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MYCOTAXON
ISSN (print) 0093-4666 (online) 2154-8889 © 2016. Mycotaxon, Ltd.
January–March 2016—Volume 131, pp. 61–85
http://dx.doi.org/10.5248/131.61
Some stromatic pyrenomycetous fungi from northern Thailand —
2. Annulohypoxylon and Ustulina
L N. V
*
, S L. S
K D. H
,
1
Institute of Biology & Soil Science, Far East Branch of the Russian Academy of Sciences,
Vladivostok 690022, Russia
2
Department of Biological Sciences, University of Arkansas, Fayetteville, Arkansas 72701, USA
3
Institute of Excellence in Fungal Research &
4
School of Science, Mae Fah Lung University,
Chiang Rai 57100, ailand
*
C : vasilyeva@biosoil.ru
A—Fourteen species of Annulohypoxylon and one species of Ustulina collected
in northern ailand are discussed. Only three of the species (Annulohypoxylon
bahnphadengense, A. bogoriense comb. nov., A. urceolatum) have been reported previously
from southeastern Asia. e other twelve species (A. chiangmaiense, A. derelictum,
A. dipterocarpi, A. maesaeense, A. microbovei, A. morisuspectum, A. neglectum,
A. paratruncatum, A. planodiscum, A. pseudonitens, A. sordidum, Ustulina pseudozonata) are
proposed as spp. nov., described, and illustrated.
K —Ascomycota, taxonomy, Xylariaceae
Introduction
In our rst paper in this series (Vasilyeva et al. 2012), we emphasized that
ailand is the portion of the Indo-Malayan center of biodiversity characterized
by a unique assemblage of vascular plants and, consequently, many associated
species of fungi. As numerous xylariaceous fungi are host-specic (Rogers
1979, Whalley 1985, 1996, Stadler 2011) and those within the Indo-Malayan
center of biodiversity would be expected to be likewise, it can be dicult to
trace host-fungus associations in tropical forests, where dead branches are
mixed together on the forest oor and the tree (or trees) from which they have
fallen are not easy to identify. Nonetheless, we anticipate that future research
will reveal denite substrate preferences for southeastern Asian species for
which associations are now unknown.
62 ... Vasilyeva, Stephenson & Hyde
In attempting to follow a biogeographical approach to taxonomy, we made
a preliminary assessment of Annulohypoxylon species that might be found in
ailand. Several species and varieties are known only from southeastern Asia
(Ju & Rogers 1996, Ju et al. 2004, Fournier et al. 2010), including Annulohypoxylon
bahnphadengense (ailand), A. bovei var. microsporum (J.H. Mill.) Y.M. Ju et al.
(China, Japan, Philippines, Taiwan), A. discophorum (Penz. & Sacc.) Y.M.
Ju et al. (Indonesia), A. elevatidiscum (Y.M. Ju et al.) Y.M. Ju et al. (Taiwan),
A. ilanense (Y.M. Ju & J.D. Rogers) Y.M. Ju et al. (Taiwan), A. maeteangense
J. Fourn. & M. Stadler (ailand), A. microcarpum (Penz. & Sacc.) Y.M. Ju
et al. (Indonesia), A. moriforme var. microdiscum (Y.M. Ju & J.D. Rogers)
Y.M. Ju et al. (Taiwan), and A. urceolatum (Philippines, Sri Lanka, and Taiwan).
ese were the rst taxa checked when identifying our specimens collected in
ailand. Although we recorded only two species (A. bahnphadengense and
A. urceolatum) again in 2011, we expect others will be represented by additional
collections in the future.
ere are a few other species reported from both southeastern Asia and the
tropics of the western hemisphere, but their types and many other specimens
are mostly from the eastern hemisphere. For example, Annulohypoxylon nitens
(Ces.) Y.M. Ju et al., described originally from Malaysia, has been found also in
China, Philippines, Sri Lanka, and Taiwan. Its occurrence in ailand has been
reported by Suwannasai et al. (2002, 2005), ienhirun et al. (2003), ienhirun
& Whalley (2004), Phosri et al. (2008), and Okane et al. (2008). Records of
A. nitens from Mexico and Puerto Rico (Ju & Rogers 1996), Argentina (Hladki
& Romero 2009), and Brazil (Pereira et al. 2010b) may represent a dierent
taxonomic entity (cf. Vasilyeva et al. 2012, Vasilyeva & Stephenson 2014). In
contrast to the southeastern Asian concentration of A. nitens, the distribution
of A. stygium (Lév.) Y.M. Ju et al. is centered primarily in the Caribbean Basin.
Specimens reported from Indonesia and Malaysia as Hypoxylon stygium
(e.g., Whalley & Whalley 2007) might be assigned to other species (such as
Hypoxylon bogoriense and H. stigmoideum Ces.) described previously from
these countries, and the unication of the three entities (H. bogoriense,
H. stigmoideum, H. stygium) into a single species (Ju & Rogers 1996) might
need further consideration. Collections from ailand identied as H. stygium
(Whalley et al. 1995, Suwannasai et al. 2002, 2005, ienhirun et al. 2003, Tang
et al. 2007, Phosri et al. 2008) might also represent dierent species.
Materials & methods
e specimens considered herein were collected in two provinces in northern
ailand: Chiang Mai (Mae Taeng District near the villages of Pha Deng and Mae
Sae; and the Doi Suthep temple in the vicinity of the city of Chiang Mai) and Chiang
Annulohypoxylon & Ustulina spp. nov. (ailand) ... 63
Rai (Doi Pui Mountain in the Muang District). Microscopic analyses were carried out
using standard techniques. Material was mounted in 10% KOH for examination of
stromatal pigments and dehiscence of the perispore, and Melzer’s iodine reagent was
used for examination of the ascal apical ring. Color designations follow Rayner (1970).
Photographs of stromata and associated structures were taken using a Nikon D40x
digital camera. Measurements were obtained from about 15 ascospores and 5 glomerate
stromata to represent the range of variation. e material from ailand is deposited in
the herbarium of Mae Fah Luang University, Chiang Rai, ailand (MFLU).
