REVISION OF THE NIVICOLOUS SPECIES OF THE GENUS LEPIDODERMA

Abstract

This work is a morphological study of the nivicolous species within the genus Lepidoderma and, wherever possible, the type material has been studied. Both macro- and microphotographs are provided.
The ornamentation of the spores was studied under SEM with the critical point technique,
so we can provide a plate with the most distinctive features of each species. They are compared with Diderma fallax, because the latter sometimes shows lime scales on the peridium. At the end of the paper we present a dichotomous key for the genus Lepidoderma, including as well the nonnivicolous
species.

Full text

Bol. Soc. Micol. Madrid 42. 2018 39
REVISION OF THE NIVICOLOUS SPECIES
OF THE GENUS LEPIDODERMA
* G. MORENO1 , A. SÁNCHEZ1 , M. MEYER2 , Á. LÓPEZ-VILLALBA1 & A. CASTILLO1
1Dpto. de Ciencias de la Vida (Botánica), Facultad de Biología, Universidad de Alcalá,
28805 Alcalá de Henares, Madrid, Spain
Correspondence to *: gabriel.moreno@uah.es
2Le Bayet, 73730 Rognaix, France
marianne.meyer2@wanadoo.fr
Summary. MORENO, G., A. SÁNCHEZ, M. MEYER, Á. LÓPEZ-VILLALBA & A. CASTILLO
(2018). Revision of the nivicolous species of the genus Lepidoderma. Bol. Soc. Micol. Madrid 42:
39–77.
This work is a morphological study of the nivicolous species within the genus Lepidoderma and,
wherever possible, the type material has been studied. Both macro- and microphotographs are pro-
vided. The ornamentation of the spores was studied under SEM with the critical point technique,
so we can provide a plate with the most distinctive features of each species. They are compared
with Diderma fallax, because the latter sometimes shows lime scales on the peridium. At the end
of the paper we present a dichotomous key for the genus Lepidoderma, including as well the non-
nivicolous species.
Key words: Amoebozoa, myxobiota, slime moulds, taxonomy.
Resumen. MORENO, G., A. SÁNCHEZ, M. MEYER, Á. LÓPEZ-VILLALBA & A. CASTILLO
(2018). Revisión de las especies nivales del género Lepidoderma. Bol. Soc. Micol. Madrid 42:
39–77.
En este trabajo realizamos un estudio morfológico de las especies nivales del género Lepidoderma,
y siempre que nos ha sido posible revisamos el material tipo. De todas ellas realizamos fotografías
macro y microscópicas, estudiamos la ornamentación esporal al SEM con la técnica del punto críti-
co, y aportamos una plancha con sus caracteres más importantes. Se comparan con Diderma fallax,
que a veces también presenta placas cristalinas en el peridio. Finalmente incorporamos una clave
dicotómica de todas las especies descritas de Lepidoderma incluyendo las no nivales.
Palabras clave: Amoebozoa, hongos mucilaginosos, mixobiota, taxonomía.
INTRODUCTION
The genus Lepidoderma de Bary was pub-
lished by ROSTAFINSKI (1873). MARTIN &
ALEXOPOULOS (1969) described it as “sporan-
giate or plasmodiocarpous. Peridium cartilagi-
nous or membranous, covered with crystalline
scales, these usually conspicuous but sometimes
united into a nearly or quite continuous crust
forming a distinct outer wall, sometimes into a
nearly or quite continuous crust forming a distinct
outer wall, sometimes composed of rather loosely
compacted crystals. Capillitium as in Didymium,
typically limeless or in one species with large and
expanded nodes which are often vesicular and
may enclose clusters of lime crystals”. The pres-

G. MORENO, A. SÁNCHEZ, M. MEYER, Á. LÓPEZ-VILLALBA & A. CASTILLO
40 Bol. Soc. Micol. Madrid 42. 2018
ence of crystalline scales allows members of the
genus to be differentiated, but sometimes these
crystalline scales are also present in the genus
Diderma, possibly because of a process of recrys-
tallization of the calcium carbonate. However, the
genus Diderma always shows abundant globular
calcium carbonate. For example, it is common
to find samples of Diderma fallax (Rostaf.) E.
Sheld. with crystalline scales, frequently over
the outer peridium. The taxonomic position of
Diderma fallax is currently a problem, because
recent molecular studies by SHCHEPIN & al.
(2016) showed that it belongs to the same clade as
Lepidoderma peyerimhoffii. These authors sug-
gested the hypothesis that both species might rep-
resent the same unique and polymorphic species.
Nevertheless, our morphological studies (and re-
cent molecular studies [SHCHEPIN & al., not
published]) suggest otherwise.
Photographic plates of all the studied species
are provided. These show the most important
morphological features and are useful to com-
pare the species considered herein with other
described species.
MATERIALS & METHODS
The studied samples were recollected mainly
in France and Spain, occasionally in Austria,
Germany, and Italy, and more rarely in the United
States and Slovenia. The new proposed species
has been recollected only in the south of Spain,
in the Penibética mountain range.
All the studied material is preserved in the
herbarium of the Department of Plant Biology
of Alcalá de Henares (AH). The specimens
were mounted in Hoyer’s medium according
to MORENO & MANJÓN (2010). Spores were
measured under an oil immersion lens of a Nikon
microscope (Nikon Eclipse 80i) equipped with an
automatic photographic system (Nikon DS–5M).
The measurements provided herein included the
surface structures of the spores, such as spines
or warts.
Scanning electron microscopy (SEM) micro-
graphs were obtained at the University of Alcalá
de Henares using a Zeiss DSM–950 device. Only
a small amount of material is needed for this
type of study; only one or a small piece of a
single sporocarp (most rarely two sporocarps)
was taken from the sample and put into a 2 ×
2 cm square of cellulose filter paper Whatman
nº 1. Then the sporocarp was packed inside the
paper and the latter was stapled, to avoid losing
the sample. Then the sporocarp was rehydrated
in concentrated ammonium hydroxide (28–30%)
for 30 minutes, dehydrated in aqueous ethanol
(70%) for 30 minutes, fixed for two hours in pure
ethylene glycol dimethyl ether (= 1,2–dimethoxy-
methane), and finally immersed in pure acetone
for at least two hours, followed by critical point
drying and sputtering with gold-palladium.
TAXONOMY
Diderma fallax (Rostaf.) E. Sheld., Minnesota
Bot. Stud. 1: 477 (1895) Fig. 1a‒k, Fig. 2a‒l, Fig.
3a‒i
≡ Chondrioderma fallax Rostaf., Sluzowce
monogr. 171 (1874)
= C. lyallii Massee, Monogr. Myxogastr. 201
(1892)
= D. lyallii (Massee) Kuntze, Revis. gen. pl. 3(3):
466 (1898)
= C. niveum var. lyallii (Massee) Meyl., Bull. Soc.
Vaud. Sci. Nat. 44: 290 (1908)
= D. niveum subsp. lyallii (Massee) G. Lister, in
Lister, Monogr. mycetozoa, ed. 2, 105 (1911)
Material examined: Austria: Stanzach, Fallerscheinalpe,
1302 m, on living moss beside the snow, 12–VIII–2000, leg.
H. Singer, AH 19622. Hahntennjoch, near Anhalter Hütte,
2036 m, on living grass beside snow, 12–VIII–2000, leg.
H. Singer, AH 19623. France: Bonneval, Lachat, Savoie,
1800 m, woody debris, leg. M. Meyer, 15–V–1996, Meyer
16839 in AH 49085. Fontaine–le–Puits, Savoie, 1411 m,
herbes sèches, leg. F. Meyer & M. Meyer, 20–IV–2016,
Meyer 47575 duplo in AH 49084. Germany: Bavaria,
Wettersteingebirge: Garmisch-Partenkirchen, 1876 m, lit-
ter of grasses, on the ground, shallow slope, alpine mats
with Pinus mugo upper station Längenfelder Bahn, near up-
per station cableway Längenfelder Bahn, leg. M. Schnittler
& M. Borg Dahl, 5–V–2015, collect. M. Schnittler 28458.
Bavaria, Wettersteingebirge: Garmisch-Partenkirchen,
1690 m, open, shrubby gavelly slopes, alder, shrub pine
and meadows, 725 m SSW Kreuzalm cableway (upper

Bol. Soc. Micol. Madrid 42. 2018 41
REVISION OF THE NIVICOLOUS SPECIES OF THE GENUS LEPIDODERMA
Fig. 1.– Diderma fallax Schnittler 29242, a. Sporocarps, b. Details of the pseudocolumella and the double peridium, c. Outer and inner layers
of the peridium, d. Capillitium, e. Spores, f. Outer layer of the peridium consisting of globular calcium carbonate, g. Capillitium with nodules,
h. Details of the smooth and nodular capillitium, i–j. Spores, k. Details of the spore ornamentation. Scale bars: a = 1 mm, b–c = 0.25 mm, d =
100 µm, e = 10 µm, f, h= 5 µm, g = 20 µm, i–j = 2 µm, k = 1 µm.

G. MORENO, A. SÁNCHEZ, M. MEYER, Á. LÓPEZ-VILLALBA & A. CASTILLO
42 Bol. Soc. Micol. Madrid 42. 2018
Fig. 2.– Diderma fallax Cainelli 9052411, a. Sporocarps, b. Details of the sporocarps and the calcium carbonate recrystallizations
forming scales, c. Details of the pseudocolumella and the peridium, d. Threads of the capillitium, e. Capillitium with nodules, f.
Spinose spores, g. Capillitium, h. Peridium with recrystallizations of calcium carbonate forming scales and globular carbonate
calcium backwards, i–k. Spores, l. Details of the spore ornamentation. Scale bars: a–c = 1 mm, d = 100 µm, e = 50 µm, f = 10
µm, g–h = 10 µm, i–k = 5 µm, l = 1 µm.

Bol. Soc. Micol. Madrid 42. 2018 43
REVISION OF THE NIVICOLOUS SPECIES OF THE GENUS LEPIDODERMA
Fig. 3.– Diderma fallax Cainelli 9051507, a. Sporocarps with the typical tessellate shape, b–c. Peridium with recrystallizations of calcium
carbonate forming scales, d. Capillitium with widenings, e. Spores, f. Outer layer of the peridium consisting of globular calcium carbonate, g–h.
Spores, i. Details of the spore ornamentation. Scale bars: a = 0.5 mm, b–c = 0.1 mm, d = 100 µm, e = 10 µm, f = 5 µm, g–h = 2 µm, i = 1 µm.

