Studies in the Truncospora ohiensis – T. ochroleuca group (Polyporales, Basidiomycota)

Abstract

Based on both morphological and phylogenetic analyses, 7 species are recognized in the Truncospora ohiensis group and 6 of them are described as new. T. ohiensis s.str. occurs in the American North-East; its closest relatives are T. arizonica, the long-spored species
from the South-West USA, and T. ornata from East Asia. T. atlantica is another long-spored member of the group, distributed in Macaronesia and the Iberian Peninsula. The dwarf-sized and pale-coloured T. mexicana is described from the western coast of the Gulf of Mexico,
and its look-alikes, T. floridana and T. tropicalis, are from the Caribbean. The identification key for the T. ohiensis – ochroleuca group is provided.

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Germany. DOI: 10.1127/nova_hedwigia/2014/0221 0029-5035/2014/0221 $ 4.25
Nova Hedwigia Vol. 100 (2015) Issue 1–2, 159–175
published online October 22, 2014; published in print February 2015 Article
C
Studies in the Truncospora ohiensis – T. ochroleuca group
(Polyporales, Basidiomycota)
Viacheslav Spirin1*, Jiří Kout2 and Josef Vlasák3
1 Botanical Unit (Mycology), Finnish Museum of Natural History, P.O. Box 7, FI-00014
University of Helsinki, Finland
2 Department of Biology, Geosciences and Environmental Education, Faculty of Edu-
cation, University of West Bohemia, Klatovská 51, Pilsen, CZ-30619, Czech Republic
3 Biol. Centre of the Academy of Sciences of the Czech Republic, Branišovská 31, České
Budějovice, CZ-37005, Czech Republic
With 4 gures and 1 table
Abstract: Based on both morphological and phylogenetic analyses, 7 species are recognized in
the Truncospora ohiensis group and 6 of them are described as new. T. ohiensis s.str. occurs in the
American North-East; its closest relatives are T. arizonica, the long-spored species from the South-
West USA, and T. ornata from East Asia. T. atlantica is another long-spored member of the group,
distributed in Macaronesia and the Iberian Peninsula. The dwarf-sized and pale-coloured T. mexicana
is described from the western coast of the Gulf of Mexico, and its look-alikes, T. floridana and
T. tropicalis, are from the Caribbean. The identification key for the T. ohiensis – ochroleuca group
is provided.
Key words: polypores, taxonomy, Perenniporia.
Introduction
The genus Truncospora was established by Pilát (1953) with two species, T. ochroleuca
(Berk.) Pilát (the genus type) and T. ohiensis (Berk.) Pilát, and later it was placed
into the synonyms of Perenniporia (Ryvarden 1972). Recent DNA-based studies
indicated that Truncospora should be separated from other genera in the so called
"core polyporoid clade" (Robledo et al. 2009, Zhao & Cui 2013). Decock & Ryvarden
(1999) and Decock (2011) emphasized some morphological features (i.e., large truncate
basidiospores with a germ pore and skeletal hyphae variably branched at their distal
parts) as unique to this genus.
*Corresponding author: viacheslav.spirin@helsinki.fi
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Zhao & Cui (2013) described a new species, T. macrospora, and provided a key to the
species of Truncospora worldwide; T. ohiensis was included as a species occurring
in both North America and East Asia. However, DNA data and morphological study
of specimens collected in North America, Europe and East Asia and labelled as
T. ohiensis or T. ochroleuca showed that 7 closely related species exist. Their
descriptions are presented below.
Materials and methods
The specimens from mycological herbaria of Botanical Museum, University of Helsinki (H), New
York Botanical Garden (NY), Center for Forest Mycology Research (CFMR), Mycological Herbarium
of the Department of Biology, University of West Bohemia (KBI), and Institute of Biology and Soil
Science, Russian Academy of Sciences (VLAM), as well as from a private herbarium of the author
JV, were studied. Herbarium acronyms are given according to Index Herbariorum (Thiers 2014). Type
specimens are deposited in H. Morphological techniques of this study follow Miettinen et al. (2006,
2012). Microscopic structures were studied and measured with Leitz Diaplan microscope (×1250
magnification). Measurements were done in Cotton Blue using phase contrast illumination and oil
immersion (with a subjective accuracy of 0.1 µm – Miettinen et al. 2006). The next abbreviations
are used in the species descriptions: d/p – dissepiments thickness/pore diameter ratio (a quotient
counted from the measurements of 10 subjectively chosen tube dissepiments and equal number of
pores), L – mean spore length, W – mean spore width, Q’ – spore length/width ratio, Q – mean spore
length/width ratio (counted for each specimen measured).
