Identifi cation by Reid, fi rst to give the species a valid taxonomic Once we exclude arrhenias with a cap over 2 cm description, who stated that his selected type collection came from a “burnt patch of soil amongst wide, those that grow on wood or Sphagnum, those 5 with a primitive lateral or no stem, the colourful Sphagnum”, i.e. an area with a goodly supply of ones and the non-translucent opaque ones, those carbonaceous organic matter able to support with cystidia, those with consistent bisporigerous other multicellular organisms. Kühner, who fi rst basidia and characteristic spore shapes, and those described this species (but illegitimately), said that the mushroom is found on bare ground or among moss, without clamps (Lichenomphalia), the macroscopic and 6 microscopic descriptions for about half a dozen little sometimes on rock (parfois sur les rochers). He did brown Arrhenia species are virtually interchangeable. not elaborate, so depending on how one interprets If you go on an excursion through the mountains and those impassive words, this can be said to fi t with fi nd a single sample of our mushroom, any pretense at our observations, at least at times. Descriptions its confi dent identifi cation based on macroscopic and by authorities like Lange, Clémençon, Lamoure microscopic morphology seems risible. Look over all and others, use the same understated and laconic the pictures again to get an idea of the wide range of language without stressing our mushroom’s most variation in the macroscopic appearance of any part startling microhabitat. Watling virtually dismissed it in Greenland with the terse, “appears to be a common of it you wish to consider. The section on microscopy 7 made the same point for its microscopic appearance. rather northerly agaric on base-poor soils.” If we cannot tell them apart by looks, we need some A little more comfort for our identifi cation is provided other character for identifi cation. by Gminder, who states that Arrhenia obscurata is one of a group of pioneer mushrooms which grows on The most arresting aspect of this beautiful mushroom virgin soil (Rohböden), but adds that the species also is its rigid adherence to a very inhospitable can be found in nourishment-poor rather sandy soil.8 microhabitat. Like many beauties, it is a master of the To a soul, a very phlegmatic group, these mycologists: grand entrance: after all, it is very diffi cult to be more it is diffi cult to correlate words like “sandy and loamy dramatic than to spring forth from solid rock! Surely soil” with the dramatic setting we observed. this attention-getting behaviour would be a strong Apart from Gminder, the only description that identifying feature? Yet, one of the things that made 9 10 us hesitant about the identifi cation was the lack of struck a chord was by Peck. Bigelow believes that emphasis authorities gave to what to us was the most Peck’s Agaricus (Omphalia) montanus may be the fi rst noteworthy aspect of this organism. The experts were description of this species. As with most of Peck’s underwhelmed and blasé. laconic descriptions, at fi rst glance it seems hopeless to guess what organism he described. If we include The most current description of the omphalinoid the title, treat single-letter abbreviations and numerals mushrooms can be found in Steen Elborne’s as full words, and each side of a hyphenated word treatment of the group in the 2012 edition of as separate, then Peck accords this new species a Funga Nordica.1 Both macro- and microscopic total of 37 words! However, for us only eight were descriptions of our fi nd fi t best with Elborne’s enough: “Thin soil covering rocks. Summit of Mt. OMPHALINA 19 Marcy.” These eight words, more than any number in have seen in the same microhabitat in 14 years, was most other treatments of what is purported to be facilitated by a special situation: a caribou dropped this same entity, convinced us that they described some concentrated heavenly organic manna into a the organism we observed. Peck’s name for it was small furrow like a gift from the gods. illegitimate, because Agaricus montanus had been If ours were a mycorrhizal mushroom, associated with used previously for the beautiful mushroom we now a tree, as are the Inocybe and Lactarius from