G. MORENO, A. SÁNCHEZ, M. MEYER, Á. LÓPEZ-VILLALBA & A. CASTILLO
44 Bol. Soc. Micol. Madrid 42. 2018
station), E–exp. slopes above the road from Kreuzeck
(branch serpentine trail) to Hochalm, below a rock crest,
leg. M. Schnittler & M. Borg Dahl, 8–V–2016, collect. M.
Schnittler 29134. Bavaria, Wettersteingebirge: Garmisch-
Partenkirchen, 1760 m, pastures with Pinus mugo thickets,
240 m SW Hochalm (restaurant), valley between Hochalm
hill and Osterfelder Kopf, leg. M. Schnittler & M. Borg
Dahl, 11–V–2016, collect. M. Schnittler 29178. Bavaria,
Wettersteingebirge: Garmisch-Partenkirchen, 1870 m,
dry rocky meadow in Pinus mugo shrubbery, 160 m
SSW Laengenfelder Bahn (upper station), along the trail
over the ridge, near the outlook, other side, leg. M. Borg
Dahl, 3–VI–2016, collect. M. Schnittler 29239. Idem, col-
lect. M. Schnittler 29242. Slovenia: Snevznik, 1700 m,
15‒V‒2009, leg. R. Cainelli, Cainelli 9051507 duplo in
AH 49126. Spain: Granada, Güéjar Sierra, Dehesa de San
Juan, 2583 m, leg. J.F. Moreno, 6–V–2014, JA–CUSSTA
8018. Huesca, Jaca, Hoz de Jaca, road to Ibón de los Asnos,
leg. J. Hernanz, 16–V–2014, AH 50466. Huesca, Sallent de
Gállego, Portalet, leg. J. Hernanz, 31–V–2014, AH 50473.
Navarra, Isaba, Belagua, leg. M. Tapia, 15–VI–2018, AH
50413. USA: California, Montecito sequoia lodge, 2225
m, 24‒V‒2009, leg. R. Cainelli, Cainelli 9052411 in AH
49127.
Remarks ‒ Diderma fallax is characterised
by sporangiate sporocarps 1.3–2 mm in diam.,
occurring in groups, globose to subglobose.
Peridium double, with an outer layer of globular
calcium carbonate, thick, white to light straw-
yellow, which breaks into polygonal plates,
with the centre creamy-white to brownish. The
inner layer membranous and grey-iridescent.
Pseudocolumella obvious, elongated to cylindri-
cal, with a widened apex (club-shaped), creamy
to straw-yellow. Capillitium filamentous, dark,
with abundant nodules, transversal unions, paler
tips, and 2–3(–4) µm diam. threads. Spores dark
brown in transmitted light, globose to subglo-
bose, 14–17 µm diam., with dense and irregularly
distributed spines. Under SEM the nodules of the
capillitium are easy to see along the threads, and
the ornamentation of the spores consists of elon-
gated baculae 1–2 µm in length.
In some samples (Fig. 2a–b), in addition to
the globular calcium carbonate of the outer pe-
ridium, recrystallizations can be observed. The
shape of the scales is similar to those of the ge-
nus Lepidoderma. We have observed the typical
globular calcium carbonate of the genus Diderma,
which can be used as the diagnostic characteristic
of the genus.
Cainelli 9051507 (Fig. 3a–i) is macroscopi-
cally similar to Diderma fallax, but the capilli-
tium shows membranous expansions at the rami-
fications and the spore ornamentation is laxer
than in the other samples and their diameter is
14–15 µm. In addition, the spores of this sample
have a paler zone and the spines are only 1 µm
in length. It also has recrystallizations, but SEM
reveals globular calcium carbonate. It is possi-
ble that these variations are due to adverse en-
vironmental conditions while fruiting, or that it
is a different taxon (variety or species). For this
molecular studies would help. We doubt about
the assignment of the specimen Cainelli 9502411
(Fig. 2a–l),, which has spores with rather dense
and faint warts. This could be something dif-
ferent, when looking on it with molecular tools,
since our experience has shown that the spore
ornamentation is quite a valuable character.
Lepidoderma aggregatum Kowalski, Mycologia
63(3): 511 (1971). Fig. 4a–i
Material examined: USA: Whatcom Co., Washington, 16
miles east of Glacier, 4000 ft, on living twigs, 16–VI–1968,
leg. D.T. Kowalski, D.T. Kowalski 8870 type (UC).
Remarks – This species is characterised by
the aggregated sporocarps, which occur in dense
and scattered groups, sessile, pulviniform to ap-
planate, 1.5–3 mm diam., with lime scales form-
ing a whitish crust. Capillitium filamentous,
threads thin, 1–2 µm diam., brownish, scarcely
branched, somewhat flexuous. Spores globose
to subglobose, 11–15 µm diam., spinose. Under
SEM the ornamentation of the spores consists of
long baculae, with an irregular distribution and
the apices with few nodules, which give them a
coralloid look.
When comparing the plates of the types of
Lepidoderma aggregatum and L. chailletii, it is
evident that the capillitium, the ornamentation
of the spores, and the size of the spores are very
similar in both species. The only difference be-
tween them are the lime scales, which always

Bol. Soc. Micol. Madrid 42. 2018 45
REVISION OF THE NIVICOLOUS SPECIES OF THE GENUS LEPIDODERMA
form a continuous layer in L. aggregatum, while
only sometimes in L. chailletii, as indicated be-
low. We think it can be due to the environmental
conditions when fruiting takes place, as indicated
in MORENO & al. (2004). Recently, SHCHEPIN
& al. (2016) indicated that the lime scales of L.
chailletii never form a continuous layer or crust.
Additionally, their molecular studies revealed
that there are three different clades of L. chail-
letii, and they suggested that possibly one of
those clades would enclose the American species
L. aggregatum. To confirm this hypothesis, it is
necessary to carry out molecular studies of the
type specimen or to collect samples at Kowalski’s
(1971) original location again (Olympic National
Park, Washington, USA). Kowalski (1971) in-
dicated that this species was very abundant, so
much that he studied almost 275 collections.
As such, after studying the morphology of the
type material of Lepidoderma aggregatum and
comparing it with L. chailletii, and supported by
the molecular results of SHCHEPIN & al. (2016),
we conclude that L. aggregatum and L. chailletii
may constitute two different sister species, but
this needs to be confirmed by molecular data
from the American samples.
Fig. 4.– Lepidoderma aggregatum D.T. Kowalski 8870 type, (UC), a. Original label, b. Sporocarps, c. Details of the peridium, d. Lime scales
of the peridium, e. Capillitium, f. Capillitium and its connections with the peridium, g‒h. Spores, i. Details of the spore ornamentation. Scale
bars: b‒c = 5 mm, d‒f = 100 µm, g‒h = 2 µm, i = 1 µm.

G. MORENO, A. SÁNCHEZ, M. MEYER, Á. LÓPEZ-VILLALBA & A. CASTILLO
46 Bol. Soc. Micol. Madrid 42. 2018
Fig. 5.– Lepidoderma alpestroides Meyer 16595 Isotype duplo in AH 49121, a–b. Plasmodiocarps, c. Insertion of the capillitium into the
peridium, d. Capillitium, e. Inner peridium and spores, f. Spores under LM, g–h. Spores under SEM (Meyer 17476 duplo in AH 31776). i.
Details of the spore ornamentation (Meyer 17476 duplo in AH 31776). Scale bars: a–b = 1mm, c–d = 50 µm, e–f = 10 µm, g–h = 2 µm, i = 1
µm.
Lepidoderma alpestroides Mar. Mey. &
Poulain, in Poulain, Meyer & Bozonnet, Bull.
Mycol. Bot. Dauphiné–Savoie 165: 9 (2002).
Fig. 5a–i
Material examined: France: Bourg-Saint-Maurice, Les
Arcs, Savoie, 1900 m, on Vaccinium mytillus, 24–V–1996,
Meyer 16595 Isotype in AH 49121. Bourg-Saint-Maurice,
Les Arcs, Savoie, 1900 m, on living shrub, mainly

Bol. Soc. Micol. Madrid 42. 2018 47
REVISION OF THE NIVICOLOUS SPECIES OF THE GENUS LEPIDODERMA
Rhododendron sp., 28–V–1997, Meyer 17476 duplo in AH
31776. Bonneval, Lachat, Savoie 1800 m, woody debris
and leaves, 31–V–2001, leg. M. Meyer, Meyer 27795 in
AH 49082.
Sporocarps whitish to yellowish, forming
lonely to gregarious plasmodiocarps, flattened
to depressed in the centre, 3–10 × 1.5–2.5 mm.
Peridium double, outer layer with white to straw
lime scales, which form a continuous and al-
most smooth crust. Inner layer of the perid-
ium membranous, hyaline, and non-iridescent.
Pseudocolumella bulky, ochraceous-white, fill-
ing half of the sporocarp. Capillitium abundant,
filamentous, non–reticulated, dichotomously
branching, emerging from the pseudocolumella
to the peridium, dark with paler tips, without
or with scanty nodules. Hypothallus underde-
veloped, silvery. Spores blackish in mass, dark
brown in transmitted light, globose to subglo-
bose, 12–15 µm diam., spinose. Under SEM the
ornamentation of the spores consists of spines
with coralloid apex.
Remarks – This species is well characterised
by its plasmodiocarpous sporocarps, the crustose
peridium formed by coalescing lime scales, the
presence of a pseudocolumella, the abundant capil-
litium, and spinose spores 12–15 µm in diam.
At the first glance it can be confused with
members of the genus Physarum, but in this
genus separable lime scales on the peridium do
not occur (although the lime can be patchy) and
the capillitium exhibits calcareous nodules con-
nected by hyaline tubules.
Recent molecular studies carried out by
SHCHEPIN & al. (2016) confirmed that
Lepidoderma alpestroides can be considered as
an independent species.
Lepidoderma carestianum (Rabenh.) Rostaf.,
Sluzowce Monogr.: 188 (1874). Fig. 6a–c, Fig.
7a–l, Fig. 8a–i
≡ Reticularia carestiana Rabenh., Fungi Eur.
Editio nova, ser. Secunda No. 436 (1862).
= Amaurochaete minor Sacc. & Ellis, in Sac-
cardo, Michelia 2: 566 (1882).
= Dermodium minus (Sacc. & Ellis) Kuntze, Re-
vis. gen. pl. 3(3): 465 (1898).
= L. carestianum f. granulosum Poulain & al., ad
inter., Les myxomycètes tome 2, fig. 398 (2011).
Fig. 6.– Lepidoderma carestianum Fungi europaei exs. No. 436, in BR as Reticularia carestiana lectotype, a‒b. Plasmodiocarps, c. Details of
the brown lime nodules. Scale bars: a‒b = 1 mm, c = 20 µm.

G. MORENO, A. SÁNCHEZ, M. MEYER, Á. LÓPEZ-VILLALBA & A. CASTILLO
48 Bol. Soc. Micol. Madrid 42. 2018
Fig. 7.– Lepidoderma carestianum AH 19504, a‒b. Plasmodiocarps, c. Details of the brown lime nodules, d. Reticulate capillitium, e‒f.
Details of the lime nodules, g. Capillitium with lime nodules, h‒i. Typical ornamentation of the lime nodules, j‒k. Spores, l. Details of the
spore ornamentation. Scale bars: a‒b = 1 mm, c = 100 µm, d = 20 µm, e‒f = 10 µm, g = 20 µ, h‒i = 5 µm, j‒k = 2 µm, i = 1 µm.

Bol. Soc. Micol. Madrid 42. 2018 49
REVISION OF THE NIVICOLOUS SPECIES OF THE GENUS LEPIDODERMA
Fig. 8.– Amaurochaete minor S.J. Harkness (Ellis collection) NYBG 5607 and NYBG 5608 lectotypus, a–b. Original labels, c. Plasmodiocarps
NYBG 5607, d. Plasmodiocarps NYBG 5608, e. Reticulate capillitium NYBG 5608, f. Spores under LM NYBG 5608, g–h. Spores NYBG
5608, i. Details of the spore ornamentation NYBG 5608. Scale bars: c‒d = 1 cm, e = 50 µm, f = 10 µm, g‒h = 2 µm, i = 1 µm.

