Lamprospora verrucispora sp. nov. (Pezizales)

Abstract

A description and illustrations of both macroscopic and microscopic characters of Lamprospora verrucispora sp. nov. are provided. Differences from other bryophilous Pezizales with globose ascospores ornamented with isolated warts are discussed and a dichotomous key to these species is given. Gall formation on rhizoids and chloronema of the bryophyte host Campylopus pyriformis is described and illustrated. This represents the first report of gall formation for a species of the genus Lamprospora.

Full text

163
Lamprospora verrucispora sp. nov. (Pezizales)
Marcel VEGA
Jan ECKSTEIN
Henk-Jan VAN DER KOLK
Ascomycete.org, 8 (4) : 163-171.
Juillet 2016
Mise en ligne le 08/07/2016
Abstract: A description and illustrations of both macroscopic and microscopic characters of Lamprospora
verrucispora sp. nov. are provided. Dierences from other bryophilous Pezizales with globose ascospores or-
namented with isolated warts are discussed and a dichotomous key to these species is given. Gall formation
on rhizoids and chloronema of the bryophyte host Campylopus pyriformis is described and illustrated. This
represents the rst report of gall formation for a species of the genus Lamprospora.
Keywords: Bryophilous Pezizales, Campylopus pyriformis, dichotomous key, galls, Pyronemataceae, taxo-
nomy.
Introduction
Since the website Ascofrance.com with its forum has been put
online by its founder and owner Christian Lechat in 2003, more than
1200 users from countries all over the world have decided to be-
come a member of this platform aiming at the international scien-
tic exchange of mycologists specialized in ascomycetes. There are
stories galore about contacts, cooperations and friendships arisen
from this unique website.
Ours started in January 2014 when the third author of this article
asked for help to identify a Lamprospora he could not determine
with the existing literature. First mails, then specimens went from
one to another of the authors. Based on ascospore ornamentation
and its type of infection on the bryophyte host Campylopus pyrifor-
mis (K.F. Schultz) Brid., the Lamprospora soon proved to be an un-
described species. One and a half years later, the new species was
found by the rst author in further sites in great abundance and sub-
sequently is described here as L. verrucispora.
Methods
The description of Lamprospora verrucispora is based on the re-
sults of the examination of several vital collections from ve locali-
ties: one in the Netherlands and four in Germany. Most observations
took place in tap water. Ascospore ornamentation was additionally
studied after staining with Lactophenol Cotton Blue or Methylene
Blue, and the iodine reaction of asci was examined with Lugol’s so-
lution. Ascospores’ size was measured from spore prints, which were
acquired by placing a cover glass on fresh apothecia. Additional
measurements from rehydrated material were found to be within
the range of those gained from vital material and are therefore not
stated separately. Macrographs were made with digital cameras. Mi-
crographs were taken either in tap water, Lactophenol Cotton Blue
or Methylene Blue, using digital cameras mounted on microscopes
and software for stacking pictures. Scanning electron micrographs
(SEM) were taken with a LEO-438 machine from air dryed samples.
Taxonomy
Lamprospora verrucispora M. Vega, Eckstein, Van der Kolk, sp.
nov. — MycoBank 817460.
Diagnosis: Diers from other species of Lamprospora by its per-
fectly globose ascospores with a surface densely covered with nu-
merous small isolated warts and few isolated bigger warts together
with an infection inducing galls on the rhizoids of its host Campylo-
pus pyriformis (K.F. Schultz) Brid.
Holotype: The holotype of Lamprospora verrucispora is deposited
in the Herbarium Hamburgense of the University of Hamburg (HBG–
holotypus, inv. no. 1412). The isotypes of Lamprospora verrucispora
are led under MV20151025-04 in the private herbaria of Marcel
Vega and Jan Eckstein.
Etymology: verrucispora = referring to the warted ascospores of
the species.
Macroscopic features (plate 1)
Apothecia mostly scattered, sometimes gregarious on soil bet-
ween shoots of Campylopus pyriformis, protonema and other bryo-
phytes; 1–1.8 (2.5) mm in diam., rst spherical, becoming
saucer–shaped, nally thick and discoid, sessile; with few hyaline
septate anchoring hyphae; mostly with a low and narrow mbriate
margin; hymenium yellow-orange to light orange, margin and outer
surface paler than the hymenium.
