A taxonomic evaluation of the stipitate Licea species

Abstract

A taxonomic study was made of the type specimens of 21 stipitate Licea species. Relevant characters were examined by light microscope and by SEM. Evidence of synonymy was found in five of the taxa, Licea tropica with L. bulbosa, L. cristallifera with L. eleanorae, L. longa and L. capitata with L. floriformis var. aureospora, and L. tanzanica with L. scyphoides. One taxon Licea capitatoides var. fujiokana is recombined as a variety of a different species, Licea rugosa var. fujiokana. A new name Licea verrucospora, and a new status was given to Licea scyphoides var. reticulata. One species, Licea takahashii was excluded as it is an immature form of another genus. Detailed standardized descriptions are made of each species examined, with comments on the most relevant taxonomic characters. Light and scanning electron micrographs of relevant morphological details are included. Nomenclatural information is given for each taxon. A key to the stipitate Licea species is proposed.

Full text

Fungal Diversity
A taxonomic evaluation of the stipitate Licea species
D. Wrigley de Basanta
and C. Lado
∗
Real Jardín Botánico de Madrid, CSIC. Plaza de Murillo, 2, 28014 Madrid, Spain
Wrigley de Basanta, D. and Lado, C. (2005). A taxonomic evaluation of the stipitate Licea
species. Fungal Diversity 20: 261-314.
A taxonomic study was made of the type specimens of 21 stipitate Licea species. Relevant
characters were examined by light microscope and by SEM. Evidence of synonymy was found
in five of the taxa, Licea tropica with L. bulbosa, L. cristallifera with L. eleanorae, L. longa
and L. capitata with L. floriformis var. aureospora, and L. tanzanica with L. scyphoides. One
taxon Licea capitatoides var. fujiokana is recombined as a variety of a different species, Licea
rugosa var. fujiokana. A new name Licea verrucospora, and a new status was given to Licea
scyphoides var. reticulata. One species, Licea takahashii was excluded as it is an immature
form of another genus. Detailed standardized descriptions are made of each species examined,
with comments on the most relevant taxonomic characters. Light and scanning electron
micrographs of relevant morphological details are included. Nomenclatural information is
given for each taxon. A key to the stipitate Licea species is proposed.
Key words: distribution, Liceales, Myxomycetes, nomenclature, taxonomy, type collections.
Introduction
The genus Licea (order Liceales, Myxomycetes) currently encompasses
more than 65 species of worldwide distribution (Lado, 2001). The original
description of the genus Licea was by Schrader at the end of the 18
th
century.
In his Nova Genera Plantarum, Schrader (1797: 16) established the “character
essentialis” and the “character generalis” of the genus and included
descriptions of and comments on four species, L. variabilis Schrad. and L.
pusilla Schrad., now accepted as autonomous species, and L. tubulina Schrad.
and L. clavata Schrad., which are now considered synonyms of Tubulifera
arachnoidea Jacq. [= Tubifera ferruginosa (Batsch) J.F. Gmel.] (Lado, 2001).
Since then the genus has been maintained, although it is an unnatural genus
(Gilert, 1994) since it is defined mainly by the lack of one of the most
informative characters of myxomycetes, the capillitium.
The taxonomic treatment of the genus has been changed over the years.
The type of dehiscence of the peridium and the presence or absence of a stalk
∗
Corresponding author: C. Lado; e-mail: lado@ma-rjb.csic.es
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have been used in the past as the most relevant characters, and served to
segregate the genus Orcadella Wingate (1889), based on sporocarps which
were stalked and dehisced by a membranous lid (Lister, 1925), and the genus
Hymenobolina Zukal (1893), based on sessile sporocarps opening by a
membranous lid (Lister, 1925). Even these characters, are of questionable
taxonomic relevance, as Lado and Pando (1997: 109) recognized, and
characters which are important at first glance, and used occasionally to
distinguish species, can be unstable within taxa or even within fructifications.
Nannenga-Bremekamp (1965), in a revision of the genus also
emphasized the type of dehiscence and recognized within the genus Licea three
subgenera, the subgen. Orcadella (Wingate) Nann.-Bremek., which dehisces
by means of a circular or almost circular lid. The subgen. Pleiomorpha Nann.-
Bremek., cracks open irregularly or by a split, and the subgen. Licea, opens
along ridges that divide the peridium in platelets. With the descriptions of
recent new species, the limits between these subgenera are unclear, and the
proposal of Nannenga-Bremekamp is now of doubtful application.
Martin and Alexopoulos (1969) recognized 19 species in the genus and
this total was increased eight years later by Keller and Brooks (1977) to 30. In
the latter classic contribution towards a monograph of the genus the authors
established the bases for a more rational study of the group, and elaborated a
detailed key of the species that the authors recognised. In this paper all the
species were placed in a single genus Licea, the subgenera proposed by
Nannenga-Bremekamp were not recognised, and only 5 truly stipitate species
were included.
Since then Lakhanpal et al. (1990) added three new stipitate species in
their paper “Notes on Licea (Myxomycetes) from India”, and at present, among
the 67 recognized species belonging to this genus (Lado, 2001), a large number
(27) of stipitate Licea have been described in the literature (Table 1).
The current concept of the genus (Lado and Pando, 1997) includes
species with plasmodiocarpic to sporocarpic, sessile or stipitate minute
sporophores, with a peridium that is membranous or coriaceous, consisting of
one or two layers. A columella, capillitium and pseudocapillitium are always
absent. On account of this, species and specimens that should be placed under
the genus Didymium (order Physarales) or Perichaena (order Trichiales) are
sometimes erroneously identified as belonging to Licea, if they lack capillitium
when the specimens develop under unfavourable conditions.
The number of recent descriptions of stipitate taxa (Table 1), some of
them practically simultaneously in press (Ing, 1999; Flatau, 2000), coupled
with the simplicity of the sporophores and lack of taxonomic characters, made
the systematics of these minute myxomycetes somewhat confused. The fact
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Table 1. Stipitate Licea described in the literature in chronological order of their description.
Name Reference
Licea operculata (Wingate) G.W. Martin, described as
Orcadella operculata Wingate, 1889
Wingate (1889); Martin (1942)
Licea pedicellata (H.C. Gilbert) H.C. Gilbert, described as
Hymenobolina pedicellata H.C. Gilbert, 1934
Gilbert (1934); Martin (1942)
Licea erecta K.S. Thind & Dhillon Thind and Dhillon (1967)
Licea scyphoides T.E. Brooks & H.W. Keller Keller and Brooks (1977)
Licea perexigua T.E. Brooks & H.W. Keller Keller and Brooks (1977)
Licea lucens Nann.-Bremek. Nannenga-Bremekamp (1981)
Licea capitata Ing & McHugh Ing (1982)
Licea atricapilla Nann.-Bremek. & Y. Yamam. Nannenga-Bremekamp and
Yamamoto (1983)
Licea erectoides Nann.-Bremek. & Y. Yamam. = Licea
erecta var. erectoides (Nann.-Bremek. & Y. Yamam.) Y.
Yamam.
Nannenga-Bremekamp and
Yamamoto (1983); Yamamoto
(1998)
Licea bulbosa Nann.-Bremek. & Y. Yamam. Nannenga-Bremekamp and
Yamamoto (1987)
Licea rugosa Nann.-Bremek. & Y. Yamam. Nannenga-Bremekamp and
Yamamoto (1987)
Licea capitatoides Nann.-Bremek. & Y. Yamam. Nannenga-Bremekamp and
Yamamoto (1990)
Licea lilacina Nann.-Bremek., T.N. Lakh. & R.K. Chopra Lakhanpal et al. (1990)
Licea scyphoides var. reticulata T.N. Lakh., Nann.-
Bremek. & R.K. Chopra
Lakhanpal et al. (1990)
Licea floriformis T.N. Lakh. & R.K. Chopra Lakhanpal et al. (1990)
Licea floriformis var. aureospora M.T.M. Willemse &
Nann.-Bremek.
Willemse and Nannenga-
Bremekamp (1994)
Licea tropica Chao H. Chung & C.H. Liu Chung and Liu (1996)
Licea tanzanica Ukkola, Härk. & Gilert Ukkola et al. (1996)
Licea poculiformis Ukkola Ukkola (1998)
Licea erddigensis Ing Ing (1999)
Licea eleanorae Ing Ing (1999)
Licea crateriformis Ing Ing (1999)
Licea cristallifera Flatau Flatau (2000)
Licea longa Flatau Flatau (2000)
Licea capitatoides var. fujiokana Y. Yamam. Yamamoto (2000)
Licea parvicapitata Y. Yamam. Yamamoto (2000)
Licea takahashii Y. Yamam. Yamamoto (2000)
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that the species of Licea are very small (about 0.1 to 1 mm in height) and
mainly the result of moist chamber culture, which means that there are often
few specimens to work with, and many aberrant forms, have further
complicated the picture. Gilert (1994) did ultrastructure studies on some
members of the genus, which clarified the relationships between some species,
but she studied mainly sessile species. She commented on the heterogeneous
nature of the genus as a whole and expressed doubt over the taxonomic
significance of the presence or absence of a capillitium, concluding that a
protoplasmodium with certain fructification morphotypes was more important.
The only stipitate species in her study was Licea operculata.
No comparative study of type material of the stipitate species, had been
done when this paper was begun, there were no keys to the numerous new
species recently described, some of the descriptions did not allow for easy
species identification, and so a taxonomic revision of these members of the
genus was deemed necessary. The contribution of Keller and Brooks (1977)
detailed most of the sessile species, of which 17 more have been described
since. The present paper attempts to clarify the similarities and differences
between the stipitate species, as a step towards further analysis of the genus
Licea as a whole.
Materials and methods
This revision is based on the study of almost 100 herbarium collections.
Type specimens of all the stipitate species in the literature (Table 1) were
requested, and 21 of the 27 listed in the table were examined simultaneously.
The remainder were requested, but were not sent. Type material was loaned by
the following herbaria BPI, BR, H, TNS and B, and material conserved in MA-
Fungi, our own collections (dwb, Lado), and those kindly lent to us by Dr. L.
Flatau (LF), Dr. B. Ing (Ing), Dr. F. Pando (Pando), Dr. M. Schnittler (sc), Ms.
M. de Haan (MdH) and Mr. A. Varela-García (VGA), were all studied in the
same manner. The same optical instruments, a Nikon SMZ-1000
stereomicroscope and a Nikon Eclipse E-600 microscope with a Nomarski
system, were used to examine the specimens. The same magnifications and the
same light intensity were used, to allow valid comparisons of such taxonomic
characters as spore and peridial colour, and ornamentation by transmitted light,
and in order to unify the terminology used for their description. Slides of most
of the type material were not included in the samples sent from the different
herbaria. Consequently slides were made by us in a standardized way by
mounting a whole sporocarp in PVA (polyvynil alcohol) and squashing it very
gently with a cover slip, just sufficient to encourage dehiscence while
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Fungal Diversity
maintaining the sporocarp whole when possible. Other diagnostic techniques
such as PCR were not possible due to the scant type material of some species
and the regulations of some herbaria. Light micrographs were made with
Nomarski optics, and SEM photographs were taken using the critical point
technique. The age and treatment of some herbarium specimens meant they did
not respond well to the critical point technique used for the SEM observations,
leaving some spores collapsed (Fig. 32). Colour notations in parentheses are
from the ISCC-NBS Color-Name Charts Illustrated with Centroid Colors
(Anonymous, 1976). The terminology used follows Lado and Pando (1997).
Results and discussion
As described by Martin and Alexopoulos (1969) and Martin et al. (1983:
41) the family Liceaceae, has a single genus, Licea. This genus includes
species with plasmodiocarpic to sporocarpic, sessile or stipitate sporophores,
with a peridium membranous or coriaceous, consisting of one or two layers,
then the external layer gelatinous when wet, drying horny, the inner always
membranous, the external surface frequently with deposits of granular material,
the peridium dehiscence can be irregular, by platelets or by lids; the columella,
the capillitium and the pseudocapillitium are always absent, and the spores are
free, globose, subglobose or ovoid, of variable colour but usually pale,
decorated or smooth, and with spore wall of uniform thickness or with a
thinner area. The nomenclatural treatment of the genus in this paper is
according to Lado (2001) and Hernández-Crespo and Lado (2005).
Licea Schrad., Nov. Gen. Pl.: 16. 1797.- Lectotype, Licea pusilla Schrad.
(designated by Martin, 1942: 700)
= Cylichnium Wallr., Fl. Crypt. Germ. 2: 267. 1833.- Type: Cylichnium operculatum
Wallr.
= Protoderma Rostaf., Sluzowce Monogr.: 90. 1874 [Nom. illeg., non Protoderma
Kütz., 1854]. Protodermium Rostaf. ex Berl. in Saccardo, Syll. Fung. 7: 328. 1888 [Nom. nov.,
based on Protoderma Rostaf.]. Protodermodium Kuntze, Revis. Gen. Pl. 2: 867. 1891 [Nom.
nov., based on Protoderma Rostaf.].- Type: Protoderma pusillum (Schrad.) Rostaf. (= Licea
pusilla Schrad.).
= Orcadella Wingate, Proc. Acad. Nat. Sci. Philadelphia 41: 280. 1889. Licea subgen.
Orcadella (Wingate) Nann.-Bremek., Acta Bot. Neerl. 14: 132. 1965.- Type: Orcadella
operculata Wingate.
= Hymenobolus Zukal, Oesterr. Bot. 43: 73. 1893 [Nom. illeg., non Hymenobolus
Durieu & Mont., 1845]. Hymenobolina Zukal, Oesterr. Bot. 43: 133. 1893 [Nom. subst., based
on Hymenobolus Zukal].- Type: Hymenobolus parasiticus Zukal.
= Kleistobolus C. Lippert, Verh. Zool.-Bot. Ges. Wien 44: Abh. 70. 1894.- Type:
Kleistobolus pusillus C. Lippert.
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= Licea subgen. Pleiomorpha Nann.-Bremek., Acta Bot. Neerl. 14: 132. 1965 [as
“Pleismorpha”].- Type: Licea variabilis Schrad.
The genus Licea has a worldwide distribution, and a total of 67 species
are now recognised (Lado, 2001). The species described in this genus as
stipitate were studied and are treated here in alphabetical order.
1. Licea bulbosa Nann.-Bremek. & Y. Yamam., Proc. Kon. Ned. Akad.
Wetensch., C 90(3): 324. 1987. TYPE: JAPAN, Kochi Pref., Aki-shi, Nabika,
developed on the bark of a living tree in a moist chamber, 28 VI – 16 V 1986,
Y.Y. 3919 (holotype: TNS; isotype NENB 15.055 now at BR!). (Figs. 1-7)
= Licea tropica Chao H. Chung & C.H. Liu, Proc. Natl. Sci. Council Republ. China, B
20(4): 140. 1996, syn. nov. TYPE: TAIWAN, Pingtung, Manchou Hsiang, Wan-li-te-shan,
about 120º50’E 22º2’N, tropical rain forest, on unidentified dead leaves of angiosperms
collected in 2 II 1996, cultured in a moist chamber from 4 II 1996-25 III 1996, fruiting bodies
appeared since 18 III 1996, C.-h. Chung M1000 (holotype: TAI).
Sporocarps scattered, stipitate, 150-280(-340) µm in height. Sporotheca
brown, subglobose, up to 160 µm diam. Hypothallus inconspicuous. Stalk up
to 180 µm in height, straight, subcylindrical, tapering slightly towards the
apex, opaque, nearly black by reflected light, especially at the base, mid-width
less than one third of the height, by transmitted light (TL) pale brown and filled
with granular material. Peridium double, except for the equatorial area where it
is single, in this area a clear ring of dehiscence is visible by reflected light
(Figs. 1-2), the top half to two thirds of the peridium lifts off as a lid with very
prominent protuberances on the inner edge, the outer layer gelatinous
continuous with the stalk, transparent in the lower half showing the spores
inside, brown by reflected light in the upper half with granular refuse material,
the inner layer membranous, smooth except for the area of dehiscence, where
there are some warts; dehiscence circumcissile and more or less equatorial.
Spores free, pale yellow or greyish green to almost hyaline (121. p. Y G-122.
gy. Y G) by TL, globose, 10-11 µm diam., smooth; spore wall of uniform
thickness (Fig. 6).
Material examined: JAPAN, Kochi Pref.: Aki-shi, Nabika, on bark of living tree in
moist chamber, 28-IV to 19-V-1986, NENB 15.055 (BR, isotype). CUBA, Sancti Spiritus,
Alturas de Banao, 21°52’39”N 79°36’03”W, on Cyathea woodwardiodes petioles in moist
chamber, 8-II-2003, dwb 2256.
Habitat: bark of living trees, angiosperm leaf litter, tree-fern petioles, ground litter.
Distribution: Australia, Cuba, Japan, Taiwan, Tanzania.
Illustrations: Nannenga-Bremekamp and Yamamoto (1987: Fig. 9); Chung and Liu
(1996: 141, Figs. 2-4) as L. tropica; Ukkola et al. (1996: 56, Figs. 2-4); Yamamoto (1998:
149); McHugh et al. (2003: 492, Fig. 2)
The sporocarps of the isotype of this species had a height and diam.
somewhat larger than the original description. The height was up to 280 µm vs.
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Fungal Diversity
Figs. 1-7. Licea bulbosa. 1-2. Habit. Note line of circumcissile dehiscence (isotype NENB
15055 BR). 3. Dehiscing sporocarp with transparent base to sporotheca and upper half lifting
as a lid (NENB 15055). 4. Whole sporocarp with transparent base to sporotheca containing
spores and band of refuse below the line of dehiscence (dwb 2256). 5. Dehisced sporocarp by
SEM (NENB 15055). 6. Spore by TL (NENB 15055). 7. Detail of outer edge of base of
sporotheca with warted edge (NENB 15055).
150-200 µm in the original description, and the diam. of the sporotheca was up
to 160 µm in the largest specimens (40-60 µm in the original description). The
stalk was light brown by TL with refuse deposits inside (Figs. 3-4). There was
a definite separation between the stalk and the sporotheca. The inner peridium
was smooth and transparent with an adhering outer layer containing refuse
particles, some of which can be crystals visible with Nomarski. It is difficult to
267