Taxonomy
Annulohypoxylon bahnphadengense J. Fourn. & M. Stadler,
Fungal Diversity 40: 30. 2010 F 1, 15a
S : THAILAND, C M P, Mae Taeng District,
Pha Deng village, Mushroom Research Centre, on wood, 19 June 2011, L. Vasilyeva
(MFLU11-1165).
N: Annhulohypoxylon bahnphadengense, characterized by eused-
applanate, dull black to shiny black stromata, yielding grayish sepia (106)
pigments in 10% KOH, ostiolar discs 0.25–0.3 mm diam., ascospores 6.5–8 ×
3–3.5 µm, with a faint straight germ slit spore-length, perispore dehiscent in
10% KOH, and described only recently by Fournier et al. (2010), was collected
again in same general locality.
Annulohypoxylon bogoriense (Höhn.) Lar.N. Vassiljeva, S.L. Stephenson &
K.D. Hyde, comb. nov. F 2, 15b
MB MB
≡ Hypoxylon bogoriense Höhn., Sitz. K. Akad. Wiss. Math.-Nat. Kl., Abt. 1, 118: 341. 1909.
S eused, rather plane, with mostly inconspicuous (sometimes
conspicuous) perithecial mounds at the margins, surface brown or dark-
reddish-brown to the naked eye but pinkish between the black ostiolar discs
in direct light and under low magnication, shiny black around the individual
perithecia, with citrine (13) to olivaceous bu (89) KOH-extractable pigments.
P spherical, 0.3–0.4 mm diam., ostioles papillate, encircled with a
slightly concave or at ostiolar discs 0.2–0.25 mm diam. A not observed.
A light brown, ellipsoid-inequilateral, 5–7 × 2.4–3 µm, with a
straight germ slit shorter than the spore length; perispore dehiscent in 10 %
KOH.
S : THAILAND, C M P, Mae Taeng District,
Pha Deng village, Mushroom Research Centre, on wood, 24 June 2011, L. Vasilyeva
(MFLU11-1166).
N: Hypoxylon bogoriense was listed as a synonym of Hypoxylon stygium
by Ju & Rogers (1996). Our illustration (F. 2) of the specimen from ailand
64 ... Vasilyeva, Stephenson & Hyde
shows the pinkish stromatal surface between the black ostiolar discs, which is
similar to A. atroroseum (J.D. Rogers) Y.M. Ju et al. Annulohypoxylon stygium
and A. atroroseum share many similar features, but mature stromata are usually
shiny black in A. stygium, whereas those of A. atroroseum are rose-colored with
a black ostiolar region (Ju & Rogers 1996). However, the mature stromata in
our specimen from ailand are not rose-colored; this tint is apparent only
in direct light and under low magnication when photographed, and the
actual color is dark brown or in accordance with Höhnel’s (1909) description
of Hypoxylon bogoriense as being violet-brown and darkening with the age.
Moreover, Rogers (1981) described H. atroroseum as having conspicuously
sunken ostiolar discs, whereas those in our specimen are slightly concave and
almost at, thus slightly elevated above the surface. As Höhnel indicated, the
at-convex tops of the perithecia, always at the margins but rarely in the middle
parts of the stromata, stick out (“Scheitel der randständigen Perithecien stets,
seltener auch der mittelständigen achkonvex vorragend;” Höhnel 1909, p. 341),
and this condition corresponds to our specimen. Although H. atroroseum
has been reported from ailand (Suwannasai et al. 2005, Phosri et al. 2008),
it was described originally from the west of central Africa (Gabon), and all
specimens from ailand might represent H. bogoriense, originally described
from Indonesia.
Annulohypoxylon chiangmaiense Lar.N. Vassiljeva, S.L. Stephenson & K.D. Hyde,
sp. nov. F 3, 15c
MB MB
Diers from Annulohypoxylon bahnphadengense by its larger ascospores and its citrine
to olivaceous bu KOH-extractable pigments
T: ailand, Chiang Mai Province, Mae Taeng District, Pha Deng village, Mushroom
Research Centre, on wood, 22 June 2011, L. Vasilyeva (Holotype, MFLU11-1167).
E: refers to the province where the fungus was collected.
S eused-pulvinate, with perithecial mounds 1/2 to 1/4 exposed,
surface at rst brownish-olivaceous, then dark brown, with black granules
immediately beneath surface and citrine (13) to olivaceous bu (89) KOH-
extractable pigments. P spherical, 0.4–0.5 mm diam., ostioles
papillate, encircled with at ostiolar discs 0.3–0.35 mm diam. A in the spore-
bearing portions 70–80 × 4.5–5 µm, the stipes ≤60 µm, with a tiny apical ring
bluing in Melzer’s iodine reagent. A brown, ellipsoid-inequilateral,
8.5–9.5 × 4-5 µm, with a straight germ slit slightly shorter than the length of the
spore; perispore dehiscent in 10 % KOH.
N: Annulohypoxylon chiangmaiense is supercially quite similar to
A. bahnphadengense, also collected from the same site. However, the pigments
Annulohypoxylon & Ustulina spp. nov. (ailand) ... 65
F. 1–3. Stromata: 1. Annulohypoxylon bahnphadengense. 2. A. bogoriense. 3. A. chiangmaiense.
Scale bars: 1, 2, 3a = 0.4 mm; 3b = 1.7 mm.
66 ... Vasilyeva, Stephenson & Hyde
of A. bahnphadengense are grayish sepia and its light-brown ascospores
(F. 15a) are 6.5–8.4 µm long (7.5 µm in average). It is also noteworthy that
young portions of the stromata in A. chiangmaiense are brownish-olivaceous.
Annulohypoxylon derelictum
Lar.N. Vassiljeva, S.L. Stephenson & K.D. Hyde,
sp. nov. F. 4
MB MB
Diers from other Annulohypoxylon species in lacking KOH-extractable pigments.