Microscopic features (plate 2a-d)
Asci (160) 200–370 (420) × 16–24 (28) µm, cylindrical, 8-spored,
operculate, inamyloid; shortly bifurcate at the base, arising from per-
forated croziers. Paraphyses liform, same length as the asci; contai-
ning, at least in the upper half, many vacuoles 1–3 µm diam. with
orange pigment turning cyan to olivaceous in Lugol’s solution;
straight, pluriseptate, apically inated, terminal cell 30–75 (100) µm
× 4–8 µm, cells 2–4 µm broad near base, rarely clavate. Ascospores
(only ascospores from sporeprint considered): (13) 14–17 µm (orna-
mentation included), hyaline, globose, always with a large lipid drop,
diameter 9–11 (12) µm, uniseriate. Ascospore surface densely cove-
red with numerous isolated warts that are mostly (0.2) 0.4–0.9 µm
broad and 0.4–0.8 µm high, occasionally with one bigger wart which
is 1–2 µm broad and 0.8–1.6 µm high. Warts are mostly hemisphe-
rical though some can be pear-shaped, elongated or conuent, oc-
casionally two warts can be connected by a thin and low ridge.
Many warts have a punctiform cavity, best visible after staining in
Lactophenol Cotton Blue. The bigger warts are predominantly he-
mispherical and not elongated or conuent. The surface of the warts
of some ascospores appears to be nely rugose in SEM. Subhyme-
nium merged with the medullary excipulum, of textura angularis
with hyaline and thin-walled hyphae, walls becoming thicker at the
base, 140 µm thick. Ectal excipulum of textura intricata of elongated
cells, up to 120 µm thick. Margo of textura prismatica, elongated hy-
phae, septate, 15–50 × 5–14 µm, some protruding.

Infection (plate 2e–f & gure 1)
L. verrucispora infects rhizoid and chloronema (the above ground,
green part of protonema) cells of the moss Campylopus pyriformis.
The terminal cells of weak rhizoids or chloronema strands are aec-
ted the most. Infected cells swell to a spherical, one-celled gall
which is many times the width of neighboring cells. Occasionally,
the infected rhizoid or chloronema strand keeps growing, due to
which the gall becomes sub-terminal. Shortly after the infection the
surface of the gall is only partly covered with hyphae (g. 1). Soon,
however, the gall has a complete sheathing of connate hyphae,
which is one to two cell layers thick (plate 2e–f). The hyphae-cove-
red galls can reach up to 180 µm in diameter where the galls mea-
sure up to 150 µm in diameter and the hyphal sheaths are 8-15 µm
thick. The infection structure consists of appressoria, infection pegs
and haustoria. Only one infection structure per gall is developed.
Appressoria are 5–8 µm wide and if at all only slightly wider than
adjacent cells (g. 1). Because of their weak dierentiation, they are
discernable in early stages of gall development only. From the ap-
pressoria an infection peg, 2–3 µm wide, penetrates the rhizoid wall.
The infection pegs are surrounded by a tube of host cell wall mate-
rial. Haustoria are thin walled, strongly ramied and ll the gall up
to half of its volume. No septa were seen within the haustoria and
they do not grow through cross walls of rhizoid or chloronema
strands. The infection does not weaken the host discernably.
Habitat (plate 1, gs. a–c)
Dutch collection: The Dreijen Arboretum is a botanical garden in
Wageningen in the province of Gelderland. A large variety of both
deciduous and coniferous bushes and trees are grown in the Arbo-
retum. The locality is at an exposed patch of about 2 m next to a
low boxwood hedge. The top soil consists of humus and is partly
covered with decaying wood chips. The moss vegetation on the soil
surface is dominated by Campylopus pyriformis.
German collections: The forest Göhrde is a mixed woodland in the
north-eastern part of Lower Saxony, extending over 75 square kilo-
metres. The four localities of L. verrucispora are situated within a ra-
dius of circa two kilometres in the western part near the hamlet
Boitze. The Göhrde in this part is dominated by Pinus sylvestris L. on
sandy soil and, being a working forest, is crossed by many regularly
used tracks. The road tracks have an upper humose layer on which
Campylopus pyriformis ourishes in large extent. Lamprospora cam-
pylopodis W.D. Buckley, also associated with C. pyriformis is ubiqui-
tous in this area and was present in three out of four localities of
L. verrucispora. The locality of the holotype of L. verrucispora was
chosen both because of the very rich collection of apothecia of
L. verrucispora growing on a forest track over a 12 metres length,
and the absence of L. campylopodis. Only one other species of bryo-
philous Pezizales could be found in the surroundings of the four lo-
calities: Lamprospora rugensis Benkert on Pohlia annotina (Hedw.)
Lindb. in a puddle fallen dry.
In general, Campylopus pyriformis grows mostly on acidic sandy,
humous or peaty soils and rotten wood on bare and disturbed
places, such as peat cuttings, roadsides, forest trails and banks
(ATHERTON et al., 2010). It is common in Western Europe, and inciden-
tally reported from other locations in the temperate zone throu-
ghout the whole Northern Hemisphere (OCHYRA et al., 2008).