G. MORENO, A. SÁNCHEZ, M. MEYER, Á. LÓPEZ-VILLALBA & A. CASTILLO
50 Bol. Soc. Micol. Madrid 42. 2018
Material examined: France: Bourg-Saint-Maurice, Les
Arcs, Savoie, on woody twigs, 28–V–1994, Herb. Meyer
14542 duplo in AH 49124. Ibidem, on twigs of Rhododen-
dron sp., 3–VI–2004, 2047 m, Herb. Meyer 24142, duplo in
AH 46500. Italy: Riva, Valsesia, ramulus fruticum varium,
spring 1861, leg. Ab. Carestia Rabenhorst Fungi europaei
exs. No. 436, in BR as Reticularia carestiana (proposed
as lectotype). Spain: Madrid, Puerto de Navacerrada, on
twigs of Cryptogramma crispa, 1–V–1997, 2100 m, leg.
A. Sánchez, AH 19504. Ibidem, Bola del Mundo, on dead
twigs of Senecio pyrenaicus, 27–V–1999, 2150 m, leg. A.
Sánchez, AH 19398. Segovia, Puerto de Navacerrada, on
twigs of Senecio pyrenaicus, 30–V–2000, 1950 m, leg. A.
Sánchez, AH 33547. Puerto de Navacerrada, on twigs of
Senecio pyrenaicus, 3–VI–2001, 2175 m, leg. A. Sánchez,
AH 28548. USA: Utah, S.J. Harkness, (Ellis collection),
NYBG 5607 and NYBG 5608 typus, both as Amaurocha-
ete minor.
Elongated plasmodiocarps, rarely branched,
20–60 × 1–5 mm, convex, sometimes flattened,
white to greyish-white to red-brown. Hypothal-
lus scanty, membranous, dark brown. Peridium
simple, dark brown to blackish-brown, it can be
easily seen when the lime scales are lacking; lime
scales white to slightly yellow. Apical and irregu-
lar dehiscence. Pseudocolumella lacking. Capil-
litium filamentous, abundant, with hyaline to
slightly brownish threads, 2–4 µm diam., reticu-
late, with plentiful to scanty lime nodules onto
the threads, which are yellow to brown and elon-
gated or globular, measuring 30–50 × 20–32 µm.
Spores violaceous in mass, violaceous-brown in
transmitted light, with a paler zone, globose to
subglobose to ovoid, 15–20 × 14–16 µm, with
clustered spines that sometimes coalesce into
short crests. Under SEM not only the nodules of
the capillitium, but also the threads of the capil-
litium are ornamented with delicate, longitudinal
to sinuous crests. This character is only visible
under SEM, and occurs consistently in all stud-
ied specimens. The ornamentation of the spores
consists of baculae which become curved easily
at the apex, and sometimes they show small un-
ion lines which gives the spore a subreticulated
look.
Remarks – Lepidoderma carestianum is char-
acterised by elongated and plasmodiocarpous
sporocarps, a simple peridium, filamentous and
anastomosed capillitium with lime nodules, and
spinose spores under LM. Under SEM the orna-
mentation of the capillitium and its nodules is
characteristic.
The first problem noted for this species is the
confusion expressed by RABENHORST (1862),
because he distributed both Lepidoderma cares-
tianum and Reticularia carestiana Rabenh., [Rab-
enhorst 436 Fungi europaei from Riva (Valsesia),
spring 1862, ad ramulos fruticum varium, leg.
Ab. Carestia] under the same name. One of the
two taxa has a reticulate capillitium with lime
nodules (Fig. 6a–c) and the other one has an al-
most non-reticulate and non-branched capillitium
consisting of straight, hyaline to slightly coloured
threads without lime nodules (Fig. 9a–f). Pos-
sibly this confusion has subsequently caused a
wrong interpretation of the species Lepidoderma
carestianum.
LISTER & G. LISTER (1911) sketched Lepi-
doderma carestianum from Switzerland (plate
115, figures a and d) with a reticulate capillitium
and with abundant lime nodules, while they drew
L. carestianum var. granuliferum from Califor-
nia (plate 115, figures b and c) with a reticulate
capillitium and globular lime nodules covered by
an asteriform membrane. They described L. car-
estianum var. chailletii as characterized by well-
developed sporangiate sporocarps with a obvious
pseudocolumella, spores 10–13 µm diam., and a
filamentous and scanty branching capillitium
(plate 116, figures a–d). Later, these authors con-
sidered this variety as the independent species L.
chailletii.
The first monograph relating to the genus
Lepidoderma was written by KOWALSKI (1971),
where he recognised six different species. He
studied the type material of five of these species,
apart from Lepidoderma tigrinum. In any case,
he was able to sample and study all six species,
including the latter one.
Some authors, such as KOWALSKI (1971) and
NEUBERT & al. (1995) considered Lepidoderma
carestianum as a synonym of L. chailletii. They
did not consider the different type of the sporo-
carps, the differences in the capillitium and in
the spore ornamentation as having taxonomic

Bol. Soc. Micol. Madrid 42. 2018 51
REVISION OF THE NIVICOLOUS SPECIES OF THE GENUS LEPIDODERMA
value. Lepidoderma carestianum has plasmodio-
carpous sporocarps, while L. chailletii virtually
always develops discernible sporangiate sporo-
carps. KOWALSKI (1971) described the capil-
litium of L. carestianum as “abundant, weakly
attached to the peridium and columella, com-
posed of straight or, more commonly, f lexuose
threads, threads of uniform diameter, commonly
branching and anastomosing to form an intricate,
wide-meshed net, dark purple- brown through-
out, becoming hyaline only at the extremities,
not noticeably tapering towards the tips, smooth
but often bearing spherical to fusiform enlarge-
ments”. This description is applicable to the cur-
rent circumscription of L. chailletii.
POULAIN & al. (2002) studied one of the
samples designed as typus by Rabenhorst, from
the BR herbarium. They concluded that Lepi-
doderma carestianum was a single species,
typically plasmodiocarpous, and they sketched
it as showing a reticulate capillitium without
lime nodules. While when we studied the same
material, it was found to have the characteristic
lime nodules in the capillitium, although scanty.
Afterwards, POULAIN & al. (2011) made a new
interpretation of this species and sketched L.
carestianum f. granulosum ad inter. in the plate
398, which we understand as the true L. cares-
tianum.
LADO (2005–2018) followed LISTER & G.
LISTER (1911) while considering as Lepidoder-
ma carestianum the taxon with nodules in the
capillitium. He synonymized L. granuliferum
with L. carestianum, which is as well a differ-
ent species, as we will show below.
SHCHEPIN & al. (2016) studied only a single
specimen of Lepidoderma granuliferum (sc26564),
which was placed within the clade of L. cares-
tianum, and recommended the need to study more
material to ascertain its relationship to L. cares-
tianum. Their phylogeny of partial SSU sequences
does not exclude the possibility that Lepidoderma
carestianum and L. chailletii are two different spe-
cies. We agree with this conclusion because the
two taxa differ morphologically (compare plates
of both species in this study).
Lepidoderma carestianum can be confused
with other plasmodiocarpous species such as L.
perforatum, L. neoperforatum, and L. granu-
liferum. Lepidoderma perforatum and L. neop-
erforatum both show perforations in the peridium
under LM. These are formed at the points where
the threads of the capillitium are inserted into
the peridium; both species have a dark-brown
capillitium and differ in spore ornamentation
(see comments to these species). Lepidoderma.
granuliferum possesses a reticulate capillitium
which forms a dense net with abundant asteri-
form nodules, smooth under SEM, and its spores
(15–18 µm in diam.) are larger than those of L.
carestianum.
In this work, and after revising the type ma-
terial of Lepidoderma carestianum (Fig. 6a–c),
we have decided to follow the interpretation of
LISTER & G. LISTER (1925) for this species.
For us, they were the first authors that established
a proper species circumscription.
We have checked the type of Amaurochaete
minor (Fig. 8a–i) and it shows a reticulated capil-
litium, forming a dense net with abundant lime
nodules, and the spores are globose to ovoid,
14‒18 × 13‒15 µm, so we consider this species
to be a synonym of Lepidoderma carestianum.
The record of Lepidoderma granuliferum
from Sierra de Guadarrama (Segovia, Spain)
published by SÁNCHEZ & al. (2002) was re-
vised and found to be L. carestianum. The latter
species is not yet recorded for our country, so the
specimens mentioned herein are the first records
for Spain.
Lepidoderma carestianum var. pseudocares-
tianum G. Moreno, Ant. Sánchez, Mar. Mey.,
López-Vill. & A. Castillo var. nov. Fig. 9a–f,
Fig. 10a–j
= L. carestianum sensu Poulain, Meyer & Bozon-
net, Les myxomycètes tome 2, fig. 399 (2011).
MycoBank: MB 827961
Etymology: “pseudocarestianum” refers to its
resemble to Lepidoderma carestianum.
Typus. Spain: Segovia, Riaza, Pico del Lobo, on
leaves of Festuca indigesta, 14–V–2003, 2000 m,
leg. A. Sánchez, AH 32515 Holotype

G. MORENO, A. SÁNCHEZ, M. MEYER, Á. LÓPEZ-VILLALBA & A. CASTILLO
52 Bol. Soc. Micol. Madrid 42. 2018
Material examined: Italy: Riva, Valsesia, ramulus fruticum
varium, spring 1861, leg. Ab. Carestia, Rabenhorst, Fungi
europaei exs. No. 436, in B, BPI and WSRL as Reticularia
carestiana. Pian del Regina, unidentified fistulous twigs,
8–V–2004, 1500 m, Herb. M. Meyer nº 23823, duplo in
AH 46501. Spain: Segovia, Riaza, Pico del Lobo, vegetal
debris, 1950 m, AH 32530. Pto. de Navacerrada, on twigs
of Senecio pyrenaicus, 30–V–2000, 1950 m, AH 33547.
Sporocarps occur as elongated plasmodio-
carps, grey to coppery-brown, 5–12 × 1.5–2.0 mm.
Hypothallus membranous, brownish. Peridium
simple, membranous to subcartilaginous, with
abundant calcium carbonate lime scales, white
to straw-white. Dehiscence irregular and usually
apical. Capillitium abundant, hyaline to slightly
brown, threads parallel, 1–3 µm diam., sparsely
anastomosing, not forming an evident reticulum,
without or (rarely) with only a few lime nodules,
tips of the threads funnel-shaped or Y-shaped at
the reunion with the peridium, sometimes retain-
ing a piece of it. Spores dark brown in mass, vio-
laceous-brown under transmitted light, globose to
subglobose, seldom widely ellipsoid, 14–18 µm
diam., spinose. Under SEM the threads of the
capillitium do not show nodules and they have
fragile, membranous remainders. The spores are
spinose with the apex typically curved.
Remarks – Lepidoderma carestianum var.
pseudocarestianum is characterised by the elon-
gated, greyish-white to coppery-brown, and plas-
modiocarpous sporocarps, the simple peridium,
a filamentous, hyaline capillitium with threads
Fig. 9.– Lepidoderma carestianum var. pseudocarestianum Fungi europaei exs. No. 436, in BPI as Reticularia carestiana, a. Plasmodiocarp,
b. Capillitium and tips of the threads without lime nodules, d‒e. Spores, f. Details of the spore ornamentation. Scale bars: a = 1 mm, b‒c = 20
µm, d‒e = 2 µm, f = 1 µm.

Bol. Soc. Micol. Madrid 42. 2018 53
REVISION OF THE NIVICOLOUS SPECIES OF THE GENUS LEPIDODERMA
rarely anastomosing, virtually without lime nod-
ules, with funnel-shaped tips, and spinose spores.
As mentioned in the comments to L. carestianum,
the specimens distributed by RABENHORST
(1862) as Reticularia carestiana represented two
different taxa.
Lepidoderma carestianum var. pseudocares-
tianum can be confused with other species of
Lepidoderma, such as L. carestianum, L. perfo-
ratum, L. neoperforatum and L. granuliferum.
Lepidoderma carestianum usually has a reticu-
late capillitium with abundant lime nodules with
variable shapes, ranging from fusiform to glo-
bose. Under SEM both the capillitium and the
lime nodules are ornamented with short crests,
as indicated above.
Under LM and SEM Lepidoderma perfora-
tum and L. neoperforatum show perforations
in the peridium and the ornamentation of the
spores is different (see observations in both spe-
cies).
Lepidoderma granuliferum shows intermediate
sporocarps, ranging from sessile and subglobose
sporangiate forms to sinuous and short plasmodio-
carps. The capillitium is filamentous, with very
reticulate threads, and numerous membranous ap-
pendixes which cover the groups of the smooth
and globular lime nodules. The membrane and the
nodules are smooth even under SEM.
The three species Lepidoderma carestianum,
L. chailletii, and L. granuliferum have been con-
fused in many studies, and the reasons for this
Fig. 10.– Lepidoderma carestianum var. pseudocarestianum AH 32515 holotypus, a. Plasmodiocarp, b‒c. Details of the crystalline scales and
capillitium, d. Capillitium without lime nodules, e‒f. Widened tips of the capillitium, g‒h. Capillitium, i. Spore, j. Details of the spore ornamen-
tation. Scale bars: a = 1 mm, b = 2 mm, c = 100 µm, d = 50 µm, e = 20 µm, f = 10 µ, g‒h = 5 µm, i = 2 µm, j = 1 µm.