distinguish the two layers. The upper part of the peridium lifts off in one piece
almost like a lid (Figs. 3-4) with very prominent protuberances on the edge
(Fig. 7). The cup that is left has few refuse particles included (Fig. 5) and
appeared totally transparent.
We were not able to examine the type of Licea tropica, which was found
on unidentified dead angiosperm leaves in Wan-li-te-shan, Manchou Hsiang,
Pingtung, Taiwan (Chung and Liu, 1996). The material was requested, but not
sent. However according to the description, comments and illustrations by
Chung and Liu (1996: 141), the constant and very clear characters of the
transparent base to the sporotheca and the darkened brownish band below the
area of dehiscence (Fig. 4), were observed in material recently isolated from
moist chamber cultures of tree fern petioles from Cuba. The separation
between the outer peridial layer of the top half and the bottom half of the
sporotheca are other clear features. All these characters, except the brown
stalk, concur with the isotype of L. bulbosa, and so we agree with McHugh et
al. (2003: 491) and believe these to be the same species. These authors made
no comments in their synonymy of the species, but their photograph of
Australian material shows all the above-mentioned characters. In the
description of L. tropica, Chung and Liu (1996: 142) recognized the similarity
of their species with L. bulbosa, but found height and diam. differences of the
sporocarps. We found, in our examination of the isotype of L. bulbosa, that
there is an overlap in the measurements.
In the Cuban material in moist chamber, the equatorial thinner band was
observed as a light line in developing sporocarps as soon as the sporotheca
formed as a transparent bulb at the top of the stalk, before spore formation and
darkening of the upper half. It was as if the developing outer peridium is
stretched forming two halves, which are separated in the middle.
This species is also like L. scyphoides and L. eleanorae in its
circumcissile dehiscence, but differs from the former in its almost hyaline
smaller spores (10-11 µm vs. 11-14 µm in L. scyphoides), and from both in the
smooth spores, smooth inner peridium, and smooth transparent base to the
sporotheca.
2. Licea capitatoides Nann.-Bremek. & Y. Yamam., Proc. Kon. Ned. Akad.
Wetensch. 93(3): 269. 1990. TYPE: JAPAN, Tokushima Pref., Tokushima-shi,
Kamo, developed in a moist chamber on bark of a living Aphananthe aspera,
11-28 V 1988, Y. Yamamoto 6387 (holotype: TNS!, isotype NENB 16.136,
now at BR!). (Figs. 8-11)
Sporocarps scattered, stipitate, 200-400 µm in height. Sporothecae dark
grey-brown, globose or subglobose, 100-120 µm diam. Stalk 100-200(-280)
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Fungal Diversity
Figs. 8-11. Licea capitatoides (NENB 16.136). 8. Habit. 9. Inner surface of peridium stippled
with dispersed warts. 10. Dehisced sporocarp with v-shaped calyculus. 11. Spore by SEM with
dense even warts.
µm in height, concolourous and continuous with sporotheca, cylindrical,
expanded at the base, full of refuse, mid-width less than half the height.
Peridium double, outer gelatinous layer with granular refuse in patches, the
inner layer membranous yellowish grey (93. y. Gray) by TL, inner surface
stippled with very obvious warts; dehiscence by irregular fracture, into “wavy-
edged” fragments of the upper sporotheca, leaving a calyculus with a clear
inner separation from stalk (Fig. 10). Spores free, greyish yellow to pale yellow
(90. gy. Y-89. p. Y) by TL, 8-10 µm diam., smooth by TL, closely warted by
SEM (Fig. 11); spore wall with an obvious paler area.
Material examined: JAPAN, Tokushima Pref.: Tokushima-shi, Kamo, on bark of living
Aphananthe aspera in moist chamber, 11 to 28-V-1988, Y. Yamamoto 6387 (TNS, holotype)
as Licea capitatoides; also NENB 16.136 (BR, isotype)
Habitat: bark of living Aphananthe aspera.
Distribution: Known only from the type locality.
269

Illustrations: Nannenga-Bremekamp and Yamamoto (1990: 269, Figs. 3A-C);
Yamamoto (1998: 139).
This species is known only from the type locality. It is most similar in
habit to L. pedicellata, but differs from it in the ornamentation and smaller size
of the spores. By transmitted light the spores appear to be smooth, but by SEM
(Fig. 11) very dense close warts are visible which are smaller and denser than
the warts in L. pedicellata. The L. capitatoides also has paler spores which are
slightly smaller than those of L. pedicellata (8-10 vs. 11-13 µm diam.).
Another difference is that in L. capitatoides only the top part of the sporotheca,
not the whole sporotheca, breaks into platelets on dehiscence. This leaves a
large v-shaped calyculus showing the separation of the inner peridium from the
stalk. The separation visible in the L. pedicellata is a small flat disc shape. The
densely stippled inner surface of the peridium however, is very similar by SEM
(Fig. 9) in the two species (Fig. 68).
Licea capitatoides can be distinguished from L. scyphoides and L.
tanzanica on the basis of the mode of dehiscence (platelets vs. circumcissile),
the different spore ornamentation and darker smaller spores of L. capitatoides.
3. Licea eleanorae Ing, Myxomycetes Britain and Ireland: 50. 1999. TYPE:
SWITZERLAND, Ticino, Lugano, Cantine di Gandria, in cortice vivo
Platanus × hispanicae in camera humida, September 1997, B. Ing (holotype:
Hb. Ing No. 97097!) (Figs. 12-20)
= Licea cristallifera Flatau, Stapfia 73: 65. 2000., syn. nov. TYPE: GERMANY,
Hessen, Kassel, Borke, liegender Fagus-Stamm (F. sylvatica), (2), 170 m über NN., 3.-27.7
1998, LF 3148 (holotype: B!).
Sporocarps scattered, solitary stipitate, 360-480 µm in height. Sporotheca
shiny golden, 175-220 µm diam. Stalk 185-260 µm in height, dark brown by
reflected light, straight, sub-cylindrical, full of crystals and refuse matter,
continuous with outer layer of sporotheca, mid-width one third of the height.
Peridium double, outer layer dark and full of refuse, birefringent crystals on the
outer upper surface (Fig. 13), inner layer translucent and finely warted (Fig.
19); dehiscence by means of a circular apical split (Figs. 15-16). Spores free,
pale yellow (89. p. Y) by TL, subglobose 8-11 µm diam., smooth by TL (Fig.
14), with square or rhomboid ornamentation by SEM (Fig. 20); spore wall of
uniform thickness (Fig. 18).
Figs. 12-20. Licea eleanorae. 12. Habit (B. Ing 98136). 13. Dehiscing sporocarp showing
crystals on upper surface of sporotheca (B. Ing 97097). 14. Spores by TL with wall of uniform
diam. (B. Ing 97097). 15. Whole sporocarp (LF 3148 as Licea cristallifera Flatau). 16.
Dehiscing sporocarp showing inner peridium separate from stalk (LF 3148). 17. Dehisced
sporocarp and spores by SEM (LF 3148). 18. Spores by TL with wall of uniform diam. (LF
3242 Licea cristallifera Flatau). 19. Inner surface of peridium (LF 3148). 20. Spore
ornamentation by SEM showing rhomboid granules (LF 3148).
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271

Material examined: SWITZERLAND, Ticino: Lugano, Cantine di Gandria, on bark of
living Platanus × hispanicae in moist chamber, IX-1997, Ing 97097 (slide of the holotype).
ENGLAND, London: Holland Park, bark of Tilia in moist chamber, XII-1998, Ing 98136.
GERMANY, Hessen, Kassel, Fuldatal, Nähe Kragenhofbrücke, on bark of Fagus sylvatica, 3
to 27-VII-1998, LF 3148 (1) (B, isotype of L. cristallifera); LF 3148 (18) (isotype of L.
cristallifera); 8 to 20-VII-1999 LF 3242 (all specimens as L. cristallifera).
Habitat: bark of living trees.
Distribution: England, Germany, Scotland, Switzerland.
Illustrations: Ing (1999: 49, Fig. 24); Flatau (2000: 65, Fig. 1A-E as L. cristallifera)
The type material shows shiny sporocarps somewhat vase shaped with a
slightly flattened top to the sporotheca (Fig. 12). The most distinguishing
character apart from the crystals in the peridium and stalk is the inner peridium
holding the spores. By TL this looks like a clear bubble full of spores (Figs. 13,
16), surrounded by the outer crystalline layer which is continuous with the
stalk (Fig. 17). The appearance and characters of the type material of Licea
cristallifera, as well as the abundant material kindly sent to us by L. Flatau, are
identical to those of L. eleanorae. A range of sizes was seen in the sporocarps
of L. cristallifera. The former species was being described while the
description of L. eleanorae was in press which may account for the double
description. We synonymize these two species here.
Licea eleanorae is similar in its method of dehiscence to L. bulbosa, L.
tanzanica and L. scyphoides. It differs from them and from all the other
stipitate Licea species in having the upper half of the peridium covered with
birefringent crystals and the tiny rhomboid or square ornamentation of the
spores by SEM (Fig. 20).
4. Licea erddigensis Ing, Myxomycetes Britain and Ireland: 48. 1999. TYPE:
UNITED KINGDOM, Gallia septentrionalis, Wrexham, Erddig Park, in cortice
vivo Aceris pseudoplatanis in camera humida, March 1999, B. Ing (holotype:
Hb. B. Ing No. 99006). (Figs. 21-25)
Sporocarps solitary scattered, stipitate, 500-800 µm in height. Sporotheca
subglobose, brown, 250-500 µm diam. Hypothallus inconspicuous. Stalk black
by refelected light, furrowed, straight, cylindrical but narrower at the apex, full
of debris, mid-width one third of height, continuous with the sporotheca.
Peridium single, dark greyish yellow to mid yellow brown (91. d. gy. Y-77. m.
y Br), stippled with very faint flattened warts on the inner surface (Fig. 24);
dehiscence by irregular apical split. Spores free, very pale yellow to almost
colourless (89. p. Y-92. y White) by TL, subglobose, smooth, 10-12 µm diam.;
spore wall of uniform thickness (Fig. 25).
Material examined: UNITED KINGDOM, Wales: Wrexham, Erddig Park, on bark of
living Acer pseudoplatanus in moist chamber, II-1999, Ing 99005 (paratype).
Habitat: bark of living sycamore tree.
Distribution: Known only from the type locality.
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Fungal Diversity
Figs. 21-25. Licea erddigensis (B. Ing 99005). 21. Habit (photo D.W. Mitchell with
permission). 22. Sporotheca by TL showing irregular dehiscence. 23. Thin transparent
peridium. 24. Detail peridial surface with faint flat warts. 25. Spores with uniform thick wall
inside peridium.
Illustrations: Ing (1999: 49, Fig. 23)
This species has a very wide sporotheca and the stalk is gathered below it
(Fig. 22). This, the membranous peridium (Fig. 23), which tends to roll back
on itself, with very faint warts on the inner surface, and the large size of the
sporocarp make it very easy to distinguish this taxon from the other stipitate
species examined. The reddish-brown colour of the spores, described by Ing
(1999: 49) was not visible in the paratype examined, which had pale yellow to
colourless spores. The author suggests that this is a sign of immaturity.
This species looks very similar to the description of L. floriformis var.
floriformis (see below), which we requested, but were not able to obtain. In
comparison to the original description of the latter species (Lakhanpal et al.,
1990), it is different in its persistent dark peridium and mode of dehiscence.
Ing (1999) also distinguishes it from other stipitate species by the orange
colour of the sporocarp during development.
273