Specimens examined
1. The Netherlands, Arboretum De Dreijen Wageningen (Gelder-
land), 51°58’04.3’’ N, 5°40’40.5’’ E, 32 m asl, besides a low boxwood
hedge, leg. Henk-Jan van der Kolk, January 23rd, 2014. Bryophyte
host: Campylopus pyriformis, accompanying mosses: Ceratodon pur-
pureus (Hedw.) Brid., Campylopus introexus (Hedw.) Brid., Atrichum
undulatum (Hedw.) P. Beauv., Hypnum cupressiforme Hedw., Bryum
barnesii J.B.Wood ex Schimp.; pers. herb. VDK fungi # 0194, duplicate
pers. herb. MV20140202-01, duplicate pers. herb. JE.
Additional collection at this site: February 11th 2014, pers. herb.
VDK fungi # 0195.
2. Germany, Forest Göhrde near Boitze (Lower Saxony), 53°07’47’’
N, 10°46’17.4’’ E, 75 m asl, in a rut on a forest track crossing a pine-
wood, leg. Marcel Vega, October 3rd 2015. Host: C. pyriformis, accom-
panying moss: Hypnum cupressiforme; soc.: Lamprospora
campylopodis, pers. herb. MV20151003-01.
Additional collections at this site: October 9th 2015, pers. herb.
MV20151009-01 , October 25th 2015, pers. herb. MV20151025-01.
3. Germany, Forest Göhrde near Boitze (Lower Saxony),
53°07’37.2’’ N, 10°45’14.9’’ E, 71 m asl, besides a forest track in a pine-
wood, leg. Marcel Vega, October 9th 2015. Host: C. pyriformis; soc.:
L. campylopodis, pers. herb. MV20151009-02.
Additional collection at this site: October 25th 2015, pers. herb.
MV20151025-02.
4. Germany, Forest Göhrde near Boitze (Lower Saxony), 53°07’49’’
N, 10°45’10.5’’ E, 63 m asl, in a rut on a forest track in a pine-wood,
leg. Marcel Vega October 9th 2015. Host: C. pyriformis; soc.: L. cam-
pylopodis, pers. herb. MV20151009-03.
Additional collections at this site: October 25th 2015, pers. herb.
MV20151025-03, December 5th 2015, pers. herb. MV20151205-03.
5. Germany, Forest Göhrde near Boitze (Lower Saxony), 53°07’36’’
N, 10°45’53.5’’ E, 79 m asl, in a rut on a forest track passing through
a pine-wood, leg. Marcel Vega, October 25th 2015. Host: C. pyriformis,
accompanying moss: H. cupressiforme. Holotype (HBG–holotypus,
inv. no. 1412), isotypes: pers. herb. MV20151025-04 and pers. her-
barium JE.
Additional collection at this site: November 7th 2015, pers. herb.
MV20151107-01, duplicate pers. herb. VDK fungi # 0246.
Discussion
To compare the new L. verrucispora with similar species of bryo-
philous Pezizales, we consider here a) bryophilous Pezizales known
to parasitise species of the moss genus Campylopus on one hand
and b) bryophilous Pezizales with similar ascospores to those of
L. verrucispora.
In a) there are Lamprospora campylopodis, L. areolata Seaver,
L. australis (Mc Lennan & Cookson) Rifai and Octospora meslinii (Le
Gal) Svrček & Kubička and in b) L. ecksteinii Benkert, L. lutziana Boud.,
L. lubicensis Benkert, Octospora svrcekii Benkert and O. wrightii (Berk.
& M.A. Curtis) J. Moravec.
164 Ascomycete.org
Figure 1 – Young gall in optical section. Scale bar: 40 µm.
Drawing: Jan Eckstein

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Plate 1 – Lamprospora verrucispora, macroscopic pictures
a & b: Locality D-Forest Göhrde, holotype; c: Locality NL-Wageningen Arboretum De Dreijen; d: Young apothecia, NL-Wageningen; e: Young
apothecium, D-Göhrde; f–h: Mature apothecia, D-Göhrde. Pictures: a,b, f–h: Marcel Vega; c & d: Henk-Jan van der Kolk; e: Jan Eckstein.

166 Ascomycete.org
Plate 2 – Lamprospora verrucispora, microscopic pictures
a: Ascospores in ascus stained with methylene blue; b: Paraphyses in tap water; c: Free ascospores stained with cotton blue: d: Free
ascospores stained with methylene blue; e: Galls in tap water; f: Gall stained with cotton blue. Scale bars: a–d = 10 µm: e = 100 µm; f =
25 µm. All micrographs taken from the holotype collection. Pictures: a & d: Raúl Tena Lahoz; b: Rubén Martínez Gil; c, e & f: Jan Eckstein.