G. MORENO, A. SÁNCHEZ, M. MEYER, Á. LÓPEZ-VILLALBA & A. CASTILLO
54 Bol. Soc. Micol. Madrid 42. 2018
confusion are mentioned above in the remarks on
L. carestianum.
Lepidoderma chailletii Rostaf., Sluzowce Mon.:
189 (1874). Fig. 11 a–i, Fig. 12a–h, Fig. 13a–k
Material examined: Spain: Madrid, Puerto de Cotos,
Valdesquí, on bark of Pinus sylvestris, 23–III–1997, 1775
m, leg. A. Sánchez, AH 19214. Ibidem, on branches, 23–
IV–1997, 1875 m, AH 19575. Ibidem, on bark, 11–IV–1999,
1825 m, AH 19467. Ibidem, 14–IV–1999, 1850 m, AH
19393, AH 19399. Ibidem, 1925 m, AH 19474. Ibidem, 23–
IV–1999, 1850 m, AH 19389. Ibidem, 25–IV–1999, 1800 m,
AH 19384, AH 19465. Ibidem, 1825 m, AH 19466. Ibidem,
on branches of Juniperus communis subsp. nana, 1925 m,
AH 19473. Ibidem, on bark of fallen branches of Pinus
sylvestris, 1–V–1999, 1850 m, AH 26328. Ibidem, on living
branch of Cytisus oromediterraneus, 2–V–1999, 1900 m,
AH 26324. Ibidem, on branches, 1–IV–2001, AH 33058.
Puerto de Navacerrada, on Festuca indigesta, 16–III–2000,
2125 m, leg. A. Sánchez, AH 25650. Ibidem, 2150 m, AH
26317. Ibidem, on leaves and twigs of Digitalis purpurea,
AH 26319. Ibidem, on twigs, 2100 m, AH 19216. Ibidem,
on stems of Poaceae, 2150 m, AH 26326. Ibidem, on twigs
of Gentiana lutea, 8–VI–2000, 2000 m, AH 25905. Ibidem,
on twigs of Senecio pyrenaicus, 2100 m, AH 25897. Ibidem,
2150 m, AH 25933. Ibidem, on twigs of Rumex suffructi-
cosus, 17–VI–2000, 2150 m, AH 25520. Segovia, Pico del
Lobo, Riaza, on Festuca indigesta, 14–V–2003, 2000 m,
leg. A. Sánchez, AH 32515. Ibidem, on living branches
of Cytisus oromediterraneus, 1925 m, AH 32516. Ibidem,
2000 m, AH 32514, AH 32517. Puerto de Navacerrada, on
branches of Cytisus oromediterraneus, 20–IV–1997, 1925
m, leg. A. Sánchez, AH 26231, AH 33054. Ibidem, 1950
m, AH 19215. Ibidem, on twigs of Rumex suffructicosus,
1900 m, AH 19211. AH 19213. Ibidem, 1925 m, AH 26320.
Ibidem, 25–IV–1997, 1875 m, AH 26325. Ibidem, on fronds
of Cryptogamma crispa, 20–V–1997, 2000 m, AH 19218.
Ibidem, on stems of Poaceae, 2025 m, AH 19219. Ibidem,
on twigs of Senecio pyrenaicus, 24–V–1997, 1925 m, AH
19217. Ibidem, on processed wood, 23–V–1998, 2000 m,
AH 18951. Ibidem, on twigs of Rumex suffructicosus, 2–
IV–1999, 1925 m, AH 19212. Ibidem, on living needles of
Pinus sylvestris, 2–V–1999, 1750 m, AH 26327. Ibidem, on
leaves of Gentiana lutea, 1950 m, AH 26322. Ibidem, on
stems of Poaceae, 1925 m, AH 33050. Ibidem, on twigs
of Digitalis purpurea, 5–V–1999, AH 19385. Ibidem,
on twigs of Rumex suffructicosus, 1950 m, AH 19383.
Ibidem, on stems of Poaceae, AH 19396. Ibidem, 1975
m, AH 18390. Ibidem, on fronds of Cryptogamma crispa,
7–V–1999, AH 26329. Ibidem, on stems of Poaceae, 1950
m, AH 33051. Ibidem, on Festuca indigesta, 9–V–1999, AH
19386. Ibidem, on leaves of Juniperus communis subsp.
nana, AH 19391. Ibidem, on stems of Poaceae, 11–V–1999,
1925 m, AH 33048. Ibidem, on fronds of Cryptogamma
crispa, 15–V–1999, 2000 m, AH 26330. Ibidem, on twigs
of Senecio pyrenaicus, 1975 m, AH 19387. Ibidem, on
twigs of Gentiana lutea, 28–V–1999, AH 26337. Ibidem,
on twig of Gentiana lutea, 30–V–1999, AH 19382. Ibidem,
on stems of Poaceae, 1950 m, AH 19470. Ibidem, 1975
m, AH 19381. Ibidem, on twigs of Senecio pyrenaicus,
16–III–2000, 2175 m, AH 25656. Ibidem, 17–III–2000,
2150 m, AH 25651, AH 25653. Ibidem, 2175 m, AH 25652.
Ibidem, on stems of Poaceae, 14–V–2000, 1900 m, AH
26334. Ibidem, 18–V–2000, 1925 m, AH 25954. Ibidem,
19–V–2000, AH 26335. Ibidem, 1950 m, AH 25884. Ibidem,
28–V–2000, AH 26336. Ibidem, on branches of Juniperus
communis subsp. nana, 6–VI–2000, AH 28723. Ibidem,
on twigs of Senecio pyrenaicus, 15–VI–2000, 2200 m, AH
25818. Ibidem, on processed wood, 10–VIII–2000, 2100
m, AH 28729. Ibidem, on fronds of Cryptogamma crispa,
1–IV–2001, 1975 m, AH 18931. Ibidem, on twigs of Rubus
idaeus, 4–IV–2001, 1725 m, AH 33057. Ibidem, on living
leaves of Erica arborea, 12–IV–2001, 1975 m, AH 33055.
Ibidem, on fronds of Cryptogamma crispa, 18–IV–2001,
AH 33053. Ibidem, on twigs of Senecio pyrenaicus, 1900
m, AH 33047. Ibidem, on twigs of Rubus idaeus, 23–IV–
2001, AH 33056. Ibidem, on stems of Poaceae, AH 33049.
Ibidem, on branches of Pinus sylvestris, 24–IV–2001, 1800
m, AH 33045. Ibidem, on branches of Cytisus oromediter-
raneus, 25–IV–2001, 1975 m, AH 33052. Ibidem, on proc-
essed wood, 25–VII–2001, 2150 m, AH 25864. Ibidem, on
stems of Poaceae, 11–V–2003, 1975 m, AH 32610. Puerto
de Navafría, on branch of Pinus sylvestris, 22–III–1997,
1800 m, leg. A. Sánchez, AH 33046. Ibidem, on twigs of
Rubus ulmifolius, 25–III–2001, 1675 m, AH 26401. Ibidem,
1700 m, AH 18534. Granada, Sierra Nevada, Hoya de la
Mora, on stems of Poaceae, 11–V–2001, 2600 m, leg. A.
Sánchez, AH 29047.
Sporocarps very variable in shape, reach-
ing from sporangiate forms to short plasmodio-
carps, 1–3 mm in length, in large and crowded
groups, whitish to greyish-white. Peridium sim-
ple, pale brown to dark brown, covered by white
to cream-white lime scales, very variable in size
and shape, sometimes covering the entire perid-

Bol. Soc. Micol. Madrid 42. 2018 55
REVISION OF THE NIVICOLOUS SPECIES OF THE GENUS LEPIDODERMA
Fig. 11.– Lepidoderma chailletii AH 19383, a. Sporocarps, b. Details of the sporocarps and the crystalline scales, c‒d. Capillitium, e.
Spores, f. Capillitium with a granulose cover, g‒h. Spores, i. Details of the spore ornamentation. Scale bars: a‒b = 1 mm, c‒d = 50 µm,
e = 10 µm, g‒h = 2 µm, i = 1 µm.

G. MORENO, A. SÁNCHEZ, M. MEYER, Á. LÓPEZ-VILLALBA & A. CASTILLO
56 Bol. Soc. Micol. Madrid 42. 2018
Fig. 12.– Lepidoderma chailletii AH 19391, a. Sporocarps, b. Details of sporocaps, c‒d. Capillitium and peridium, e. Details of the capillitium
with small nodules, f‒g. Spore, h. Details of the spore ornamentation. Scale bars: a‒b = 1 mm, c‒d = 100 µm, e = 50 µm, f‒g= 2 µm, h = 1
µm.
ium and sometimes leaving gaps between them,
which reveal the membraneous layer of the pe-
ridium. Pseudocolumella pulvinulate to subglo-
bose, whitish to yellowish, sometimes lacking.
Capillitium filamentous, brownish, paler at the
tips of the threads, variable, but normally rigid,
smooth, poorly branched, and barely reticulated
(AH 19383, AH 19391), 1–1.5 µm diam. Other

Bol. Soc. Micol. Madrid 42. 2018 57
REVISION OF THE NIVICOLOUS SPECIES OF THE GENUS LEPIDODERMA
Fig. 13.– Lepidoderma chailletii AH 25821, a. Sporocarps, b. Crystalline scales, c‒d. Details of the outer reticulate capillitium,
e. Tips of the capillitium, f. Capillitium reticulate at the tips, g‒h. Capillitium with a granulose cover, i‒j. Spores, k. Details of the
spore ornamentation. Scale bars: a = 1 mm, b‒d = 50 µm, e‒f = 20 µm, g‒h = 5 µm, i‒j = 2 µm, k = 1 µm.

G. MORENO, A. SÁNCHEZ, M. MEYER, Á. LÓPEZ-VILLALBA & A. CASTILLO
58 Bol. Soc. Micol. Madrid 42. 2018
samples show abundant and small nodules (AH
26320), or rarely a reticulated pattern at the tips
of the threads (AH 25821). Spores globose to sub-
globose, 12–14 µm diam., spinose. Under SEM,
the threads of the capillitium show a cover of
amorphous material which entirely covers these
structures, and the ornamentation of the spores
consists of irregularly distributed baculae with
small nodules at the apices, which gives them a
coralloid look.
Remarks – Lepidoderma chailletii is charac-
terised by sporangiate to short plasmodiocarpous
sporocarps, a whitish to yellowish pseudocolu-
mella, the peridium with calcium carbonate lime
scales which vary in size and shape, sometimes
stablishing a continuous layer and sometimes ap-
pearing isolated, the filamentous, rigid, slightly
branched, dark brown capillitium with paler
tips and threads of 1–1.5 µm diam., and spinose
spores, 12–14 µm diam.
As indicated above, some authors such as
KOWALSKI (1971) and NEUBERT & al. (1995)
have misinterpreted Lepidoderma chailletii.
Their concept merged specimens with plasmo-
diocarpous and sporangiate sporocarps into a
single variable species, and synonymized it with
L. carestianum. LISTER & G. LISTER (1911,
1925) considered L. chailettii to be an independ-
ent taxon, just like POULAIN & al. (2011). We
agree and follow this most recent interpretation
of L. chailletii, so we conclude that L. chailletii
and L. carestianum are two different and well
defined species.
In the molecular studies carried out by SH-
CHEPIN & al. (2016), specimens determined
as Lepidoderma chailletii segregated into three
different clades, which are likely to constitute
different species, but these authors did not find
clear morphological differences to separate them
properly, so they considered them “cryptic spe-
cies”. They concluded that further studies using
scanning electron microscopy to visualize spore
ornamentation patterns would be necessary to
separate them. While studying our samples un-
der SEM, we have not observed significant dif-
ferences in the ornamentation of the spores, but
we did see variations in the size of the spores
and in the capillitium. Sometimes, the threads
of the capillitium are flexuous while other times
they are rigid. In addition, the threads were
smooth in some samples, while in others they
had nodules. This variability is illustrated in the
plates.
Lepidoderma crustaceum Kowalski, Mycologia
59(1): 167 (1967). Fig. 14a–h
Material examined: USA: Butte Co., California, 4 miles E
of Stirling, 4000 ft, on decaying twig, 4–VI–1966, leg. D.T.
Kowalski, D.T. Kowalski 2643 type (MICH).
Remarks – This species is characterised by
the short stalk, pale violaceous-brown sporo-
carps 1–1.5 mm in diam., occurring separat-
ed or in small groups, and a double peridium
with lime scales forming a continuous layer. A
pseudocolumella is often present. The capilli-
tium consists of filamentous, dark brown, rigid,
scarcely branched, and anastomosing threads.
The spores are globose to subglobose, 11–13 µm
diam., prominently and densely spinose. Under
SEM the ornamentation of the spores consists
of dense baculae.
KOWALSKI (1971) commented that Lepido-
derma crustaceum is a “extremely rare, perhaps
the rarest member of the genus”. After the revi-
sion of Kowalski’s type material carried out by
MORENO & al. (2004), the authors concluded
that this was a well defined and rare species in
the genus.
SHCHEPIN & al. (2016) could obtain only one
sequence of Lepidoderma crustaceum from Gen-
Bank, and their study included it within the clade
of Diderma fallax and L. peyerimhoffii. Their
conclusion is that the sequence is taxonomically
wrong determined, because the morphological
differences between the species are not signifi-
cant. We think that more molecular studies are
needed, especially using the American material,
to ascertain its taxonomic position.
Lepidoderma didermoides Kowalski, Mycologia
63(3): 503 (1971). Fig. 15a–j, Fig. 16a–h
Material examined: USA: Whatcom Co., Washington, 18
miles east of Glacier, 4050 ft, on living twigs, 22–VI–1970,
leg. D.T. Kowalski, D.T. Kowalski 10580 type, (UC).
Remarks – This rare species described by

