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Phaeomarasmius siquieri (Agaricoid clade, Tubariaceae), a new mediterranean
resupinate species found in Formentera (Balearic Islands, Spain)
Article · January 2011
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Micol. Veget. Medit., 26 (1): 29-36. 2011
Manoscritto ricevuto il 18.05.2011
Accettato per la pubblicazione il 07.06.2011
PHAEOMARASMIUS SIQUIERII
(AGARICOID CLADE, TUBARIACEAE),
A NEW MEDITERRANEAN RESUPINATE SPECIES FOUND
IN FORMENTERA (BALEARIC ISLANDS, SPAIN)
Joan Carles Salom
Conselleria de Medi Ambient i Mobilitat,- Carrer Gremi Corredors, 10, 07009
Palma de Mallorca (Mallorca, Illes Balears).
E-mail: joancarles.salom@gmail.com
Fernando Esteve-Raventós
Departamento de Biologia Vegetal (Botánica), Universidad de Alcalá, 28871
Alcalá de Henares (Madrid, Spain). E-mail: fernando.esteve@uah.es
Salom J. C. & Esteve-Raventós F., 2011: Phaeomarasmius siquierii (Agaricoid clade,
Tubariaceae), una nuova specie resupinata mediterranea trovata a Formentera (Isole
Baleari, Spagna).
Key words: Basidiomycota, agaricoid clade, Agaricales s.l., taxonomy, ecology.
Abstract: Phaeomarasmius siquierii is described and illustrated as a new species, based
on morphological and ecological characters. It is characterized by its peculiar small
resupinate habit, long, thick-walled, brown cystidia and habitat on Juniperus phoenicea
ritidomes. It has been found several times at one locality in Formentera, Balearic Islands.
The new taxon honours Josep Ll. Siquier for his long dedication to the study of the
macromycetes in the Balearic Islands.
Riassunto: Phaeomarasmius siquierii vienne descritta ed illustrata come nuova specie.
sulla base di caratteri morfologici ed ecologici. Il nuovo taxon, ritrovato varie volte in una
località di Formentera (Isole Baleari), è caratterizzato dai piccoli basidiomi resupinati, i
cistidi bruni a parete spessa e l’habitat su ritidoma di Juniperus phoenicea e viene dedicato a Josep Ll. Siquier in riconoscimento della sua lunga dedizione allo studio dei macromiceti delle Isole Baleari.
INTRODUCTION
Some weeks after heavy rainfall, unusual in Formentera, we visited the Island on 9
December 2008 to enlarge its mycofloristic catalogue. Among a large diversity of fungal
species, some minute basidiomata attracted our attention, especially by their habitat,
because growing on Juniperus phoenicea L. (Cupressaceae) soaked ritidomes, a mediterranean tree that is called “savina” in the territory. Though very tiny, we could
30
Salom J. C. & Esteve-Raventós F.
Fig. 1 - Phaeomarasmius siquierii Salom & Esteve-Rav. (JCS-816 B).
Fig. 2 - Phaeomarasmius siquierii Salom & Esteve-Rav. (JCS-816 B).
Phaeomarasmius siquierii
31
Fig. 3 - Phaeomarasmius siquierii Salom & Esteve-Rav.: A) Basidiospores (JCS-816 B),
B) Basidia (JCS-1067 B- Holotypus), C) Cheilocystidia (JCS-816 B), D)
Cheilocystidia (JCS-1067 B- Holotypus), E) Pileipellis (JCS-1067 B- Holotypus),
F) Pileipellis (JCS-1067B- Holotypus), G) Hymenophoral trama (JCS-1067 BHolotypus), H) Clamp connections (JCS-1067 B- Holotypus).