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a) Bryophilous Pezizales known to parasitise species of the
moss genus Campylopus
BUCKLEY (1923: 44-45) described Lamprospora campylopodis from
Campylopus fragilis (Brid.) Bruch & Schimp. However, all but one of
the later collections of this species have been reported from C. pyri-
formis (pers. observations; BENKERT, 1987: 213-214; RUBIO et al., 2002:
69-70; ECKSTEIN & ECKSTEIN, 2013: 57-58). The one other collection had
the host Campylopus oerstedianus (Müll. Hall.) Mitt. (BENKERT, 2008:
15). Lamprospora campylopodis clearly diers from L. verrucispora in
its ascospores which have an areolate reticulum consisting of mostly
penta– or hexagonal meshes reminiscent of the surface of a football,
thus confusion with L. verrucispora is out of the question. Since L.
campylopodis was found frequently growing together with L. verru-
cispora, macroscopic dierences became apparent. With some ex-
perience both species can be separated in the eld: the apothecia
of L. campylopodis are — at least when mature — rather deeply
orange and have a distinctive membranaceous margin diering
from the yellowish-orange apothecia with a small and indistict mar-
gin of L. verrucispora.
According to BENKERT (1987: 210-211), all exsiccata of Lamprospora
australis (Mc Lennan & Cookson) Rifai had Campylopus introexus as
their host. The ascospores of L. australis dier markedly from those
Plate 3 – Lamprospora verrucispora, SEM-Pictures of ascospores
Pictures on the left from the holotype collection MV20151025-04. Pictures on the right from the Wageningen collection VDK fungi # 0194.
Scale bar: 10 m. Pictures: Jan Eckstein.

168 Ascomycete.org
of L. verrucispora in having an ornamentation consisting of very high
and narrow ridges forming a high reticulum. So far, L. australis is only
known from Australia.
LEGAL (1939: 137-139) attributed to Lamprospora areolata a col-
lection which BENKERT (1987: 207-208) reported to be associated with
the genus Campylopus. However, he doubted the identity of this
specimen with L. areolata, but could not refer it to L. campylopodis
or L. australis either. As the ascospore ornamentation of L. areolata
is similar to that of L. australis in consisting of a reticulum of high
and narrow ridges, this species could not be mistaken for L. verru-
cispora.
Finally, VIVANT (1998: 22-24) reports a nd of Octospora meslinii on
Campylopus subulatus Schimp. ex Milde. This species of Octospora
has distinctively ellipsoid spores and therefore it cannot be mixed
up with L. verrucispora.
b) Bryophilous Pezizales with similar ascospores (plate 4)
We consider species with globose or subglobose ascospores and
an ornamention consisting of isolated warts on average less than
1 µm wide as similar to L. verrucispora. There are ve such species
and each of them is known from only one host, whereas all the hosts
are not related to the genus Campylopus. Information on the species
has been taken from literature and from personal observations.
Both Octospora svrcekii and O. wrightii share one special feature
with L. verrucispora which had not yet been evidently demonstrated
for any Lamprospora species: They induce galls on the rhizoids of
their hosts (DÖBBELER, 1979: 844–846; BENKERT, 1998: 26–27 and 31).
The size of ascospore warts is, although slightly larger, similar to
L. verrucispora, with warts of O. svrcekii being 0.5–1 µm broad and
0.5–1 µm high and those of O. wrightii being 0.5–1 (2) µm broad and
0.5–1 (2) µm high. However, O. svrcekii parasitises Cratoneuron lici-
Plate 4 – Comparative Plate
a: Lamprospora verrucispora; b: L. lubicensis; c: L. lutziana; d: L. ecksteinii; e: Octospora svrcekii; f: O. wrightii. Spores stained with cotton blue.
Scale bars: a–f = 10 µm. Leg.: a: Marcel Vega; b: Jan Eckstein; c: Elisabeth Stöckli; d: Günter Eckstein; e: Lukáš Janošík; f: Torsten Richter.
Pictures: a, b & d: Jan Eckstein; c & e: Lukáš Janošík; f: Hartmut Schubert.

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num (Hedw.) Spruce, a pleurocarpic moss typical for wet or moist
habitats like streams and wet rocks (BENKERT, 1998: 26–27). Octospora
wrightii occurs on Amblystegium serpens (Hedw.) Schimp., also a
pleurocarpic moss occuring on many substrates including stone,
wood or soil. Furthermore, both species of Octospora have predo-
minantely subglobose ascospores and have not as many warts as
L. verrucispora (Plate 4e–f; BOUDIER, 1917: plate 399; SEM-pics in BEN-
KERT, 1998: 27 and 32). In these respects, they clearly dier from
L. verrucispora with its globose ascospores ornamented with many
ne warts.
