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521
Nova Hedwigia 85 3—4 521—539 Stuttgart, November 2007
DOI: 10.1127/0029-5035/2007/0085-0521 0029-5035/07/0085-0521 $ 4.75
© 2007 J. Cramer in der Gebrüder Borntraeger
Verlagsbuchhandlung, D-14129 Berlin · D-70176 Stuttgart
Type studies of three tomentelloid species
(Basidiomycota, Thelephorales): Tomentella radiosa,
Tomentella cinereoumbrina and Tomentella punicea
by
Nourou Soulemane Yorou
1,2*
and Reinhard Agerer
1
1
Department Biology I, Organismic Biology: Mycology, University of München,
Menzinger Str. 67, D-80638, München, Germany
2
Laboratoire d´Ecologie Appliquée, Faculté des Sciences Agronomiques,
Université d´Abomey-Calavi, 01 BP 526, Cotonou, Benin
With 14 figures and 1 table
Yorou, N.S. & R. Agerer (2007): Type studies of three tomentelloid species (Basidiomycota,
Thelephorales): Tomentella radiosa, Tomentella cinereoumbrina and Tomentella punicea. Nova
Hedwigia 85: 521-539.
Abstract: This paper presents additional anatomical descriptions of T. radiosa, T. cenereoumbrina
and T. punicea based on their holotype. For each species, original iconographies and anatomical
features are presented and discussed in details. Similarities and differences between closest species
are highlighted. ITS rDNA sequences´ similarities and phylogenetic relationship between T. radiosa
and Thelephora terrestris are reported. This paper confirms the presence of skeletals and binding-
like hyphae on the rhizomorphs of T. punicea.
Key words: Anatomy, rhizomorphal structures, Tomentella radiosa, T. cinereoumbrina, T. punicea.
Introduction
Many species of resupinate Thelephorales have been considered as conspecific,
resulting in the adoption of a wide species concept, at least for some species (e.g.,
T. radiosa (P.Karst.) Rick in Kõljalg 1996), whilst on the over hand same species
has been repeatedly described as new, resulting in widespread synonymy (comp.
Larsen 1965, 1966, 1968, 1970, 1974, Kõljalg 1996, Wakefield 1960). To avoid
such confusion, it is advisable to retrieve detailed informative anatomical data from
type specimens. Such comparative analyses have already helped discriminate
1
* Corresponding author, Email: n.s.yorou@lrz.uni-muenchen.de
522
tomentelloid species previously considered as synonymous, as in the case of
Tomentella lilacinogrisea Wakef. and Tomentella neobourdotti M.J.Larsen (Martini
and Hentic 2005). Detailed anatomical descriptions highlighting notable features,
coupled with reliable, informative and faithful illustrations are invaluable for the
correct species delimitation. The tropical African thelephoroid fungi can be particularly
difficult to identify due to a lack of easily available, scientifically reliable literature.
Anatomical comparisons with type species have enabled the identification of new
species (Yorou & Agerer 2007a,b). Herewith, we aim at providing additional
anatomical characters of previously published descriptions and to highlight important
discriminative features for each type.
Materials and methods
Measurements and drawings were made from microscopical preparations. Fine sections of fruit
bodies were mounted in water and afterwards in 2.5% KOH, in Congo Red, in Cotton Blue and in
Melzer’s reagent (Kreisel & Schauer 1987), respectively. Microscopic studies were performed using
a light microscope Leica DM LB2. Measurements were made at magnification X 1000 and did not
include the apiculus, ornamentation of basidiospores and sterigmata. Line drawings were made at
magnification X 1000 using a drawing tube. Colour codes of dried basidiocaps are given according
to Kornerup and Wanscher (1978). Descriptions follow criteria compiled by Kõljalg (1996). Herbarium
abbreviations follow Holmgren et al (1990). We refer to Yorou & Agerer (2007a, b) for protocols of
molecular and phylogenetic studies.
Results
Tomentella radiosa (P.Karst.) Rick, Broteria 2 (Ser. 3): 79. 1934. Figs 1-5
BASIDIOCARP resupinate, adherent to the substratum, pelliculose, continuous, 0.5- 0.8
mm thick. Hymenophore crustose to granulose, brown (6E5), continuous, subiculum
pale to yellow brown, sterile margin determinate, fimbriate, consisting of agglutinated
parallel hyphae, whitish to concolorous with subiculum.