the same know as Psilocybe montana. Too bad, because from our area, then the tree’s cycle determines when enough limited experience, montanus is more appropriate to food can be given the fungus to produce fruiting this organism than to the Psilocybe species. bodies. However, the absence of obvious mycorrhizal Back to identifi cation—if this is a species limited to hosts suggests that ours is not mycorrhizal. A barren rock in a arctoalpine setting, its daramatic basidiolichen might fruit all season long. The fi rst lifestyle should serve to identify it, provided the Lichenomphalia umbellifera fruitbody, a well-known algal other fi ndings fi t. If this species is ubiquitous, and may associate, was seen on Mt. Ignoble on May 27, just a be found on sand dunes, loam, burns in Sphagnum, few weeks after the snow disappeared. Why did our and many other places, usually in moss, then the Arrhenia obscurata not fruit until the end of July? identifi cation becomes diffi cult to hopeless. Questions: The answer is probably related to its choice of 1. Is it possible that several species are included in microhabitat. Whereas Lichenomphalia, Inocybe and these descriptions? (Our guess is yes.) Lactarius live in turf, where their mycelium can 2. Is it possible that it behaves differently on the probably overwinter, it is quite likely that our fungus mountaintop than in the valley? (Our guess is no.) lives so exposed and in such shallow and superfi cial cracks and crevasses, that it freezes completely in Range the winter, killing the mycelium. There is no soil in Is it possible that the same species fruits not only its preferred cracks between foliation plates. Where in arcto-alpine regions, but also in lowlands? Most fruiting bodies appeared on crumbled granite “soil”, its organisms tend to specialize for one major climatic measured depth was seldom over 1 cm. The fungus setting and tend to be intolerant of major differences. probably reappears as the result of new mycelial Therefore, the possibility of the taxon containing more growth from sporulation into its substrate of choice. than one species is real. On the other hand, perhaps Once things thaw out and warm up, it takes several the species is unable to compete in a biodiverse months for spores to develop into mycelia capable of environment, and its only requirement is pioneer soil, producing fruiting bodies. The ability to survive early devoid of suffi cient nutritional content to support frost and ephemeral snow in the fall (in fact producing competing species. In that case it is conceivable more sporocarps at that time) may be part of its that it can thrive in submarine Dutch littoral sand strategy to drop a lot of spores with the falling and bars equally well as on arcto-alpine rocks. But Reid’s melting snow right into the same areas that it likes, to carbonaceous Sphagnum? be ready to grow the next year. Season Drought may also contribute to the late onset of fruiting. We have 30% more rain and more cloudy Some observers report two fruitings during the days in Aug-Oct than May-Jul. In an unprotected year. Climate makes this unlikely in most arctoalpine setting exposed to constant sun and wind, this may be situations. But if it really also lives in regions with little suffi cient to prevent viable fruiting bodies in the earlier, or inconsistent frost and snow, like The Netherlands sunnier and drier quarter. where double fruiting was reported, this is theoretically possible. Lifestyle We did not observe double fruitings. But why How does Arrhenia obscurata produce such substantial did it not fruit before the end of July? If it were a beauty from its self-imposed austerity? As mentioned, saprobe, presumably it could fruit any time that the the life of a traditional saprobe or mycorrhiza are temperature and moisture were right—but there unlikely. Among potential smaller partners we found is not enough organic matter to sustain a saprobe. moss, liverwort and alga. Many arrhenias are reputed Microglossum atropurpureum, the only saprobe we to be moss associates. Two observations argue against