Lamprospora ecksteinii has an ornamentation of warts which can
be elongated and conuent like those of L. verrucispora, but the
warts are on average slightly larger, measuring 0.5–1 (1.5) µm broad
and 0.5–1 (1.5) µm high. In contrast to L. verrucispora, the ascospores
of L. ecksteinii are slightly subglobose and their surface is less den-
sely covered with warts (Plate 4d and SEM–pics in BENKERT, 2009: 52-
53). Furthermore, L. ecksteinii does not induce galls on the rhizoids
of its host Microbryum curvicollum (Hedw.) R.H.Zander. The moss
M. curvicollum occurs in sun–exposed pioneer sites in grassland,
quarries or arable elds and therefore in habitats quite dierent
from those of Campylopus pyriformis, the host of L. verrucispora.
Lamprospora lutziana, a species with perfectly globose
ascospores, diers from L. verrucispora in having an ascospore orna-
mentation with fewer warts (Plate 4c and SEM-pics in SCHUMACHER,
1993: 321). The warts in L. lutziana are 0.3–1.4 µm broad and 0.6–
1.6 µm high diering from the more or less isodiametric and lower
warts of L. verrucispora. Also, L. lutziana does not induce galls on the
rhizoids of its host Philonotis fontana (Hedw.) Brid. which occurs in
fairly wet habitats such as marshes, ditches, by streams or lakes as
well as on wet rocks.
Lamprospora lubicensis is the most similar species to L. verrucis-
pora regarding ascospore shape, size and ornamentation. However,
L. lubicensis diers from L. verrucispora by having an ascospore or-
namentation of, on average, slightly larger and less numerous warts
and a stronger tendency for elongated and/or conuent warts (Plate
4b and SEM-pics in BENKERT, 1994: 198, 2009: 55). Occasionally, the
warts of L. lubicensis can be hook-shaped and in extreme cases they
can even indicate an irregular reticulum (pers. observation). Regular
warts in L. lubicensis are 0.5–1.5 µm broad and 0.5–1.5 µm high, and
when elongated they are 2–3 (5) µm long. However, L. lubicensis
does not induce galls on the rhizoids of its host Hennediella heimii
(Hedw.) R.H.Zander which is restricted to salt inuenced habitats i.e.
coastal areas, salt works and salted verges of roads (ATHERTON et al.,
2010: 491). Macroscopically, L. lubicensis diers from L. verrucispora
in the more orange-red coloured apothecia with a much more pro-
minent margin compared with the yellow-orange and rather indis-
tinctively marginate apothecia of L. verrucispora.
Gall formation on mosses induced by bryophilous Pezizales
Gall formation is documented for several bryophilous Pezizales. It
is a characteristic feature dening the section Wrightoideae (BENKERT,
1998, 2006) within the genus Octospora encompassing the species
O. americana Benkert, O. erzbergeri Benkert, O. hygrohypnophila Dis-
sing & Sivertsen, O. orthotrichi (Cooke & Ellis) K.B. Khare & V.P. Tewari,
O. svrcekii, O. texensis Benkert and O. wrightii. Spherical, unicellular
galls are induced on rhizoid end-cells of their hosts, whereas the gall
surface is completely covered with a layer of hyphae (DÖBBELER, 1979:
845, g. 8). These galls are very similar in all members of the section
Wrightoideae and also highly resemble the galls observed in L. ver-
rucispora. However there is a dierence in the number of appresso-
ria per gall: In L. verrucispora there is only one appressorium per gall,
whereas DÖBBELER (1979: 844) reported up to six appressoria per gall
in O. wrightii. Admittedly, the appressoria in mature galls are rather
dicult to see and L. verrucispora could possibly have more than
one per gall.
Gall formation is also reported from O. meslinii associated with
Grimmia pulvinata (Hedw.) Sm. (ITZEROTT & DÖBBELER, 1982: 202, g. 1)
and in O. pseudoampezzana (Svrček) Caillet & Moyne associated with
Schistidium crassipilum H.H. Blom (ECKSTEIN & ECKSTEIN, 2009: 225). In
both cases the infectious structures are very similar to those of the
section Wrightoideae. In contrast, O. rubens (Boud.) M.M. Moser
forms unicellular galls on rhizoids of Ceratodon purpureus which in
general do not develop a complete cover of hyphae, most of the
gall’s surface remaining free (ITZEROTT & DÖBBELER, 1982: 205-206, g.