T: ailand, Chiang Mai Province, Muang District, T. asai, on dead branch
of Dipterocarpus (?), in a dipterocarp forest, 30 June 2011, L. Vasilyeva (Holotype,
MFLU11-1168).
E: from the Latin derelictus, meaning ‘abandoned’ or ‘neglected’.
S eused-pulvinate, with perithecial mounds 1/4 exposed or
inconspicuous, surface dull black, blackish granules immediately beneath
surface, without apparent KOH-extractable pigments. P spherical,
0.3–0.5 mm diam., ostioles papillate and encircled with a at or slightly concave
disc 0.2–0.3 mm diam. A not observed. A brown to light brown,
ellipsoid-inequilateral, 8–10.5 × 4.5–5.2 µm, with a straight germ slit almost the
length of the spore; perispore dehiscent in 10% KOH.
N: e most prominent feature of A. derelictum is the lack of KOH-
extractable pigments, a condition not previously recorded for Annulohypoxylon.
Annulohypoxylon dipterocarpi Lar.N. Vassiljeva, S.L. Stephenson & K.D. Hyde,
sp. nov. Fs 5, 15g
MB MB
Diers from Annulohypoxylon purpureonitens by its smaller ostiolar discs and larger
ascospores.
T: ailand, Chiang Mai Province, Muang District, T. asai, on dead branch
of Dipterocarpus (?), in a dipterocarp forest, 30 June 2011, L. Vasilyeva (Holotype,
MFLU11-1169).
E: refers to the genus of the probable host plant, with the specimen collected
in an almost pure dipterocarp forest.
S eused-pulvinate, with perithecial mounds 1/2 to 1/4 exposed or
inconspicuous, surface black to the naked eye but brownish under direct light,
blackish granules immediately beneath the surface, KOH-extractable pigments
vinaceous purple (101). P spherical, 0.3–0.5 mm diam., ostioles
papillate and encircled with a small concave disc about 0.1–0.2 mm diam. A
70–80 × 5–6 µm long in the spore-bearing portions, the stipes 30–35 µm long,
with an apical part not bluing in Melzer’s iodine reagent. A brown to
Annulohypoxylon & Ustulina spp. nov. (ailand) ... 67
light brown, ellipsoid-inequilateral, (8–)9–11(–12) × 4.5–5 µm, with a straight
germ slit the length of the spore; perispore dehiscent in 10% KOH.
N: Annulohypoxylon dipterocarpi has vinaceous purple KOH-extractable
pigments and is characterized by eused pulvinate stromata similar to A. purpureo-
nitens (Y.M. Ju & J.D. Rogers) Y.M. Ju et al., known only from Brazil and Mexico
(Ju & Rogers 1996); A. purpureonitens diers in having larger ostiolar discs
(0.2–0.5 mm diam.) and smaller ascospores (6.5–10 µm). e lack of a positive
Melzer’s iodine reaction in the asci is also diagnostic for A. dipterocarpi. When
the perithecia are crashed in a mount, a mass of tiny ellipsoid spores 1.5–3
µm long appears together with asci and ascospores, suggesting a conidial state
within the stromata but probably instead a hyperparasitic anamorphic fungus.
Annulohypoxylon maesaeense Lar.N. Vassiljeva, S.L. Stephenson & K.D. Hyde,
sp. nov. F 6a, 15h
MB MB
Diers from Annulohypoxylon moriforme by its larger ascospores.
T: ailand, Chiang Mai Province, Mae Taeng district, near Mae Sae village, on
the bark of an unidentied tree, 23 June 2011, S. Stephenson & T. Ko Ko (Holotype,
MFLU11-1170).
E: from the Mae Sae village where the specimen was collected.
S glomerate, 3–5 mm diam., oen conuent, with perithecial mounds
1/2 to 1/4 exposed, surface dark brown, blackish granules immediately beneath
surface, with pale olivaceous but mostly slightly hazel (88) tinted KOH-
extractable pigments. P spherical, ≤1 mm diam., ostioles papillate
and encircled with a concave disc ≤0.3–0.35 mm diam. A not observed.
A brown, ellipsoid-inequilateral, (8–)10–12(–12.5) × 4.5–5.5 µm,
with a straight or oblique germ slit extending the length of the spore or slightly
shorter; perispore dehiscent in 10% KOH.
N: Based on the key to taxa of Hypoxylon sect. Annulata (Ju & Rogers 1996),
of the species characterized by having ascospores with a dehiscent perispore,
ostiolar discs 0.2–0.5 mm diam, and dull brownish mature stromata), only two
resemble A. maesaeense—A. moriforme (Henn.) Y.M. Ju et al. and A. truncatum
(Schwein.) Y.M. Ju et al., both also reported from ailand (Whalley et al. 1995,
Suwannasai et al. 2002, ienhirun et al. 2003, Phosri et al. 2008, Okane et al.
2008).
Annulohypoxylon moriforme, said to occur in the tropics and subtropics
of both hemispheres (Ju & Rogers 1996), is distinguished from A. truncatum
and A. maesaeense by its smaller ascospores (6–9 × 2.5–4 µm). One variety,
A. moriforme var. microdiscum from Taiwan, has larger ascospores comparable
68 ... Vasilyeva, Stephenson & Hyde
F. 4–6. Stromata: 4. Annulohypoxylon derelictum. 5. A. dipterocarpi. 6a. A. maesaeense.
6b. A. truncatum on Quercus (from Ouachita Mountains, Arkansas, VLA P-2156).
Scale bars: 4, 5 = 0.4 mm; 6a = 0.8 mm; 6b = 1 mm.
Annulohypoxylon & Ustulina spp. nov. (ailand) ... 69
in size to those of A. maesaeense, but diers by having ostiolar discs smaller
than 0.2 mm diam.
Annulohypoxylon truncatum (F. 6b) diers from A. maesaeense in
having greenish olivaceous (90) to dull green (70) KOH-extractable pigments.