DNA isolAtioN AND sequeNciNg: 0.02–0.2 g of the context tissue was disintegrated for 30 s with a
steel ball mixer mill MM301 RETSCH at room temperature. DNA was isolated using CTAB/NaCl
extraction buffer as described by Murray & Thompson (1980), followed by two times repeated
extraction with chloroform, and isopropanol precipitation. Crude DNA was dissolved in 100 mL of
sterile water and further purified using Wizard Clean Up kit PROMEGA. 0.5 ul of resulting DNA
solution of 50 mL was used as a template for amplification with ITS5 and ITS4 primers (White et al.
1990), or EF983F and EF2218R primers (Matheny et al. 2007) in 25 mL reaction mixture using 55°
C annealing temperature. With ITS primers, 30 cycles were used and 0.3 mL of crude amplified DNA
was directly applied for sequencing. With EF primers, 35 cycles PCR were used and the amplified
DNA band was purified from agarose gel using Machery-Nagel NucleoSpin kit. Sequencing from
ITS1 and ITS4 primers, resp. from EF983F, EF2218R and EF1567R primers was performed in
the Genomics laboratory of Biology Centre, Academy of Sciences of the Czech Republic, České
Budějovice, on ABI 3730xl DNA analyzer, using BigDye Terminator 3.1 kit.
PhylogeNetic ANAlysis We prepared two datasets for analysis: ITS dataset containing our sequences
of 18 Truncospora specimens (two of them yielded 2 haplotypes), a sequence of Perenniporia
medulla-panis (GenBank KJ410710) for rooting the phylogeny, and four similar sequences of
P. ohiensis/ochroleuca from GenBank. Tef1 dataset contained only newly generated sequences and
P. medulla-panis sequence (GenBank KJ410711). After alignment in MEGA5 (Tamura et al. 2011),
ITS dataset yielded 634 characters of which 126 variable and 76 parsimony informative. In the tef1
dataset, unaligned 3' tail was removed first (about 30 bp), yielding 1152 aligned characters, 258
variable and 159 parsimony informative. Evolutionary analyses were conducted in MEGA5 using
the Maximum Likelihood (ML) method.
Results
The species of the T. ohiensis group share all principal features characterizing the genus
in general. Skeletal hyphae are dextrinoid and cyanophilous (although those reactions
vary a bit from species to species), unbranched along their most length, except distal
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parts located predominantly at the dissepiment edges or in upper layer of tube trama.
Basidia in all species studied are rather uniform, clavate-pedunculate, with strongly
inflated epibasidial part, 25–35 × 10–14 µm, and basidiospores are unevenly thick-
walled, with a germ pore and usually with a large oil-drop. The spore wall thickness
is an important character in this group; however, the wall changes depending on age,
partly degrading in mature and, especially, in old spores.
Analysis of ITS sequences of T. ohiensis coll. from East Asia, Macaronesia, USA,
Mexico and Puerto-Rico showed very distinct sequence differences between
geographical samples, and ITS based phylogeny produced 7 strongly supported terminal
clades (Fig. 1). Dataset of tef1 sequences prepared from the same samples also showed
considerable sequence differences of ecologically-determined samples and nearly
the same topology (Fig. 2), strongly suggesting that 7 species are involved. This was
Fig. 1. Phylogenetic relationships of 22 Truncospora specimens inferred with ITS rRNA sequences.
Perenniporia medulla-panis was used to root the tree. Topology from maximum likelihood (ML)
analysis. Support values along branches from ML bootstrap (1000 replicates). The tree is drawn to
scale, with branch lengths measured in the number of substitutions per site. GenBank numbers with
assigned names indicate sequences retrieved from GenBank. Other GenBank numbers in "Specimens
examined".
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confirmed by painstaking micro- and macroscopic study of many new collections and
older herbarium specimens (see below).