R
HIZOMORPHS present in subiculum, common at the margins, colourless, pale yellow
to brown under dissection microscope, pale yellow to pale brown in water and in
2.5% KOH, 20-70 µm wide, undifferentiated, uniform loose (Agerer 1999), of type
A (Agerer 1987-2006), margin smooth, individual hyphae uniform and loosely
arranged (Fig. 1), 4-6(7) µm diameter, clamped, simple septa also present, rarely
inflated, thin to thick-walled (0.5-1 µm), colourless, sometimes pale yellow to pale
brown in water and in 2.5% KOH.
S
UBICULAR HYPHAE broadly clamped, simple septa infrequent, clamps commonly thick-
walled (2-3 µm), mostly with a large (1-2 µm diam.) hole, hyphae 4-7(8) µm wide,
sometimes inflated, then up to 11 µm diam., exceptionally with a balloon-like inflation
(20 µm) (Fig. 2), thick-walled (0.5-1.5 µm), walls light yellow, cross-shaped
ramifications absent, anastomosis common (Fig. 3), elbow-like outgrowths or short
side-branches common (Fig. 4). Subicular hyphae light brown to brown in water
and in 2.5% KOH. Hyphae from the sterile margin loosely arranged and parallel,
never isolated, clamped, simple septa common, thin-walled (0.2-0.6 µm), without
523
encrustations, colourless to very pale yellow in water and in 2.5% KOH, congophilous,
cyanophilous, rarely patchily amyloid especially at septa and tips of the hyphae.
S
UBHYMENIAL HYPHAE clamped, 5-8 µm wide, often short and inflated (Fig. 5), thin
to thick-walled (0.5-1 µm), always thick-walled at their base, colourless to pale
brown in water and in 2.5% KOH, strongly congophilous, cyanophilous, not amyloid.
CYSTIDIA absent.
Fig. 1: Tomentella radiosa. Optical section through the rhizomorph (from the holotype). Scale
bar = 10 µm.
Fig. 2: T. radiosa. Optical section of subicular hyphae showing balloon-like inflation (from the
holotype). Scale bar = 10 µm.
524
BASIDIA 40-55 µm long, 8-10 µm at apex, 6-8 µm at base, clavate, clamped at base,
not stalked, not sinuous, colourless to rarely light yellow in water and in 2.5%
KOH, 4-sterigmate, sterigmata 4-7 µm long and 1-1.5 µm at base.
B
ASIDIOSPORES of two types, first type (6)6.5-7.5 × 6-8 µm in frontal face, 6-7.5 ×
6-8 µm in lateral face, ellipsoid to lobed in frontal view, ellipsoid in lateral view,
second type 7.5-11.5 (13) × 7-9 µm in frontal face, 7.5-10 (12) × 7.5-9 µm in lateral
face, triangular in frontal view, ellipsoid to reniform in lateral view, echinulate,
aculei very short (0.2-0.4 µm), dense and irregular, pale brown in water and in 2.5%
KOH, slightly congophilous, cyanophilous, not amyloid.
C
HLAMYDOSPORES absent.
MATERIAL STUDIED: HOLOTYPE: Hypochnus fuscus (Pers.: Fr.) P.Karst. var. radiosus P.Karst., Medd.
Soc. Fauna Fl. Fenn. 9: 71. 1882: Finland, Helsingfors, on dead wood, W.Nylander, X 1858 (H).
REMARKS: The presences of large, broadly clamped, thick-walled subicular hyphae
with large holes at clamps and thickened septa, as well as triangular and reniform
pale brown basidiospores with very short aculei, are typical anatomical features for
Fig. 3: T. radiosa. Optical section of subicular hyphae showing the H-like anastomosis (from the
holotype). Scale bar = 10 µm.
525
T. radiosa. Previous descriptions of T. radiosa (Agerer & Bougher 2001, Dämmrich
2006, Kõljalg 1996, Melo et al. 2003) already reported a wide variability of
basidiospore shapes and size. Tomentella ellisii (Sacc.) Jülich & Stalpers present
also somewhat reniform basidiospores in lateral view (Dämmrich 2006). Both species
present basidiospores with size ranging between 6-13 µm (Dämmrich 2006, Kõljalg,
1996, Melo et al. 2003). Furthermore, T. ellisii reveals subicular and subhymenial
hyphae with the same size and inflations as those of T. radiosa. However, T. ellisii
is characterised by a cyanescent reaction of its basidia in KOH (a character that
T. radiosa lacks) and the more prominent basidiospores ornaments of up to 1 µm
long aculei.