5. Licea erecta var. erectoides (Nann.-Bremek. & Y. Yamam.) Y. Yamam.,
Myxomycete Biota Japan: 130. 1998. Licea erectoides Nann.-Bremek. & Y.
Yamam., Proc. Kon. Ned. Akad. Wetensch., C 86(2): 209. 1983. TYPE:
JAPAN, Kochi Pref., Motoyama-cho, cult. on bark of an unidentified tree, 1-19
VII 1980 (holotype: NENB 13.043 now at BR!, isotype Y.Y. 640 p.p.)
(Figs. 26-32)
Sporocarps scattered, stipitate, 800 -1300 µm in height. Sporothecae dark
brown, ovoid 200-400 µm diam. Hypothallus inconspicuous. Stalk up to 700
µm long, concolourous with sporotheca, brownish orange (54. br O), straight,
sub-cylindrical, tapering towards the top, furrowed, mid-width one tenth of the
height. Peridium double, although appearing single, inner layer yellowish grey
to colourless (93. y Gray) by TL, covered with a gelatinous layer with granular
refuse material forming longitudinal striations continuous with the outer layer
of the stalk, the inner surface prominently warted by TL (Fig. 29); dehiscence
into elongated platelets in upper sporotheca leaving a short lobed calyculus.
Spores free, whitish in mass, almost hyaline to pale yellow green (121. p. Y G)
by TL, subglobose, 10-12 µm diam., smooth by TL, evenly and densely warted
by SEM (Fig. 32); spore wall of uniform thickness.
Material examined: JAPAN, Kochi Pref.: Motoyama-cho, bark of unidentified tree, in
moist chamber, 1 to 19-VII-1980, NENB 13.043 (BR, holotype). COSTA RICA, Puntarenas
Prov: Monteverde, nr St. Elena village, 10°19’21”N 84°46’05”W, on a decayed frond of
Chamaedorea tepijiote, field collection. 11-VI-1999, M. Schnittler & Y. Novozhilov, sc
14529. CUBA, Sancti Spiritus: Alturas de Banao, 21°52’39”N 79°37’03”W, on dead liana in
moist chamber, 6-II-2003, dwb 2258.
Habitat: bark of trees and vines.
Distribution: Costa Rica, Cuba, Japan,
Illustrations: Nannenga-Bremekamp and Yamamoto (1983: 210, Fig. 2a-d); Yamamoto
(1998: 131).
The large closed tulip-shaped sporocarps (Figs. 26-28) were over 800 µm
tall. When viewed with TL the sporotheca can be seen as almost transparent
and distinct from the brown stalk. The stalk has longitudinal striations of
granular material within an outer covering giving the appearance of a stalk
within a stalk. The inner surface of the peridium is covered with large warts
(Fig. 31) especially in the upper half. Between these warts or baculae by SEM,
at very high magnification, smaller flatter warts are visible (Fig. 30).
Dehiscence is by means of platelets which break away and open leaving a
lobed cup. The spores are whitish in mass, almost hyaline by TL, smooth and
thin-walled, 10-12 µm diam. (Fig. 29). The neotropical specimens (sc 14529
and dwb 2258) have shorter peridial warts.
This species was published as Licea erectoides, but later combined as a
variety of L. erecta by one of the authors (Yamamoto, 1998), who stated that
the only difference from L. erecta was the spore size. We requested the type of
274

Fungal Diversity
Figs. 26-32. Licea erecta var. erectoides. 26. Habit (dwb 2258). 27. Tulip-shaped sporotheca
by SEM with granular striations (NENB 13.043 as Licea erectoides). 28. Habit (NENB
13.043). 29 Warted inner peridial surface and spores with wall of uniform diam. by TL (dwb
2258). 30. Detail of inner peridial ornamentation with smallest surface warts x9000 by SEM.
31. Larger warts of inner peridial surface x6000 by SEM. 32. Detail of spore surface by SEM.
L. erecta var. erecta but had no reply and so have been unable to study it. De
Haan (2002) SEM photographs shows a spore of L. erecta var. erecta from the
collection of Nannenga- Bremekamp, showing it is smooth. We studied this
material [BR-Myc 067670,61 from the collection Nannenga- Bremekamp
275

14807 (ex Yamamoto Y. 2912)] and doubt the rather small sporocarps belong
to L. erecta var. erecta. The sporocarps measured 288 µm in height, and the
diam. of the sporotheca was 115 µm. The box is labeled “Licea erecta Thind ?
& Dhillon”. It did not fit the description of L. erecta var. erecta (see below),
which exactly matches our observations of the holotype of L. erecta var.
erectoides, with the exception of the spore size and ornamentation.
Our SEM and microscopic examination of L. floriformis var. aureospora
(see below) shows similarities between these two species. The inner surface of
the peridium is warted in both, they have almost colourless spores by TL of
similar size range, and the ornamentation of the spores by SEM is also similar.
They are different, however, when viewed by TL, as the inner layer of the
peridium of Licea erecta var. erectoides appears as a flat base to the sporotheca
and the outer layer of refuse appears as ribs outside along the length of the
sporotheca, continuous with the ribbed stalk. The inner layer of the peridium of
L. floriformis var. aureospora ends in a pointed calyculus, protruding down
into the stalk and the outer refuse appears as a thin layer covering only the
basal portion of the sporotheca. Macroscopically they are also different, with L.
floriformis var. aureospora having a rounded sporotheca with a yellow spore
mass showing through the top half, which fractures into small irregular
platelets. The sporotheca of Licea erecta var. erectoides is tulip shaped, denser
and all one colour, since none of the whitish spore mass shows through. The
platelets are also slightly elongated.
Nannenga-Bremekamp and Yamamoto (1983) and de Haan (2002) both
commented on specimen YY 101 (NENB 13044), as being different from the
type of L. erecta var. erectoides. The spores by SEM are very similar to those
of L. floriformis var. aureospora, and it seems probable that it belongs to that
taxon.
6. Licea floriformis var. aureospora M.T.M. Willemse & Nann.-Bremek.,
Proc. Kon. Ned. Akad. Wetensch. 97(1): 137. 1994. TYPE: THE
NETHERLANDS, in the rural estate of Hoekelum near Ede, on bark of living
Sambucus nigra, on 16 XII 1986 (holotype: 104 L in the collection of M.T.M.
Willemse 104a, isotype NENB 16.075 now at BR!). (Figs. 33-46)
= Licea longa Flatau, Stapfia 73: 67. 2000, syn. nov. TYPE: GERMANY, Kassel,
Borke von Populus spec., (1), 150 m über NN, 6.-26.10.1998, LF 3180 (holotype: B!).
Figs. 33-39. Licea floriformis var. aureospora. 33-34. Habit (NENB 16.705). 35. Whole
sporocarp by SEM. Dehiscing sporotheca and stalk with multiple strands (M. de Haan
001014). 36. Dehisced sporocarp by TL (M. de Haan 001014). 37. Detail of spores by TL (M.
de Haan 001014). 38. Inner peridial surface with warts (M.de Haan 001014). 39. Spore by
SEM (Lado 11715).
276

Fungal Diversity
277

= Licea capitata Ing & McHugh in Ing, Trans. Brit. Mycol. Soc. 78(3): 439. 1982, syn.
nov. TYPE: UNITED KINGDOM, Scotia, Dumbarton, Balloch Park, reperta in vitro in cortice
ablato ab arboris vivae Aceris pseudoplatani, 4 Oct. 1980, B. Ing (holotype: K!).
Sporocarps scattered, stipitate, 450 -1200 µm in height. Sporotheca light
olive brown, subglobose, 200-400(-700) µm diam. Stalk olive brown (94. l. Ol
Br) concolourous with the sporotheca, 250-800 µm long, straight,
subcylindrical, tapering towards the top, plicate, filled with algae by TL,
continuous with the sporotheca, mid-width a quarter of the height. Peridium
double, covered below with gelatinous layer with granular refuse material
forming patches or longitudinal striations continuous with the outer layer of the
stalk, the inner layer membranous, greyish yellow (90. gy. Y- 91. d. gy. Y)
with no inclusions, its inner surface warted to almost spiny by TL and by SEM
(Fig. 38); dehiscence irregular in upper sporotheca into platelets, which are
spiny at edge, leaving a short calyculus. Spores free, yellow gold in mass, pale
yellow green (121. p. Y G) to colourless by TL, smooth, densely warted by
SEM, 10-12,5 µm diam.; spore wall of uniform thickness (Fig. 37).
Material examined: THE NETHERLANDS, Ede: Hoekelum, on bark of living
Sambucus nigra in moist chamber, 16-XII-1986, NENB 16.705 (BR, isotype); BELGIUM,
Zillebeke: Palingbeek, on logs of broad-leaved trees, 28-X-2001, MdH 001014; MEXICO,
Veracruz: Catemaco, Los Tuxtlas Biological Reserve, 18°35’04”N 95°04’30”W, 200 m, on
bark of living tree and bryophytes, 2-XII-1999, Lado11715. GERMANY, Hessen, Kassel-
Wolfsanger, bark of living Populus sp. in moist chamber, 6 to 26-X-1998, LF 3180 (B,
holotype of Licea longa); 7 to 26-X-1998, Hessen, Kassel, Fuldatal, bark of living Populus sp.
in moist chamber, 17-IX to 26-X-1998, LF 3169; 10-XII-1998, LF 3194 as Licea longa;
Hessen, Kassel-Wolfsanger, bark of living Populus sp. in moist chamber, 21-XII-1999 to 3-II-
2000, LF 3277 as Licea longa. UNITED KINGDOM, Scotland, Dumbartonshire, Dumbarton,
Balloch Park, on bark of Acer pseudoplatanus, 4-X-1980, B. Ing [K(M): 129676 holotype of
Licea capitata]. Wales, Coed Aber, SH665 718, on bark of Quercus petraea in moist chamber,
2-III-1993, dwb 1128.
Habitat: bark of living trees.
Distribution: Belgium, Germany, Ireland, Japan, Mexico, The Netherlands, United
Kingdom.
Illustrations: Willemse and Nannenga Bremekamp (1994: 138, 139, Figs. 1-2); Flatau
(2000: 67, Fig. 3 as Licea longa); de Haan (2002: 30, Plate 1; 32, Figs. 3-4)
The type specimens were very big with a habit that is more like a Trichia
than a Licea (Figs. 33-34) but without capillitium. The sporocarps were up to
1200 µm in height with the largest sporotheca 700 µm in diam. The peridium
was dull pale brown by reflected light with a mottled appearance at the top
with the yellow spore mass showing through the thinner outer peridial layer in
places. The inner layer adhered to the outer layer and was densely warted on
the inner surface (Fig. 38). The dark, obviously plicate stalk was made of
overlapping strands visible by SEM (Figs. 35, 42) and filled with algae. The
stalk was continuous with the sporotheca. Dehiscence was by platelets which
had few patchy areas with refuse inclusions from the outer layer showing
278