In DNA terms, the T. ohiensis group is not a monophyletic unity. The members of the
T. ochroleuca complex split it at least in two subgroups (see Figs 1, 2). One of them
consists of T. arizonica, T. ohiensis and T. ornata and thus it encompasses temperate
species, while another subgroup includes the subtropical or tropical T. floridana,
T. mexicana and T. tropicalis. The Macaronesian T. atlantica position is somewhat
ambiguous in our phylogenies, quite close to the Australian T. ochroleuca; more samples
are needed to reveal its phylogenetic relations.
The closely related T. ochroleuca group differs in having distinctly wider skeletal
hyphae (4–6 µm) in both context and trama; it seems to include many species, too.
This group is not yet studied properly, and many older names should be checked for
establishing modern species concepts therein.
Truncospora arizonica Spirin & Vlasák, sp. nov. Fig. 4
Pileus perennis, consoliformis, superficies nigra, rimosa, poris 3–4 per mm. Systema hypharum
dimiticum. Basidiosporae crassitunicatae, ellipsoideae, truncatae, 11.7–15.6 × 7.1–9.5 µm,
dextrinoideae, cyanophilae.
holotyPe: USA. Arizona: Cochise Co., Portal Area, Chiricahua Mts., Quercus arizonica, 6.IX.2012
Vlasák 1209/69 (H).
Mycobank no. MB808453
Fig. 2. Phylogenetic relationships of 16 Truncospora taxa inferred with tef1sequences. Other data
as in Fig. 1.
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BAsiDiocArPs perennial, sessile, dimidiate, solitary, 20–30 mm long, 10–20 mm
wide. Upper surface first ochraceous to pale brown, in mature basidiocarps black,
matt, indistinctly striate, with several unclear annual zones, sometimes longitudinally
cracking. Margin sharp but rather thick, even, fertile. Pore surface first pale cream-
colored, later pale ochraceous; pores regular, circular, 3–4 per mm, dissepiments
normally rather thin, entire (d/p = 1), pubescent under lens. Section: pileal crust brown
to black, 0.2–0.8 mm thick; context 1–3 mm thick, ochraceous to brownish; tubes
indistinctly stratified, 6–10 mm thick, pale ochraceous to brownish.
hyPhAl structure di-trimitic; generative hyphae clamped, skeletal hyphae strongly
dextrinoid, distinctly cyanophilous (reactions more or less uniform in all parts of
basidiocarps).
coNtext: Hyphal structure dimitic; skeletal hyphae dominating, irregularly arranged
and occasionally branched, subsolid in upper part, having a distinct lumen in lower
layer, (3.0–)3.1–4.9(–5.2) µm in diam. (n = 35/2).
tuBes: Hyphal structure di-trimitic; trama irregular. Skeletal hyphae densely inter-
woven, thick-walled (wall normally not exceeding 1 µm thick), in some parts easily
collapsing, lumen distinct to wide, (2.9–)3.0–4.1(–4.2) µm in diam. (n = 40/2),
occasionally branched (terminal branches 1.5–2 µm in diam.), some with adventive
septa; generative hyphae thin-walled, 2–3 µm in diam.
BAsiDiosPores thick-walled (spore wall 1–1.5 µm thick), narrowly ellipsoid to ellipsoid,
mostly regularly but sometimes obliquely truncate, (11.3–)11.7–15.6(–15.8) × (6.5–)
Fig. 3. Hymenium and trama of T. ohiensis (isotype).
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7.1–9.5(–10.3) µm, L = 13.70, W = 8.11, Q' = (1.3–)1.4–1.9(–2.1), Q = 1.64–1.75
(n = 60/2), strongly dextrinoid, cyanophilous.
remArks: T. arizonica is a long-spored species in the T. ohiensis complex, along with
T. atlantica (Table 1). According to DNA data, it is closely related to T. ohiensis s.str.
Fig. 4. Basidiospores in the T. ohiensis group: a. T. arizonica (holotype), b. T. atlantica (holotype),
c. T. floridana (holotype), d. T. mexicana (holotype), e. T. ohiensis (JV 0108/116), f. T. ornata
(holotype), g. T. tropicalis (holotype).
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Table 1. Spore dimensions in the Truncspora ohiensis – T. ochroleuca group.