Moreover, A. truncatum has been reported to inhabit Quercus in temperate
regions, but species records actually have been indicated only from Mexico
and the southeastern United States (North Carolina, Georgia, Louisiana,
Massachusetts, Pennsylvania; Ju & Rogers 1996) as well as from Tennessee
(Vasilyeva et al. 2007) and Arkansas and Texas (our unpublished data).
Annulohypoxylon truncatum has also been recorded (as Hypoxylon truncatum)
from Japan (Abe 1990, Abe & Doi 2000) and Korea (Velmurugan et al. 2007),
not unexpected given that an appreciable number of pyrenomycetous fungi
occur in both eastern North America and temperate regions of eastern Asia
(Vasilyeva & Stephenson 2010). However, the eastern Asian material of
A. truncatum needs to be reconsidered, in light of the redetermination of a
specimen identied as “A. truncatum” from the Russian Far East (Vasilyeva
1998) to A. orientale Lar.N. Vassiljeva & S.L. Stephenson (Vasilyeva et al. 2013).
Annulohypoxylon microbovei
Lar.N. Vassiljeva, S.L. Stephenson & K.D. Hyde,
sp. nov. F 7, 15d
MB MB
Diers from other species of the Annulohypoxylon bovei complex by its very small
ostiolar discs.
T: ailand, Chiang Mai Province, vicinity of Chiang Mai city, near Doi Suthep
temple, Huai Kok Ma, on dead branches of unidentied tree, 21 June 2011, L. Vasilyeva
(Holotype, MFLU11-1171).
E: refers to the similarity of this fungus to the Annulohypoxylon bovei
complex, albeit having smaller ostiolar discs.
S glomerate or eused-pulvinate, mostly conuent, with perithecial
mounds 1/4 to 3/4 exposed, surface dark brown, blackish granules immediately
beneath surface, with citrine (13) to hazel (88) tinted KOH-extractable
pigments. P spherical, 0.5–0.7 mm diam., ostioles papillate and
encircled with a slightly concave or at disc 0.2–0.3 mm diam. A not
observed. A brown, ellipsoid-inequilateral, 8–11 × 4.5–5.5 µm, with
a straight germ slit spore length; perispore dehiscent in 10% KOH.
N: e stromata and ostiolar discs in A. microbovei are similar to those
in A. bovei (Speg.) Y.M. Ju et al., especially as illustrated by Miller (1961,
Figs 158, 159). Annulohypoxylon bovei var. microsporum, widely distributed in
southeastern Asia, is also known from China, Japan, Philippines, and Taiwan
70 ... Vasilyeva, Stephenson & Hyde
(Ju & Rogers 1996) and shares a number of features (including ascospore size)
with A. microbovei but diers in its larger (0.3–0.7 mm diam) ostiolar discs.
Annulohypoxylon morisuspectum Lar.N. Vassiljeva, S.L. Stephenson & K.D. Hyde,
sp. nov. F 8, 15e
MB MB
Diers from Annulohypoxylon moriforme by its at ostiolar discs and its brownish
stromatal surface.
T: ailand, Chiang Mai Province, vicinity of Chiang Mai city, near Doi Suthep
temple, Huai Kok Ma, on dead branches of unidentied tree, 21 June 2011, L. Vasilyeva
(Holotype, MFLU11-1172).
E: as compared with A. moriforme, with which it shares a number of similar
features.
S glomerate, 1.5–2.5 mm diam., oen conuent, with perithecial
mounds 1/2 to 1/4 exposed, surface dark brown, blackish granules immediately
beneath surface, with greenish to citrine (13) KOH-extractable pigments.
P spherical, 0.4–0.5 mm diam., ostioles nely papillate and encircled
with a at disc 0.2–0.3 mm diam. A 60–70 × 4–5 µm long in the spore-
bearing portions, the stipes 35–50 µm long, with an apical ring bluing in
Melzer’s iodine reagent, discoid, 1.5–2 µm broad. A brown to light
brown, ellipsoid-inequilateral, 7.5–10 × 3–4.5 µm, with a straight germ slit
slightly shorter than the length of the spore; perispore of at least some spores
dehiscent in 10% KOH, but rarely.
N: Annulohypoxylon morisuspectum resembles A. moriforme and
A. truncatum, which are distinguished by their stromata with concave ostiolar
discs of the truncatum-type; A. moriforme is further distinguished by the
olivaceous shades of its stromatal surface (Ju & Rogers 1996).
Annulohypoxylon neglectum Lar.N. Vassiljeva, S.L. Stephenson & K.D. Hyde,
sp. nov. F 9, 15i
MB MB
Diers from Annulohypoxylon purpureopigmentum by its smaller ascospores.
T: ailand, Chiang Mai Province, vicinity of Chiang Mai city, near Doi Suthep
temple, Huai Kok Ma, on dead branches of an unidentied tree, 21 June 2011,
L. Vasilyeva (Holotype, MFLU11-1173).
E: refers to the small and inconspicuous, easily overlooked stromata.
S small, glomerate, 1.5–2 mm diam., oen conuent, with perithecial
mounds 1/3 to 1/4 exposed, surface dark brown, almost black to the naked
eye, with a purplish tint under the dissecting microscope, blackish granules
immediately beneath surface, with dilute bay (6) to purplish KOH-extractable
Annulohypoxylon & Ustulina spp. nov. (ailand) ... 71
F. 7–9. Stromata: 7. Annulohypoxylon microbovei. 8. A. morisuspectum. 9. A. neglectum.
Scale bars: 7 = 0.5 mm; 8 = 0.6 mm; 9 = 0.7 mm.
72 ... Vasilyeva, Stephenson & Hyde
pigments. P spherical, 0.2–0.3 mm diam., ostioles rather coarsely
papillate and encircled with a concave disc 0.15–0.2 mm diam. A 40–50 ×
3.5–4 µm long in the spore-bearing portions, the stipes 15–20 µm long, with an
apical ring bluing in Melzer’s iodine reagent, discoid, about 0.5–0.7 µm broad.
