The Lichenologist 42(4): 397–403 (2010)
doi:10.1017/S0024282910000150
© British Lichen Society, 2010
A new species of Stigmidium sensu stricto on Thelenella
muscorum
Sergio PÉREZ-ORTEGA, M. Gökhan HALICI, Kerry KNUDSEN
and Mehmet CANDAN
Abstract: Stigmidium humidum is described from Spain and Turkey on Thelenella muscorum (Fr.) Vain.
(syn Chromatochlamys muscorum (Fr.) H. Mayrhofer & Poelt). The species is the first Stigmidium s. str.
species on a host lichen of the family Thelenellaceae and is compared with other species with ‘Type a’
pseudoparaphyses.
Key words: Ascomycota, lichenicolous fungi, periphysoids, ‘Type a’ pseudoparaphyses
Introduction
Stigmidium Trevis. is a genus of ascomycete
fungi, comprising mostly lichenicolous taxa,
currently included in the Mycosphaerellaceae
(Lumbsch & Huhndorf 2007). The generic
type S. schaereri (A. Massal.) Trevis., is
lichenicolous on the sterile thalli of a species
of Solorina. Stigmidium s. str. and was revised
by Roux & Triebel (1994). The genus is
distinguished by perithecioid ascomata with
punctiform ostioles with a hamathecium of
external periphyses (sensu Roux & Triebel
1994), with periphysoids, lacking interascal
filaments, and by hyaline 1-septate ascospores (rarely turning brown in a few species)
(Roux & Triebel 1994; Calatayud & Triebel
2003). The periphysoids originate from the
wall of the upper cavity of the pseudothecia
are never attached to the bottom of the cavity
and are referred to as pseudoparaphyses of
which two types are recognized (Roux &
S. Pérez-Ortega: Instituto de Recursos Naturales,
Centro de Ciencias Medioambientales CSIC, E-28006
Madrid, Spain. Email: sperezortega@ccma.csic.es
M. G. Halici: Erciyes Üniversitesi, Fen Edebiyat Fakültesi, Biyoloji Bölümü, 38039 Kayseri, Turkey.
K. Knudsen: The Herbarium, Department of Botany
and Plant Sciences, University of California, Riverside,
California 92521, USA.
M. Candan: Anadolu University, Faculty of Science,
Department of Biology, Eskişehir, Turkey.
Triebel 1994). Stigmidium s. str. has ‘Type a’
pseudoparaphyses which are narrow and
two-celled. It currently comprises 22 species
including the species to be described in this
paper (Roux & Triebel 1994; Roux &
Navarro-Rosinés 1994; Roux et al. 1995;
Calatayud & Triebel 2001, 2003; Triebel &
Cáceres 2004; Zhurbenko & Triebel 2008).
A second group of species currently included
in Stigmidium differ from Stigmidium s. str. in
having ‘Type b’ pseudoparaphyses sensu
Roux & Triebel (1994) which are long and
relatively large, made up of 3–5 more or less
equal cells, and comprise the S. placynthii
group which currently includes five species
(Roux & Triebel 1994 & 2005; Roux &
Navarro-Rosinés 1994; Kocourková &
Knudsen 2009a, b).
The genus Stigmidium includes at least 90
binominals (Etayo 2008; Mycobank 2009;
Zhurbenko 2009; Kocourková & Knudsen
2009a, b). The majority of the species, 65 or
more depending on validity of synonymies,
make up Stigimidium s. lat. and are a heterogeneous group. Species have been transferred or described in Stigmidium using
genera concepts differing from those in Roux
& Triebel (1994). Some species have interascal filaments (see Triebel 1989, Calatayud &
Triebel 2003), others have a hamathecium
not described in detail using the same
concepts as Roux’s & Triebel’s taxonomy
398
THE LICHENOLOGIST
(see Halici & Hawksworth 2007), or a
hamathecium has not been observed (see
Zhurbenko 2009), or with observations
uncertain because of limited type material
(see Etayo 2008). Kocourková & Knudsen
(2009b) summarize the situation: “Due to a
lack of standardized morphological nomenclature coupled with differences in taxonomic descriptions and generic concepts of
Stigmidium by different authors, many Stigmidium species have overlapping ascospore
and ascomata size, forcing keys to be structured according to host (Halici 2008; Ihlen &
Wedin 2008) rather than by individual morphological characters”. We expect that some
species in Stigmidium s. lat. may actually belong in Stigmidium s. str. or in the closely
related S. placynthii group with ‘Type b’
pseudoparaphyses. But other species will
need to be segregated into other genera.