The presence of elbow-like outgrowths of subicular hyphae and the rarely slightly
amyloid hyphae in T. radiosa have been reported only recently by Agerer & Bougher
(2001). Amyloidy is currently a rarely documented character within tomentelloid
fungi. Amyloidy occurs patchily and only very slightly either on the subicular hyphae
of a few species such as Tomentella subamyloidea Agerer and T. radiosa (Agerer &
Bougher 2001), or on the apiculus of basidiospores of Tomentella lapida (Pers.)
Stalpers and Tomentella bryophila (Pers.) M. J. Larsen (Dämmrich 2006, Yorou &
Agerer 2007a).
According to Larsen (1974), T. radiosa presents two kinds of subicular hyphae. He
reports that some hyphae are 3-5.5 µm in diameter, frequently clamped, thick-walled,
and pale to medium brown, and that other hyphae are 5.5-7 µm in diameter, simple
septate, mostly thin-walled, and pale yellowish to brown. The holotype we examined
Fig. 4: T. radiosa. Optical section of subicular hyphae showing elbow-like outgrowths (from the
holotype). Scale bar = 10 µm.
526
reveals clamped hyphae of similar size ranges without a distinct hiatus in dimensions.
Differences are found only in the colour, thickness and the frequency of clamps,
rather than their width. Subicular hyphae are thick-walled (0.5-1.5 µm), light brown
to brown, regularly clamped, whilst hyphae from the sterile margin of fruit body are
Fig. 5: Tomentella radiosa. a. Section thought the basidiocarp, b. Basidiospores in frontal view,
c. Basidiospores in lateral view (from the holotype). Scale bar = 10 µm.
527
rather thin-walled (0.2-0.6 µm), colourless to light yellow, clamped and simple
septate. Clampless hyphae as mentioned by Larsen (1974) are absent.
Kõljalg (1996) adopted a broad species concept for T. radiosa, including specimens
with quite resupinate fruit bodies and those with clavaroid processes at their margins.
According to Kõljalg (1996), T. radiosa, T. oligofibula M.J.Larsen, T. purpurea
Wakefield, and T. carbonaria M.J.Larsen are conspecific. He highlighted great
similarities between T. radiosa and Thelephora terrestris Ehrhart, a species with pileate
to sub-sessile, infundibuliform to imbricate fruit bodies (Corner 1968). Anatomical
differences between T. radiosa and Th. terrestris refer to basidiospore shape and
ornaments, and as a consequence, both species are considered by Kõljalg (1996) as
different, thus applying a narrow species concept. In addition to basidiospores shape
and ornaments, rhizomorphal structure shows important dissimilarities between both
Fig. 6. One of the most parsimonious trees showing the grouping of Tomentella radiosa and
Thelephora terrestris in the same clade. Bootstrap values are shown above branches. The Genbank
(NCBI, EMBL or UNITE) codes as well as origin of the vouchers are indicated after species names.
528
Th. terrestris Th. terrestris Th. terrestris
(AF272923) (DQ068970) (AF272921)
T. radiosa 98.91 99.82 99.82
(UDB000964)
Th. terrestris 99.1 100
(AF272921)
species. Like T. radiosa, Th. terrestris has undifferentiated rhizomorphs (Agerer 1988)
with loosely arranged uniform hyphae. However, they differ from those of T. radiosa
by having cystidia (Agerer & Weiß 1989). Contrary to Kõljalg (1996), Dämmrich
(2006) adopted a broad species concept and suggested T. radiosa as the resupinate
form of Th. terrestris (= Thelephora terrestris f. resupinata (Bourdot & Galzin)
Donk). We didn’t examine the holotype of Th. terrestris. However, in recent molecular
investigations (Yorou & Agerer 2007a, b, see also Fig. 6), a specimen identified as
T. radiosa presents almost identical sequences with those of Th. terrestris. Genetic
distance between T. radiosa and Th. terrestris ranges from 0.18 to 1.09% regarding
ITS rDNA sequences (Table 1). Phylogenetically, T. radiosa (accession number
UDB000964, Genbank UNITE) clusters together with Th. terrestris s. str. (accession
numbers AF272921, AF272923 and DQ068970, Genbank NCBI) with a very strong
bootstrap support (100%). Based on cited specimens, molecular data support the
synonymy suggested by Dämmrich (2006). However, to make reliable conclusions
about taxonomical relationships between both species, molecular investigations should
be undertaken using all specimens described as T. radiosa (Agerer & Bougher 2001,
Dämmrich 2006, Kõljalg 1996, Melo et al, 2003, Larsen, 1965, 1974, Wakefield
1966, 1969) and compared to Th. terrestris s. str. It is likely that specimens described
as T. radiosa encompass representatives of Th. terrestris f. resupinata (sensu Dämmrich
2006) and those of T. radiosa (sensu Kõljalg 1996).