A light brown, ellipsoid-inequilateral, 4–6.3 × 2.5–3 µm, with a
straight germ slit, perispore dehiscent in 10% KOH
N: Despite its inconspicuous habit, A. neglectum possesses features
uncommonly encountered in Annulohypoxylon, and their combination
in the same taxon is noteworthy. Very small ascospores were reported
for A. atroroseum, A. ilanense and A. stygium, but these species dier from
A. neglectum by their widely eused stromata; in addition, A. ilanense diers
by its brick colored KOH-extractable pigments (Ju & Rogers 1999), and
A. atroroseum and A. stygium by their greenish olivaceous (90) to dull green
(70) pigments (Ju & Rogers 1996).
Purplish KOH-extractable pigments are present in A. purpureonitens and
A. urceolatum; the pigments in A. neglectum are similar but lack a vinaceous
tinge (101 or 116). Annulohypoxylon minutellum (Syd. & P. Syd.) Y.M. Ju et al.
[= Hypoxylon cohaerens var. microsporum J.D. Rogers & Cand.] has vinaceous
to rusty KOH-extractable pigments (Hsieh et al. 2005), but this species does
not have annulate ostioles. Annulohypoxylon purpureopigmentum Jad. Pereira
et al. (Pereira et al. 2010a, Fig. 16) is characterized by purple vinaceous KOH-
extractable pigments, but the ascospores (6.5–8(–10) µm long) are larger than
those of A. neglectum. e appearance of glomerate stromata with large coarse
papillae that almost ll up the concave discs is typical for A. neglectum and
for A. purpureopigmentum, which seems the most similar western hemisphere
species to A. neglectum.
Annulohypoxylon paratruncatum Lar.N. Vassiljeva, S.L. Stephenson & K.D. Hyde,
sp. nov. F 10a, 15f
MB MB
Diers from Annulohypoxylon truncatum by its smaller ostiolar discs and its citrine to
olivaceous-bu KOH-extractable pigments.
T: ailand, Chiang Mai Province, Mae Taeng District, Pha Deng village, Mushroom
Research Centre, on the bark, 19 June 2011, L. Vasilyeva (Holotype, MFLU11-1174).
E: refers to the close similarity of the new fungus to A. truncatum.
S glomerate to eused-pulvinate, with perithecial mounds 1/2 to 1/4
exposed, surface dull black to the naked eye, dark brown under the dissecting
microscope, blackish granules immediately beneath surface, with citrine (13)
to olivaceous-bu (89) KOH-extractable pigments. P spherical,
0.3–0.4 mm diam., ostioles papillate and encircled with a concave disc about 0.2
Annulohypoxylon & Ustulina spp. nov. (ailand) ... 73
F. 10–11. Stromata: 10a. Annulohypoxylon paratruncatum.
10b. A. truncatum (from Ouachita Mountains, Arkansas, VLA P-2156). 11. A. planodiscum.
Scale bars: 10a = 1 mm; 10b = 0.9 mm; 11 = 1.3 mm.
(–0.25) mm diam. A 60–80 × 4.5–5 µm long in the spore-bearing portions,
the stipes 15–20 µm long, with an apical ring bluing in Melzer’s iodine reagent,
discoid, about 0.5 µm broad. A brown to light brown, ellipsoid-
inequilateral, 8–10.5 × 4–4.6 µm, with a straight germ slit shorter than length
of the spore; perispore dehiscent in 10% KOH.
N: Images of stromata show that A. paratruncatum (F. 10a) and
A. truncatum (F. 10b) are supercially very similar; however, A. truncatum,
which appears to have much larger perithecia and ostiolar discs, also diers by
its greenish olivaceous (90) to dull green (70) KOH-extractable pigments.
74 ... Vasilyeva, Stephenson & Hyde
Annulohypoxylon planodiscum Lar.N. Vassiljeva, S.L. Stephenson & K.D. Hyde,
sp. nov. F 11, 15j
MB MB
Diers from Annulohypoxylon elevatidiscum by its glomerate stromata and at ostiolar
discs.
T: ailand, Chiang Mai Province, Muang District, Doi Pui Mountain, on the trunk
of an unidentied tree, 30 June 2011, L. Vasilyeva (Holotype, MFLU11-1175).
E: refers to the mostly at ostiolar discs.
S glomerate (seemingly eused-pulvinate when conuent), with
perithecial mounds 1/2 to 1/4 exposed, surface dull black (sometimes overlaid
with a white substance, blackish granules immediately beneath the surface),
with greenish-olivaceous (90) KOH-extractable pigments. P
spherical, 0.3–0.4 mm diam., ostioles papillate and encircled with a at disc
0.2–0.3 mm diam. A 60–80 × 4.5–5 µm in the spore-bearing portions, the
stipes 15–20 µm long, with an apical ring bluing in Melzer’s iodine reagent,
discoid, about 0.5 µm broad. A brown to light brown, ellipsoid-
inequilateral, 9–11.5 × 4–5 µm, with a straight germ slit spore extending the
length of the convex side of the spore; perispore dehiscent in 10% KOH.
N:
e combination of several key characters in the type of
A. planodiscum—greenish olivaceous KOH-extractable pigments, a 0.2–0.3 mm
diam ostiolar disc, a straight germ slit on the convex side of the ascospores, and
a perispore dehiscent in 10% KOH—are also found in A. elevatidiscum (Ju et al.
2004), which diers by its slightly larger ascospores and regularly eused-
pulvinate stromata. Although some A. planodiscum discs appear to be slightly
convex, the majority are at, with its stromata initially glomerate and only later
conuent, like those of A. truncatum.
Annulohypoxylon pseudonitens Lar.N. Vassiljeva, S.L. Stephenson & K.D. Hyde,
sp. nov. F 12, 15l
MB MB
Diers from Annulohypoxylon nitens by its smaller ascospores and its smaller ostiolar
discs.
T: ailand, Chiang Mai Province, vicinity of Chiang Mai city, near Doi Suthep
temple, Huai Kok Ma, on dead branches of an unidentied tree, 21 June 2011, L.
Vasilyeva (Holotype, MFLU11-1176).