A further problem in Stigmidium, as in
other lichenicolous genera, is that many
species concepts were influenced by a hostbased taxonomy. This problem was recognized already before the work of Roux &
Triebel (1994). Hawksworth (1986) wrote
“Species concepts in the genus Stigmidium
Trevisan require a thorough re-appraisal as
the taxonomy is currently largely hostbased”. Stigmidium species appear to be narrowly host specific to particular species,
genera, or families. On some host genera one
species of Stigmidium s. lat. may comprise
two or more species differing, for instance, in
having ‘Type a’ or ‘Type b’ pseudoparaphyses, but with statistically narrow ascospore
differences.
In this paper we describe a new species
of Stigmidium on a new host species and
genus, Thelenella muscorum (Fr.) Vain. (syn.
Chromatochlamys
muscorum
(Fr.)
H.
Mayrhofer & Poelt), in a new host family
Thelenellaceae O. E. Erikss. ex H. Mayrhofer.
The species is a member of Stigmidium s. str.
and is compared with other species with
‘Type a’ pseudoparaphyses.
Material and Methods
Specimens were examined using a Nikon SMZ 600
stereomicroscope. Hand-cut sections of ascomata were
Vol. 42
studied in distilled water, lactophenol cotton blue and
KOH. The amyloid reactions were tested in iodinepotassium iodide solution (1%) (=Lugol’s iodine solution), directly (I) and after a KOH pre-treatment (K/I).
A 1 % solution of Brilliant Cresyl blue (BCr) was used
for testing metachromatic reactions of the endoascus,
epispore and vegetative hyphae. Samples were observed
under a Nikon Eclipse 80i microscope fitted with
‘Nomarski’ differential interference constrast and a
Nikon DS-Fi1 digital image system. All measurements
were made in water, and taken by means of Nikon Image
Analyze System®. Microscopic pictures were taken by
mean of this system; habitus picture was taken by mean
of a Nikon Image System® fitted to a Leica dissecting
microscope. For ascospore and ascus size measurements, the highest and lowest values are given within
parentheses, with the average together with ± standard
deviation in italics; values are approximated to the
nearest 0·5 µm.
The Species
Stigmidium humidum Pérez-Ortega,
Halici & K. Knudsen sp. nov.
MycoBank No.: MB 513453
Fungus lichenicola in thallo Thelenella muscorum parasiticus. Ascomata perithecioidea, subglobosa, atra, 75–
140 µm diametro, semiimmersa. Peridium olivaceum,
textura angulari, inaequaliter pigmentatum, K+
brunneo-caeruleus. Gelatina hymenialis I−, K/I−. Filamenta interascalia observata: pseudoparaphyses ‘typo
a’. Asci saccati vel clavati (24–)41 ± 6(–54) × (9–)
14·5 ± 2(–17) µm magni, 8-spori, endoascus pariete
BCr+ cyanea, I−, K/I−. Ascosporae incoloratae, anguste
pediformes, ad septum non vel laeviter constrictae,
halonatae, (12–)14·5 ± 1·5(–18) × (3.5–)5 ± 0·5(–6·5)
[cellula supera] × (3–)4 ± 0·5(–5) [cellula infera] µm
magnae, longitudinis/latitudinis [cellulae superae]
ratio: (2·5–) 3 ± 0·5(–4), epiplasma BCr+ cyanea.
Conidiomata non observata.
Typus: Turkey, Afyon, Sandıklı, Akdağ Tabiat Parkı,
Başalan Yaylası, 38° 20$ 37$ N, 30° 01# 24$ E, on
Thelenella muscorum, on mosses, alt. 1477 m, 03 June
2008, M. Candan (ANES 12281—holotypus).
(Fig. 1)
Vegetative hyphae hyaline to pale brown,
composed of elongate to rectangular cells,
c. 5 µm diam., immersed in the host tissues,
BCr−.
Ascomata perithecioid, black, ± shiny, in
dense groups, sessile or more often 1/3 to 2/3
immersed in the host thallus, 75–140 µm
(n = 11). In transverse section, globose to
subglobose, frequently with a depression
2010
Stigmidium humidum—Pérez-Ortega et al.
399
F. 1. Stigmidium humidum (MAF 16095). A, habitus; B, ascomata in cross section; C, detail of hamathecial
filaments: ‘Type a’ pseudoparaphyses; D, young ascus; E, mature asci with 8 ascospores; F. detail of vegetative
hyphae (arrow) in host thallus; G, ascomatal wall, detail of cells in the lower part of the ascomata; H, young halonate
ascospores, arrows pointing to halo; I, mature ascospore. B & F in lactophenol blue; D & E in Lugol’s; C, G, H &
I in water. Scales: A = 250 µm; B = 50 µm; C– G = 10 µm; H & I = 5 µm.