Bol. Soc. Micol. Madrid 42. 2018 59
REVISION OF THE NIVICOLOUS SPECIES OF THE GENUS LEPIDODERMA
Fig. 14.– Lepidoderma crustaceum D.T. Kowalski 2643 type (MICH), a. Original label, b. Sporocarps, c. Details of the crystalline scales of
the peridium, d. Capillitium, e. Tips of the capillitium, f‒g. Spores, h. Details of the spore ornamentation. Scale bars: b = 5 mm, c = 100 µm,
d‒e = 50 µm, f‒g = 2 µm, h = 1 µm.
KOWALSKI (1971) is characterised by dispersed
to grouped sporocarps which are sessile, globose
to subglobose, reaching 1–1.5 mm in diam.; a
simple, cartilaginous, and brownish peridium
covered only sparsely with whitish lime scales;
and an abundant, rigid, dark brown, anasto-
mosed, and scantly branched capillitium, with
threads around 1 µm diam. and numerous dark
nodules. Spores are globose to subglobose, 14–17
µm diam., spinose.
Lepidoderma didermoides can be distin-
guished from L. chailletii by its cartilaginous
peridium, few lime scales on the peridium, and
its larger spores. However, the capillitium and
the ornamentation of the spores resemble those
of L. chailletii. The presence of large spores in
nivicolous myxomycetes is not unusual and may
be triggered by unfavourable environmental con-
ditions during development, which influences as
well the size and shape of peridial lime scales.
Consequently, we think that this is a poorly de-
fined species, and consider it to be a synonym
of L. chailletii, as indicated by MORENO & al.
(2004).
After studying the type material of Diache-
opsis spinosifila M.L. Farr & R.L. Critchf., we

G. MORENO, A. SÁNCHEZ, M. MEYER, Á. LÓPEZ-VILLALBA & A. CASTILLO
60 Bol. Soc. Micol. Madrid 42. 2018
think it is possibly a Lepidoderma with very few
lime scales. It has been treated as a synonym
of L. didermoides by MORENO & al. (2003)
and, as mentioned above, L. didermoides has
been considered a synonym of L. chailletii by
MORENO & al. (2004). In this work we agree
with these previous conclusions, considering D.
spinosifila a synonym of L. didermoides, and L.
didermoides a synonym of D. chailletii.
Fig. 15.– Lepidoderma didermoides D.T. Kowalski 10580 type (UC), a. Original label, b. Sporocarps, c. Sporocarps with few crystalline scales
and capillitium, e. Capillitium, f‒g. Details of the nodules of the capillitium, h‒i = Spores, j = Details of the spore ornamentation. Scale bars:
b = 5 mm, c‒d = 1 mm, e–f = 50 µm, g = 10 µm, h‒i = 2 µm, j = 1 µm.

Bol. Soc. Micol. Madrid 42. 2018 61
REVISION OF THE NIVICOLOUS SPECIES OF THE GENUS LEPIDODERMA
Fig. 16.– Diacheopsis spinosifila BPI 818265 holotypus, a. Original label, b. Sporocarps, c. Capillitium, d‒e. Details of the nodules of the
capillitium, f. Nodules of the capillitium, g. Spore, h. Details of the spore ornamentation. Scale bars: b = 1 mm, c–e = 50 µm, f = 5 µm, g = 2
µm, h = 1 µm.
Lepidoderma echinosporum G. Moreno, López-
Vill. & S.L. Stephenson, in Crous & al., Persoo-
nia 37: 231 (2016). Fig. 17a‒l
Material examined: India: Himachal Pradesh Province,
50 km N of Manali and S of Rohtang Pass, 32°20’23.9”
N and 77°13’08.2” E, elevation 3240 m, on dead semi–
woody stems near the margin of a melting snowbank along
National Highway 21, 19 May 2006, S. L. Stephenson 21862
(holotype AH 46061, isotype in BPI).
Remarks – Lepidoderma echinosporum has
been recently described in CROUS & al. (2016).
It is characterised by sporocarps with a stellate
dehiscence, a cylindrical columella which occu-

G. MORENO, A. SÁNCHEZ, M. MEYER, Á. LÓPEZ-VILLALBA & A. CASTILLO
62 Bol. Soc. Micol. Madrid 42. 2018
Fig. 17.– Lepidoderma echinosporum AH 46061 holotype, a‒b. Details of the sporocarps, dehiscence, cylindric pseudocolumella, and
crystalline scales, c. Crystalline scales of the peridium, d. Capillitium, e. Spores and capillitium, f. Crystalline scales of the peridium, g.
Inside a crystalline scale of the peridium, h. Capillitium, i–k. Spores, l. Details of the spore ornamentation. Scale bars: a‒b = 1 mm, c‒d
= 100 µm, e = 10 µm, f = 50 µm, g‒k = 5 µm, l = 2 µm.

Bol. Soc. Micol. Madrid 42. 2018 63
REVISION OF THE NIVICOLOUS SPECIES OF THE GENUS LEPIDODERMA
pies almost the entire sporotheca, an inner layer
of the peridium formed by prismatic and thick
plates of calcium carbonate, and dark spores that
are globose to subglobose, 15–17.7(–18) × 14–17
μm in size, including the prominent spines. Un-
der SEM the spore ornamentation consists of
long baculae which sometimes are fused into
short ridges and with coralloid nodules at the
apices.
Lepidoderma nevadense is macroscopically
similar but differs from L. echinosporum by the
elastic and reticulated capillitium, which forms
a dense, tight net, and smaller [13–14(–15) µm
diam.] and warted spores.
After studying the type of Lepidoderma pey-
erimhoffii, we found it to differ from L. echi-
nosporum by the darker, more rigid and more
fragile capillitium, widened at the edges where
threads merge into each other, and by shortly
spinose spores.
Lepidoderma granuliferum (W. Philips) R.E. Fr.,
Ark. Bot. 6(7): 3 (1906). Fig. 18a‒k
≡ Didymium granuliferum W. Phillips, Grevillea
5: 114 (1877).
≡ Badhamia granulifera (W. Phillips) Massee,
Monogr. Myxogastr. 321 (1892).
≡ L. carestianum var. granuliferum (W. Phillips)
G. Lister, in Schinz, Mitt. Naturwiss. Ges. Win-
terthur 6: 63 (1906).
Material examined: Spain: Madrid, Puerto de Navacerrada,
on twigs of Senecio pyrenaicus, 17–VI–1999, 2050 m, leg.
A. Sánchez, AH 19388. Ibidem, on twigs of Digitalis pur-
purea, 17–III–2000, 2025 m, AH 25654. Ibidem, 18–III–
2000, 2100 m, AH 32592. Ibidem, on twigs of Gentiana
lutea, 8–VI–2000, 2175 m, AH 25928. Ibidem, on leaves
of Gentiana lutea, 17–VI–2000, AH 25822. Ibidem, on
twigs, 3–VI–2001, 2125 m, AH 28547. Ibidem, on twigs
of Digitalis purpurea, 21–VI–2001, 2000 m, AH 26623, AH
33059. Ibidem, 30–VI–2001, 2200 m, AH 19464. Ibidem,
on twigs of Senecio pyrenaicus, 2150 m, AH 28579. Ibidem,
on twigs of Digitalis purpurea, 20–VII–2001, 2050 m, AH
26323. Ibidem, on living branches of Cytisus oromediter-
raneus, 18–V–2003, 2125 m, AH 32529. Segovia, Pico del
Lobo, Riaza, on vegetal debris, 14–V–2003, 1975 m, leg. A.
Sánchez, AH 32530. Ibidem, on stems of Poaceae, 1950 m,
AH 32513. Puerto de Navacerrada, on stems of Poaceae,
1–V–1997, 2100 m, leg. A. Sánchez, AH 19504. Ibidem,
on living branches of Cytisus oromediterraneus, 31–V–
1998, AH 32574. Ibidem, on twigs of Senecio pyrenaicus,
27–V–1999, 2150 m, AH 19398. Ibidem, 7–VI–1999, 2000
m, AH 19379. Ibidem, 2050 m, AH 19311. Ibidem, 2100
m, AH 19380. Ibidem, 8–VI–2000, 2125 m, AH 25899.
Ibidem, 2150 m, AH 25901. Ibidem, 11–VI–2000, 2050 m,
AH 25931, AH 25935, AH 32543. Ibidem, 2075 m, AH
32575. Ibidem, on living branches of Cytisus oromediter-
raneus, 14–VI–2000, 2175 m, AH 32538. Ibidem, on fronds
of Cryptogamma crispa, 16–III–2001, 2150 m, AH 25649.
Ibidem, on twigs of Rubus idaeus, 23–IV–2001, 1925 m,
AH 32573. Ibidem, on branches of Cytisus oromediter-
raneus, 6–VI–2001, 2100 m, AH 32564. Granada, Sierra
Nevada, on processed wood, 13–VIII–2001, 2700 m, leg. A.
Sánchez, AH 29292. Ibidem, Hoya de la Mora, on Carduus
carlinoides, 14–VIII–2001, 2750 m, AH 29289. Ibidem,
2600 m, AH 29290. Ibidem, 2750 m, AH 29288, AH 29291.
Ibidem, Carihuela, on vegetal debris, 5–VII–2016, 3100 m,
leg. G. Moreno, A. Sánchez & L. Monje, AH 46328, AH
46329, AH 46329.
Sporocarps variable, usually sinuous plasmo-
diocarps but more rarely sporangiate forms in
dense groups, 5–14 × 1–3 mm, cream-white to
greyish-white. Peridium double, the inner layer
membranous, transparent, and iridescent; and
the outer layer brown, covering the membrane
as a crust of whitish to cream-white lime scales,
variable in size and shape, with sometimes a few
interruptions revealing the inner membraneous
layer of the peridium. Capillitium abundant,
filamentous, threads of the capillitium pale to
slightly yellow, paler at the tips, with a lot of
membranous and asteriform expansions with
globose lime nodules inside. Spores violaceous-
brown in transmitted light, globose to subglo-
bose, 15–18 µm diam., with dense and short
spinules. Under SEM, the asteriform expansions
and the lime nodules of the threads of the capil-
litium are smooth, and the ornamentation of the
spores consists of short baculae.
Remarks – Lepidoderma granuliferum is char-
acterised by sinuous plasmodiocarps occurring
in small groups, a double peridium, a filamen-
tous capillitium with threads creating an intricate
net with abundant globular nodules covered by
an asteriform membrane, which is smooth under

G. MORENO, A. SÁNCHEZ, M. MEYER, Á. LÓPEZ-VILLALBA & A. CASTILLO
64 Bol. Soc. Micol. Madrid 42. 2018
Fig. 18.– Lepidoderma granuliferum AH 29288, a. Plasmodiocarps, b. Details of the lime nodules of the capillitium, c‒e. Smooth and as-
teriform nodules of the capillitium under LM, f. Plentiful nodules of the capillitium (AH 32592). g‒h. Smooth and asteriform nodules of the
capillitium under SEM (AH 32592), i‒j. Spores, k. Details of the spore ornamentation. Scale bars: a = 1 mm, b = 100 µm, c‒e = 20 µm, f = 20
µm, g = 10 µm, h = 5 µm, i‒j = 2 µm, k = 1 µm.