Fungal Diversity
through, and were warted to almost spiny and irregularly shaped (Figs. 35-36).
An irregular very shallow calyculus was left. The spores were yellow gold in
mass, hyaline to pale yellow green (121. p. Y G) and smooth by TL but evenly
and densely warted with some fused warts by SEM (Fig. 39).
We also examined several collections of L. longa Flatau, kindly sent to
us by the author, and the type material of this species (LF 3180). The habit,
height (1000-1200 µm) and diam. (400-720 µm) of the specimens match those
of the isotype of L. floriformis var. aureospora (Figs. 35, 42). The dehiscence
was by irregular platelets which had plates with zip-like junctions and spines
on the edge (Fig. 44) and similar patches of refuse to L. floriformis var.
aureospora and which we think may be the remains of the outer peridium. The
inner peridium surface has the same ornamentation of closely packed warts and
the stalk was continuous with sporotheca SEM (Fig. 43). The spores were 10-
12 µm yellow gold and smooth by TL but evenly and densely warted with
some fused warts by SEM (Figs. 45-46). In the description Flatau (2000)
commented that these species were differentiated by the apparently single (L.
longa) versus double peridium, and the opaque stalk of L. floriformis var.
aureospora. We found the stalks and the peridium to be identical and we
believe these to be the same species, on the basis of the detailed comparisons
of the morphological features made of the type material of each and the SEM
photographs. We therefore synonymize them here.
Comparison by SEM of this species with material from Mexico
published under Licea sp. 1 by Lado et al. (2003) has allowed us to include this
specimen in L. floriformis var. aureospora. De Haan (2002), in her SEM
photographs shows the spores of the collection YY 101 (NENB 13044),
tentatively identified as L. erecta var. erectoides, to be very similar to those of
L. floriformis var. aureospora, and we agree that the specimen should be
included in this species. Spore ornamentation viewed by SEM has been shown
by Gilert (1994) to be a stable and important taxonomic character.
Ing (1999: 55) included under Licea operculata inoperculate forms
previously described as L. capitata by Ing (1982), which he stated were the
result of conditions of moisture during fructification. We requested the type
specimen of L. capitata from Kew and compared it with the isotype of L.
floriformis var. aureospora, and found that the size and ornamentation of the
spores were the same, as were the characters of the stalk and the peridium.
There were also comments on inoperculate sporocarps placed in L. operculata
by Mitchell and McHugh (2000), but the photograph they publish with refuse
material from the stalk continuing up the sides of the sporotheca to the middle,
and the description of its irregular dehiscence at the apex, suggest that these
specimens also belong to L. floriformis var. aureospora.
279

280

Fungal Diversity
This variety was also recently reported from Asahi Town, Fukui
Prefecture, Japan (Yamamoto et al., 2004), extending its geographical
distribution, previously limited to Europe and America.
The authors state that L. floriformis var. aureospora differs from L.
floriformis var. floriformis in having a longer stalk and the colour of the spores
in mass are yellow not black and colourless by TL not pale pink. As we were
unable to study material of the latter variety we can not confirm this.
7. Licea lucens Nann-Bremek., Proc. Kon. Ned. Akad. Wetecsch., C 84(3):
285. 1981. TYPE: FRANCE, Dept. Doubs, near Gigot, on moses and
liverworts on bark taken from a living Acer, developed in a moist chamber:
bark collected on 10 VI 1978, moistened 12 II 1979, the sporangia ripened
from 15 III to 30 IV 1979 (holotype: NENB 11.194, now at BR!). (Figs. 47-51)
Sporocarps scattered or gregarious, stipitate, up to 150 µm in height.
Sporotheca red brown when moist, rosy when dry, elliptical, 35-45 µm width,
70-100 µm height. Hypothallus inconspicuous. Stalk very short up to 30 µm
long, hyaline by transmitted light, straight, cylindrical, with little included
granular material, mid-width approximately equal to the height. Peridium
single, transparent, hyaline, smooth and glossy on both sides; dehiscence by
irregular fracture of the top, leaving a deep cup. Spores free, red brown to rosy
in mass, pale yellow to light yellow by TL (89. p. Y-86. l. Y), subglobose, 8-10
µm diam., warted by TL, very prominent warts by SEM, spore wall uniformly
thin.
Material examined: FRANCE, Dept. Doubs, Nr Gigot, on mosses and liverworts on
bark of living Acer sp. in moist chamber, 15-III-1979 to 30-IV-1979, NENB 11.194 (BR,
holotype)
Habitat: on mosses and liverworts on bark of living Acer.
Distribution: Known only from the type locality.
Illustrations: Nannenga-Bremekamp (1981: 286, Fig. 1).
The type material shows many beautiful tiny sporocarps. They look like
miniature glass beads full of rosy golden spores (Fig. 47). They appear to be
sessile but close examination shows they are on a very short (30 µm) stalk
which is just an extension of the peridium (Fig. 48). The peridium is
completely hyaline and smooth outside and inside by transmitted light. By
SEM
however the inner surface is very sparsely and irregularly warted (Fig.
Figs. 40-46. Licea floriformis var. aureospora (as Licea longa Flatau). 40. Dehiscing
sporocarp and spores by TL (LF 3169). 41. Fragments of peridium by TL (LF 3180). 42.
Whole sporocarp by SEM. Dehiscing sporotheca and stalk with multiple strands (LF 3169). 43.
Detail of sporotheca by SEM (LF 3169). 44. Detail of inner peridial surface and edge of
platelet by TL (LF 3180). 45. Group of spores by SEM (LF 3169). 46. Surface detail of spore
by SEM (LF 3169).
281

Figs. 47-51. Licea lucens (NENB 11.194). 47. Habit. 48. Intact sporocarp with spores by TL.
49. Group of spores by TL. 50. Smooth inner surface of peridium by SEM. 51. Two spores
showing verrucate ornamentation.
50). The spores show very clear dense ornamentation by TL (Fig. 49) which is
prominent, and of the echinulate to verrucate type (Rammeloo, 1974, 1975) by
SEM (Fig. 51).
This combination of characters is completely unique among the stipitate
Licea species we have examined. The size and initial habit of shiny sporocarps
is similar to L. perexigua (see below), but the shape, spore colour and
ornamentation, and the mode of dehiscence are all different. This species is
only known from the type locality, and its minute dimensions have probably
caused it to be overlooked, or confused with algae or other bark epiphytes.
8. Licea operculata (Wingate) G.W. Martin, Mycologia 34(6): 702.
1942. Orcadella operculata Wingate, Proc. Acad. Nat. Sci. Philadelphia 41:
280. 1889. TYPE: U.S.A., Pennsylvania: Philadelphia, Fairmount Park or
282

Fungal Diversity
Chestnut Hill, on trunks of living Quercus rubra L. (holotype: PH, now at
BPI!). (Figs. 52-58)
Sporocarps scattered to loosely gregarious, stipitate, 500-1000 µm in
total height. Sporotheca dull brown, usually urn-shaped, but rarely ovoid or
almost globose, 200-500 µm diam. Stalk 300-640 µm in height, nearly black,
slightly tapering towards the apex, furrowed, mid-width less than one fifth of
height, filled with granular refuse material. Peridium single, thin, warted on the
inner surface (Fig. 54) except for the lid, with ornamentation particularly
visible at the edge of dehiscence (Fig. 55); dehiscence by a distinct shiny gold
lid with a smooth inner surface by TL; the lid minutely punctate by SEM.
Spores free, almost colourless to pale greenish yellow (104. p. g Y-121. p. Y
G), smooth by TL (Fig. 54), 8-11µm diam., minutely punctate by SEM (Fig.
58); spore wall of uniform thickness.
Material examined: USA, Philadelphia: Fairmount Park, [On bark living red-oak trees
(Quercus rubra) Wingate 1889] (Orcadella Wingate 1275) ex state coll. (BPI 826296, type).
ECUADOR, Prov. Orellana: Amazonian basin, Yasuni National Park, Lago Agrio, 0°40’30”S
76°23’45”W ± 600 m, bark of deciduous tree in moist chamber, sc17779; 0°40’09”S
76°24’03”W bark of Cedrelinga catenaeformis in moist chamber 28-V-2000, dwb 1833;
0°40’31”S 76°23’47”W, bark of deciduous tree in moist chamber, sc17808. Pichincha Prov.:
Western Andean Slopes, Macquipucuna Reserve; Calacalí, 0°5’40”N 78°37’0”W, bark of dead
liana in moist chamber, sc 13587. PERU, Mazan: 3°28'S 74°55'W, on dead liana in moist
chamber, 22-VI-2002, dwb 2225. PUERTO RICO, Fajardo: El Verde, 18°20’30”N
65°49’30”W bark of living deciduous trees in moist chamber, sc 17091; sc 17336. Fajardo: on
a ca. 1 km ESE San Vicente, 18°21’05”N 65°43’57”W, bark of a living deciduous tree in
moist chamber, sc 17103. MEXICO, Veracruz: Catemaco, Los Tuxtlas Biological Reserve,
18°35’04”N 95°04’30”W, on bark of Porteroni viridescens in moist chamber, 31-V-2000, dwb
1836. Queretaro: Sta María del Mexicano, 20°49’25”N 100°03’24”W, on bark of Yucca sp. in
moist chamber, 17-II-2004, dwb 2378.
Habitat: Bark of living trees.
Distribution: Ecuador, Mexico, Peru, Puerto Rico, Tanzania, USA.
Illustrations: Wingate (1889: 280); Lister (1925: pl. 149d-f, as Orcadella operculata);
Nannenga-Bremekamp (1965: 134, 1974: 72); Martin and Alexopoulos (1969: Fig. 9); Emoto
(1977: pl. 7, Figs. 5-8); Neubert et al. (1993: pl. 6, Figs. 3-4); Gilert (1994: Figs. 47-70); Lado
and Pando (1997: 124, Fig. 24); Yamamoto (1998: 152).
The presence of variable morphotypes in this species was commented on
by Wingate, in his original description (Wingate, 1889). He gave the range in
height from 375-1250 µm. The sporocarps of the type material examined were
> 800 µm in total height. The sporotheca were 300-500 µm in diam., urn-
shaped, dull brown, and with a distinct shiny gold lid (Fig. 52) with a smooth
inner surface by TL. The nearly black furrowed stipe had 3 or 4 intertwined
strands inside it visible by TL. The type material had only a few (6) sporocarps
left so we were not able to do SEMs of it. Our SEM pictures are of Schnittler’s
collection (sc17103) from Puerto Rico, and there are further SEM pictures in
Gilert (1994).
283

Figs. 52-58. Licea operculata. 52. Habit (BPI 826296). 53. Urn-shaped sporotheca by TL (sc
17103). 54. Inner peridial surface and spores by TL (sc 17103). 55. Detail of rim of sporotheca
by SEM (sc 17103). 56. Urn-shaped sporotheca by TL (dwb 1833). 57. Outer surface of
sporotheca by SEM (sc 17103). 58. Spore by SEM (sc 17103).
Typical large urn-shaped sporothecae with a distinct lid like the type
specimens are unmistakable (Figs. 53, 56). Many of the smaller particularly
tropical forms from moist chamber cultures we have examined are not as easily
determined. They are variable in size from 500-700 µm in total height and the
stalk is sometimes light brown and not furrowed. In addition in some
collections the lid appears almost smooth on the inner surface, whereas in
others some ornamentation (baculate processes see Gilert, 1994) is easily
visible at 400x by TL. We have included in this taxon any specimens larger
than 500 µm with a distinct lid, tapering stalk and smooth spores by TL within
the size range and colour of this species.
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Fungal Diversity
This species is similar in habit and spore ornamentation to L. floriformis
var. aureospora and L. erecta var. erectoides (see above), but it can be
distinguished easily by its mode of dehiscence with a lid, and the edge of the
sporotheca where the lid separates which has clear warts (Fig. 56). The spores
of L. operculata are also smaller although there is overlap in the extremes of
the sizes. The differences between this species and L. poculiformis Ukkola, the
only other operculate stipitate Licea, are commented upon under that species
below.
After comparing some specimens from Mexico published as Licea sp. 1
by Lado et al. (2003), with a visible lid, similar spores and peridium, we also
include them in L. operculata, although the sporocarps are smaller. Ing (1999:
55) included inoperculate forms previously described as L. capitata but we
have found that the type of L. capitata belongs to L. floriformis var.
aureospora (see comments under this taxon). Licea operculata was also
reported from Tanzania (Ukkola, 1998) on Araucaria bark in moist chamber
culture. The author mentions that some specimens had poorly developed lids.
The brief description of Orcadella operculata var. sessile G. Lister
(Monogr. Mycetozoa, ed. 3: 186. 1925) suggests a completely different sessile
species.
9. Licea parvicapitata Y. Yamam., Bull. Natl. Sci. Mus., Tokyo, B 26(3): 113.
2000. TYPE: JAPAN, Chiba Pref., Sawara-shi, Sawara-ho, Atago-jinja, on
bark of living Quercus acuta, in culture, 18 V 1982, MT-2664 (holotype:
TNS!). (Figs. 59-65)
Sporocarps scattered, solitary, stipitate, 480-700 µm in height.
Sporotheca golden, globose, 160 µm diam., with large warts on the outer
surface held up by a distinct thin darker stalk. Stalk 200-400 µm in height,
black (Fig. 59), straight, subcylindrical, tapering slightly towards the apex,
filled with refuse matter (Fig. 61), opaque, mid-width up to a quarter of its
height. Peridium single membranous, almost hyaline to pale grey, warted on its
inner surface (Figs. 63-64), covered with refuse material discontinuous with
the stalk, and irregular darker warts scattered on outer surface (Fig. 62);
dehiscence circumcissile by a line very close to the base of the sporotheca
leaving a very small calyculus, the remainder of the peridium breaks into small
irregular platelets. Spores free, pale greenish yellow (121. p. Y G-104. p. g Y)
to colourless, subglobose, smooth (Fig. 65), 7-8 µm diam.; spore wall thin,
with thickened area on one side.
Material examined: JAPAN, Chiba Pref.: Sawara-shi, Sawara-ho, Atago-jinja, on bark
of living Quercus acuta, 18-V-1982, MT 2664 (TNS, holotype)
Habitat: bark of Quercus acuta.
Distribution: Known only from the type locality.
285