Species/specimen L' L W' W Q' Q n
T. arizonica (11.3)11.7–15.6(15.8) 13.70 (6.5)7.1–9.5(10.3) 8.11 (1.3)1.4–1.9(2.1) 1.70 60
Vlasák 1209/69 (12.7)13.1–15.6(15.8) 14.48 (6.7)7.3–9.5(10.3) 8.31 (1.5)1.6–1.9(2.1) 1.75 30
Vlasák 1307/11bJ (11.3)11.7–14.3(15.1) 12.91 (6.5)7.1–9.2(9.4) 7.91 (1.3)1.4–1.8(1.9) 1.64 30
T. atlantica (12.1)12.6–16.0(16.1) 14.19 (7.2)7.4–10.2(10.3) 8.86 (1.4)1.5–1.9(2.1) 1.61 60
Alanko 128700 (12.1)12.6–16.0(16.1) 14.21 (7.3)8.0–10.2(10.3) 9.03 (1.4)1.5–1.8(2.0) 1.58 30
Väisälä 22 (12.3)12.8–15.6(16.1) 14.17 (7.2)7.4–9.8(10.0) 8.69 (1.4)1.5–1.9(2.1) 1.64 30
T. floridana (9.8)9.9–12.0(12.1) 11.03 (6.0)6.2–8.9(9.0) 7.19 (1.3)1.4–1.7(1.8) 1.54 90
Vlasák 1008/78 (10.0)10.1–11.8(12.1) 11.05 (6.1)6.3–8.3(8.4) 7.25 (1.3)1.4–1.7(1.8) 1.53 30
Vlasák 1008/86 (9.8)9.9–12.0(12.1) 10.76 (6.0)6.2–7.7(8.1) 7.02 (1.3)1.4–1.6(1.7) 1.54 30
West 1936 (10.3)10.6–12.0(12.1) 11.29 (6.2)6.3–8.9(9.0) 7.30 (1.3)1.4–1.7(1.8) 1.56 30
T. mexicana (10.4)10.6–13.7(14.0) 11.90 (6.5)6.6–9.2(9.3) 7.68 (1.3)1.4–1.7(1.8) 1.56 90
JV 0610/U-4 Kout (11.2)12.0–13.7(14.0) 12.79 (7.2)7.4–9.2(9.3) 8.11 (1.3)1.4–1.7(1.8) 1.58 30
Murrill 183 (10.8)10.9–13.3(13.8) 11.76 (6.5)6.6–8.6(8.9) 7.61 (1.3)1.4–1.7(1.8) 1.55 30
Vlasák 1109/83 (10.4)10.6–12.2(13.6) 11.13 (6.7)6.8–8.1(8.2) 7.32 (1.3)1.4–1.6(1.7) 1.53 30
T. ochroleuca coll. - - - - - - -
JV0610/7B-Kout (10.9)11.2–13.2(13.3) 12.39 (6.7)6.8–8.8(9.2) 7.52 (1.3)1.4–1.8(1.9) 1.66 30
Lowe 13123 (12.0)12.2–15.7(15.8) 13.70 (7.1)7.3–10.2(10.3) 8.62 (1.3)1.4–1.8(1.9) 1.81 30
Niemelä 1178 (14.4)14.9–21.8(23.4) 18.46 (8.0)8.4–11.8(12.2) 10.24 (1.5)1.6–2.2(2.4) 1.81 30
Niemelä 5268 (14.6)14.8–17.2(17.6) 15.86 (6.5)7.0–9.0(9.1) 8.03 (1.6)1.7–2.2(2.4) 1.99 30
Niemelä 5647 (10.2)10.8–13.6(14.3) 12.36 (6.3)6.6–9.1(9.7) 7.71 (1.4)1.5–1.9(2.0) 1.61 30
T. ohiensis (9.3)9.6–13.2(13.4) 11.14 (6.2)6.4–9.3(9.8) 7.47 (1.2)1.3–1.8(1.9) 1.50 150
Lea (isolectotype) (9.5)9.6–11.7(12.2) 10.75 (6.4)6.7–8.5(8.8) 7.41 (1.2)1.3–1.6(1.7) 1.46 30
Vlasák 0108/116c (9.8)10.0–12.0(12.1) 10.86 (6.2)6.4–8.7(8.9) 7.29 (1.2)1.3–1.7(1.8) 1.50 30
Vlasák 0309/114 (9.3)10.1–12.1(12.2) 10.99 (6.2)6.5–8.7(9.2) 7.32 (1.2)1.3–1.6(1.7) 1.51 30
Vlasák 0509/64 (10.3)10.7–13.2(13.4) 11.92 (7.2)7.3–9.3(9.8) 8.14 (1.2)1.3–1.6(1.7) 1.47 30
Vlasák 1208/21a (9.7)10.0–12.8(13.2) 11.19 (6.3)6.5–8.0(8.1) 7.20 (1.3)1.4–1.8(1.9) 1.56 30
T. ornata (10.1)10.2–12.8(13.0) 11.17 (6.7)6.8–9.1(9.3) 7.73 (1.2)1.3–1.6(1.7) 1.45 60
Spirin 6672 (10.2)10.3–11.8(12.1) 11.02 (6.7)6.8–9.1(9.3) 7.69 (1.2)1.3–1.6(1.7) 1.44 30
Dai 1644 (10.1)10.2–12.8(13.0) 11.32 (6.9)7.0–9.1(9.2) 7.76 (1.3)1.4–1.6(1.7) 1.46 30