Tomentella cinereoumbrina (Bres.) Stalpers, Stud. Mycol. 35: 96. 1993. Figs 7-8
BASIONYM: Hypochnus cinnereoumbrina Bres., Stud. Trent. (Ser. 2, Sci. Nat. Econom. ) 7: 62. 1926.
BASIDIOCARP resupinate, adherent to the substratum, arachnoid to crustose, continuous,
hymenophore grey to light brown (7D4), smooth to granular, sterile margin indeterminate.
R
HIZOMORPHS absent.
S
UBICULAR HYPHAE mostly simple septate (Fig. 7), 3-6(6.5) µm, thick-walled (0.5-1 µm),
walls yellowish, sometimes tortuous, lateral protuberances common, without
encrustations, colourless to brown in water and in 2.5% KOH, rarely cyanescent,
congophilous, cyanophilous, not amyloid.
S
UBHYMENIAL HYPHAE simple septate, clamps present, more common than in subicular
hyphae, 3-6 µm wide, thin-walled, without encrustations, colourless to light brown in
water and in 2.5% KOH, sometimes cyanescent, congophilous, cyanophilous, not amyloid.
Table 1. ITS rDNA sequences’ similarities (%) between Tomentella radiosa and Thelephora terrestris.
529
Fig. 7: Tomentella cinereoumbrina. a. Section through the basidiocarp; b. Basidiospores in frontal
view; c. Basidiospores in lateral view (from the lectotype). Scale bar = 10 µm.
530
Fig. 8: T. cinereoumbrina. Young basidia showing distal apice (from the lectotype). Scale bar = 10 µm.
CYSTIDIA absent.
B
ASIDIA 35-70 µm long, 8-11(12) µm at apex, 4-8 µm at base, clamped at base,
clavate to utriform, sinuous, not stalked, colourless to very light brown in water and
in 2.5% KOH, sometimes cyanescent, young basidia with distinctive distal apex
(Fig. 8), 4-sterigmate, sterigmata 6-10 × 2-4 µm, basidia congophilous, cyanophilous,
not amyloid.
B
ASIDIOSPORES (8.5)9-11(12) × (7.5)8-9(11) µm in frontal face, (7.5)8-10(11) ×
(7)8-9 µm in lateral face, irregularly ellipsoid to broadly ellipsoid in frontal view,
broadly ellipsoid in lateral view, echinulate, aculei dense and very short, 0.4-1 µm,
pale brown in water and in 2.5% KOH, very slightly congophilous, not cyanophilous,
not amyloid.
C
HLAMYDOSPORES absent.
MATERIAL STUDIED: LECTOTYPE: Italy, ad truncos arb. frond., Gocciadoro, J.Bresadola, VI 1901 (BPI)!
REMARKS: T. cinereoumbrina is easily identifiable by the mostly simple septate subicular
and subhymenial hyphae, the young basidia with distinct distal apex and the broadly
ellipsoid pale brown basidiospores with very short aculei. Simple septate subicular
hyphae are known from only few Tomentella species including Tomentella fibrosa
(Berk. & M.A.Curtis) Kõljalg, Tomentella fuscocinerea (Pers.) Donk, Tomentella badia
(Link) Stalpers (Kõljalg 1996, Melo et al. 2000, Dämmrich 2006) and Tomentella
furcata Yorou & Agerer (Yorou & Agerer 2007a). Infrequent clamps on subicular
hyphae have been also reported for T. oligofibula. The first three species are completely
clampless whilst clamps occur occasionally on subicular and more frequently on
subhymenial hyphae of T. cinereoumbrina, T. oligofibula and T. furcata. From this
531
last group, T. furcata is easily recognised by its distinctly yellow subglobose to globose
basidiospores that bear simple, and/or distinctly forked spines, and by the presence of
rather short suburniform basidia with infrequent transverse septa and/or intra-basidial
hyphae. T. cinereoumbrina differs from T. oligofibula by its broadly ellipsoid
basidiospores with rather short spines. Basidiospores of T. oligofibula sometimes have
bifurcate ornaments and are reminiscent of those of T. radiosa (Larsen et al. 1994,
Melo et al 2003, Kõljalg 1996).