Figs. 59-65. Licea parvicapitata (MT 2664). 59. Habit. 60. Whole sporocarp showing line of
dehiscence and spotted surface. 61. dehisced sporocarp by TL. 62. Outer surface of peridium
with refuse and darker irregular spots. 63-64. Inner surface of hyaline peridium with warts. 65.
Spores by TL.
Illustrations: Yamamoto (2000: 113)
This species is only known from type material which showed stipitate
sporocarps, most with a spotty appearance due to the peridial warts, although
these were not evident on all sporocarps. The stalk was very thin and fragile
and fractured on mounting. The combination of the dark thin stalk, the spotted
golden brown sporotheca and the small spores make this species easily
distinguished from the others. There were crystals among the refuse material of
some specimens when viewed with Nomarski, but the longer thinner stalk,
larger sporocarp size (480-700 µm vs. 360-480 in L. eleanorae), dehiscence
near the base not the apex, and lighter spores (121. p. Y G-104. p. g Y vs. 89.
p. Y) of L. parvicapitata differentiate it from L. eleanorae.
286

Fungal Diversity
10. Licea pedicellata (H.C. Gilbert) H.C. Gilbert in Martin, Mycologia 34(6):
702. 1942. Hymenobolina pedicellata H.C. Gilbert, Stud. Nat. Hist. Iowa Univ.
16(2): 153. 1934. TYPE: U.S.A., Iowa, Milford, on bark of living Ulmus, 16
July, 1932, H.C. Gilbert 2117 (holotype: BPI!). (Figs. 66-74)
Sporocarps scattered to gregarious, stipitate, 200-500 µm in height.
Sporotheca dark brown or black, subglobose, rounded even when dry, 75-175
µm diam. Stalk 125-325 µm in height, thick, straight, cylindrical, furrowed,
continuous with outer peridium, filled with granular refuse material, mid-width
less than half the height. Peridium double, but apparently single, with an
adhering outer layer of refuse material, membranous, dull brown; dehiscence
into irregular platelets 20-30 µm wide, inner layer warted (Fig. 68). Spores
free, dark brown in mass, medium yellow to greyish yellow (87. m. Y-90. gy.
Y) by TL, subglobose, (10-)11-13(-14) µm diam., minutely warted by TL (Fig.
70) and densely warted by SEM (Fig. 72); spore wall with a paler thinner area.
Material examined: USA, Iowa, Milford [coll W. Okoboji acc. to Gilberts letter Dec. 7,
1934 on box] on bark of elm (in moist chamber), 16-VII-1932, H.C. Gilbert 2117 (BPI 826385,
holotype) as Hymenobolina pedicellata; (BPI 657294, isotype) as Licea pedicellata. Iowa:
Milford [on bark of living Ulmus, see Gilbert (1934: 154)], 16-VII-1932, coll. W. Okaboji [HC
Gilbert 2117] (BPI 826368, isotype) as Licea pedicellata. Iowa city on dead Ulmus sp., 00-XII-
1961, coll. G.W. Martin (BPI 805746). Virginia, Mt. Lake Biol. Station, on bark of Quercus
sp. in moist chamber, 12-VIII-1969, leg. & det. C.J. Alexopoulos (BPI 826366). Texas: Austin,
Hancock Center, on bark of living Ulmus crassifolia in moist chamber, 5-VIII-1974, leg. &
det. C.J. Alexopoulos (BPI 737641).
Habitat: Bark of living trees.
Distribution: Zaire?, USA.
Illustrations: Gilbert (1934: 154, Fig. 1); Martin and Alexopoulos (1969: 481, pl. 1 Fig.
11a-c); Gilert (1994: 17, Fig. 47); Yamamoto (1998: 142).
In the original description of the species, Gilbert (1934) states that the
spores are “smooth or faintly and finely warted”. Examination of all the BPI
collections of type material, both by TL and SEM, showed all the spores to be
warted, and none were smooth as Gilbert suggested. This is the main
distinguishing character of this species from other similar species like L.
rugosa (Table 2). The faint warts are discernible by TL at 400x (Fig. 70), and
SEM examination revealed dispersed warts (Fig. 72), which were irregular in
size. At great magnification (Fig. 74) they were seen to have a warted surface
themselves.
Another clear characteristic visible by TL is that the interior of the
peridium is warted, but the warts are dispersed and separate (Fig. 73). In
addition almost the entire peridium breaks into platelets leaving a very shallow
structure attached to the stalk (Figs. 69, 71). We were not able to see clear
ridges or lines of dehiscence in any of the specimens, and all the specimens
retained their rounded shape even when dry (Fig. 67). Martin and Alexopoulos
(1969: 48) also commented that they “rarely observed platelets in the
287

Figs. 66-74. Licea pedicellata. 66. Habit (BPI 826368). 67. Whole sporocarp by SEM (BPI
737641). 68. Irregular platelets with warted inner surface (BPI 826385). 69. Dehisced
sporocarp by TL with shallow flat remains of peridium (BPI 826385). 70. Spore by TL (BPI
826385). 71. Dehisced sporocarp with flat remains of peridium by SEM (BPI 657294). 72. Spore
by SEM (BPI 657294). 73. Group of spores and detail of inner peridial surface by SEM (BPI
657294). 74. Highly magnified
(× 20,000) detail of spore ornamentation by SEM (BPI
657294).
288

Fungal Diversity
Table 2. Characters differentiating L. pedicellata, L. rugosa var. rugosa and L. rugosa var.
fujiokana.
Name Dehiscence Peridium Spore Other
L. pedicellata
Irregular
fragments;
flat base of
sporotheca left
Dispersed
warts on inner
surface
Yellowish brown;
warted; (10-)11-13(-14)
µm diam.
Sporotheca
round
when dry
L. rugosa var.
rugosa
Irregular
fragments
Prominent
dense warts on
inner surface
Olive-brown; smooth;
14-16 µm diam.
Sporotheca
wrinkled
when dry
L. rugosa var.
fujiokana
Irregular
fragments
Prominent
dense warts on
inner surface
Greyish yellow brown;
smooth; 10-12 µm diam.
Sporotheca
wrinkled
when dry
peridium”. We did a SEM study of material cited by Pando and Oltra (2001)
from Mogente, Spain [MA-Fungi 40549 (Oltra 2519)] but found it to have
smooth spores and a different ornamentation of the inner peridium and we
conclude that it does not belong to this taxon. Of the other material examined
in our herbarium that had been assigned to this species (DWM 3313 from
Mexico on red cedar bark in moist chamber, and TEB 3704 from Virginia-
Baskerville cemetery, USA, cedar tree #3, 28-VIII-1970), none had warted
spores except another sample (DWM 4478) from Zaire, but the single slide of
this was insufficient for us to positively confirm its identity. The SEM picture
in Gilert (1994) shows a rugose sporocarp with apparent platelets, which casts
doubt on its identity as L. pedicellata. It is possible that this species has been
confused with L. rugosa, and that its distribution is not as widespread as Martin
and Alexopoulos (1969: 48) and other authors (Yamamoto, 1998; Ing, 1999)
believed, since they mention also Australia, Austria, Great Britain, Greece,
India, Japan, Mexico, Tunisia and Turkey. The similarities between L.
pedicellata, L. rugosa and the varieties of each are discussed below.
It is worth noting that the three boxes of dried type material lent to us by
the BPI herbarium have slight differences in the labels. One of them (BPI
826385) lists the locality as Milford Iowa, as does Gilbert´s description (1934),
with a note “coll. W. Okoboji acc. to Gilberts letter Dec. 7, 1934”, another
(BPI 657294) lists Okoboji, Iowa, both in script. Collection BPI 826368 lists
Milford and coll. W. Okaboji (sic). We note for clarification that West Okoboji
is a town slightly West of Milford, both in Dickinson County Iowa. It seems
that “coll.” here refers to the place of collection, not who it was collected by,
the accepted meaning of this abbreviation (Stearn, 1996).
289

Figs. 75-80. Licea perexigua. 75. Habit (dwb 1936). 76. Inner surface of peridium densely
papillate by TL (HWK 1166). 77. Inner surface of dehisced sporotheca with spores by SEM
(dwb 1936). 78. Inner surface of peridium densely papillate by TL (HWK 1166). 79-80. Detail
of spore surface ornamentation by SEM (dwb1936).
290

Fungal Diversity
11. Licea perexigua T.E. Brooks & H.W. Keller in Keller and Brooks,
Mycologia 69(4): 674. 1977. TYPE: U.S.A., Arkansas, Crawford County, on
the bark of living Juniperus virginiana, October 27, 1964, T.E. Brooks 2747
(holotype: BPI!). (Figs. 75-80)
Sporocarps scattered to gregarious, stipitate or sessile on a narrowed
base, up to 100 µm in height. Sporotheca golden, iridescent, subglobose, 40-
100 µm diam. Hypothallus inconspicuous. The stalk when present pale yellow
by TL, less than half the total height of sporocarp, filled with granular refuse
material. Peridium single, membranous, colourless by TL, with scant granular
refuse matter evenly distributed over the surface, inner surface closely
papillate; dehiscence circumcissile low down at the base of the sporotheca
leaving a calyculus. Spores free, pale to greyish green yellow (121. p. Y G-
122. gy. Y G), from 8.5-10.5 µm diam., smooth by TL, minutely ornamented
with unevenly scattered warts by SEM (Fig. 79); spore wall thin, with a paler
thinner area.
Material examined: USA, Arkansas: Crawford Co., [on bark of living Juniperus
virginiana in moist chamber (Keller and Brooks, 1977)], 27-X-1964, coll. TEB 2747 (BPI
826404, holotype). Mississippi: Mississippi co., moist chamber, coll. HWK 1166 (slide).
MEXICO, Tlaxcala: Cuapiaxtla, 19°19’40”N 97°41’44”W, on bark of living Juniperus
deppeana in moist chamber, 18-I-2001, dwb 1936. Hidalgo: Progreso, La Cruz, 20°16’02”N
99°10’48”W, on bark of living tree Opuntia sp. in moist chamber, 26-III-2001, dwb 2021.
SPAIN, Cuenca: Saceda-Trassierra, on bark of Quercus ilex in moist chamber, Pando 1185,
21-III-1990, MA-Fungi 31964.
Habitat: bark of living trees.
Distribution: Mexico, Spain, USA.
Illustrations: Keller and Brooks (1977: 675, Figs. 13-17); Lado and Pando (1997: 127,
Fig. 35); Novozhilov et al. (2003) (as L. cf. perexigua).
The type material showed very tiny, almost sessile, golden irridescent
sporocarps. The specific epithet gives the most obvious character of
exceedingly small and with a stalk so short that even the stipitate sporocarps
appear sessile. Many calyculi of dehisced sporocarps were visible on the bark
surface in the type specimen, the stalk remaining with a tiny cup-shaped
sporothecal base on top. The single peridium in some specimens is completely
hyaline, and refuse material is very sparce even when present. The
ornamentation on the inner surface is visible at 400x (Figs. 76, 78), appearing
to be in a reticulate pattern as described by the authors (Keller and Brooks,
1977), but by SEM (Fig. 77) it can be seen that this pattern is not a true
reticulum but is the result of the uneven distribution of the papillae and there
are some smooth areas without ornamentation. By TL the spores seem smooth
and have thin walls with a paler thinner area at which they have a tendency to
fold (Fig. 78), but by SEM irregularly distributed rhomboid warts are visible
(Figs. 79-80) with some so close together as to appear fussed, and others with
spaces between them.
291

Figs. 81-86. Licea poculiformis. 81. Habit (Ukkola 319A). 82. Whole sporocarp showing
goblet-shaped sporotheca and pale lid (Ukkola 319A). 83. Dehisced sporocarp (dwb 1759). 84.
Deep cup of dehisced sporotheca by TL (dwb 1759). 85. Spores by TL (dwb 1759). 86. Inner
surface of peridium by TL (dwb 1759).
The unique pattern of ornamentation on the spores distinguishes this
species from the other stipitate species. In addition it is smaller than all of them
except L. lucens (see above). The transparent peridium is another clear
character. Keller and Brooks (1977) differentiate L. perexigua from L. tenera
by the broad base of the sessile sporocarps in the latter species, its different
292