20 OMPHALINA our mushroom’s being one of those. If it were a moss the fi rst mushroom—an omphalinoid very similar to partner, then one might expect to fi nd it preferentially ours—to appear on Surtsey, an island off the coast among moss. Our mushroom preferred bare “soil” of Iceland that arose from the sea due to volcanic and avoided furrows where low moss had established activity. The island is a protected nature reserve, set itself. Secondly, the microscope did not reveal any aside exclusively to document the successive stages hyphae associated with moss. of colonization of pioneer soil. The story of that mushroom seems so parallel to our observations— Liverworts might be a little more likely, because at even to a later discovery of green granules at its least we saw a few hyphae near them. However, these base—that we suspect that the fi rst mushroom were very few in number, and macroscopic liverworts identifi ed on Surtsey must be the same species. were relatively uncommon around these mushrooms. Although we have studied our mushroom enough Several reasons argue that this mushroom might be a to almost fi ll this journal issue, we actually know very closet basidiolichen, i.e. an undeclared algal partner: little about it. Even its identity is not certain. We can 1. With their bare soil and botrydinal globules these speculate, advance our theories with chest thumping mushrooms reminded us of Multiclavula vernalis, and brave posturing, but really, we have more another basidiolichen denizen of pioneer soil. questions than answers. Greg Thorn made a brave 2. Based on what we have seen, algal globules were attempt to extract DNA from the hyphae around the the one consistent potential partner around these algal granules, but unfortunately these were reluctant mushrooms—even on bare rock! to reveal their secrets. 3. Macroscopic connecting threads could be seen Stay tuned as we learn more of this saxifragous between mushrooms and algae. Phoenix, arising anew each summer from cracks in the 4. Microscopically the algae were all inside hyphal rock of Mt. Ignoble. containers, connected by copious hyphae. 5. No other basidiolichen was seen in the vicinity References to account for these mycelial algae farms. 1. Elborne SA: Arrhenia. In Knudsen H, Vesterholt J, eds.: Funga (Lichenomphalia umbellifera grew on the mountain, Nordica. Nordsvamp, København. 2012. but not in these thin cracks and furrows.) 2. Larson D: Springtails on mushrooms. The Osprey, 45(1):19-21. 2014. 6. Sherlock Holmes states that if you discard 3. Oberwinkler F: Basidiolichens, in Hock B, ed: Fungal explanations that do not fi t, what is left—no Associations, 2nd edition, The Mycota IX, Springer Verlag, matter how unlikely—is the solution: here, only Berlin. 2012. algae are left. 4. Kuyper TW: Omphalina. In Bas C, Kuyper TW, Noordeloos ME, Vellinga EC, eds: Flora Agarica Neerlandica, vol 3:78-88. 7. Throughout their evolution, fungi have turned A. A. Balkema, Rotterdam. 1995. to the lichenized lifestyle whenever it suited 5. Reid DA: New or interesting records of British them, with no respect to any classifi cation we Hymenomycetes II. Transactions of the British Mycological may impose upon them, so when it comes to Society 41:419-445. 1958. omphalinoids, surely Lichenomphalia does not hold 6. Kühner W: Omphalia obscurata spec. nov. Annales de l’Université de Lyon, Ser. 3, Sci., Sect. C (Sci. nat.) 6:130-131. the patent to the use of algae? 1949. 8. The hyphae going to and from the granules, 7. Watling R: Larger cold-climate fungi. Sydowia 26:308-325. possibly even some surrounding them, seem to 1983. have clamp connections; better magnifi cation and 8. Gminder A: Omphalina. In Kreiglsteiner GJ, ed.: Die Grosspilze resolution is required to confi rm this with absolute Baden-Württebergs, Band 3, Ständerpilze: Blätterpilze I:476-488. Verlag Eugen Ulmer, Stuttgart. 2003. certainty. 9. Peck CH: Report of the New Your State botanist for 1873. 9. And the most overwhelming reason: it goes Annual Report of the New York State Museum of Natural against conventional wisdom. History, 27: 73-116. 1875. 10. Bigelow HE: North American species of Clitocybe. Part II. One of us was privileged to pen a few words Beihefte zur Nova Hedwigia. Heft 81. J. Cramer, Berlin. 1985. on basidiolichens for an upcoming lichen issue of 11. Voitk A: The basidiolichens of Surtsey and other places. FUNGI.11 That article also makes a comparison with FUNGI, 7(2-3):59-62. 2014.