Tomentella punicea (Alb. & Schwein.) J.Schröt. In Cohn, Krypt.-Fl. Schles. 3:
420. 1889. Figs 9-14
BASIONYM: Thelephora punicea Alb. & Schwein.: Fr. Albertini J.B. & Schweinitz L.D.Consp. Fung.
Lusat. 278. 1805
BASIDIOCARP resupinate, separable from the substrate, arachnoid to membranous,
continuous, up to 1 mm thick, hymenophore olive brown (4E4) to yellow-brown
(5E4 to 5E5), granulose, subiculum yellow brown, sterile margin determinate, byssoid
to fimbriate, brownish yellow.
R
HIZOMORPHS present in subiculum and at margins, dark brown under the dissection
microscope, thick and visible already at X6, yellow brown in water and in 2.5%
KOH, dimitic to trimitic (Figs. 9-10), slightly differentiated (type C or thelephoroid
rhizomorph type according to Agerer 1987-2006, 1999), individual hyphae colourless,
sometimes yellowish to very pale brown in water and in 2.5% KOH, strongly
congophilous, slightly cyanophilous, not amyloid, central hyphae wide (Fig. 11),
6-10 µm diam., thin-walled, of two kinds, some usually clamped, but simple septa
common, some completely clampless, then tortuous; below surface hyphae similar
to subicular ones, 3-4.5 µm wide, clamped, simple septa common, resulting in short
cells, surface hyphae consisting mostly of skeletals that commonly end in binding-
like hyphae; skeletals (Fig. 12) 2-4 µm wide, thick-walled (0.5-2 µm), clampless,
simple septa rare, rarely branched, tortuous and commonly bent; yellowish, sometimes
filled with yellow-brown content; binding-like hyphae (Fig. 13) emerging from
skeletals and below surface generative hyphae, 1-2(2.5) µm diam., thin-walled,
dendroid, tortuous, clampless, simple septa rare.
S
UBICULAR HYPHAE clamped, 2-4.5(5) µm diam., thin-walled, with shiny reddish to
yellow-brown particles/granules (in water) (Fig. 14) that rapidly dissolve in 2.5%
KOH resulting in a citrine (yellow-green to yellow-brown) solution (observable
even with naked eyes), colourless to pale yellow in water and in 2.5% KOH,
congophilous, slightly cyanophilous, not amyloid;
S
UBHYMENIAL HYPHAE clamped, 3-4 µm wide, thin-walled, with yellow-brown to red
shiny particles/granules (observable in water), particles rapidly dissolving in 2.5%
KOH, colourless to pale yellow in water and in 2.5% KOH, congophilous, slightly
cyanophilous, not amyloid,
C
YSTIDIA absent.
B
ASIDIA (35)40-60 µm long, 5.5-7.5 µm wide at apex and 3.5-5 µm wide at base,
clamped at base, clavate, sinuous, sometimes with transverse septa, sometimes with
532
yellow-brown to red-brown particles (observable only in water) on their surface,
colourless to pale yellow, congophilous, slightly cyanophilous, not amyloid,
4-sterigmate, sterigmata 4.5-7 µm long and 1-1.5 µm wide at base.
BASIDIOSPORES (6)6.5-8(8.5) × 6(6.5)8 µm in frontal face, (6)6.5-8(8.5) × 6(6.5)8 µm
in lateral face, triangular with widened proximal part (proximal part as wide as the
Fig. 9: Tomentella punicea. Views of an old rhizomorph. a. Surface view; b. Optical section (from
the lectotype). Scale bar = 10 µm.
533
length of the spores) to lobed in frontal view, ellipsoid in lateral view, thick-walled
(0.5-1 µm), pale yellow to pale brown, slightly congophilous, cyanophilous, not
amyloid, commonly with oil drops, oils drops turning dark-brown in Melzer’s reagent,
Fig. 10: T. punicea. Optical section through an old rhizomorph. Note the presence of distinctive
skeletals that ending into binding-like hyphae. Scale bar = 10 µm.
534
Fig. 11: T. punicea. Surface view of central hyphae of a rhizomorph. a. Hyphae simple septate; b.
Hyphae clamped and simple septate. Scale bar = 10 µm.
Fig. 12: T. punicea. Details of skeletals on rhizomorphs (from the lectotype). Scale bar = 10 µm.