Fungal Diversity
peridial colour and ornamentation, and its larger (12 µm vs. 8.5-10.5 µm
diam.) spores. The authors also differentiate it from L. scyphoides by its almost
sessile habit, its smaller [11-13.5(-14) µm diam. in the latter species] slightly
darker spores (121. p. Y G-122. gy. Y G vs. 90. gy. Y-104. p. g Y), and its
manner of dehiscence (irregular vs. circumcissile in L. scyphoides). Our
observations concur with theirs.
12. Licea poculiformis Ukkola, Acta Bot. Fennica 160: 5. 1998. TYPE:
TANZANIA, Tanga Province, Lushoto District, East Usambara Mts., on
mosses growing on bark of Cupressus sp., in moist chamber culture,
12.XII.1995, Ukkola 319A (holotype: H!). (Figs. 81-86)
Sporocarps scattered, stipitate, 150-200(-400) µm in height. Sporotheca
treacle-brown, goblet-shaped, 100-150 µm diam., with a shiny pale lid (Fig.
81). Stalk 50-150 µm in height, furrowed, mid-width half the stalk height.
Peridium double, outer layer coriaceous, shiny, inner layer membranous;
dehiscence by a pale shiny lid with a clear edge and warts on the margin,
leaving a deep cup (Fig. 84). Spores free, pale greenish to greyish yellow (90.
gy. Y - 104. p. g Y) by TL, 8.5-10 µm diam., smooth by TL (Figs. 85-86) but
warted by SEM (Ukkola, 1998: 6, Fig. 7); spore wall with a slightly thinner
paler area.
Material examined: TANZANIA, Tanga Province, Lushoto District, East Usambra
Mountains, Amani Forest Reserve, submontane forest, alt. 900 m, on mosses growing (sec.
Ukkola, 1998) on bark of living Cupressus sp. in moist chamber culture, 13-IX-1995 (on the
label), Ukkola 319A (H, holotype). MEXICO, Quintana Roo, El Eden, bark of living
Hematoxylon campechianum in moist chamber, 10-II-2000, dwb 1759 [Lado et al., 2003: 88].
Habitat: moss and bark of living trees.
Distribution: Mexico, Tanzania.
Illustrations: Ukkola (1998: 6, Figs. 2-7).
The sporotheca and stalk of this small species are concolourous, treacle-
brown. The abundant sporocarps in the type specimen were mostly projecting
from the tips of moss leaflets. The Mexican material is much larger (up to 400
µm see Fig. 83) but otherwise agrees with the type. This species has been
reported recently from Okayama Prefecture, Japan on the bark of living Prunus
sp. (Yamamoto and Fujioka, 2004) and the Japanese specimens described are
also larger than the type. This species is closest to L. operculata, and shows
similarity in the habit, and also the inner surface of the peridium by SEM, but
the type specimens are different in their deep brown colour, have a large
difference in size (150-400 µm in height vs. 500-1000 µm in L. operculata),
and a double peridium vs. single in L. operculata. The lid of L. poculiformis is
at the broadest diam. of the sporotheca, and wider than that of L. operculata,
has a clear edge and is distinctly papillose by TL, whereas the lid of the L.
operculata appears smooth by TL and is at the narrowing top portion of its
293

more globose sporotheca. Finally, the ratio of stalk width to height is different
with L. poculiformis much shorter and squatter and with a straight wide stalk
(Fig. 82) and with the outer peridium continuous with the sporotheca, not long
and tapered and separate from the sporotheca like L. operculata.
13. Licea rugosa Nann.-Bremek. & Y. Yamam., Proc. Kon. Ned. Akad.
Wetensch., C 90(3): 326. 1987. var. rugosa. TYPE: JAPAN, Kochi Pref.,
Agawa-mura, Mnt. Nakatsu, on the bark of a living Aesculus turbinate Blume,
VII 1984, Y.Y. 2300 (holotype: TNS, isotype NENB 14.476 now at BR!).
(Figs. 87-93)
Sporocarps scattered, stipitate, up to 150-500 µm in height. Sporotheca
dark brown to almost black, subglobose up to 150 µm diam. Stalk up to 180
µm in height, straight, cylindrical, furrowed (Fig. 88), stout, continuous with
the outer peridium, mid-width half of the stalk height. Peridium single,
membranous, wrinkled with darkened ridges visible when dry, prominently
warted on the inner surface; dehiscence irregular fracture into fragments in the
upper part leaving a calyculus (Figs. 88, 92) Spores free, dark brown in mass,
olive brown (94. l. Ol Br) by TL, subglobose, 14-16 µm diam., totally smooth;
spore wall with a thinner area (Fig. 89).
Material examined: JAPAN, Kochi Pref.: Agawa-mura, Mnt. Nakatsu, bark of living
Aesculus turbinate, VII 1984, coll. NENB 14.476 [ex YY 2300] (BR, isotype). MEXICO,
Hidalgo: Progreso, La Cruz, 20°16’02”N 99°10’48”W, on bark of living Prosopis laevigata in
moist chamber, 22-III-2001, dwb 2005; on bark of living Schinus molle in moist chamber, 16-
III-2001, dwb 2006. Puebla: Emilio Portes Gil, 19°17’40”N 97°30’22”W, on bark of living
Yucca filifera in moist chamber, 25-I-2001, dwb 1955. Zapotitlán de las Salinas, 18°20’00”N
97°27’45”W, on bark of living Beaucarnea gracilis in moist chamber, 11-IX-2003, dwb 2311;
ibidem, 13-IX-2003, dwb 2319. Tehuacán, Santiago Nopala, 18°25’57”N 97°36’46”W, on
bark of living Yucca periculosa in moist chamber, 11-XI-2003, dwb 2332; ibidem, bark of
living Beaucarnea gracilis in moist chamber, 22-XI-2003, dwb 2335. Queretaro: Peña Miller,
Camargo, 21°06’06”N 99°43’29”W, on bark of living Prosopis laevigata in moist chamber,
15-II-2002, dwb 2154.
Habitat: bark of living trees.
Distribution: Japan, Mexico.
Illustrations: Nannenga-Bremekamp and Yamamoto (1987: 326); Yamamoto (1998:
144).
Figs. 87-93. Licea rugosa var. rugosa. 87. Habit (NENB 14.476). 88. Dehisced sporocarp by
SEM showing calyculus and furrowed stalk (NENB 14.476). 89. Dark spores with wall thinner
in one area by TL (dwb 2006). 90. Inner surface of peridium with prominent dense warts
(NENB 14.476). 91. Smooth spore by SEM (NENB 14.476). 92. Calyculus of dehisced
sporocarp containing spores (dwb 2006). 93. Group of smooth spores by SEM (dwb 2319).
294

Fungal Diversity
295

The abundant Mexican material was identical to the type both by
microscopic examination and by SEM. The most obvious differentiating
features of this species are the size and rather dark colour of the spores. The
sporocarps examined were of varied height from almost sessile to 500 µm. The
sporotheca are globose when wet but very wrinled when dry, with prominent
ridges. The spores by TL look like the spores of L. parasitica (Zukal) Martin
[11-13(-16) µm diam.], but the thinner area is less pronounced, and the spores
were completely smooth even by SEM (Figs. 91, 93). The wrinkled sporotheca
breaking into platelets instead of the lid of L. parasitica easily avoids any
confusion even with the sessile specimens. The plasmodium, visible in moist
chamber cultures, is thick dull brown and appears as a continuous sludge on
the bark surface like many large protoplasmodia combined.
In the original description of L. rugosa Nannenga Bremekamp &
Yamamoto (1987) differentiated it from L. pedicellata on the basis of the spore
colour being dark brown and the smooth spores. The spores are also slightly
bigger in L. rugosa and the smooth surface vs. the warted surface of L.
pedicellata spores (Figs. 68-70) is confirmed by SEM. The peridial
ornamentation is also different in density and distribution (Figs. 64, 90) as
revealed by SEM examination, and some of the warts are fused in L. rugosa
(Fig. 90). However, L. rugosa and its var. fujiokana (see below) are very like
L. pedicellata but with the definable differences listed (Table 2). DNA
sequencing techniques may demonstrate that they are in fact a continuum of
the same species, but at present pending further studies, we prefer to maintain
them as separate taxa.
14. Licea rugosa var. fujiokana (Y. Yamam.) D. Wrigley & Lado, comb. nov.
(Figs. 94-99)
≡ Licea capitatoides var. fujiokana Y. Yamam., Bull. Natl. Sci. Mus., Tokyo, B 26(3):
112. 2000. TYPE: JAPAN, Tokyo Pref., Chiyoda-ku, on the premises of Imperial Palace, on
bark of living Acer buergerianum, 3 VI 1999, Y. Yamamoto, 99TK-29 (holotype: TNS!;
isotype 99TK-26 in TNS).
Sporocarps scattered, stipitate, 150-320 µm in height. Sporothecae dark
brown, globose or subglobose, 80-200 µm diam. Stalk 70-160 µm in height,
slightly tapering towards the top, filled with refuse material, mid-width half the
height. Peridium double, wrinkled, outer layer gelatinous with granular refuse
material, inner layer membranous, yellow-grey (93. y Gray) with a densely
warted surface; dehiscence by irregular fracture into fragments in upper
peridium leaving a calyculus. Spores free, greyish yellow brown (80. gy. Y
Br), subglobose, (9-)10-11 µm diam., smooth by TL and SEM; spore wall with
an obvious paler area about half the diam. of the spore (Figs. 96-97).
296

Fungal Diversity
Figs. 94-99. Licea rugosa var. fujiokana (99TK29 as Licea capitatoides var. fujiokana Y.
Yamam.). 94. Habit. 95. Dehiscing sporocarp by TL. 96. Spores by TL with paler area. 97.
Spore by SEM. 98. Inner surface of peridium by TL. 99. Group of spores and inner surface of
peridium by SEM.
Material examined: JAPAN, Tokyo Pref.: Chiyoda-ku, bark of living Acer
buergerianum in moist chamber, 3-VI-1999, leg. Y. Yamamoto, 99TK-29 (TNS, holotype).
MEXICO, Tlaxcala: Calpulalpan, El Peñón, on Abies religiosa in moist chamber, 19-II-1998,
VGA 671. Hidalgo: El Cardonal, 20°38’56”N 99°00’11”W, on bark of Prosopis juliflora in
moist chamber, 9-X1-2000, dwb 1851. Metzquititlan, 20°29’48”N 98°40’03”W, on bark of
Acacia sp. in moist chamber, 5-X-2001, dwb 2076, idem, 23-XI-2001, dwb 2067. Puebla:
Emilio Portes Gil, 19°17’40”N 97°30’22”W, on bark of living Nolina parviflora in moist
chamber, 18-I-2001, dwb 1932; on bark of living Yucca filifera in moist chamber, 25-I-2001,
dwb 1955; 3-II-2001, dwb 1979. Zapotitlán de las Salinas, 18°19’22”N 97°29’57”W, on bark
of living Beaucarnea gracilis in moist chamber, 1-X-2001, dwb 2077. S. Martin Esperilla,
18°44’45”N 97°31’44”W, on bark of living Yucca periculosa in moist chamber, 22-X1-2003,
dwb 2348; idem, 29-X1-2003, dwb 2344. Veracruz: Totalco, 19°28’56”N 97°22’00”W, on
bark of living Juniperus deppeana in moist chamber, 18-I-2001, dwb 1966. San Luis Potosi:
297

Xilitla, Reten, 21°18’04”N 99°05’07”W, on bark of living Liquidamber styraciflua in moist
chamber, 26-X1-2001, dwb 2104. Oaxaca: Tepelmeme, La Unión, 18°06’33”N 97°20’31”W,
on bark of living Prosopis laevigata in moist chamber, 15-XI-2003, dwb 2316; Tepelmeme,
Mex-135 highway, Km. 109, 18°00’35”N 97°21’19”W, on bark of living Yucca periculosa in
moist chamber, 29-X1-2003, dwb 2349.
Habitat: bark of living trees.
Distribution: Japan, Mexico.
Illustrations: Yamamoto (2000: 112, Fig. 6).
When this species was originally described by Yamamoto as L.
capitatoides var. fujiokana he stated that it differed from L. capitatoides by the
colour and size of the spores (greyish yellow and 10-12 µm diam. in the
former, and pale yellow, 8-10 µm diam. in the latter). However comparison of
the types showed additional differences. Licea rugosa var. fujiokana has a
roughened sporotheca when dry, a lighter stalk by TL and darker spores (Fig.
95). SEM examination showed the spores to be totally smooth (Fig. 97) like L.
rugosa var. rugosa, although smaller. The peridium fractured into fragments of
approximately 30 µm in diam. (Fig. 98), and the inner peridium by SEM (Fig.
99) was covered with warts with some fused warts. This ornamentation is the
same as that of the inner peridium of L. rugosa var. rugosa, and different in
both density and distribution from L. capitatoides. There are also patches of
refuse material on the outer surface of the peridium, which suggest the remains
of a gelatinous layer. The Mexican material was examined by SEM, and is
identical to the type.
The type of var. fujiokana is therefore exactly the same as the type of var.
rugosa in all other characters except its double peridium, spore size (10-12 µm
diam. in var. fujiokana and 14-16 µm diam. in var. rugosa) and the yellowish
brown spore colour of var. fujiokana (80. gy. y Br) instead of olive brown in
var. rugosa (94. l. Ol Br). For this reason the new combination as a variety of
L. rugosa is justifiable. The two taxa can be separated under the light
microscope on the basis of the darker, larger spores of L. rugosa var. rugosa
(Table 2).
15. Licea scyphoides T.E. Brooks & H.W. Keller in Keller & Brooks,
Mycologia 69(4): 679. 1977. TYPE: U.S.A., Ohio, Greene County, next to the
swimming pool at John Bryan State Park, from the bark of living Juniperus
virginiana, August 18, 1976, H.W. Keller 1945 (holotype: BPI!).(Figs. 100-109)
= Licea tanzanica Ukkola, Härk. & Gilert in Ukkola, Härkönen & Saarimäki, Karstenia
36(2): 57. 1996, syn. nov. TYPE: TANZANIA, Tanga Prov., Lushoto Distr., W Usambara
Mts., on Azadirachta indica in moist chamber cultura, 12.XII.1988, Härkönen 3693 (holotype:
H!).
Sporocarps scattered to gregarious, stipitate, 150-280(-400) µm in height.
Sporotheca globose, golden brown, shiny, 80-160(-220) µm diam. Stalk 70-
298

Fungal Diversity
299
Figs. 100-105. Licea scyphoides (HWK 1945). 100. Habit. 101. Base of sporotheca from above
containing spores by SEM. 102. Inner surface of peridium by SEM. 103. Upper half of
sporotheca showing clear area of dehiscence by TL. 104. Spore with closely punctate
ornamentation by SEM. 105. Group of spores appearing roughened.