E: refers to the similarity of this species with Annulohypoxylon nitens in a
number of features.
S eused-pulvinate, with perithecial mounds 1/2 to 1/4 exposed,
surface dull to shiny black in some parts, blackish granules immediately
beneath the surface, with citrine (13) KOH-extractable pigments. P
Annulohypoxylon & Ustulina spp. nov. (ailand) ... 75
spherical, 0.3–0.5 mm diam., ostioles papillate and encircled with a concave
disc about 0.2 mm diam. A 60–65 × 4–4.5 µm in the spore-bearing part,
stalks 20–25 µm long, with an apical ring bluing in Melzer’s iodine reagent,
discoid, about 0.5 µm broad. A brown to light brown, ellipsoid-
inequilateral, 5–7.5 × 2.5–3 µm, with a straight or slightly oblique germ slit
shorter than the length of the spore; perispore dehiscent in 10% KOH.
N: Annulohypoxylon pseudonitens resembles A. nitens in having shiny
black mature stromata, somewhat exposed perithecial mounds, smaller
ascospores, and greenish tinged KOH-extractable pigments. However, the
ascospore length in A. pseudonitens does not exceed 7.5 µm, overlapping
with the lower range cited for A. nitens (6.5–10(–11) µm long; Ju & Rogers
1996). As A. nitens has been reported previously from ailand (Suwannasai
et al. 2002, 2005, ienhirun et al. 2003, Okane et al. 2008, Phosri et al. 2008),
we rst assigned our specimen to A. nitens. However, the type specimen of
A. nitens [≡ Rosellinia nitens Ces.], treated as a synonym of Hypoxylon truncatum
(Schwein.) J. Mill. (Miller 1961), is cited as having 8–12 µm long ascospores (cf.
Saccardo 1882, Miller 1961). In addition Hladki & Romero (2009) cite longer
(8–9 µm) ascospores for a specimen of A. nitens from Argentina. e material
of A. nitens from China and Philippines, identied previously as Hypoxylon
bovei var. microsporum, also has 8–10 µm long ascospores (Miller 1961, Ju &
Rogers 1996), exceeding the spore length of A. pseudonitens.
Annulohypoxylon bovei and its variety dier from A. pseudonitens by their
mainly glomerate stromata.
Annulohypoxylon sordidum
Lar.N. Vassiljeva, S.L. Stephenson & K.D. Hyde,
sp. nov. F 13, 15k
MB MB
Diers from Annulohypoxylon moriforme by its euse stromata and pale fawn KOH-
extractable pigments.
T: ailand, Chiang Mai Province, Mae Taeng District, Pha Deng village, Mushroom
Research Centre, on wood, 20 June 2011, L. Vasilyeva (Holotype, MFLU11-1177).
E: derived from the Latin sordidus, meaning ‘dirty-looking’.
S eused-pulvinate, with perithecial mounds 1/2 to 1/4 exposed,
surface dull, dark brown, blackish granules immediately beneath surface, with
dilute fawn (87) KOH-extractable pigments. P spherical, 0.3–0.4
mm diam., ostioles papillate and encircled with a tiny concave disc up to 0.1
mm diam. A 80–85 × 4.5–5 µm in the spore-bearing part, stalks 15–20 µm
long, with an apical ring bluing in Melzer’s reagent, discoid, about 0.5–0.7 µm
broad. A brown to light brown, ellipsoid-inequilateral, 9–11(–12) ×
76 ... Vasilyeva, Stephenson & Hyde
F. 12–14. Stromata: 12. Annulohypoxylon pseudonitens. 13. A. sordidum. 14. A. urceolatum.
Scale bars: 12, 13 = 0.6 mm; 14 = 1.4 mm.
4–5.5 µm, with a straight germ slit slightly shorter than the length of the spore;
perispore infrequently dehiscent in 10% KOH.
N: Annulohypoxylon sordidum has very small ostiolar discs similar to
those of A. moriforme var. microdiscum, but both A. moriforme varieties are
characterized by basically glomerate and blackish stromata with olivaceous
shades, as well as greenish olivaceous (90) to dull green (70) KOH-extractable
pigments.
Annulohypoxylon & Ustulina spp. nov. (ailand) ... 77
e stroma of A. sordidum illustrated in F. 13 is somewhat similar to that
of A. pyriforme (Y.M. Ju & J.D. Rogers) Y.M. Ju et al. (Ju & Rogers 1996,
Fig. 14G) in having rather prominent perithecial mounds and small ostiolar
discs. Annulohypoxylon pyriforme diers in its slightly smaller (8–10 × 3.5–4
µm) pyriform ascospores, a perispore that is always indehiscent in 10% KOH,
and dull green to dark green KOH-extractable pigments. Annulohypoxylon
sordidum may represent a western hemisphere counterpart of A. pyriforme,
known only from Venezuela (Cerro de la Neblina).
Annulohypoxylon urceolatum
(Rehm) Y.M. Ju, J.D. Rogers & H.M. Hsieh,
Mycologia 97: 861. 2005. F. 14
S : ailand, Chiang Mai Province, Muang District, Doi Pui
Mountain, on the branch of an unidentied tree, 30 June 2011, L. Vasilyeva (Holotype,
MFLU11-1178).
N: Annulohypoxylon urceolatum is described in Ju & Rogers (1996, as
Hypoxylon urceolatum). As indicated earlier, this species has been found only
in southeastern Asia, but our record is the rst for ailand. We observed all
the main features of this species such as the vinaceous purple KOH-extractable
pigments, ascospores that are 9–14 × 3.5–4.5 µm, and a perispore that in our
specimen is indehiscent in 10% KOH.