around the ostiole. Outer part of the pseudothecium with an irregular rugose appearance. Wall pseudoparenchymatous, of
textura angularis, brown, olive greenish in K,
c. 10–20 µm wide, usually thinner at the base
than near the ostiole, with 2–6 layers of cells;
inner part light brown to hyaline, outer part
dark brown. Cells of the ascomatal wall ±
rectangular and compressed in transverse
sections of the ascomata, and ± roundedpolygonal in superficial view, mostly 4–8 µm
diam., I−; cells in the outer part of the
ascomata forming an uneven surface. External periphyses poorly developed. Hymenial
gel I−, K/I−. Ostiolar filaments present,
4–6 µm long, hamathecium composed of
400
THE LICHENOLOGIST
pseudoparaphyses of ‘type a’ (Roux &
Triebel 1994) 5–10 µm long. Asci bitunicate,
sessile to distinctly stalked at the base, clavate
to saccate, thickened at the apex, 8-spored,
ascospores often arranged distichiously,
(24–)41 ± 6(–54) × (9–)14·5 ± 2(–17) µm (n
= 28), endoascus BCr+ violet. Ascospores
1-septate, constricted at the septum, obovate, lower cell narrower and longer than the
upper one, hyaline, halonate, surface
smooth, not pseudotetrablastic (with a large
oil droplet in each cell), inner ascospore wall
BCr+ violet, (12–)14·5 ± 1·5(–18) × (3·5–)
5 ± 0·5(–6·5) (upper cell) × (3–)4 ± 0·5(–5)
(lower cell) µm [n = 61; length/breadth ratio
upper cell (2·5–)3 ± 0·5(–4)].
Etymology. The name refers to the occurrence of the species in moist microhabitats on
Thelenella muscorum among bryophytes and it
may eventually be discovered on related host
genera that also occur in these microhabitats.
Ecology and distribution. The species is
known from three localities in central Spain
and one in western Turkey. The species
seems to be relatively common on T. muscorum (syn. Chromatochlamys muscorum, see
Fryday & Coppins 2004 for a recent discussion of the taxonomy of the host) thalli in
higher elevation areas of Sistema Central and
Sistema Ibérico. The species does not produce clear necrosis or decolouration in host
thalli, so we assume a parasymbiotic state for
the species. However, thalli infected with S.
humidum show a smaller number of host
perithecia than those thalli not infected,
which might indicate a negative effect of this
taxon on its host. More careful observations
are needed to clarify this question.
Additional specimens examined. Spain: Castilla y León:
Segovia, Honrubia de la Cuesta, siliceous outcrops close
to the village, 41° 30# 45$ N, 3° 42# 8$ W, on Thelenella
muscorum on mosses, alt. 968 m, 06 August 2007, S.
Pérez-Ortega (MAF 16094); Burgos, Barbadillo de Pez,
siliceous outcrops close to the road, 5 km N Bardadillo,
42° 8# 13·2$ N, 3° 11# 33$ W, alt. 1122 m, 22 May
2006, S. Pérez-Ortega (MAF 16095).
Discussion
Stigmidium humidum is the first species in the
genus to be recognized on the lichen host
Vol. 42
genus Thelenella and belongs to Stigmidium s.
str., having ‘type A’ periphysoids or pseudoparaphyses (sensu Roux & Triebel 1994).
Within the genus 22 species are known with
‘type a’ periphysoids including S. humidum,
and several characters of these species are
compared in Table 1.
Stigmidium humidum is most similar to S.
bellemerei Cl. Roux & Nav.-Ros., S. degelii R.
Sant., S. leucophlebiae Cl. Roux & Triebel, S.
mycobilimbiae Cl. Roux, Triebel & Etayo, S.