T. tropicalis (9.2)9.3–12.2(12.4) 10.75 (6.1)6.2–9.1(9.2) 7.40 (1.1)1.3–1.7(1.8) 1.47 60
Vlasák 1112/18J (9.2)10.0–12.2(12.4) 11.21 (6.4)6.8–9.1(9.2) 7.88 (1.2)1.3–1.7(1.8) 1.43 30
Vlasák 1008/45 (9.2)9.3–12.0(12.1) 10.28 (6.1)6.2–8.4(8.8) 6.92 (1.1)1.3–1.7(1.8) 1.50 30
Truncospora sp. (10.2)10.3–12.2(12.3) 11.19 (6.6)7.1–8.3(8.8) 7.75 (1.3)1.4–1.6(1.7) 1.45 30
Ryvarden 5042 (10.2)10.3–12.2(12.3) 11.19 (6.6)7.1–8.3(8.8) 7.75 (1.3)1.4–1.6(1.7) 1.45 30
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from the American North-East and T. ornata from East Asia. T. arizonica differs from
those species in having of thick black crust, rimose in older basidiocarps, and larger
pores. Context is also darker than in other members of the group, and contextual skeletal
hyphae are wider.
T. arizonica is so far known only from Arizona inhabiting oak wood. Both description
and illustration of Perenniporia ohiensis given by Gilberston & Ryvarden (1987) refer
to this species.
Truncospora atlantica Spirin & Vlasák, sp. nov. Fig. 4
Pileus perennis, dimidiatus, superficies cremea vel ochracea, laevis, poris 3–4 per mm. Systema
hypharum dimiticum. Basidiosporae crassitunicatae, ellipsoideae, truncatae, 12.6–16.0 × 7.4–10.2
µm, dextrinoideae, cyanophilae.
holotyPe: Spain. Canary Islands: Tenerife, Anaga, Erica arborea, 21.XII.2005 Alanko 128700 (H).
Mycobank no. MB808454
BAsiDiocArPs perennial, sessile, dimidiate, solitary, 15–45 mm long, 10–25 mm wide.
Upper surface first pale ochraceous, in mature basidiocarps ochraceous to brownish,
matt, indistinctly striate, with several unclear annual zones. Margin sharp or blunt, even,
fertile. Pore surface first pale cream-colored, later pale ochraceous, sometimes with
brownish stains; pores regular, circular, 3–4 per mm, dissepiments thick, entire (d/p =
0.5–1.5), pubescent under lens. Section: no distinct pileal crust; context 1–3 mm thick,
cream- to pale wood-colored; tubes one-layered, 4–15 mm thick, cream-colored to
pale ochraceous.
hyPhAl structure di-trimitic; generative hyphae clamped, skeletal hyphae dextrinoid
(reaction distinct), moderately to strongly cyanophilous.
coNtext: Hyphal structure dimitic; skeletal hyphae dominating, irregularly arranged
to subparallel, occasionally branched, (2.7–)3.1–5.0(–5.1) µm in diam. (n = 40/2).
tuBes: Hyphal structure di-trimitic; trama irregular. Skeletal hyphae densely
interwoven, thick-walled (wall up to 2 µm thick), lumen distinct to capillary, (2.7–)
2.8–4.0(–4.1) µm in diam. (n = 40/2), occasionally branched (terminal branches 1.5–2
µm in diam.); generative hyphae thin-walled, 1.5–3 µm in diam.