Bol. Soc. Micol. Madrid 42. 2018 65
REVISION OF THE NIVICOLOUS SPECIES OF THE GENUS LEPIDODERMA
LM and SEM, and spores that are globose to sub-
globose, 15–18 µm diam. and densely spinose.
KOWALSKI (1971) described and under-
stood this species correctly. However, he included
Amaurochaete minor Sacc. & Ellis as a synonym.
For us Lepidoderma granuliferum and A. minor are
different species, and we consider the latter as a
synonym of L. carestianum, as mentioned above.
The most common mistake in the interpreta-
tion of Lepidoderma carestianum is to use it as
a prioritary name for L. granuliferum (LADO
& RONIKIER, 2008; LADO, 2005–2018), but
these two taxa represent different species. We
consider them as different since L. carestianum
has nodules in the capillitium, its threads are not
as intricate as in L. granuliferum, and the nodules
are more irregular than in L. granuliferum, they
are along the threads, they are not covered by
an asteriform membrane, and they have a subtle
ornamentation consisting of thin lines or small
crests visible under SEM.
LISTER & G. LISTER (1911) considered
Lepidoderma granuliferum as a variety of L. car-
estianum. NEUBERT & al. (1995) considered L.
granuliferum and L. carestianum sensu Lister as
synonyms, while on the other hand they consid-
ered L. carestianum as the prioritary name of L.
chailletii. KOWALSKI (1971) and POULAIN &
al. (2011) considered these species as independ-
ent, and we follow their point of view.
Lepidoderma neoperforatum Kuhnt, Ber. Bayer.
Bot. Ges. 87: 99 (2017). Fig. 19a‒j
Material examined: Germany, Bayern, Bayerischer
Wald, Bayerisch–Eisenstein, nahe Ruckenwies, Höhe ca.
1200 m, MTB 6854/4; Fichtenwaldrand, auf anhängen-
den, lebenden und abgestorbenen, dünnen Äst chen von
Vaccinium myrtillus, in Vergesellschaftung mit Physarum
albescens Ellis ex T. Macbr. Und Lepidoderma chailletii
Rostaf., 4˗V˗2004, leg. A. Kuhnt. Holotypus: M–0046559,
Botanische Staatssammlung München (M). Isotypen: M–
0046560, GZU 313862, Hb. W. Nowotny (Now 13034), Hb.
Mar. Meyer (MM48350), Hb. Kuhnt (HK 040504–10, duplo
in AH 49122). Norwegen, Provinz Aust–Agder, Kommune
Bykle, Hovden, Wanderpfad Richtung Hovdenuten, Höhe
ca. 900m, auf anhängenden, lebenden Ästchen von Betula
sp., 3˗VI˗2012, leg. A. Kuhnt, (HK 120603–24, duplo in
AH 49123).
Sporocarps plasmodiocarpous, greyish-white,
slightly shining or pearly. Peridium thick, dou-
ble, outer peridium membranous, thin, with lime
scales; inner peridium dark reddish-brown to
blackish with small perforations, these 3–5 µm
in diam., these marking the points where capil-
litial threads touch the peridium. Capillitium
abundant, threads rigid, attached to the base of
the plasmodiocarp and to the inner peridium,
dark brown, paler in the tips, 1–3 µm diam., al-
most smooth, and sometimes with small and dark
nodules. Spores dark brown to blackish in mass,
brownish in transmitted light, with a paler side
and with a thinner germination wall, globose to
oval, 12–14.5 µm diam. or 14–15 × 12–13 µm,
spinose. Under SEM the ornamentation of the
spores consists of dense and tight spines.
Remarks – According to KUHNT (2017)
Lepidoderma neoperforatum is characterised
by robust whitish-grey plasmodiocarps with an
outer peridium consisting of small, compact,
and non-crystalline calcareous lime scales that
do not detach. Capillitium and spores are dark
brown, and the most remarkable feature are the
perforations 2.5–5 µm in diam. that the capil-
litial threads leave when detaching from the pe-
ridium. Revising the isotype of this species (HK
040504–10, duplo in AH 49122) we confirmed by
LM and SEM the size of these perforations.
This species can be confused with other plas-
modiocarpous members of the genus Lepido-
derma such as L. aggregatum, L. carestianum
or L. perforatum. In their study, MORENO &
al. (2004) mentioned that L. aggregatum does
not form elongated plasmodiocarps and that the
inner membranous peridium is hyaline to irides-
cent.
Lepidoderma carestianum shows scanty lime
scales on the outer peridium which do not form
a crust. The inner peridium is dark brown as in
L. neoperforatum but lacks perforations.
Finally, Lepidoderma perforatum is macro-
scopically the most similar species, showing a
paler, greyish to yellowish-violaceous brown in-
ner peridium; the lime scales of the outer perid-
ium are larger and not pearl-white. Nevertheless,
the diameter of the spores and the peridial per-
forations of L. perforatum and L. neoperforatum

G. MORENO, A. SÁNCHEZ, M. MEYER, Á. LÓPEZ-VILLALBA & A. CASTILLO
66 Bol. Soc. Micol. Madrid 42. 2018
Fig. 19.– Lepidoderma neoperforatum HK04504–10 isotype, a. Plasmodiocarps, b. Details of the small boreholes of the peridium, c.
Capillitium, d. Spores, e. Capillitium under SEM, f. Details of the boreholes of the peridium under SEM, g. Remainders of the capil-
litium in the peridium, h‒i. Spores, j. Details of the spore ornamentation. Scale bars: a = 1cm, b‒c = 50 µm, d‒g = 10 µm, h‒i = 2 µm,
j = 1 µm.

Bol. Soc. Micol. Madrid 42. 2018 67
REVISION OF THE NIVICOLOUS SPECIES OF THE GENUS LEPIDODERMA
are quite similar. Under SEM the ornamentation
of the spores differs, being spinose in L. neop-
erforatum and warted with coralloid apices in
L. perforatum. In summary, we judge the dif-
ferences between these two species as difficult
to comprehend. As such, molecular studies are
needed to make a decision if these are indeed
separate species.
Lepidoderma nevadense G. Moreno, Ant.
Sánchez, Mar. Mey., López-Vill. & A. Castillo,
sp. nov. Fig. 20a‒n
MycoBank: MB 827960
Etymology: Latin nevadense, referring to its
collection in the National Park of Sierra Nevada
from Spain.
Typus. Spain: Granada, Parque Nacional Sierra Nevada,
Jardín Botánico Universitario, Albergue Universitario, 2500
m., on debris of endemic plants (herbaceous and shrubs),
leg. A. Sánchez, A. Castillo & G. Moreno, 20–VI–2013, AH
46324 Holotypus, Isotypus MM 48610 and BPI.
Other material examined: Spain: Granada, Parque Nacional
Sierra Nevada, Jardín Botánico Universitario, Albergue
Universitario, 2500 m., on debris of endemic plants (her-
baceous and shrubs), leg. A. Sánchez, A. Castillo & G.
Moreno, 20–VI–2013, AH 46320, AH 46321, AH 46322,
AH 46323, AH 46325. Ibidem, on Helianthemum panno-
sum, leg. Á. López-Villalba & J.F. Moreno, 4–VI–2018, AH
50263 (2 boxes).
Sessile sporocarps, rarely with a short and
blackish stalk, in large groups. Sporotheca 1.2–2
mm diam., globose to subglobose, with plates
which give it a tessellate appearance. Dehiscence
apical and irregular, which sometimes leaves a
star-like structure. Peridium brittle, easily de-
stroyed after dehiscence, triple, outer perid-
ium consisting of a reddish-brown to yellow-
ish-brown cover of calcium carbonate scales,
inner peridium consists of a white to cream-
white cover of calcium carbonate scales 30–45
× 25–37 µm in size. The intermediate layer is
thin, membranous, hyaline, iridescent, and in
contrast to Lepidoderma peyerimhoffii is visible
only in immature sporocarps. Pseudocolumella
cylindrical to club-shaped, not always present,
cream-white. Hypothallus membranous and
pale brown. Capillitium filamentous, abundant,
brown, quickly falling apart when manipulated
(in this aspect it resembles that of the genus
Arcyria), reticulate, threads of the capillitium
forming a dense and crowded net, 1–4 µm diam.,
with fusiform (8–12 × 4–6 µm), triangular (18–
20 × 5–8 µm), or globose (5–7 µm) dark brown
nodules. The tips of the threads of the capillitium
are long, sharp, and slightly coloured. Spores
dark violaceous-brown in transmitted light with
a paler zone, globose to subglobose, 12–14(–15)
µm diam., with small and dense warts. Under
SEM the scales of the peridium are polyhedric,
and globular calcium carbonate does not appear;
the threads of the capillitium are smooth and re-
ticulated; and the spores show abundant, dense,
and short baculae which sometimes become at-
tached in short crests.
Remarks – Lepidoderma nevadense is recog-
nisable by the sessile sporocarps, with evident
peridial plates and a tessellate dehiscence, the
capillitium detaches totally from the sporothe-
ca and forms a very dense net with abundant
meshes; and the spores show flattened and dense
warts under LM.
Lepidoderma peyerimhoffii is a closely re-
lated species which can be easily distinguished
from L. nevadense by the filamentous and
slightly reticulate capillitium, and the spinose
spores. Under SEM the ornamentation of the
spores consists of longer baculae.
Lepidoderma echinosporum, a recently de-
scribed species from India (CROUSS & al., 2016)
is macroscopically similar to L. nevadense, but
L. echinosporum has very characteristic spores
with an echinulate ornamentation.
Lepidoderma perforatum Mar. Mey. &
Poulain, in Poulain, Meyer & Bozonnet, Bull.
Mycol. Bot. Dauphiné–Savoie 165: 6 (2002).
Fig. 21a‒k, Fig. 22l‒q
Material examined: France: La Bathie, Savoie, on Vac-
cinium myrtillus, 1300 m, 22–IV–1987, leg. M. Meyer,
Meyer 2696, duplo in AH 31773. La Bathie, Savoie, on
twigs, 1850 m, 22–V–2000, leg. M. Meyer & al., M. Meyer
21092, duplo in AH 49083.

G. MORENO, A. SÁNCHEZ, M. MEYER, Á. LÓPEZ-VILLALBA & A. CASTILLO
68 Bol. Soc. Micol. Madrid 42. 2018
Fig. 20.– Lepidoderma nevadense AH 46324 holotypus, a. Sporocarps, b. Dehiscence of the sporocarps, c. Crystalline scales of the
peridium, d. Crystalline scales of the peridium under LM, e. Reticulated capillitium, f. Nodules of the capillitium, g. Filamentous
tips of the capillitium, h. Spores. i. Crystalline scales of the peridium under SEM, j. Inside a crystalline scale of the peridium, k.
Reticulate capillitium, l‒m. Spores, n. Details of the spore ornamentation. Scale bars: a‒b = 1 mm, c = 0.2 mm, d = 50 µm, e‒g = 20
µm, h‒j = 10 µm, k = 5 µm, l‒m = 2 µm, n = 1 µm.

Bol. Soc. Micol. Madrid 42. 2018 69
REVISION OF THE NIVICOLOUS SPECIES OF THE GENUS LEPIDODERMA
Fig. 21.– Lepidoderma perforatum Meyer 2696, a. Plasmodiocarps, b. Details of the small boreholes of the peridium, c. Insertion
in Y-shape of the capillitium into the peridium, d. Insertion of the capillitium into the peridium, e. A tip of the capillitium uproots a
piece of the peridium, f. Spores, g. Details of the peridium and the capillitium, h. Details of the insertion of the capillitium into the
peridium, i–j. Peridium enclosing the tip of the capillitium, k. The capillitium is smooth under the covering peridium. Scale bars: a
= 1 cm, b–e = 50 µm, f = 10 µm, g = 50 µm, h = 10 µm, i–j = 5 µm, k = 1 µm.