schaereri (A. Massal.) Trevis and S. stereocaulorum Zhurb. & Triebel, but, apart from a
different unrelated lichen host, also differs in
some other characters. Stigmidium bellemerei,
a species recognized on Lecania nylanderiana
has relatively longer ascospores [(13·5–)15–
18(–19·5) × (3·5–)4–5·5 µm] which are more
ellipsoid and pseudotetrablastic (having
2 large oil droplets per ascospore cell) and
relatively smaller ascomata [70–100 × 65–
100 µm] (Roux et al. 1998). Stigmidium
degelii, a species recognized on Degelia
plumbea, has ascospores rather similar in size
to those of S. humidum [(12–)13–17
(–19·5) × (3–)4–5(–6)], but they are pseudotetrablastic, and the ascomata are relatively smaller [55–106 × 50–90] (Roux &
Triebel 1994). Stigmidium leucophlebiae, a
species recognized on Peltigera leucophlebia,
also has rather similar sized but pseudotetrablastic
ascospores
[(12–)13·5–15
(–15·5) × 3·5–5 (–5·5) µm], but this species
has immersed ascomata in the dying parts of
the host thallus which are relatively smaller
[55–70 × 50–65 µm], while S. humidum has
semi-immersed to superficial ascomata
(Roux & Triebel 1994). Stigmidium mycobilimbiae, a species recognized on muscicole
Mycobilimbia spp., has relatively shorter
and narrower ascospores than S. humidum
and they are also pseudotetrablastic and
non-halonate [(12–)13·5–15(–15·5) × 3·5–5
(–5·5) µm]. This species also has smaller
ascomata [60–90 × 50–75 µm] (Roux &
Triebel 1994). Stigmidium schaereri, a species
recognized on Solorina sp., has rather similar
sized ascospores to S. humidum [(10–)12–
15(–18) × (3–)4–5 µm] which are pseudotetrablastic and non-halonate, but the endoascus and inner perispore of S. schaereri give a
2010
T 1. Comparison of Stigmidium species showing ‘Type a’ pseudoparaphyses
Species
Ascus size
(µm)
Ascospores
(µm)
S. humidum
75–140
S. acetabuli
50–85
(24–)41 ±6(–54) ×
(9–)14·5 ±2(–17)
20–30(–35) × 9–11
S. bellemerei
70–100 × 65–100
41–49 × 12–16
S. cartilagineae
S. cerinae
S. collematis
60–100(–140)
50–70 × 45–70
30–45
28–32 × 11–13
27–34 × 10–11.5
21–23 × 8–10
S. congestum
40–85 × 30–77
24–37 × 10–18
S. croceae
47–62 × 43–60
22–34 × 9–11
S. degelii
55–106 × 50–90
30–57 × 11–17
S. epixanthum
40–60 × 12–15
S. fuscatae
S. joergensenii
S. lecidellae
S. leucophlebiae
100–140 ×
120–180
50–100
30–45
60–90 × 42–75
55–70 × 50–65
(12–)14·5 ±1·5(–18) ×
(3·5–)5 ±0·5(–6·5)
(7·5–)8–10·5(–11) ×
3–4(–4·5)
(13·5–)15–18(–19·5) ×
(3·5–)4–5·5
8–12(–14) × 3–4·5(–5)
(9–)10–12·5(–13) × 3–4
(7–)7·5–9·5(–10·5) ×
(2–)2·5–3
(11–)12–14(–16) ×
(2·5–)3–4
(8·5–)9–13(–14·5) ×
(2·5–)3–3·5(–4·5)
(12–)13–17(–19·5) ×
(3–)4–5(–6)
11–17 × 5–7
30–40 × 10–15
19–27 × 11–13
22–36 × 8–13
30–39 × 13–17
S. mitchelii
55–65 × 50–60
28–35 × 10–14
S. mycobilimbiae
60–90 × 55–75
29–38 × 10–15
S. peltidae
50–70 × 45–65
27–29 × 9–11.5
90–100
60–125 × 57–115
55–75
24–35 × 11–16
29–43 × 12–17
22–35 × 9–13
50–80
30–34 × 11–13
S. pseudopeltidae
S. schaereri
S. solorinarium
S. tabacinae
(8–)10–12 × 4–5(–5·5)
(8–)9–12 × 3·5–4·5
(11–)12–15(–16) × 3–4
(12–)13·5–15(–15·5) ×
3·5–5(–5·5)
(10·5–)12–14(–14·5) ×
3–4(–5)
(10·5–)11·5–16(–16·5) ×
(2·5–)3– 4(–4·5)
(8·5–)9–11·5(–12) ×
(2·5–)3–3·5(–4·5)
(10–)10·5–14 × 3·5–5·5(–6)
(10–)12–15(–18) × (3–)4–5
(10–)10·5–13·5(–14) ×
3–4(–4·5)
10–14 × 3–4
Host
lichen
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Stigmidium humidum—Pérez-Ortega et al.
Ascomatum
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401
402
THE LICHENOLOGIST
negative reaction in BCr, whereas S. humidum shows violet reactions in the endoascus
and inner perispore (Roux & Triebel 1994).
Stigmidium stereocaulorum, a species recognized on Stereocaulon spp., also has rather
similar sized ascospores to S. humidum
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known from Turkey and Spain (Halici 2008;
Halici et al. 2010). It differs from Stigmidium
humidum in having dark brown setae arising
from the upper part of ascomata and 3-5septate brown ascospores.
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Mayrhofer & Poelt in the family Protothelenellaceae Vĕzda, H. Mayrhofer & Poelt, which is
related to the Thelenellaceae. The type species
of P. haesitans as well as the sample collected
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to Stigmidium s. str. as they show K/I+ blue
hymenial gel, long periphysoids with 3–5
cells, and ascospores larger than previously
published and will be revised separately by
Kocourková and Knudsen.
We thank Harrie Sipman for checking our Latin. SPO
is very obliged to the project ANT-COMPLEX
(CGL2006-12179-CO2-01, MEC, Spain) for support.
The studies of MC were financially supported by
TÜBÌTAK (107T605 coded project).
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