2-3, ut O. libussae Svrček & Kubička). The galls of Octospora humosa
(Fr.) Dennis in species of the Polytrichaceae are somewhat dierent
in being very large, up to 350 µm in diameter, and thick walled. The
surface is almost free with only a few hyphae clustered around the
appressorium at the base of the gall (DÖBBELER & ITZEROTT, 1981: 130-
133, g. 3). The gall inducing species O. meslinii, O. pseudoampez-
zana, O. rubens and O. humosa clearly dier from L. verrucispora
having broadly ellipsoid to ellipsoid spores.
There was a report of possible gall formation for Lamprospora tu-
berculata by EGERTOVÁ et al. (2015: 121-122) but so far this has not
been conrmed in other collections of that species (pers. observa-
tion). Therefore, Lamprospora verrucispora is the rst Lamprospora
species convincingly known as inducing galls on its host. However,
detailed studies of infections are still lacking for many species and
it seems likely that gall formation will also be discovered in other
Lamprospora species in the future.
Key
With several new species described in recent years the keys avai-
lable so far are outdated. Therefore, a new key is provided below to
facilitate identication of all species of bryophilous Pezizales with
globose or nearly globose ascospores and an ascospore ornamen-
tation consisting solely of isolated warts. In addition to the similar
species mentioned above, the key includes Lamprospora esterlech-
nerae Benkert, L. maireana Seaver, L. rehmii Benkert, L. spinulosa Sea-
ver, L. tuberculata Seaver and L. tuberculatella Seaver, which all have
an ascospore ornamentation of rather large warts or tubercles. Spe-
cimens of all species mentioned were studied by at least one of the
authors except for L. esterlechnerae which is currently known only
from the type. Additional information has been gathered from BEN-
KERT (1987, 2002, 2006, 2011), CAILLET & MOYNE (1980, 1991), ECKSTEIN
(2014), ECKSTEIN et al. (2014), EGERTOVÁ et al. (2015), GRAF & MÜHLEBACH
(2014), KULLMAN (1997), SEAVER (1914, 1928); WANG & KIMBROUGH (1992).
Acknowledgements
We would like to thank the following for sharing information
and/or providing us with samples and photos: Raúl Tena Lahoz and
Rubén Martínez Gil (Spain), Lukáš Janošík (Slovakia), Elisabeth
Stöckli and Ueli Graf (Switzerland), Maren Kamke and Hartmut Schu-
bert (Germany), and Gilbert Moyne (France). Our most profound
thanks go to Brian Spooner (UK) for revising the manuscript. We are
very grateful to the sta of the Johann-Friedrich-Blumenbach-Ins-
titut für Zoologie und Anthropologie Göttingen for allowing us to
take SEM-pics by means of their equipment. Thanks go to Uwe Lin-
demann, Hans-Otto Baral, Jürgen Hechler and Emiel Brouwer for
helpful advice. Last but not least our thanks go to Christian Lechat
for starting ascofrance.com and for sticking to it.
References
ATHERTON I., BOSANQUET S., & LAWLEY M. (eds.). 2010. — Mosses and li-
verworts of Britain and Ireland: a eld guide. Plymouth, British Bryo-
logical Society, 856 p.
BENKERT D. 1987. — Beiträge zur Taxonomie der Gattung Lampros-
pora (Pezizales). Zeitschrift für Mykologie, 53 (2): 195-271.
BENKERT D. 1994. — Beiträge zur Kenntnis bryophiler Pezizales-Arten.
1. Lamprospora lubicensis, eine neue Art aus Norddeutschland.
Zeitschrift für Mykologie, 60 (1): 195-198.

BENKERT D. 1998. — Beiträge zur Kenntnis bryophiler Pezizales-Arten.
6. Wrightoideae, eine neue Sektion der Gattung Octospora. Zeit-
schrift für Mykologie, 64 (1): 17-40.
BENKERT D. 2002. — Beiträge zur Kenntnis bryophiler Pezizales. 10.
Variabilität und Verbreitung von Lamprospora maireana Seaver
und Lamprospora tuberculatella Seaver. Feddes Repertorium, 113
(1-2), 80-95.
BENKERT D. 2006. — Octospora erzbergeri (Pezizales, Ascomycota), eine
neue Art aus Serbien-Montenegro. Österreichische Zeitschrift für
Pilzkunde, 15: 1-6.
BENKERT D. 2008 [2007]. — Zur Kenntnis des Vorkommens bryophiler
Pezizales (Ascomycota) in Südost-Europa. Mycologia Montene-
grina, 10: 7-21.
BENKERT D. 2009. — Zwei neue Arten bryophiler Pezizales (Ascomy-
cota) aus der Bundesrepublik Deutschland und Auistung der aus
Deutschland bisher nachgewiesenen Arten mit Kurzdiagnostik.
Zeitschrift für Mykologie, 75 (1): 51-68.