BAsiDiosPores: thick-walled (spore wall 1–2 µm thick), narrowly ellipsoid to ellipsoid,
often regularly truncate, (12.1–)12.6–16.0(–16.1) × (7.2–)7.4–10.2(–10.3) µm, L =
14.19, W = 8.86, Q' = (1.4–)1.5–1.9(–2.1), Q = 1.58–1.64 (n = 60/2), moderately to
strongly dextrinoid, cyanophilous (staining more or less even in all spores).
remArks: T. atlantica was first described by Torrend (1910) as Polyporus ochroleucus
var. lusitanica. Later it was reported as Perenniporia ochroleuca from several European
countries (Jahn 1972, Ryvarden & Gilbertson 1994). However, it differs from T. ochroleuca
coll. in having distinctly narrower tramal and contextual skeletal hyphae.
T. atlantica is distributed in Macaronesia (Canary Islands and Madeira) and the Iberian
Peninsula, and it is probably the only species of Truncospora occurring in Europe. It
inhabits branches of trees and shrubs, as well as woody fences, poles etc.
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Truncospora floridana Vlasák & Spirin, sp. nov. Fig. 4
Pileus perennis, perpusillus, superficies cremea vel pallide ferruginea, laevis, poris 6–7 per mm.
Systema hypharum dimiticum. Basidiosporae crassitunicatae, ellipsoideae, truncatae, 9.9–12.0 ×
6.2–8.3 µm, dextrinoideae, cyanophilae.
holotyPe: USA. Florida: Key Largo, John Pennekamp St. Pk., hardwood, 29.VIII.2010 Vlasák
1008/78 (H).
Mycobank no. MB808455
BAsiDiocArPs perennial, sessile, dimidiate, solitary, 12–25 mm long, 8–20 mm wide.
Upper surface first almost white to pale cream-colored, in very old basidiocarps
brownish to rusty brown, matt, indistinctly striate, azonate. Margin sharp but rather
thick, even, fertile. Pore surface first white, later pale cream-colored; pores regular,
circular, later elongated, almost invisible by naked eye, (5) 6–7 per mm, dissepiments
very thick, entire (d/p = 1.5–2), pubescent under lens. Section: pileal crust first
indistinct, in older basidiocarps very thin, dark-brown; context 2–3 mm thick, cream-
to pale wood-colored; tubes one-layered, 1–3 mm thick, cream- to pale wood-colored.
hyPhAl structure di-trimitic; generative hyphae clamped, skeletal hyphae dextrinoid
(reaction weak in context, weak to moderate in tube trama), variably cyanophilous.
coNtext: Hyphal structure dimitic; skeletal hyphae dominating, irregularly arranged
and occasionally branched, (3.0–)3.1–3.9(–4.1) µm in diam. (n = 40/2).
tuBes: Hyphal structure di-trimitic; trama irregular. Skeletal hyphae densely interwoven,
thick-walled (wall normally not exceeding 1 µm thick), lumen distinct, (2.3–)2.4–3.6
(–3.9) µm in diam. (n = 60/3), occasionally branched (terminal branches 1.5–2 µm in
diam.), some with adventive septa; generative hyphae thin-walled, 2–3.3 µm in diam.
BAsiDiosPores thick-walled (spore wall 0.8–1.5(2) µm thick), ellipsoid to broadly
ellipsoid, often regularly truncate, (9.8–)9.9–12.0(–12.1) × (6.0–)6.2–8.3(–8.4) µm,
L = 11.03, W = 7.19, Q' = (1.3–)1.4–1.7(–1.8), Q = 1.53–1.56 (n = 90/3), moderately
to strongly dextrinoid, cyanophilous (coloration more or less even in all spores).
remArks: T. floridana produces small-sized basidiocarps. Its pores are smallest in the whole
group, poorly visible by the naked eye due to exceptionally thick dissepiments. Moreover,
contextual layer of T. floridana is as thick as tubes or thicker; this feature distinguishes it
from the similarly looking T. mexicana and T. tropicalis. The latter species, also occurring
in South Florida, has wider skeletal hyphae in both context and tubes; in addition, its pores
are wider and tube dissepiments are thinner (see below).
T. floridana is so far found only in Florida.
Truncospora mexicana Vlasák, Spirin & Kout, sp. nov. Fig. 4
Pileus perennis, perpusillus, superficies cremea vel pallide ochracea, laevis, poris 5–7 per mm.