Figs. 106-109. Licea scyphoides (MH 3693 as Licea tanzanica). 106. Habit. 107. Dehisced
sporocarp leaving calyculus. 108. Inner surface of peridium. 109. Spore by SEM with closely
punctate ornamentation.
180 µm in height, dark brown by reflected light, 150 µm in height, thick, erect,
furrowed, straight, cylindrical, mid-width half of the height. Peridium single,
membranous, with refuse matter on outside, inner surface of the peridium
clearly punctate (Fig. 102); dehiscence circumcissile at a transparent equatorial
band, visible by TL, leaving a basal calyculus (Fig. 101). Spores free,
yellowish (90. gy. Y-104. p. g Y) by TL, globose, 11-13.5(-14) µm diam.,
roughened to smooth by TL and low magnification SEM (Figs. 101, 105), and
closely punctate by at higher magnification SEM (Fig. 104); spore wall with a
paler thinner area.
Material examined: USA, Ohio: Greene county, John Bryan State Park near swimming
pool, on bark surface of red cedar, 18-XIII-1976, coll. Keller 1945 (BPI 826487, holotype).
CUBA, Sancti Spiritus: Alturas de Banao, 21°52’39”N 79°37’03”W, on dead liana in moist
300

Fungal Diversity
chamber, 6-II-2003, dwb 2255. MEXICO, Hidalgo: El Cardonal, 20°38’57”N 99°00’11”W,
bark of living Prosopis juliflora in moist chamber, 9-X1-2000, dwb 1865. Progreso, La Cruz,
20°16’02”N 99°10’48”W, bark of living Schinus molle in moist chamber, 13-III-2001, dwb
1989; ibidem, bark of living Prosopis laevigata in moist chamber, 13-III-2001, dwb 1992, dwb
1993; ibidem, bark of living Opuntia sp., 16-III-2001, dwb 2004. PERU. Mazan: 3°28’S
74°55’W, dead liana in moist chamber, 27-V-2002, dwb 2193; ibidem, 4-VI-2002, dwb 2198;
ibidem, 3-VI-2002, dwb 2232. PUERTO RICO, Fajardo, Dry Coastal Forest on rocks, Las
Cabezas de San Juan Nature Reserve, near ‘lands end’, 18°23’13”N 65°37’29”W, bark of
living Tamarindus indica in moist chamber, sc 16930. SPAIN. Cuenca: Saceda-Trasierra, bark
of living Quercus ilex, in moist chamber, 21-III-1990, 1184-1 Pando, MA-Fungi 31963.
UNITED KINGDOM, Wales: Aber, bark of living Quercus petraea, in moist chamber, 31-I-
1993, dwb 1130a. TANZANIA, Tanga Province: Lusotho District, West Usambra Mts.,
Mombo, town centre, yard of the restaurant, 0438CD, on Azadirachta indica in moist chamber,
12-XII-1988, MH 3693 (H holotype); idem, MH 3591, MH 3592. Arusha (Northern) Province:
Moshi District. Moshi International school park, 0337AD, on Jacaranda in moist chamber, 23-
V-1988, MH 3576 as Licea tanzanica.
Habitat: Bark of living trees and living and dead vines.
Distribution: Mexico, Peru, Spain, Tanzania, USA, Wales.
Illustrations: Keller and Brooks (1977: 680, Figs. 23-28); Ukkola et al. (1996: 56, 58);
Lado and Pando (1997: 132, Fig. 38); Flatau (2000: 70, Fig. 5); Yamamoto (1998: 145); de
Haan (2001: 16, pl. 1).
The sporocarps examined were from 150-280(-450) µm in total height
and 84-160(-200) µm diam., with globose, golden brown shiny sporotheca. The
peridium contained refuse matter which thins towards the middle of the
sporotheca leaving a transparent equatorial band, very obvious on both sides of
the line of dehiscence when specimens are mounted and viewed by TL (Fig.
103). The warts of the inner suface of the peridium are visible in the
transparent area of dehiscence by TL at 400x. In this species as with several
other stipitate Licea, some sporocarps in dry material appear sessile if they are
fallen against the substrate and the stalk blends with the substrate refuse
material.
We examined the type material and several other specimens of L.
tanzanica, which we found to be very similar to this species. The SEM
examination showed the spore ornamentation (Figs. 104, 109) and the
ornamentation of the inner peridium (Figs. 102, 108) of the two species to be
identical. There are no differences in the spore sizes (L. scyphoides 11-14 µm
diam. and L. tanzanica 12-13 µm diam.). The types show L. tanzanica
somewhat larger, reaching 400 µm in height and 200 µm in diam., but the sizes
overlap. The ratio of stalk width to height is also different with L. scyphoides
shorter and squatter. In L. scyphoides a cup is left after dehiscence but L.
tanzanica dehisces lower down and leaves only the base of the sporotheca. The
spore colour is also subtly different. Licea tanzanica spores are olivaceous and
L. scyphoides are pale yellow without the darker tints. The transparent border
visible at the area of circumcissile dehiscence is a constant feature of both and
301

measures up to 70 µm. The authors of each species recognized that they are
very close, but believed them to be different (Ukkola et al., 1996). We feel the
combination of small differences may represent geographic differences, but are
not sufficient to maintain these as separate species, and we synonymize them
here. Perhaps these small differences indicate a continuum of a variable
species.
Licea bulbosa (see above) is similar to L. scyphoides in habit, but its
smaller (10-11 µm diam.) almost hyaline smooth spores, its longer thinner
stalk, and the smooth inner peridium with a totally transparent basal half
differentiate it.
16. Licea verrucospora (T.N. Lakh., Nann.-Bremek. & R.K. Chopra) D.
Wrigley & Lado, nom. nov. et status nov. (Figs. 110-115)
≡ Licea scyphoides var. reticulata T.N. Lakh., Nann.-Bremek. & R.K. Chopra, Proc.
Kon. Ned. Akad. Wetensch. 93(3): 261. 1990 [basion.], non. L. reticulata Berk. & Broome,
1873. TYPE: INDIA, Himalchal Pradesh, North Western Himalaya mountains, Shimla,
Summer Hill area, 2067 m, developed in a moist chamber on bark of Quercus incana from 0-9
m above the ground, 6 IV 1981 (holotype HPUB 15405, paratype NENB 13.877 now at BR!).
Sporocarps scattered to gregarious, stipitate, 250-500(-700) µm in height.
Sporothecae dark ochraceous-brown, shiny, subglobose, 100-200 µm diam.
Stalk 150-350(-500) µm in height, long, dark, straight, subcylindrical, rugose
with several strands visible by TL within the main stalk (Fig. 110), mid-width a
quarter of stalk height. Peridium single, membranous, covered with refuse
matter, smoky grey, densely and prominently warted; dehiscence irregular into
platelets formed by fragmentation of the whole peridium. Spores free, pale
yellow to greyish yellow (89. p. Y - 90. gy. Y) by TL, globose, 10-12(-13) µm
diam., very minutely warted but warts in irregular patches on the spore surface,
visible by TL and also by SEM (Fig. 115); spore wall with a pale thinner area
(Fig. 111).
Material examined: INDIA, Himalchal Pradesh: Shimla, Summer Hill, on bark of living
Quercus incana in moist chamber, 14-IV-1981, NENB 13.877 (BR, paratype of Licea
scyphoides var. reticulata)
Habitat: bark of living Quercus incana.
Distribution: Known only from the type locality.
Illustrations: Lakhanpal et al. (1990: 261, Fig. 7).
We raise its rank here to species level because on comparison of the
material with L. scyphoides we noticed many differences, sufficient to consider
it to be a totally different species. The original description of L. scyphoides var.
reticulata notes that dehiscence takes place above an equatorial line to leave a
cup with a smooth margin. We found no such line, and in fact the platelet
margins are visible extending to the base of the sporotheca (Figs. 110, 113).
The base, when remaining after dehiscence, is a thin vestige of peridium more
302

Fungal Diversity
303
Figs. 110-115. Licea verrucospora (NENB 13.877 as L. scyphoides var. reticulata). 110.
Habit. 111. Dehisced sporocarp by TL. 112. Detail of inner surface of peridium by SEM with
flattened reticulate markings (arrow). 113. Fragmented peridium by SEM. 114. Stalk and
fragmented remains of sporotheca. 115. Spore showing irregular verrucose ornamentation.

like a collar than a cup (Fig. 114), whereas in L. scyphoides a clear cup is left
after dehiscence. Licea scyphoides has a transparent equatorial band at the area
of dehiscence which is not present in L. verrucospora. The inner surface of the
peridium in L. scyphoides is minutely warted (Fig. 102), but the inner surface
of L. verrucospora is densely marked with protruding warts of various sizes,
and the lines of the outer reticulate pattern show through (Fig. 112). The spores
in L. verrucospora have a unique ornamentation of rounded raised warts of
different sizes, dispersed irregularly over the spore surface, different from L.
scyphoides even by TL, and more so by SEM (Fig. 115). This pattern is also
different from any other stipitate Licea described so far. These differences are
in addition to those noted by the authors of this species which were the larger
size of the sporocarps but they overlap, and its reticulately-ridged peridium.
The surface of the sporothecae of the type material is crossed with a network of
shiny lines on ridges, marking the surface, but not the platelets by which it
dehisces (Fig. 112).
The epithet verrucospora was chosen to denote the character of the spore
surface, since the variety name has already been used for a different species
(Lado, 2001).
Licea verrucospora is different from L. tanzanica in spore size (10-12
µm diam. vs. 12-15 µm diam.), ornamentation, and in its mode of dehiscence
(platelets vs. circumcissile). Its yellowish, irregularly ornamented spores, also
separate it from L. pedicellata with regular ornamentation and L. rugosa which
has completely smooth spores.
Excluded species: (6)
Licea atricapilla Nann.-Bremek. & Y. Yamam., Proc. Kon. Ned. Akad.
Wetensch., C 86(2): 208. 1983. TYPE: JAPAN, Kochi Pref., Susaki-shi,
Hachimangu, cult. on bark of living Cinnamomum camphora (L.) Presl, 7-31 X
1981 (holotype: Y.Y. 1277, isotype NENB 12.984).
We were unable to examine this species. The following description is
adapted from Nannenga-Bremekamp and Yamamoto (1983).
Sporocarps scattered, stipitate, 260 µm in total height. Sporotheca silvery
irridescent with a dark apical operculum, subglobose, 160 µm diam. Stalk
nearly black, furrowed, filled with granular refuse material. Peridium
membranous, separate from the stalk, densely and minutely warted on the inner
surface. Dehiscence circumscissile along edge of the apical disc. Spores,
olivaceous brown with a large paler area and a thinner wall, 12-13 µm diam.,
smooth by TL.
Habitat: bark of living Cinnamomum camphora.
Distribution: only known from type locality.
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Fungal Diversity
Illustrations: Nannenga-Bremekamp and Yamamoto (1983: 208); Yamamoto (1998:
136).
The authors place this species in the subgenus Orcadella on account of
its black operculum. They differentiate it from L. operculata which has a
glossy operculum, and by its larger darker spores (olivaceous brown and 12-13
µm diam. vs. 8-11 µm diam. and almost colourless in L. operculata). They also
comment that L. scyphoides differs from it by spore colour (greyish yellow in
the latter species), “a more densely warted peridium” and “a larger lid, which
occupies half of the sporangium”. The description of this species is unlike any
we have examined.
Licea crateriformis B. Ing, Myxomycetes Britain and Ireland 46. 1999. TYPE:
UNITED KINGDOM, Scotland, Isle of Arran, Kings Cross Point, in thallo
vivo Radulae complanatae in rame vivo Corylus avellanae, 2 September 1998,
B. Ing (holotype: Hb. B. Ing 98068).
The following description is adapted from the original publication by Ing
(1999) since we were unable to obtain the type material of this species for
examination.
Sporocarps short stipitate, 800 µm in height. Sporothecae 700-750 µm
diam. shaped like a Craterium. Stalk dark, furrowed, from a discoid
hypothallus. Peridium single, dark brown to chestnut with a distinct golden-
bronze lid; dehiscence by lid. Spores colourless by TL, 7-9.5 µm diam.,
subglobose, smooth; spore wall of uniform thickness.
Habitat: on liverworts on living branches.
Distribution: Known only from the type locality.
Illustrations: Ing (1999: 47).
According to the author´s description, the shape and dehiscence of this
species are similar to Licea poculiformis, but that species is much smaller [150-
200(-400) µm in height] and the spores have a wall with a thinner area.
Licea erecta var. erecta K.S. Thind & Dhillon, Mycologia 59(3): 463. 1967.
TYPE: INDIA, Eastern Himalayas, Darjeeling, Tiger Hill, on decaying
bamboo twigs, Septembar 23, 1964, No. 451 (holotype: PAN).
We were unable to obtain the type material of this species for
examination. The following is adapted from the original description by Thind
and Dhillon (1967).
Sporocarps scattered, stipitate, up to 1500 µm in height. Sporothecae
dark brown, oblong or ovate, up to 600 × 400 µm. Stalk up to 900 µm in
height, 200 µm wide, long, erect, ridged, concolourous. Peridium tough, thick,
with dark granular inclusions, marked with ridges from the top of the stalk and
305