Key to the species of Annulohypoxylon considered in this paper
(New species indicated in bold font)
1. Stromata without apparent KOH-extractable pigments, eused-pulvinate;
ascospores 8–10.5 µm long .................................. A. derelictum
1. Stromata with KOH-extractable pigments ................................... 2
2. KOH-extractable pigments grayish-sepia, stromata widely eused,
ascospores 6.5–8.4 µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . A. bahnphadengense
2. KOH-extractable pigments of other colors ................................... 3
3. KOH-extractable pigments purplish tinged .................................. 4
3. KOH-extractable pigments of other colors ................................... 8
4. Stromata glomerate ...................................................... 5
4. Stromata widely eused ................................................... 6
5. Ascospores 4–6.3 µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. neglectum
5. Ascospores 6.5–8 µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. purpureopigmentum
6. Ascospores 8–12 µm long ................................................. 7
6. Ascospores up to 14(–17) µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. urceolatum
7. Ostiolar discs 0.1–0.2 mm diam ................................ A. dipterocarpi
7. Ostiolar discs larger, 0.2–0.5 mm diam ........................ A. purpureonitens
78 ... Vasilyeva, Stephenson & Hyde
8. KOH-extractable pigments greenish-olivaceous, dull green, citrine,
olivaceous bu, or hazel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
8. KOH-extractable pigments brick colored ............................ A. ilanense
9. Stromata glomerate, oen conuent and then becoming eused-pulvinate ...... 10
9. Stromata at and widely eused .......................................... 18
10. Ostiolar discs of the bovei type ........................................... 11
10. Ostiolar discs of the truncatum type . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
11. Ostiolar discs 0.2–0.3 mm diam .......................................... 12
11. Ostiolar discs 0.3–0.7 mm diam ...................... A. bovei var. microsporum
12. Ascospores narrower (3–4.5 µm), perispore almost indehiscent in 10% KOH
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .A. morisuspectum
12.
Ascospores wider (4.5–5.5 µm), perispore dehiscent in 10% KOH . . . A. microbovei
13. Ascospores 6–7(–9) µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. moriforme
13. Ascospores 8–12 µm long ............................................... 14
14. Ostiolar discs do not exceed 0.2 mm diam ................................. 15
14. Ostiolar discs 0.2–0.4 mm diam .......................................... 16
15. KOH-extractable pigments citrine to olivaceous-bu,
ascospores 8–10 µm long ............................... A. paratruncatum
15. KOH-extractable pigments greenish olivaceous to dull green,
ascospores 10–12 µm long . . . . . . . . . . . . . . . . . . . A. moriforme var. microdiscum
16. KOH-extractable pigments close to hazel tint ................... A. maesaeense
16. KOH-extractable pigments greenish-olivaceous to dull green . . . . . . . . . . . . . . . . 17
17. Ostiolar discs concave ......................................... A. truncatum
17. Ostiolar discs at . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. planodiscum
18. Ostiolar discs of bovei-type ......................................... A. nitens
18. Ostiolar discs of truncatum type ......................................... 19
19. Ostiolar discs up to 0.1 mm diam . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. sordidum
19. Ostiolar discs larger .................................................... 20
20. Ostiolar disc 0.2 mm diam . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. pyriforme
20. Ostiolar disc 0.3–0.4 mm diam . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
21. Ascospores 5–7 µm long ................................................ 22
21. Ascospores 8.5–9.5 µm long ................................ A. chiangmaiense
22. Stromatal surface pinkish between black ostioles . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
22. Stromatal surface uniformly black ........................................ 24
23. KOH-extractable pigments citrine to olivaceous-bu, ostiolar discs slightly concave
or almost at ............................................... A. bogoriense
23. KOH-extractable pigments greenish olivaceous to dull green, ostiolar discs
conspicuously sunken ...................................... A. atroroseum
Annulohypoxylon & Ustulina spp. nov. (ailand) ... 79
F. 15. Ascospores: a. Annulohypoxylon bahnphadengense. b. A. bogoriense. c. A. chiangmaiense.
d. A. microbovei. e. A. morisuspectum. f. A. paratruncatum. g. A. dipterocarpi. h. A. maesaeense.
i. A. neglectum. j. A. planodiscum. k. A. sordidum. l. A. pseudonitens. Scale bars: a = 7.5 µm;
b–d, f, i = 4 µm; e = 3 µm; g, h, j–l = 5 µm.
24. KOH-extractable pigments citrine . . . . . . . . . . . . . . . . . . . . . . . . . . . A. pseudonitens
24. KOH-extractable pigments greenish olivaceous to dull green . . . . . . . . . A. stygium
80 ... Vasilyeva, Stephenson & Hyde
Ustulina pseudozonata Lar.N. Vassiljeva, S.L. Stephenson & K.D. Hyde,
sp. nov. F. 16
MB MB
Diers from Ustulina zonata by its larger ascospores and light-brownish stromata; and
from U. deusta by its wider ascospores, longer germ slit, and light-brownish stromata.
T: ailand, Chiang Mai Province, Mae Taeng district, near Mae Sae village, on the
bark of an unidentied tree, 23 June 2011, L. Vasilyeva (Holotype, MFLU11-1179).
E: refers to some similarity with Ustulina zonata.
S eused-pulvinate with the appearance of a at elongated cake,
2.5–5 × 1.5–2.5 cm, light-brownish, with wavy and greyish margins, surface
undulate, irregularly cracked, and very nely papillate, ostioles almost
umbilicate. P tubular, 1–1.2 mm long, 0.4–0.6 mm wide. A in the
spore-bearing portions 180–200 × 12–14 µm, stipes up to 200 µm long, with
an apical ring bluing in Melzer’s iodine reagent, urn-shaped, 7–7.5 × 3–4 µm.
A brown, unicellular, fusoid-inequilateral, smooth, (27.5–)30–35
(–37.5) × 8–10(–12) µm, with a straight germ slit extending almost the length
of the spore.
N: e name Ustulina is currently treated as a synonym of Kretzschmaria
(Martin 1970, Læssøe 1994, Rogers & Ju 1998), but within Kretzschmaria sensu
lato two taxonomic complexes can be clearly distinguished, the kretzschmarioid
complex and the ustulinoid complex (Rogers & Ju 1998). Both complexes were
accepted within a single genus based primarily on similar cultural features
(Rogers & Ju 1998), but many other groups of xylariaceous genera (e.g.,
Annulohypoxylon, Daldinia, and Hypoxylon in one instance and Entoleuca,
Nemania, and Rosellinia in another) known to share cultural features are
nevertheless accepted as independent taxonomic entities (Stadler 2011).