32
Salom J. C. & Esteve-Raventós F.
collect dozens of fruitbodies at their peak of fructification period. In this particular habitat we could collect several interesting and very small species like Patellaria atrata
(Hedw.) Fr. (= Lecanidion atratum (Hedw.) Endl), Crepidotus pallidus (Berk. & Broome)
G. Petersen, H. Kundsen & Seberg (= Pellidiscus pallidus (Berk. & Broome) Donk),
Lachnella sp., Clitopilus hobsonii (Berk. & Broome) P.D. Orton and Mycenella margaritifera (Maire) Maas Geest., the last one constituting the first record for the Balearic Islands
(SIQUIER et al., 2009). In this contribution we propose and describe a new species of the
genus Phaeomarasmius Scherff., mainly characterized by the peculiar (“crepidotoid”)
aspect of the basidiomata, the long and sinuose cylindrical cystidia and the particular habitat on “savina”. A comparison with both the well-known European species of this genus,
e.g. P. erinaceus (Fr.) Scherff. ex Romagn. and P. rimulincola (Rabenh.) P.D. Orton is
made, and the possible relationship of some representatives of reduced or “cyphelloid”
genera, such as Phaeosolenia Speg. and Chromocyphella De Toni & Levi, with the new
taxon is discussed.
MATERIAL AND METHODS
All the material collected is kept in the personal herbarium of one of us (JCS), and
duplicates have been deposited at AH (Herbarium of University of Alcalá, Madrid). The
microscopical description has been made on the basis of the study of fresh material, using
an Olympus BX-51 microscope, observing the samples in distilled water, Congo Red and
5% ammonia solution, 5% and Melzer’s reagent. Microphotographs were taken with a digital compact camera Olympus C-7070. References to color codes follow MUNSELL® (1994).
RESULTS
Phaeomarasmius siquierii Salom & Esteve-Rav. spec. nov.
Basidiomata minuta, resupinata, cupulata. Pileus 1.5-3 mm latus, semiorbicularis, convexus, siccus, furfuraceus vel velutino-sericeus, brunneo-tabacinus vel brunneo-argillaceus,
margine saepe undulato, non striato. Lamellae latae (L= 3-7), pallide brunneae, albofimbriatae. Stipes nullus. Caro tenuis, odore inconspicuo. Basidiosporae 9-12(-13) x 6-7,5(8) µm, (sub)ovatae vel late ellipsoideae, interdum leviter angulateae, laeves, crassitunicatae, ochraceae sub lente, guttula oleosa praeditae, cum poro germinativo minuto (0,50,7 µm). Basidia tetraspora, clavata, sterigmatibus longis usque ad 4-7 µm.
Cheilocystidia conspicuae, brunnea, elongata, 40-110 x 3-6 µm, filiformia, subcylindracea, anguste sublageniformia, flexuosa, submoniliformia vel substrangulata.
Pleurocystidia nulla. Pileipellis ex cute hyphis cilindraceis, haud gelatinis, elementis terminalibus in subtrichodermatae sat aerifero dispositis, pigmento parietalibus brunneo
constituta. Hyphae fibulatae. Ad lignum vivum in silvis Junipero phoenicii.Holotypus:
Hispania, Illes Balears, Formentera, Es Ram-torrent dels Arbocers, 8- XII-2010- herb.
JCS-1067B. Isotypus in AH 40201.
Basidiomata (Figs. 1, 2) gregarious, resupinate, cupulate or shell-like (“crepidotoid”),
without a stem.
Phaeomarasmius siquierii
33
Pileus 1,5-3 mm diam. in wet condition and when fully developed; margin undulate,
especially with age, not striate; color buff brown (7.5YR 5/3-5) to brown or tobaccobrown (7.5YR 4/3-6), depending on water content (so the pileus can be considered as
slightly hygrophanous); surface furfuraceous to shaggy, mostly showing a covering of
whitish hairs (canescent to sericeous), reaching the margin, but more dense at the centre
or the insertion area; under the lens the surface is scurfy, covered with minute crystal granulations.
Himenophore rather simple, formed by 3-7 lamellae, with no or few lamellulae (l= 0-2),
free on hardly reaching the centre of the pileus, 1-2 mm wide, concolorous; edge irregular, finely fimbriate to serrulate, sterile, whitish or paler than the surfaces; under the lens,
the edge shows numerous packed hairs which is the microscopic correlate of the fimbriate aspect; the faces often show brownish spots due to spore accumulation.
Flesh very inconspicuous, smell absent.