BENKERT D. 2011. — Lamprospora bavarica und L. esterlechnerae (Pe-
zizales), zwei neue Arten aus dem Nationalpark Bayerischer Wald
(Deutschland, Bayern). Zeitschrift für Mykologie, 77 (2): 149-155.
BOUDIER E. 1917. — Dernières étincelles mycologiques. Bulletin de la
Société mycologique de France, 33 (1-2): 7-22.
BUCKLEY W.D. 1923. — New British Discomycetes. Transactions of the
British Mycological Society, 9 (1-2): 43-47.
CAILLET M. & MOYNE G. 1980. — Contribution à l’étude du genre Oc-
tospora Hedw. ex S.F. Gray emend. Le Gal. Espèces à spores orne-
mentées, globuleuses ou subglobuleuses. Bulletin de la Société
mycologique de France, 96 (2): 175-211.
CAILLET M. & MOYNE G. 1991. — Clé de détermination du genre Oc-
tospora et des genres voisins. Bulletin de la Société d’Histoire natu-
relle du Doubs, 84: 9-24.
DÖBBELER P. 1979. — Untersuchungen an moosparasitischen Pezizales
aus der Verwandtschaft von Octospora. Nova Hedwigia, 31 (4):
817-864.
170 Ascomycete.org
Key for bryophilous Pezizales with globose or subglobose (L/B < 1.1) and warty ascospores
1 Ascospores slightly subglobose.............................................................................................................................................................................................. 2
1 Ascospores perfectly globose.................................................................................................................................................................................................. 4
2 Ascospores (11) 12–14 × 11–13 (13.5) µm, Q: 1.08–1.12, not growing in markedly humid habitats, host Amblystegium serpens
......................................................................................................................................................................................................................................... O. wrightii
2 Ascospores bigger........................................................................................................................................................................................................................ 3
3 Ascospores 14–15 (16) × 13–14.5 µm, Q: 1.07–1.09, growing in dry, sun-exposed habitats, host Microbryum curvicollum
....................................................................................................................................................................................................................................... L. ecksteinii
3 Ascospores (13.5) 14–16 × 13–15 (15.5) µm, Q: 1.05–1.07, growing in humid to wet places, host Cratoneuron licinum .. O. svrcekii
4 Ornamentation consisting of warts higher than broad, warts truncate, 0.8–1.2 µm broad and 1–2 µm high, ascospores
15.2–17.5 (20) µm ................................................................................................................................................................................................... L. spinulosa
4 Ornamentation consisting of warts of more or less same height and breadth..................................................................................................... 5
5 Species growing in wet or humid places on Philonotis fontana, ascospores (12.8) 14–18 (19) µm, warts 0.3–1.4 µm broad and 0.5–
1.6 µm high.................................................................................................................................................................................................................. L. lutziana
5 Species not growing in markedly wet or humid places, not on Philonotis.............................................................................................................. 6
6 Ornamentation consisting of coarse warts or tubercles often > 1.5 µm in diam.................................................................................................. 7
6 Ornamentation consisting of smaller warts all or nearly all < 1.5 µm in diam. ....................................................................................................11
7 Ornamentation consisting of large, hemispherical or broadly rounded tubercles with thin walls and many internal cavities, giving
the warts a foamy appearance, 2–6 µm broad, ascospores 19–25 µm, hosts Archidium alternifolium (Hedw.) Mitt. or Fossombronia
....................................................................................................................................................................................................................................... L. maireana
7 Warts solid or only with tiny cavities..................................................................................................................................................................................... 8
8 Ascospores 19–24 µm in diameter.......................................................................................................................................................................... L. rehmii
8 Ascospores 14–19 µm in diameter......................................................................................................................................................................................... 9
9 Growing on rotting wood or similar substrata, host Dicranodontium denudatum (Brid.) E. Britton, ascospores 14–16 µm
............................................................................................................................................................................................................................ L. esterlechnerae
9 Growing on soil in more or less dry habitats, host dierent ...................................................................................................................................... 10
10 Ornamentation consisting of uneven sized warts (0.5) 1–3 (5) µm broad, of rather irregular shape, ascospores (13) 15–19 (20) µm,
host Pleuridium.................................................................................................................................................................................................... L. tuberculata
10 Ornamentation consisting of more or less even sized warts, which are regularly hemispherical, 0,5–2 µm broad, warts occasionally
connected by very low ridges, ascospores 15–18 µm, hosts Weissia or Didymodon............................................................ L. tuberculatella
11 Ornamentation consisting of small warts (0.2) 0.4–0.9 µm broad and 0.4–0.8 µm high, occasionally with one or few bigger warts
1–2 µm broad and 0.8–1.6 µm high, ascospores (13) 14–17 µm, growing on sand or acidic soil with upper humose layer, host Cam-
pylopus pyriformis ............................................................................................................................................................................................. L. verrucispora
11 Ornamentation consisting of warts on average > 0.8 µm broad, host dierent .................................................................................................12
12 Ornamentation consisting of mostly regular hemispherical warts on average > 1 µm broad, occasionally connected by very low
ridges, ascospores 15–18 µm, hosts Weissia or Didymodon ......................................................................................................... L. tuberculatella
12 Ornamentation consisting of warts 0.5–1.5 µm, ascospores 13.9–18 µm, growing on salt inuenced soil, host Hennediella heimii
....................................................................................................................................................................................................................................... L. lubicensis

DÖBBELER P. & I TZEROTT H. 1981. — Zur Biologie von Octospora libussae
und O. humosa, zwei im Moosprotonema wachsende Pezizales.