We believe that Ustulina deserves recognition as a separate genus; among
the ustulinoid taxonomic complex, the most closely related species to
U. pseudozonata are U. deusta (Hom.) Lind and U. zonata (Lév.) Sacc. Rogers
& Ju (1998) distinguish these two species in their key provided based on
U. deusta having ascospores 27–35(–38) × 7–9 µm, a germ slit much shorter
than the spore length, and a northern temperate distribution vs. U. zonata
having ascospores 21–34 × 8.5–12 µm, a germ slit that is either the length of the
spore or nearly so, and a subtropical and tropical distribution. e specimen
from northern ailand has the range in ascospores length characteristic of
U. deusta but a germ slit that ts U. zonata, and this combination is enough to
characterize it as a separate species.
Ustulina zonata, described from Java, has been mentioned repeatedly as a
pathogen that causes root decay of trees in southeastern Asia (Brooks 1915,
Annulohypoxylon & Ustulina spp. nov. (ailand) ... 81
F. 16. Ustulina pseudozonata. a. Stromata (photo by Randy Darrah). b. Ascus top with an apical
apparatus and an ascospore. c. Ascospores. Scale bars: a = 7 mm; b = 7 µm; c = 8 µm.
Petch 1928, Nandris et al. 1987, Muraleedharan & Baby 2007, Barthakur 2011).
e only color illustration found on the Internet (http://www.ava.gov.sg) shows
U. zonata as having carbonaceous fruit bodies hidden among grass and roots.
e common name of the disease (charcoal stump root) seems to emphasize the
black color of the stromata, although Petch (1928, p. 241) wrote that stromata
usually referred to as representing U. zonata are “fairly frequently reddish
on the surface, wholly or in part, but the majority of the specimens are, until
82 ... Vasilyeva, Stephenson & Hyde
weathered, grey or purple-grey; when weathered, they are, of course, black.”
He was unable to locate Léveillé’s type of U. zonata but found Léveillé’s specimen
of “U. deusta”, which did not have zones on the fragment of a stroma, but had a
reddish tinge. According to Petch, Léveillé distinguished Ustulina zonata from
U. vulgaris (= U. deusta) by its reddish color and zoned surface. None of these
features is mentioned by Rogers & Ju (1998), who indicate only that U. zonata
has stromata, perithecia, ostioles, and asci similar to those found in U. deusta.
In our specimen of U. pseudozonata from ailand, the fully mature stromata
are light brownish or sand-colored. is is the color reported for the maturing
stromata in Kretzschmaria milleri J.D. Rogers & Y.M. Ju from the Caribbean
Basin (French Guiana, Guyana, Mexico, Venezuela).
“Ustulina zonata” from southeastern Asia might represent a species
complex waiting for separation into several separate taxa. Interestingly, almost
all specimens listed under the description of U. zonata by Rogers & Ju (1998)
are from the Caribbean Basin (Cuba, French Guiana, Guadeloupe, Mexico,
Puerto Rico). Only one specimen from Taiwan was cited (Rogers & Ju 1998) for
U. zonata, which diers from U. pseudozonata by its ascospores with pinched
ends. Among the synonyms Rogers & Ju (1998) give for Kretzschmaria zonata is
Hypoxylon deustum f. madagascariensis (Henn.) Hendr., which Dennis (1963)
reported as having ascospores 20–28 × 7–10 µm and which Rogers & Ju noted
very likely represented Kretzschmaria zonata. e specimens from ailand
and Madagascar are not conspecic, primarily due to the spore size dierences
that ostensibly demonstrate the heterogeneity of the Ustulina zonata ‘complex.’
Given the economic importance of U. zonata as a pathogen of many trees and
shrubs, further taxonomic investigations are warranted.
Discussion
In this paper, we have increased the number of examples of closely
related species known from the Caribbean and Indo-Malayan centers of
biodiversity (cf. Vasilyeva et al. 2012). As noted in earlier treatments of species
from temperate latitudes of northeastern Asia and eastern North America
(Vasilyeva & Stephenson 2010, 2011), the morphological dierences between
species in the western and eastern hemispheres are rather insignicant and
sometimes pertinent to only a single character (e.g., a dierence in ascospore
size, of anamorphs, or in stromatal pigments). is enigma currently has no
biogeographic explanation but does demand an accumulation of relevant
examples to solve.
Specimens of six dierent Annulohypoxylon taxa (as Hypoxylon bovei
var. microsporum, H. moriforme, H. nitens, H. purpureonitens, H. stygium,
H. stygium var. annulatum) reported from ailand by Suwannasai et al. (2002)
Annulohypoxylon & Ustulina spp. nov. (ailand) ... 83
were said to dier from taxa described previously by Ju & Rogers (1996) due
to a high level of morphological variation in the species involved. However,
the dierences might as easily be explained by the completely dierent species
composition of the fungal assemblage found in ailand. In fact, morphological
variation within Hypoxylon and Annulohypoxylon is not very high, as both the
same characters (stromatal pigments and shapes, dehiscent or indehiscent
perispore, etc.) and the same ranges of variation (e.g., size of ascospores) repeat
themselves innitely (cf. Vasilyeva & Stephenson 2010) while combining
dierently with one another other for a particular taxon. Diversity lies not
in morphological variation per se, but rather in the combinations of features
characterizing dierent species.
Acknowledgments
e research reported herein was funded in part by a grant (OISE-1042602) from the
National Science Foundation to the University of Arkansas. e support provided by the
Global Research Network for Fungal Biology and King Saud University is also gratefully
acknowledged. anks are extended to Randy Darrah for providing the image used
for Figure 16a. We are also thankful to Liliane Petrini (Breganzona, Switzerland) and
Katherine Winsett (North Carolina State University, Raleigh, USA) for the thorough
reading of our manuscript and their valuable comments.
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