Basidiospores (Fig. 3A) 9-12 (-13) x 6-7,5 (-8) µm (Xm = 10,63 x 6,86; 9.91 x 7,10 µm
Q=1,39-1,55, Qm = 1,47), broadly ellipsoid to sub-ovoid, often sub-lentiform (slightly
compressed) in lateral view, the outline occasionally subangular or irregular, smooth,
brown ochraceous, thick walled (1 µm), with short apiculus, germ pore present but small,
central, 0,5-0,7 µm broad, sometimes rather inconspicuous and with a pseudocallus
aspect; endospore darker brown in Congo Red ammonia, with vacuolar content.
Cheilocystidia 40-110 x 3-6 µm, (sub-) cylindrical, sometimes attenuate towards the apex
or subventricose, with irregular, undulate or submoniliform outline, often bifurcate
towards the roundish, sometimes subcapitate apex; walls thick (1 µm), brown, due to
parietal pigment, frequently septate, clamped at base (Fig. 3H). Abundant microcrystals
present between cystidia or covering their walls in places, especially near the apex
(Fig. 3C, 3D).
Basidia (Fig. 3B) 22-40(-43) x (6-)9-11 µm clavate, four-spored, with long sterigmata
4-7 µm, clamped.
Pleurocystidia not observed.
Pileipellis (Fig. 3E, 3F) formed by a subtrichoderm, with filamentous and intricate
clamped hyphae in the epidermis, with numerous terminal cells similar to the
cheilocystidia. Abundant amount of microcrystals covering the hyphae, especially in the
apical endings.
Hymenophoral trama irregular, congophobic, formed by intricate clamped hyphae,
3-5 µm wide, yellowish-brown due to parietal pigment.
Material examined: Illes Balears, Formentera, Es Ram, torrent dels Arbocers, UTM
31SCC7179; alt. 5-30 msl, 9 December 2008, on ritidomes of Juniperus phoenicea
(“savina”), leg. J. Espinosa, J. Güell, T. Serra and J.C. Salom, Herb. JCS 816B, duplicate
in AH 40199. Ibidem, 8 December 2010, leg. J.C. Salom, Herb. JCS 1067B (Holotypus).
Isotypus in AH 40201.
34
Salom J. C. & Esteve-Raventós F.
COMMENTS
The number of species of the genus Phaeomarasmius in Europe is rather scarce, and
only the following taxa are known: P. erinaceus (Fr.: Fr.) Kühner, P. rimulincola (Rabenh.)
P.D.Orton, P. borealis Rald /=P. erinaceus s. A.H. Sm. & Hesler and P. gypsophilus
Esteve-Rav., Villarreal, Heykoop & E. Horak (Rald in KNUDSEN & HANSEN, 1991;
WATLING et al., 1993; LUDWIG, 2001). Both P. erinaceus and P. rimulincola are relatively
commonplace, whereas the rare Phaeomarasmius gypsophilus (ESTEVE-RAVENTÓS et al.,
1998), due to the more complex pileipellis structure, perhaps should be better placed in
Flammulaster Earle. Together with this last genus, Tubaria (W.G. Sm.) Gillet and
Phaeomarasmius include agaricoid mushrooms, many of them lignicolous, with brown
spore print, mostly small basidiomata and a simple, dry, filamentous pileipellis; in the
past, many species of these genera where included in the wide genus Naucoria Fr.
(SINGER, 1950; KÜHNER & ROMAGNESI, 1953), or interpreted as small species of Pholiota
(SMITH & HESLER, 1968). Traditionally, mycologists have considered that Tubaria species
can be easily recognized macroscopically by the (sub-)decurrent lamellae, whereas in
Flammulaster and Phaeomarasmius they are emarginate to adnate; the latter can be distinguished by the more complex pileipellis of Flammulaster, which shows more or less
isodiametric to short, wide elements or cells in the hyphae of the pileipellis (sometimes
wrongly called “sphaerocysts” or better, sphaerocytes), contrasting with the more simple,
filamentous, elongate and regular articles of Phaeomarasmius (VELLINGA, 1986);
Phaeomarasmius excentricus Scherff. /= P. rimulincola, which constitutes the type species
of the genus, differs from Flammulaster spp. by the different pileipellis structure (see
HORAK, 1968). Recent phylogenetic studies demonstrate that Flammulaster,
Phaeomarasmius and many Tubaria species group together in a “Tubaria o tubarioid
clade” (AIME et al., 2005) or an “Agaricoid clade-grade Tubarieae” (MATHENY et al., 2006,
2007), not related with Pholiota or Crepidotus and, consequently, they should not be considered as members of the Crepidotaceae. Recently, VIZZINI (2008) has established the
new family Tubariaceae for the tubarioid clade.