Systema hypharum dimiticum. Basidiosporae crassitunicatae, ellipsoideae, truncatae, 10.6–13.7 ×
6.6–9.2 µm, dextrinoideae, cyanophilae.
holotyPe: Mexico. Veracruz: Park Zamora, Casuarina equisetifolia (fallen branch), 1.X.2006 Kout
0610/U (H, isotype in KBI).
Mycobank no. MB808456
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BAsiDiocArPs perennial, sessile, dimidiate, solitary, 5–20 mm long, 3–18 mm wide.
Upper surface first almost white or pale cream-colored, in mature basidiocarps pale
ochraceous, especially at the margin, matt, indistinctly striate and zonate. Margin sharp
but rather thick or blunt, even, fertile. Pore surface first pale cream-colored, later pale
ochraceous to pale brownish; pores round to angular, later partly sinuous, 5–7 per
mm, dissepiments first entire and thick, then uneven and strongly thinning-out (d/p =
0.25–2.5), pubescent under lens. Section: no distinct pileal crust; context exceptionally
thin, 0.1–1(2) mm thick, cream- to pale wood-colored; tubes indistinctly stratified,
3–13 mm thick, cream- to wood-colored.
hyPhAl structure: di-trimitic; generative hyphae clamped, skeletal hyphae dextrinoid
(reaction weak in context, moderate in tube trama), weakly to moderately cyanophilous.
coNtext: Hyphal structure dimitic; skeletal hyphae dominating, irregularly arranged
and occasionally branched, some hyphae flexuous and unevenly inflated, (1.9–)2.0–3.5
(–3.6) µm in diam. (n = 40/2).
tuBes: Hyphal structure di-trimitic; trama irregular. Skeletal hyphae loosely interwoven,
often flexuous, with thickened walls (wall mostly not exceeding 1 µm thick) and
occasional short side branches, lumen wide, (2.2–)2.3–3.3(–3.8) µm in diam. (n =
60/3), regularly branched (terminal branches 1–2 µm in diam.), some with adventive
septa; generative hyphae thin-walled, 1.5–2 µm in diam.
BAsiDiosPores: thick-walled (spore wall 1–2 µm thick), ellipsoid to broadly ellipsoid,
often regularly truncate, (10.4–)10.6–13.7(–14.0) × (6.5–)6.6–9.2(–9.3) µm, L = 11.90,
W = 7.68, Q' = (1.3–)1.4–1.7(–1.8), Q = 1.53–1.58 (n = 90/3), moderately to strongly
dextrinoid, cyanophilous (reaction clear and even).
RemArks: T. mexicana is similar to T. floridana in having dwarf-sized basidiocarps with
rather small pores; however, its basidiocarps are more deeply colored, with ochraceous
hues, and contextual tissue is very thin. Pores of T. mexicana undergo strong changes
first being rather regular and thick-walled, and later becoming sinuous, with thinning-
out and uneven dissepiments. Moreover, T. mexicana has longer spores if compared with
T. floridana and T. tropicalis, but these differences are merely statistical (Table 1). The
latter species differs from T. mexicana due to larger pores and wider skeletal hyphae in
both context and tubes. These species seem to not coincide in their distribution areas.
T. mexicana is known from the western and southern coasts of the Gulf of Mexico and
found in South Texas (USA), Veracruz (Mexico), and the westernmost part of Cuba
(Pinar del Río Province).
Truncospora ochroleuca (Berk.) Pilát coll.
Polyporus ochroleucus Berk. was described from South-West Australia, and it represents
a large-pored (Berkeley 1845) and long-spored (Decock 2011) species. The only ITS
sequence of the Australian T. ochroleuca available in GenBank (FJ411097) shows no
total match with other species in the T. ohiensis – T. ochroleuca complex. However,
several species of the T. ochroleuca group may occur in Australia only. ITS sequences
obtained by us from some East Asian, African and Central American collections of
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T. ochroleuca coll. reveal that several species are hidden under this name (not shown
in the phylogram); further studies may elucidate this problem.
Truncospora ohiensis (Berk.) Pilát Figs 3, 4
LectotyPe: USA. Ohio: Waynesville, Lea (NY – isolectotype, studied).
BAsiDiocArPs perennial, sessile, dimidiate, solitary, 20–60 mm long, 10–40 mm wide.