535
echinulate, aculei short (0.5-1 µm), densely arranged giving an impression of
bifurcation.
C
HLAMYDOSPORES absent.
MATERIAL STUDIED: LECTOTYPE: USA, the Academy of Natural Sciences of Philadelphia, Bethlehem,
Schweinitz herbarium, herb. 676 (PH)!
REMARKS: The presence of skeletals and binding-like hyphae on the rhizomorphs is
typical for this species. In addition, shiny red-brown granules (in water) that dissolve
and produce a citrine solution in KOH make the identification of T. punicea quiet
easy.
Melo et al (2003) highlighted the presence of two kinds of generative hyphae, but
did not comments on central enlarged hyphae and whether the rhizomorphs are
differentiated. According to Agerer (1999, 1987-2006), rhizomorphs of T. punicea
belong to the thelephoroid type (rhizomorph type C or slightly differentiated).
According to Dämmrich (2006), T. punicea shows monomitic rhizomorphs. Kõljalg
(1996) mentioned the presence of both monomitic and dimitic rhizomorphs, but
provided rather scanty information about the features of the hyphae. Dimitic to
trimitic hyphal systems with binding-like hyphae have been reported for Tomentella
rubiginosa (Bres.) Maire and Tomentella atroarenicolor Nikol. (Melo et al 2003,
2006). Tomentella atroarenicolor is recognisable by the presence of hyphoid cystidia
Figure 13: T. punicea. Details of binding-like hyphae on the surface of rhizomorphs (from the
lectotype). Scale bar = 10 µm.
536
in the hymenium and on rhizomorphs. Kõljalg (1996) and Dämmrich (2006)
considered T. rubiginosa to be synonymous to T. punicea, while Melo et al (2003)
and Stalpers (1993) regarded both species as different due to the presence of longer
and narrower clavate basidia in T. rubiginosa.
Among the species we have examined so far, T. punicea is the unique tomentelloid
species that presents skeletal hyphae in the sens of Hartig (1885), Falck (1912),
Fig. 14: Tomentella punicea, a. Section through the basidiocarp, b. Basidiospores in frontal view,
c. Basidiospores in lateral view (from the lectotype). Scale bar = 10 µm.
537
Corner (1932) and Clemençon (1997). All other tomentelloid fungi reported to show
dimitic rhizomorphs, have, in addition to generative hyphae, rather exclusively very
thin (1-2 µm), thin to only slightly thick-walled, sinuous, sometimes branched and
multi-septate hyphae.
Only two Tomentelloid fungi produce a citrine (yellow-green to yellow-brown)
solution when encrustations are dissolved in KOH. To date, except T. punicea, such
a reaction is only known from Tomentella umbrinospora M.J.Larsen (Larsen 1974,
Losi 1997, Yorou & Agerer 2007b). The latter species differs from T. punicea by
the red-brown to chestnut colour (6D3-6E3) of its hymenophore and the presence of
numerous, thinner, only slightly thick-walled multi-septate skeletals and the complete
absence of binding-like hyphae (Yorou & Agerer 2007b).
Tomentella punicea is anatomically close to Tomentella ferruginea (Pers.: Pers.)
Pat. (Kõljalg 1996) and macroscopically to Tomentella botryoides (Schw.) Bourd.
& Galz. (Dämmrich 2006). A similarity between T. punicea and T. botryoides is
supported by various DNA-sequence analyses where both species form a monophyletic
group with a bootstrap support of 71% (Kõljalg et al. 2000), 82% (Yorou et al.
2007), and 95% (Yorou & Agerer 2007a). Studies by Yorou & Agerer (2007a)
indicate that T. ferruginea is a sister species to the group comprising T. punicea,
T. botryoides, Tomentella africana Yorou & Agerer and T. umbrinospora. However,
T. punicea differs from both, T. botryoides and T. ferruginea, by the lack of cyanescent
reactions in KOH and the presence of distinctive skeletals in the sense of Hartig
(1885), Falck (1912) and of binding-like hyphae.
Acknowledgements
Type materials were made available to us by curators of the herbaria H, BPI and PH to whom we
address our sincere thanks. The German Academic Exchange Service (DAAD) is also thanked for
the grant no. A/03/15106 offered to the first author. We ware much indebted to Dr. Alison Davies
(Institute for Systematic Botany, LMU-Munich) for her valuable help with linguistic checking of the
manuscript.
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Received 21 May 2007, accepted in revised version 30 June 2007.