Nova Hedwigia, 34: 127-136.
ECKSTEIN J. 2014. — Lamprospora hispanica und Lamprospora tuber-
culatella – zwei bryoparasitische Discomyceten (Ascomycota, Pe-
zizales) neu für Deutschland. Zeitschrift für Mykologie, 80 (1):
105-117.
ECKSTEIN J. & ECKSTEIN G. 2009. — Bryoparasitische Pezizales (Ascomy-
cetes) der Gattungen Lamprospora, Octospora und Neottiella im
Alten Botanischen Garten von Göttingen (Deutschland, Nieder-
sachsen). Herzogia, 22: 213-228.
ECKSTEIN J. & ECKSTEIN G. 2013. — Bemerkenswerte Funde bryoparasi-
tischer Pezizales (Ascomycota) aus Deutschland. Boletus, 34 (2):
55-66.
ECKSTEIN J., ECKSTEIN G. & VEGA M. 2014. — Bemerkenswerte Funde
bryoparasitischer Pezizales (Ascomycota) aus Deutschland II. Bo-
letus, 35 (1): 17-25.
EGERTOVÁ Z., ECKSTEIN J. & VEGA M. 2015. — Lamprospora tuberculata,
Octospora ithacaensis, O. orthotrichi and O. anis – four bryopa-
rasitic ascomycetes new to the Czech Republic. Czech Mycology,
67 (2): 119-133.
GRAF U. & MÜHLEBACH K. 2014. — Ein alpiner Moosbecherling. Schwei-
zerische Zeitschrift für Pilzkunde, 92 (3): 10-12.
ITZEROTT H. & DÖBBELER P. 1982. — Octospora meslinii und O. rubens
(Pezizales), zwei weitere bryophile Gallenbildner. Mitteilungen der
Botanischen Staatssammlung München, 18: 201-211.
KULLMAN B. 1997. — A very rare discomycete, Octospora maireana,
found in Finland. Karstenia, 37: 27-31.
LEGAL M. 1939. — Un Humaria nouveau et un Lamprospora norda-
mericain récoltés en France. Revue de mycologie, 4 (3): 133-139.
OCHYRA R., LEWIS SMITH R.I., & BEDNAREK–OCHYRA H. 2008. — The illustra-
ted moss ora of Antarctica. Cambridge University Press, 704 p.
RUBIO E., SUÁREZ A. & MIRANDA M.A. 2002. — Estudios preliminares
sobre los géneros Lamprospora De Notaris y Ramsbottomia W. D.
Buckley (Ascomycetes, Pezizales) en Asturias. Boletín de la Sociedad
Micológica de Madrid, 26: 61-82.
SCHUMACHER T. 1993. — Studies in arctic and alpine Lamprospora spe-
cies. Sydowia, 45 (2): 307-337.
SEAVER F.J. 1914. — A preliminary study of the genus Lamprospora.
Mycologia, 6 (1): 5-24.
SEAVER F.J. 1928. — The North A merican cup–fungi (Operculates). New
York, 284 p.
VIVANT J. 1998. — Octospora meslinii (Le Gal) Svrček & Kubička. Bulle-
tin de la Societé mycologique des Landes, 21: 22-24.
WANG Y.Z. & KIMBROUGH J.W. 1992. — Monographic studies of North
American species of Octospora previously ascribed to Lampros-
pora (Pezizales, Ascomycetes). Special publication of the National
Museum of Natural Science, 4: 1-68.
171
Ascomycete.org
ef
Marcel Vega
Kohlhoefen 17
20355 Hamburg
Germany
tomprodukt@web.de
Jan Eckstein
Heinrich–Heine–Strasse 9
37083 Göttingen
Germany
jan.eckstein@octospora.de
Henk–Jan van der Kolk
Hullenberglaan 9
6721 AL Bennekom
The Netherlands
henk-janvdkolk@hotmail.com