wrinkles between the segments; dehiscence irregular from above. Spores free,
hyaline by TL, 13.5-15 µm diam., minutely verrucose to almost smooth.
Habitat: decaying bamboo twig.
Distribution: India; Thailand.
Illustrations: Thind and Dhillon (1967: 464).
Licea erecta var. erecta, according to Martin and Alexopoulos (1969) is
“somewhat similar to L. operculata, from which it differs in its more robust
habit, complete lack of an operculum, and particularly in the netted peridium
which approaches that of a Cribraria, although there is no evidence that the
peridium falls away from the interstices of the net and there is no Cribraria
which approaches it in appearance or has such large spores.”
The L. erecta var. erectoides is described above. As both de Haan (2002)
and Yamamoto (1999) point out, the main difference between L. erecta var.
erecta and the var. erectoides is the spore size. One of de Haan´s SEM
photographs shows a spore of the var. erecta from the collection of Nannenga-
Bremekamp (14807). The material illustrated by de Haan (2002), and
examined by us, is a collection labeled by Nannenga-Bremekamp as a
questionable identification (Licea erecta Thind? & Dhillon). It is much smaller
than the described L. erecta and has a round sporotheca and short stalk.
Without the type to examine, it is impossible to evaluate this species.
Licea floriformis var. floriformis T.N. Lakh. & R.K. Chopra in Lakhanpal,
Nannenga-Bremekamp and Chopra, Proc. Kon. Ned. Akad. Wetensch. 93(3):
255. 1990. TYPE: INDIA, Himalchal Pradesh, North Western Himalaya
mountains, Shimla, Summer Hill area, 2067 m, developed in a moist chamber
on bark of Quercus incana taken 3-6 m the ground, harvested on 4 VIII 1981
(holotype: HPUB 76423).
The following description is adapted from Lakhanpal et al. (1990) since
we were unable to examine the type material of this species.
Sporocarps scattered, stipitate, up to 700 µm in height. Sporothecae
globose or subglobose, red-brown to nearly black, smooth or wrinkled with
minute ridges, 250-350 µm diam. Stalk 250-450 µm in height, sturdy,
subconical, nearly black. Peridium single, membranous, thin, inner surface
bearing long papillae; dehiscence by irregular fracture in the upper sporotheca
and lobes below. Spores free, smooth, smooth, black in mass, light pale pink to
colorless by TL, 11-12(-14) µm diam.
Habitat: bark of living Quercus incana.
Distribution: Known only from the type locality.
Illustrations: Lakhanpal et al. (1990: 255) [a drawing by Nannenga-Bremekamp from a
photograph].
In the description of L. floriformis var. floriformis, Lakhanpal et al.
(1990) note that it differs from L. erecta var. erecta K.S. Thind & Dhillon only
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Fungal Diversity
in its slightly smaller spores, but that they overlap in size. The spores in both
are smooth. Perhaps these are the same species, but without being able to
examine the type specimens of either species, it is impossible to be sure.
Licea lilacina Nann.-Bremek., T.N. Lakh. & R.K. Chopra in Lakhanpal,
Nannenga-Bremekamp and Chopra, Proc. Kon. Ned. Akad. Wetensch. 93(3):
256. 1990. TYPE: INDIA, Himalchal Pradesh, North Western Himalaya
mountains, Shimla, Summer Hill area, 2067 m, developed in a moist chamber
on bark from a living Quercus incana, 3 m above the ground (holotype: HPUB
11,001; isotype NENB 13.889 now at BR).
We were unable to obtain the type material of this species. The following
description is adapted from Lakhanpal et al. (1990).
Sporocarps scattered, or in pairs, stipitate or sessile, 600-700 µm in
height. Sporothecae globose or subglobose 200-350 µm diam., violet-brown or
irridescent blue/violet, sometimes sprinkled with white particles. Stalk if
present up to 300 µm long, (the description in English has an errata, reporting
the stalk up to 3 mm long. In the Latin description it is 0.3 mm which is in
keeping with other measurements) thick, opaque or ochraceous with refuse
matter, slightly tapering towards the top, width about half of the height at base.
Peridium double, the outer layer gelatinous with little granular refuse at the
base, the inner layer membranous ochraceous, smooth or minutely wrinkled;
dehiscence by irregular fracture, sometimes lobately from above leaving a cup
with an irregular margin. Spores pale lilaceous in mass, paler rosy-lilac by TL,
(8-)9-10(-12.5) µm diam., densely and minutely spinulose; spore wall
uniformely thin.
Habitat: bark of living Quercus incana and Pinus wallichiana.
Distribution: Known only from the type locality.
Illustrations: Lakhanpal et al. (1990: 256)
The authors point out that this species has lilac spores which, suggests
other orders and not the Liceales. They include it in the genus Licea because of
the absence of capillitium and a double peridium. Their reference to a
sprinkling on the peridium of white particles also suggests another group,
which is an opinion we share. Apparently plentiful material was collected (38
collections), but the second author (Nannenga-Bremekamp) states that she saw
only two slides.
Licea takahashii Y. Yamam., Bull. Natl. Sci. Mus., Tokyo, B 26(3): 114. 2000.
TYPE: JAPAN, Gifu Pref., Osaka-machi, Nigorigo-onsen, ca. 1950 m alt., on
dead wood, 8 X 1994, K. Takahashi, YY-14468 (holotype: TNS!).
(Figs. 116-117)
307

Figs. 116-117. Licea takahashii (YY 14468). 116. Habit. 117. Spiny thick-walled spores and
immature elaters (arrows).
We have examined the type material YY 14468 and we have detected
some rudimentary capillitium, made up of small, immature poorly formed
elaters (Fig. 117). It appears to be an immature Trichia instead of a Licea. This
is supported by the fact that it was found on dead wood. It also has very large
mottled sporocarps (up to 2 mm tall and 500 µm diam.), with a yellow spore
mass and spiny thick-walled yellow spores, 12-14 µm diam. We thus exclude
this material from the genus Licea. The authors of the species mentioned its
similarity in habit to Trichia or Hemitrichia but they said “it obviously lacks
capillitium”.
Discussion
More than ninety collections were examined for this evaluation. The
simultaneous study of 21 type specimens of these Licea species has enabled us
to make detailed comparisons and come to the conclusion that in five cases, L.
capitata, L. longa, L. cristallifera, L. tanzanica and L. tropica, the newly
described species actually belonged to an existing taxon. They have been
synonymized with L. floriformis var. aureospora (the first two), L. eleanorae,
L. scyphoides and L. bulbosa, respectively. In the case of L. capitatoides var.
fujiokana, it is recombined as a variety of a different species, Licea rugosa var.
fujiokana. A new name Licea verrucospora, and a new status was given to
Licea scyphoides var. reticulata, since it was found to be so different from L.
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Fungal Diversity
scyphoides that it has been raised to species level. In the case of L. takahashii,
it was excluded from the genus Licea because it is an immature form of
Trichia, a different genus.
Among the accepted 16 stipitate Licea species, there do appear to be
loose groups. There are those with dehiscence by a lid (L. operculata and L.
poculiformis), and those with circumcissile dehiscence (L. scyphoides and L.
bulbosa), although the top half of the sporotheca is not really a lid as in the
previous two species. There are also species with dehiscence by irregular
fracture of some form, or platelets separated by definite lines of dehiscence.
Where there are platelets, however, they are not as well defined as they are in
sessile species like L. minima Fr., and it is difficult to separate this form of
dehiscence from fracture into irregular fragments. Most of these Licea had not
been described in 1965, and they loosely fit only two of the subgenera
(Orcadella and Pleiomorpha) proposed by Nannenga-Bremekamp (1965). It
seems to be reasonable, therefore not to maintain her subgenera, at least in the
case of the stipitate members of the group, although the character of type of
dehiscence, when used in conjunction with others, is useful in separating
species.
Other groupings are also possible if we consider those with smooth
spores by SEM (L. rugosa and L. rugosa var. fujiokana) or those with a double
peridium (e.g. L. bulbosa, L. eleanorae, L. capitatoides or L. poculiformis), but
the use of SEM is not available to all taxonomists, and the description of the
peridium as single or double depends on the state of the material and is open to
error in dry specimens. To separate these species it is necessary to use other
characters such as the size, colour and ornamentation of the spores and
peridium, and stalk size and colour in conjunction with the type of dehiscence,
a double or single peridium and SEM examination where possible.
Using a combination of these characters, a key has been proposed below
to aid in the identification of the species discussed in this paper. The six
excluded species have been placed in the key to serve as a basis for
comparison. Characters from their original published descriptions, and a
different typeface has been used for these taxa.
In the descriptions of the material we have given reasons for considering
each of these to be different taxa of Licea. The fact that some of these species
were simultaneously in press (Table 1), coupled with the variability of
specimens from moist chamber culture and the simplicity of the sporophores,
can easily explain the taxonomic complexity. Six of the species studied have
only been found from the type locality. This may be because they are newly
described, and therefore there has not been time for other collections, or
perhaps that many environments have been little sampled until recently. The
309

Key to the stipitate Licea
1. Spores by TL dark olive, olive brown, dark brown, yellowish grey ................................. 2
1. Spores by TL very pale or colourless ............................................................................... 5
2. Spores warted .....................................................................................10. Licea pedicellata
2. Spores smooth ................................................................................................................... 3
3. Dehiscence along an apical disc ............................................................... Licea atricapilla
3. Dehiscence into irregular fragments ................................................................................. 4
4. Spores 14-16 µm diam. ......................................................... 13. Licea rugosa var. rugosa
4. Spores 9-12 µm diam. ...................................................... 14. Licea rugosa var. fujiokana
5. Outer surface of sporotheca covered with birefringent crystals ........................................ 6
5. Outer surface of sporotheca not covered with crystals ..................................................... 7
6. Sporotheca vase-shaped, without a clear line of dehiscence.................. 3. Licea eleanorae
6. Sporotheca subglobose, with a clear line of dehiscence .......................... 1. Licea bulbosa
7. Outer surface of sporotheca with distinct spots or ridges ................................................. 8
7. Outer surface of sporotheca without distinct spots or ridges ............................................ 9
8. Spotty appearance from large warts on sporotheca surface ............. 9. Licea parvicapitata
8. Sporotheca surface covered with a reticulum of ridges ................. 16. Licea verrucospora
9. Sporocarps more than 400 µm tall................................................................................... .10
9. Sporocarps up to 400 µm tall ........................................................................................... 18
10. Dehiscence by lid or circumcissile .................................................................................. 11
10. Dehiscence neither by a lid nor circumcissile .................................................................. 13
11. Dehiscence circumcissile ....................................................................15. Licea scyphoides
11. Dehiscence by a lid........................................................................................................... 12
12. Long stalk, more than half total height ................................................ 8. Licea operculata
12. Short stalk, less than half total height .................................................. Licea crateriformis
13. Peridium single ................................................................................................................ 14
13. Peridium double .............................................................................................................. 16
14. Peridium ochraceous by TL ................................................................ 4. Licea erddigensis
14. Peridium colourless by TL ............................................................................................... 15
15. Spores 10-12 µm diam. ........................................................5. Licea erecta var. erectoides
15. Spores 13-15 µm diam. .................................................................. Licea erecta var. erecta
16. Spores golden yellow in mass ................................... 6. Licea floriformis var. aureospora
16. Spores not golden yellow in mass .................................................................................... 17
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Fungal Diversity
17. Spores smooth ................................................................ Licea floriformis var. floriformis
17. Spores densely and minutely spinulose ......................................................... Licea lilacina
18. Appears almost sessile .................................................................................................... 19
18. Obviously stalked ........................................................................................................... 21
19. Dehiscence circumcissile ................................................................... 15. Licea scyphoides
19. Dehiscence irregular ........................................................................................................ 20
20. Peridium golden irridescent, with refuse material .............................. 11. Licea perexigua
20. Peridium hyaline, transparent, with no refuse covering .............................. 7. Licea lucens
21. Dehiscence into irregular fragments .................................................. 2. Licea capitatoides
21. Dehiscence by lid or circumcissile .................................................................................. 22
22. Dehiscence by a lid ......................................................................... 12. Licea poculiformis
22. Dehiscence circumcissile ................................................................................................. 23
23. Inner peridium smooth; lower half of peridium transparent ..................... 1. Licea bulbosa
23. Inner peridium warted; transparent only at line of dehiscence ...........15. Licea scyphoides
known distribution of some of the species, for example L. erecta var.
erectoides from Costa Rica, Cuba and Japan, or L. rugosa var. rugosa from
Japan and Mexico, is obviously a reflection of areas sampled rather than true
distribution, and reflects the scarcity of data for this group in some parts of the
world. Six of the taxa were found by us (Lado et al., 2003; Stephenson et al.,
2004) during recent intensive sampling in Mexico in undisturbed environments
such as Biosphere Reserves or National Parks, which had not been sampled
previously. This could be an explanation for their rarity, and is a further
compelling reason for carrying out intensive biodiversity inventories, and for
supporting future conservation of these areas. It is certainly evident that the
study of many collections is necessary to fully appreciate the variability of a
species, and to be certain that the specimens obtained from moist chamber
culture are properly mature. The presence of different sized or irregularly
shaped spores is a clue to immaturity or poor development, and mounting
moist fresh specimens that have not dried slowly will often give atypical
dehiscence. In the case of L. operculata, the dwarf forms may be an ecotype
prevalent in tropical habitats. These dwarf forms have also been seen for other
species, like Didymium squamulosum (Alb. & Schwein.) Fr. or Arcyria cinerea
(Bull.) Pers., from our moist chamber cultures.
But how many differences are sufficient to consider a species of Licea as
separate? In a group with so few characters, and the somewhat arbitrary
relative taxonomic value given to these different characters (dehiscence,
separation of the sporotheca from the stalk, ornamentation of spores and
311

peridium), coupled with apomictic life cycles (Clark, 2004), the use of
morphospecies, becomes questionable. The application of scanning electron
microscopy has certainly helped to answer some doubts and questions about
morphological characters, but further information is needed to confidently
confirm that these myxomycetes belong to separate species and are not
ecotypes or variants of fewer species. Consensus on taxonomic boundaries
informed by other techniques such as DNA sequencing, will be necessary to
corroborate these findings. It is still in its infancy in this group and until now
applied to plasmodia (Baldauf and Doolittle, 1997), mainly laboratory-
cultured, although primers specific for extraction from sporocarps have also
recently been designed (Martín et al., 2003). Spore to spore culture can be
useful, though difficult with these very small myxomycetes, to determine the
stability of the characters of new species (Keller, 1996).
The only way to be really sure that a described species is not a small
clonal population, or morphological variant of another species is, as Keller
(1996) and Clark (2004) recommended, the collection and examination of a
large amount of material for study from different and widely separated areas,
and even a study of the reproductive systems in these collections.
Acknowledgements
We would like to thank all the keepers of the following herbaria B, BPI, BR, H, MA,
TNS, and L. Flatau, M. de Hann, B. Ing, F. Pando, M. Schnittler and A. Varela-García for
supplying the material for this study. We would also like to thank Miguel Jerez of the Real
Jardín Botánico, CSIC for his valued help and skill with the SEM work. This project was
supported by the Spanish Ministry of Education and Science (REN2002-00445/GLO) and the
National Science Foundation, USA (Grant # DEB-9705464).
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