G. MORENO, A. SÁNCHEZ, M. MEYER, Á. LÓPEZ-VILLALBA & A. CASTILLO
70 Bol. Soc. Micol. Madrid 42. 2018
Sporocarps plasmodiocarpous, isolated, 10–
70 × 1–5 mm, yellowish-white. Peridium dou-
ble, outer layer consisting of a continuous crust
of white lime scales, which is supported by a
hyaline membrane. Inner peridium membranous,
dark grey, not iridescent. Pseudocolumella ab-
sent. Capillitium abundant, consisting of dark
brown, filamentous threads 1–2 µm in diam., not
forming a reticulum, without nodules or rarely
with a few dark once, emerging from the base of
the plasmodiocarp and rising towards the perid-
ium, paler towards the tips. Tips of the threads
funnel-shaped or Y-shaped at their attachment
sites to the peridium, but easily detaching from
the inner peridium, leaving perforations eas-
ily visible in LM (3–5 µm diam.). Hypothallus
slightly developed. Spores blackish-brown in
mass, dark brown in transmitted light, globose
to subglobose even ellipsoid, 13–15 µm diam.
or 13–16 × 12.5–15 µm, spinose. Under SEM
the ornamentation of the spores is warted with
coralloid apices.
Remarks – Lepidoderma perforatum shows a
spinose ornamentation by LM which is similar to
the condition in L. carestianum, L. granuliferum
and L. neoperforatum. Lepidoderma carestianum
shows elongated or globular lime nodules on the
threads of the capillitium and L. granuliferum
has an intricate capillitium with abundant and
globular nodules covered by an asteriform and
smooth membrane. Lepidoderma neoperfora-
tum is at least closely related to L. perforatum.
KUHNT (2017) emphasized that both the size of
the spores and the size of the perforations of the
peridium are larger in L. perforatum. However,
we have examined specimens of both species,
and these two characters are very similar. Under
SEM the ornamentation of the spores is different,
spinose in L. neoperforatum, and warted with
coralloid apices in L. perforatum.
Fig. 22.– Lepidoderma perforatum Meyer 2696, l. Ornamentation of the inner face of the peridium, m–n. Capillitium with a granulose cover,
o–p. Spores, q. Details of the spore ornamentation. Scale bars: l = 2 µm, m–n = 5 µm, o–p = 2 µm, q = 1 µm.

Bol. Soc. Micol. Madrid 42. 2018 71
REVISION OF THE NIVICOLOUS SPECIES OF THE GENUS LEPIDODERMA
Lepidoderma peyerimhoffii Maire & Pinoy in
Maire, Patouillard & Pinoy, Bull. Soc. Hist, Nat.
Afrique N. 17:140(1926). Fig. 23a‒i, Fig. 24a‒k,
Fig. 25a‒d
≡ Diderma peyerimhoffii (Maire & Pinoy) H.
Neubert, Nowotny & K. Baumann, Myxomyceten
3: 78 (2000).
= D. nivale (Meyl.) Nowotny, H. Neubert & K.
Baumann in Neubert, Nowotny & Baumann,
Carolinea 49: 24 (1991).
= Diderma trevelyanii var. nivale Meyl., Bull.
Soc. Vaud. Sci. Nat. 50: 8 (1914)
Material examined: Switzerland: Le Chasseron, Canton
Vaud, on decaying conifer needles, 1580 m, VI–1913,
leg. CH. Meylan, CJB 6790 lectotype. Algeria: Oriental
Djurdjura, Aït–Ouaban forest, on dead leaves of Acer ob-
tusatum W. et K., in cliffs under snow, 25–V–1915, leg.
R. Maire & De Peyerimhoff. Holotype in MPU. Austria:
Innsbruck, near Bodensteinalm, 1661 m, on twig beside
snow, 10–V–2000, leg. H. Singer, AH 19624. France: col
de la Madeleine, Savoie, 1988 m, sur bois mort d’Alnus
viridis, 18–VI–2006, leg. M. Meyer, Meyer 36552 duplo in
Fig. 23.– Lepidoderma peyerimhoffii Holotype in MPU. a. Original label, b. Original colour drawing, c. One thread of the capillitium, d.
Capillitium, e. Tips and nodules of the capillitium, f. Spores, g‒h. Spores, i. Details of the spore ornamentation. Scale bars: c‒d = 10 µm, e =
50 µm, f = 10 µm, g‒h = 2 µm, i = 1 µm.

G. MORENO, A. SÁNCHEZ, M. MEYER, Á. LÓPEZ-VILLALBA & A. CASTILLO
72 Bol. Soc. Micol. Madrid 42. 2018
Fig. 24.– Lepidoderma peyerimhoffii Cainelli 11051101, a. Sporocarps, b. Details of the triple peridium (outer and inner), c. Details of
the crystalline scales of the outer peridium, d. Capillitium, e. Details of the ramification and the nodules of the capillitium, f. Spores, g.
Details of the membrane of the peridium and of the crystalline scales, h. Capillitium, i‒j. Spores, k. Details of the spore ornamentation.
Scale bars: a = 1 µm, b‒c = 0.5 mm, d = 50 µm, e‒f = 10 µm, g = 20 µm, h = 10 µm, i–j = 5 µm, k = 1 µm

Bol. Soc. Micol. Madrid 42. 2018 73
REVISION OF THE NIVICOLOUS SPECIES OF THE GENUS LEPIDODERMA
AH 49074. Col de la Madeleine, Savoie, 1776 m, sur bois
mort d’Alnus viridis, 12–VI–2008, leg. M. Meyer, Meyer
37686 duplo in AH 49073. Col de la Salette, Isère, 1635 m,
herbes sèches, nivicole, 26–IV–2004, leg. M. Meyer, Meyer
23643 in duplo AH 49075. Val d’ Isère, Isère, 2142 m,
herbes sèches, nivicole, 4–VI–2015, leg. M. Meyer, Meyer
47410 duplo in AH 49076. Bourg-Saint-Maurice, Les Arcs,
Savoie, 2022 m, Alnus viridis vivant, 24–IV–2017, leg. M.
Meyer & F. et J. Avrit, Meyer 48200 duplo in AH 49077.
La Pierre Saint–Mar tin, Arette Ski Resort, 26‒V‒2018, leg.
M. Tapia, 180526–008 in AH 50405. Germany: Bavaria,
Wettersteingebirge: Garmisch-Partenkirchen, 1690 m, open,
shrubby gavelly slopes, alder, shrub pine and meadows, 725
m SSW Kreuzalm cableway (upper station), E–exp. slopes
above the road from Kreuzeck (branch serpentine trail)
to Hochalm, below a rock crest, leg. M. Schnittler & M.
Borg Dahl, 8–V–2016, collect. M. Schnittler 29108, 29162,
29165, 29174. Italy: Colle dell’ Agnello, Cuneo, arbustes
vivants, 12–V–2011, leg. M. Meyer, Meyer 38918 duplo in
AH 49078. Chianale vs. Colle dell’Agnello, 2176 m, 11‒
V‒2011, leg. R. Cainelli, Cainelli 11051101, duplo in AH
49125. Spain: Granada, Parque Nacional Sierra Nevada,
Cauchiles, on Carduus carlinoides, 2750 m, 26–VII–2001,
leg. A. Sánchez, AH 30032. Ibidem, 14–VIII–2001, leg.
A. Sánchez & M. Sánchez, AH 29293, AH 46502. Ibidem,
Monachil, Barranco de San Juan, 2560 m, 4–VI–2018,
leg. Á. López-Villalba & J.F. Moreno, AH 50133. Huesca,
Bielsa, Plana Fonda, vegetal debris, 2‒V‒2015, leg. J.
Hernanz, AH 50433–1, AH 50433–2.
Sessile sporocarps, more rarely short stalked,
grouped in extended or scattered colonies.
Sporotheca 1.5–2 mm diam., globose to subglo-
bose, with polyhedric and brownish plates which
give it a tessellate appearance. Peridium triple,
outer layer reddish-brown, especially towards the
centre of the plate, with whitish calcium carbon-
ate scales which are especially prominent towards
the margins of a plate. Intermediate layer whitish,
covered by calcium carbonate scales. Inner layer
membranous, iridescent; it can be easily sepa-
rated from the intermediate layer and thus can
be easily observed. Dehiscence apical, tessellate,
and irregular, never leaving a star-like structure.
Pseudocolumella prominent, cylindrical to club-
shaped, dark cream. Hypothallus brownish and
membranous. Capillitium brownish, filamentous;
it does not fall apart from the sporotheca, threads
rigid, slightly anastomosed, bifurcated and paler
towards the tips, 2–3 µm diam., with a few glo-
Figura 25.– Lepidoderma trevelyanii var. nivale CJB 6790 lectotype, a‒c. Capillitium, d. Spores. Scale bars: a‒c = 50 µm, d = 10 µm.

G. MORENO, A. SÁNCHEZ, M. MEYER, Á. LÓPEZ-VILLALBA & A. CASTILLO
74 Bol. Soc. Micol. Madrid 42. 2018
KEY TO THE SPECES OF LEPIDODERMA
1.– Nivicolous species … 2
1’.– Non-nivicolous species … 13
2.– Plasmodiocarpic species … 3
2’.– Sporangiate species … 8
3.– With large nodules in the capillitium … 4
3’.– Without wide nodules in the capillitium … 5
4.– Capillitium asteriform with globose nodules, smooth under LM and SEM
Lepidoderma granuliferum
4’.– Nodules variable in shape, appearing smooth under LM and short-crested under SEM
Lepidoderma carestianum var. carestianum
5.– Without perforations in the peridium at the points where capillitial threads are attached to the
capillitium … 6
5’.– With perforations … 7
6.– Yellowish plasmodiocarps with a crust of lime scales Lepidoderma alpestroides
6’.– Greyish or reddish-brown plasmodiocarps with discernible lime scales
Lepidoderma carestianum var. pseudocarestianum
7.– Peridium yellowish-white with well-developed lime scales, inner peridium greyish
Lepidoderma perforatum
7’.– Peridium pearl-white with weakly developed lime scales, inner peridium reddish-brown
Lepidoderma neoperforatum
bose, ellipsoid or fusiform nodules. Spores dark
violaceous-brown in transmitted light, with a
paler zone, globose to subglobose, variable in
size depending on the collection (12–15 µm diam.
or 14.5–16 µm diam.), with dense spines. Under
SEM the ornamentation of the spores consists of
long baculae.
Remarks – Lepidoderma peyerimhoffii was
described by MAIRE & al. (1926) and it is
characterised by the tessellate sporocarps, the
triple peridium, the rigid, slightly anastomo-
sed, and breakable capillitium, and the spinose
spores.
We were able to study the type material of this
species, and to prepare a plate with images of the
capillitium, the spores under LM and SEM, and
we have added to the plate of the original colour
drawing done by Maire and conserved in MPU.
We noticed some differences observed in
material from Germany (Bavarian Alps) sent by
Martin Schnittler: Schnittler 29162 shows scales
of a Lepidoderma and the spores are violaceous
in transmitted light, with a paler zone, 12–15 µm
diam., and irregularly distributed spinules; and
the capillitium is filamentous, branched, 1–2 µm
diam., and pale brown. Schnittler 29165 shows
the same spores as the previous specimen, and
a filamentous capillitium, with some nodules,
more rigid than the previous one, and transver-
sal ramifications; the threads of 1–3 µm diam.,
brown with paler tips, ending in long and sharp
apices. Schnittler 29174 has violaceous spores,
with a paler zone, 13–15 µm diam., and irregu-
larly distributed spines. Capillitium plentiful,
threads 1–3 µm diam., brown, with paler tips,
anastomosed, with small and dark nodules. Scale
limes variable in size. Schnittler 29108 has vio-
laceous spores, with the same size as the first
two samples, with irregularly distributed spines.
Capillitium filamentous, with darker nodules just
like those indicated by POULAIN & al. (2011),
and transversal ramifications; the threads of 1–2
µm diam., very pale brown with paler tips.
Lepidoderma nevadense and L. echinospo-
rum are both close to L. peyerimhoffii. The
differences between the three species are ex-
plained above, see remarks relating to L. ne-
vadense.