Phaeomarasmius siquierii is here presented as a new species, based on morphological
and ecological characters; it can be readily distinguished by its tiny basidiomata, lack of a
stipe (giving it a “crepidotoid” to “resupinatoid” habit), a very simple lamellate
hymenophore (L= 3-7), narrow and protruding cheilocystidia often with a tortuose to submoniliform outline, and a peculiar habitat on Juniperus (Cupressaceae) ritidomes. Either
P. erinaceus or P. rimulincola are very different morphologically; P. erinaceus shows a
hirsute to (sub-)squarrose pileus and a well developed stipe, whereas P. rimulincola may
superficially remind the new species by the aspect of the pileus and the small size; however, P. rimulincola shows a bigger pileus (often 4-8 mm diam.), a well developed, though
short stipe, well developed lamellae that reach the stipe, larger, amygdaliform spores
[(12-)14-18(-21) x 7-9 µm], different and non-encrusted cheilocystidia and typically
grows (usually together with some corticolous Mycena spp.) on living bark of broadleaved
trees. A revision of the literature which refers to Flammulaster or Phaeomarasmius in
America (SMITH & HESLER, loc. cit.; SINGER, 1956, 1969), as well as from different
(sub-)tropical areas of the world (HORAK, 1979, 1980; LIU, 1995) reveals that none of the
species described resemble P. siquierii, as the typical crepidotoid habit of the new species
seems not to have been noticed or found before in both genera.
The simplicity of the hymenophore in P. siquierii might call to mind the “cyphelloid”
brown-spored genera Chromocyphella and Phaeosolenia. Both include very reduced,
Phaeomarasmius siquierii
35
non-lamellate species (“cyphelloid habit”), with cup-shaped to tubular basidiomata,
growing gregariously in large numbers. On the basis of phylogenetic studies,
BODENSTEINER et al. (2004) have demonstrated that Phaeosolenia also group together with
Tubaria, Phaeomarasmius and Flammulaster, in the “Tubaria clade”, whereas data on
Chromocyphella are not available yet. Species of Phaeosolenia show smooth spores and
fructify on lax woolly stroma or subicula; Chromocyphella is a muscicolous genus with
punctate to ornamented spores (see DONK, 1959, 1962; COOKE, 1961; BODENSTEINER,
2006; SULZBACHER et al., 2009); in the Iberian Peninsula Chromocyphella muscicola and
C. pinsapinea (MORENO et al., 1985) have been recorded previously but not Phaeosolenia.
It is a fact that those little-known or unexplored mediterranean ecosystems such as
Juniperus spp. colonies (“sabinares” and “enebrales”) have turned out to be extremely
interesting when searching for macromycetes. Over the last years, in fact, many taxonomical novelties have been described growing on bark of branches or trunks of these vascular plants, e.g., Xeromphalina junipericola G. Moreno & Heykoop, Mycena juniperina
Aronsen, Marasmiellus phaeomarasmioides G. Moreno, Heykoop, Esteve-Rav. & E.
Horak, Trametes junipericola Manjón, G. Moreno & Ryvarden, etc., to name just a few.
ACKNOWLEDGEMENTS
We wish to express our sincere appreciation to our friends Joan Planas, for his help
and kindness in the composition of the photographic plates, and Toni Serra, Jaume
Espinosa and, above all, Joan Güell, for their company and help during the visit to
Formentera. Especially, we want to acknowledge our friend Josep Ll. (“Pepe”, “Pep”)
Siquier, to whom we dedicate this species, for his endless kindness and special friendship.
One of us (JCS) is proud to consider Pep his “mycological mentor”, an excellent master
and a tireless companion in numerous journeys, forays and fieldworks, in short, a great
friend whom he is very lucky to have met.
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