OMPHALINA 21 AArrheniarrhenia ggriseopallidariseopallida Andrus & Maria Voitk As mentioned when discussing the putative Arrhenia these had two, and the spores were big, 11.6-14.5 x obscurata, identifi cation of small, brown arrhenias is 6.7-7.7 μm. Two-spored basidia and big spores easily diffi cult. However, a few have distinctive characters, not keyed it out to Arrhenia griseopallida. However, before shared by others, which allows them to be separated you jubilate that fi nally you have a reliable character, from the group. One such species is Arrhenia you should be aware that four-spored forms have also griseopallida. Overall, it is a larger mushroom (cap been reported. Of course. diameter 3-25 mm, stem height 10-30 mm), whose The mushrooms illustrated were collected July 20, stem length is nearly twice the cap diameter. The cap, 2011, near a gravelly path to the lower or steep as illustrated in the title banner, is distinctly scaly, not section of the Main River. The substrate was perhaps a quite as scaly as Arrhenia sphagnicola, our logo, but bit more like disturbed soil than pioneer soil, and did reminiscent of it. The colour is darker and drabber, have more moss, with some grass and other vascular with gray rather than reddish tones. plants as well, and even some small seedling conifers Should any doubt remain, its distinct identifi cation not too far away, only a few metres from the forest character is microscopic. Whereas most of the basidia edge. That said, the soil was not rich, and one could of the other two species had four spores, all basidia of make a case for newly “created” pioneer soil.
22 OMPHALINA Andrus Voitk AArrheniarrhenia ppeltigerinaeltigerina
Photo: Roger Smith
Omphalinoid fungi live in areas of poor soil. From this directly or indirectly, is there some other mutualistic one might suspect that either they have developed relationship, or is it a saprobe specialized to dead thalli special coping mechanisms to extract nutrition form of Peltigera? this substrate, or they have developed partnerships, In size it is intermediate between A. obscurata and A. whether mutualistic or parasitic, with other organisms griseopallida: cap diameter 4-20 mm, stem length 8-30 living in these habitats. Some (Lichenomphalia) have mm; the stem again is usually twice cap width. The cap known algal associations, and other may as well. Some is smooth with a minute pebbly texture, without burr- have associations with liverworts or mosses, some like scales, like A. griseopallida. It is more translucent with lichens. and the colour is lighter than the latter. Among the latter is Arrhenia peltigerina. As the name The illustrated specimen was collected from Change suggests, it grows with the lichen Peltigera, which Island’s Squidjigger trail on September 5, 2013. The allows for relatively secure identifi cation among this lichen was identifi ed by Michele Piercy-Normore as group of similar, small, brown mushrooms. The exact P. degenii. Whether A. peltigerina associates with all nature of this relationship is unclear: is it a parasite Peltigera spp., or only some, is unknown. on the lichenized ascomycete, does it use its algae
Photo: Michael Burzynski OMPHALINA 23 AArrheniarrhenia eepichysiumpichysium Andrus Voitk
Arrhenia epichysium (above) is uncommon in NL: the mushroom illustrated is the only one I have seen in over ten years of collecting. For a little brown mushroom in a group of little brown mushrooms, it is easy to recognize because of its substrate—the only Arrhenia to grow on rotted wood. The odds of a misidentifi cation of an omphalinoid on this substrate is extremely slim. Lichenomphalia umbellifera (left) may be found on rotten wood, but it has a more crenulate or scalloped cap edge, wider gill spacing, ranges from white to yellowish tan, and if the log is bare of moss, the granular algal mat is readily seen. 24 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 Model Forest of Newfoundland and Labrador Memorial University of Newfoundland St. John’s Campus Grenfell Campus Tuckamore Lodge Quidi Vidi Brewing Company Rodrigues Winery Shorefast Foundation
OMPHALINA 2014 2014 2014 2014 2014 e 2014 2014 2014d ca 2014 2014de 2014 2014d decade 2014 2014on 2014 ec 2014 ssecond 2014 2014 he TThe GROS MORNE NATIONAL PARK A UNESCO WORLD HERITAGE SITE Headquarters: Killdevil Camp, Lomond, NL September 12-14, 2014
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