Bol. Soc. Micol. Madrid 42. 2018 75
REVISION OF THE NIVICOLOUS SPECIES OF THE GENUS LEPIDODERMA
8.– Peridium tessellate ... 9
8’– Peridium not tessellate ... 11
9.– Spores strongly spinose Lepidoderma echinosporum
9’.– Spores warted or slightly spinose ... 10
10.– Spores slightly spinose, capillitium rigid, persistent, slightly reticulated. Spores 15–17 µm diam.
Lepidoderma peyerimhoffii
10’.– Spores warted, capillitium flexuous, easily falling apart, reticulated. Spores 13–14(–15) µm
diam. Lepidoderma nevadense
11.– Stalked sporocarps, with violaceous shades, peridium covered by a crust of lime scales. Spores
11–13 µm diam. Lepidoderma crustaceum
11’.– Sessile sporocarps, without violaceous shades, peridium covered by lime scales forming or not
a crust. Spores 12-14(-16) µm diam. … 12
12.– Peridium covered by discernible lime scales not forming a crust Lepidoderma chailletii
12’.– Peridium covered by lime scales forming a crust Lepidoderma aggregatum
13.– Typically associated with bryophytes … 14
13’.– Other types of habitats … 15
14.– Spores spinose, wide and flattened lime scales … Lepidoderma tigrinum
14’.– Spores warted, small and granulose lime scales … Lepidoderma crassipes
14’’.– Spores with small crests … Lepidoderma cristatosporum
15.– On leaves, rarely in dead wood. Sporocarps short-stalked and with a star-like dehiscence
Lepidoderma trevelyanii
15’.– On dead wood. Sporocarps with a long stalk and without a star-like dehiscence. Spores with
thick scattered warts Lepidoderma stipitatum
15’’.– On dead wood. Sporocarps with a long stalk and without a star-like dehiscence. Spores spinose.
Lepidoderma tigrinum
TAXONOMIC CONCLUSIONS:
In this work we recognise the nivicolous taxa of
Lepidoderma listed below.
Lepidoderma aggregatum Kowalski, Mycologia
63(3): 511 (1971).
Lepidoderma alpestroides Mar. Mey. &
Poulain, in Poulain, Meyer & Bozonnet, Bull.
Mycol. Bot. Dauphiné-Savoie 165: 9 (2002).
Lepidoderma carestianum (Rabenh.) Rostaf.,
Sluzowce Monogr.: 188 (1874).
Lepidoderma carestianum var. pseudocaresti-
anum var. nov.
= L. carestianum sensu Poulain, Meyer & Bozon-
net, Les myxomycètes tome 2, fig. 399 (2011).
Lepidoderma chailletii Rostaf., Sluzowce Mon.:
189 (1874).
= L. didermoides Kowalski, Mycologia 63(3):
503 (1971).
Lepidoderma crustaceum Kowalski, Mycologia
59(1): 167 (1967).
Lepidoderma echinosporum G. Moreno, López-
Vill. & S.L. Stephenson, in Crous & al., Persoo-
nia 37: 231 (2016).
Lepidoderma granuliferum (W. Philips) R.E. Fr.,
Ark. Bot. 6(7): 3 (1906).
Lepidoderma neoperforatum Kuhnt, Ber. Bayer.
Bot. Ges. 87: 99 (2017).
Lepidoderma nevadense sp. nov.

G. MORENO, A. SÁNCHEZ, M. MEYER, Á. LÓPEZ-VILLALBA & A. CASTILLO
76 Bol. Soc. Micol. Madrid 42. 2018
Lepidoderma perforatum Mar. Mey. &
Poulain, in Poulain, Meyer & Bozonnet, Bull.
Mycol. Bot. Dauphiné-Savoie 165: 6 (2002).
Lepidoderma peyerimhoffii Maire & Pinoy in
Maire, Patouillard & Pinoy, Bull. Soc. Hist, Nat.
Afrique N. 17:140(1926).
= Diderma trevelyanii var. nivale Meyl., Bull.
Soc. Vaud. Sci. Nat. 50: 8 (1914).
= D. nivale (Meyl.) Nowotny, H. Neubert & K.
Baumann in Neubert, Nowotny & Baumann,
Carolinea 49: 24 (1991).
ACKNOWLEDGMENTS
We wish to express our gratitude to Steven
L. Stephenson, Martin Schnittler, and Renato
Cainelli for their revisions of the manuscript.
We acknowledge Martin Schnittler and Renato
Cainelli for providing some of the specimens,
as well as the brilliant photos taken by Renato
Cainelli. We thank Luis Monje and Ángel Pueblas
of the Department of Drawing and Scientific
Photography at the University of Alcalá for their
help in the digital preparation of the photographs.
We thank Antonio Priego and José Antonio
Pérez of the Electron Microscopy Service of the
University of Alcalá for their invaluable help
with the SEM. Thanks are extended to Jean-
Claude Malaval for the material of Lepidoderma
peyerimhoffii conserved in the herbarium MPU,
Caroline Loup, the curator responsible for the
MPU herbarium and the curators of B, BR, BPI,
NYBG, UC and WSRL herbarium, specially to
Javier Rejos, curator of the AH herbarium, for
his assistance with the specimens examined in the
present study Finally, we are thankful to “Junta
de Andalucía, Consejería de Medio Ambiente y
Ordenación del Territorio, Dirección General de
Gestión del Medio Natural y Espacios Protegidos”
and “Comunidad de Madrid, Subdirección General
de Espacios Protegidos, Dirección General
del Medio Ambiente, Consejería de Medio
Ambiente, Administración Local y Ordenación
del Territorio” for the access-permission to
“Parque Nacional de Sierra Nevada” and “Parque
Nacional de Guadarrama”, and for the license to
collect and study nivicolous Myxomycetes.
BIBLIOGRAPHY
CROUS, P.W., M.J. WINGFIELD, T.I.
BURGESS, G.E.ST.J. HARDY, C. CRANE,
S. BARRETT, J.F. CANO-LIRA, J.J. LE
ROUX, R. THANGAVEL, J. GUARRO, J.P.Z.
SIQUEIRA, C.M., C.M. SOUZA-MOTTA,
S.L. STEPHENSON, D.A. SUTTON,
N. TAMAKEAW, M.T. TELLERIA, N.
VALENZUELA-LOPEZ, A. VILJOEN, C.M.
VISAGIE, A. VIZZINI, A.M. STCHIGEL,
F. WARTCHOW, B.D., B.D. WINGFIELD,
E. YURCHENKO, J.C. ZAMORA, J.Z.
GROENEWALD, M.P. MARTÍN, D.S.
ALFREDO, P.A. BARBER, R.W. BARRETO,
I.G. BASEIA, J. CANO-CANALS, R.
CHEEWANGKOON, R.J. FERREIRA,
J. GENÉ, C. LECHAT, G. MORENO, F.
ROETS, R.G. SHIVAS, J.O. SOUSA, Y.P.
TAN, N.P. WIEDERHOLD, S.E. ABELL,
T. ACCIOLY, J.L. ALBIZU, J.L. ALVES,
Z.I. ANTONIOLLI, N. APLIN, J. ARAÚJO,
M. ARZANLOU, J.D.P. BEZERRA, J.-
P. BOUCHARA, J.R. CARLAVILLA, A.
CASTILLO, V.L. CASTROAGUDÍN, P.C.
CERESINI, G.F. CLARIDGE, G. COELHO,
P.C. CERESINI, G.F. CLARIDGE, G.
COELHO, V.R.M. COIMBRA, L.A. COSTA,
K.C. DA CUNHA, S.S. DA SILVA, R.
DANIEL, Z.W. DE BEER, M. DUEÑAS, J.
EDWARDS, P. ENWISTLE, P.O. FIUZA, J.
FOURNIER, D. GARCÍA, T.B. GIBERTONI,
S. GIRAUD, M. GUEVARA–SUAREZ, L.F.P.
GUSMÃO, S. HAITUK, M. HEYKOOP,
Y. HIROOKA, T.A. T.A. HOFMANN,
J. HOUBRAKEN, D.P. HUGHES, I.
KAUTMANOVÁ, O. KOPPEL, O. KOUKOL,
J. T.A. HOFMANN, J. HOUBRAKEN, D.P.
HUGHES, E. LARSSON, K.P.D. LATHA,
D.H. LEE, D.O. LISBOA, W.S. LISBOA,
Á. LÓPEZ-VILLALBA, J.L.N. MACIEL,
P. MANIMOHAN, J.L. MANJÓN, S.
MARINCOWITZ, T.S. MARNEY, M.
MEIJER, A.N. MILLER, I. OLARIAGA,
L.M. PAIVA, M. PIEPENBRING, J.C.
POVEDA-MOLERO, K.N.A. RAJ, H.A.
RAJA, A. ROUGERON, I. SALCEDO, R.
SAMADI, T.A.B. SANTOS, K. SCARLETT,

Bol. Soc. Micol. Madrid 42. 2018 77
REVISION OF THE NIVICOLOUS SPECIES OF THE GENUS LEPIDODERMA
K.A. SEIFERT, L.A. SHUTTLEWORTH,
G.A. SILVA, M. SILVA (2016). Fungal
Planet description sheets: 469–557. Persoonia
37: 218–403. http://dx.doi.org/10.3767/
003158516X694499
KOWALSKI, D.T. (1971). The genus Lepido-
derma. Mycologia 59(1): 167–171.
KUHNT, A. (2017). Bemerkenswerte
Myxomycetenfunde: Neue Arten, Neukom-
binationen und Nachweise seltenerArten. Ber.
Bayer. Bot. Ges. 87: 93–128.
LADO, C. (2005–2018). An on line nomenclatur-
al information system of Eumycetozoa. Real
Jardín Botánico, CSIC. Madrid, Spain. http://
www.nomen.eumycetozoa.com (22–6–2018).
LADO, C. & A. RONIKIER (2008). Nivicolous
myxomycetes from the Pyrenees: notes
on taxonomy and species diversity. Part 1.
Physarales and Trichiales. Nova Hedwigia
87(3–4): 337–360.
LISTER, A. & G. LISTER (1911). A monograph
of the Mycetozoa. A descriptive catalogue of
the species in the Herbarium of the British
Museum. Second edition. London, Witherby
& Co. 302 pp., 200 plates.
LISTER, A. & G. LISTER (1925). A monograph
of the Mycetozoa. A descriptive catalogue of
the species in the Herbarium of the British
Museum. Oxford University Press 222 p.
MAIRE, R., R. PATOUILLARD & P. PINOY
(1926). Myxomycètes de l’Afrique du Nord.
Bull. Soc. Hist. Nat. Afrique N. 17: 38–43.
MARTIN G.W. & C.J. ALEXOPOULOS (1969).
The Myxomycetes. Iowa Univ. Press. 561
pp.
MORENO, G. & J.L. MANJÓN (2010). Guía de
hongos de la Península Ibérica. Barcelona.
Ediciones Omega.
MORENO G., C. ILLANA & H. SINGER (2003).
Diacheopsis spinosifila, a synonimum of
Lepidoderma didermoides. Mycotaxon 88:
123–128.
MORENO G., H. SINGER & C. ILLANA (2004).
A taxonomic review on the nivicolous myxo-
mycetes species described by KOWALSKI. II.
Physarales and Trichiales. Österr. Z. Pilzk.
13: 61–74.
NEUBERT H., W. NOWOTNY & K. BAU-
MANN (1995). Die Myxomyceten. Band II.
Physarales. Karlheinz Baumann Verlag. Go-
maringen. 368 pp.
POULAIN, M., M. MEYER & J. BOZONNET
(2002). Deux espèces nouvelles de myxo-
mycètes: Lepidoderma alpestroides et
Lepidoderma perforatum. Bull. Mycol. Bot.
Dauphiné-Savoie 165: 5–18.
POULAIN, M., M. MEYER & J. BOZONNET
(2011). Les myxomycètes tomes 1–2. Fédération
mycologique et botanique Dauphiné-Savoie,
Sevrier, France.
RABENHORST, L. (1862). Fungi Europaei
Exsiccati. Klotzschii Herbarii, vivi Mycologici
continuation. Editio nova. Series secunda.
Centuria V. Cura Dr. L. Rabenhorst. Dresdae,
Typis Caroli Heinrich.
ROSTAFINSKI, J. (1873). Versuch eines Systems
des Mycetozoen. Inaugural-Dissertation,
Strassburg. 21 pp.
SÁNCHEZ, A., G. MORENO, C. ILLANA &
H. SINGER (2002). A study of nivicolous
Myxomycetes in Southern Europe, Sierra de
Guadarrama, Spain. Persoonia 18(1): 71–84.
SHCHEPIN, O.N., Y.K. NOVOZHILOV & M.
SCHNITTLER (2016). Disentangling the
taxonomic structure of the Lepidoderma
chailletii-carestianum species complex
(Myxogastria, Amoebozoa): genetic and mor-
phological aspects. Protistology 10(4): 117–
129. doi:10.21685/1680-0826-2016-10-4-