Upper surface first pale ochraceous to pale brown, in mature basidiocarps brown to
brownish-black, matt, with several unclear annual zones. Margin first sharpened,
in older basidiocarps blunt, cream-colored to pale ochraceous, even, fertile. Pore
surface white to pale cream-colored; pores regular, circular or elongated, 5–6 per mm,
dissepiments thick, entire (d/p = 1–2), pubescent under lens. Section: pileal crust brown
to blackish, 0.1–0.3 mm thick; context 1–3 mm thick, pale cream-colored to ochraceous;
tubes indistinctly stratified, 1–8(15) mm thick, pale cream-colored to ochraceous.
hyPhAl structure: di-trimitic; generative hyphae clamped, skeletal hyphae moderately
dextrinoid, moderately to strongly cyanophilous (reactions more or less uniform in
all parts of basidiocarps).
CoNtext: Hyphal structure dimitic; skeletal hyphae dominating, loosely interwoven
or arranged in parallel bundles, occasionally branched, lumen wide to narrow, (2.1–)
2.3–3.7(–4.0) µm in diam. (n = 70/4).
tuBes: Hyphal structure di-trimitic; trama irregular. Skeletal hyphae densely inter-
woven, thick-walled (wall normally not exceeding 1 µm thick), in some parts easily
collapsing, lumen distinct to wide, (1.8–)2.0–3.1(–3.2) µm in diam. (n = 60/3), occa-
sionally branched (terminal branches 1–1.5 µm in diam.), some with adventive septa;
generative hyphae thin-walled, 1–3.5 µm in diam.
BAsiDiosPores thick-walled to very thick-walled (spore wall 1–2.5 µm thick), narrowly
ellipsoid to ellipsoid, normally regularly truncate, (9.3–)9.6–13.2(–13.4) × (6.2–)
6.4–9.3(–9.8) µm, L = 11.14, W = 7.47, Q' = (1.2–)1.3–1.8(–1.9), Q = 1.47–1.56 (n =
150/5), strongly dextrinoid, cyanophilous (reaction more or less even).
remArks: This species was described as Trametes ohiensis based on the sole specimen
from Ohio (Berkeley 1872); we studied its part (NY), and it agrees well with other
specimens collected in the American North-East. T. ohiensis has rather big, ungulate
basidiocarps finally covered by brownish-black crust with large, unclear zones, while
margin usually stays pale-colored (concolorous with pore surface). Its closest relative,
T. arizonica, is characterized by larger pores with thinner dissepiments and distinctly
longer spores, as well as wider skeletal hyphae. Another counterpart, T. ornata from East
Asia, is microscopically almost identical with T. ohiensis, but it produces considerably
smaller, conchate basidiocarps (see below). The specimen Ryvarden 5042 from Kenya
(O, H, labeled as Perenniporia ochroleuca) is morphologically very similar to T.
ohiensis s.str., and it seems to be another, still undescribed species in this group; this
subject deserves a separate study.
T. ohiensis is a widely distributed species in the US North-East and adjacent parts of
Canada. It inhabits fallen logs and branches of deciduous trees, as well as "structural
timber" (Overholts 1914).
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Truncospora ornata Spirin & Bukharova, sp. nov. Fig. 4
Pileus perennis, conchatus, superficies pallide ochracea vel cinnamomea, laevis, poris 5–6 per mm.
Systema hypharum dimiticum.Basidiosporae crassitunicatae, ellipsoideae, truncatae, 10.2–12.8 ×
6.8–9.1 µm, dextrinoideae, cyanophilae.
HolotyPe: Russia. Khabarovsk Reg.: Khabarovsk Dist., Bol’shoi Khekhtsir Nat. Res., Fraxinus
mandshurica, 5.IX.2013 Spirin 6672 (H).
Mycobank no. MB808457
BAsiDiocArPs perennial, sessile, dimidiate, solitary, 7–25 mm long, 5–10 mm wide.
Upper surface first pale cream-colored or grayish, warted, in mature basidiocarps brown,
matt, indistinctly striate, with several unclear annual zones. Margin sharp, even, fertile.
Pore surface first white, later pale cream-colored, in senescent basidiocarps brownish;
pores regular, circular, 5–6 per mm, dissepiments entire, first think, later thinning-out
(d/p = 0.7–2). Section: pileal crust brownish, seen as a thin line ca. 0.1 mm thick; context
1–2 mm thick, cream-colored, sometimes with brownish hues; tubes one-layered or
indistinctly stratified, 1–2 mm thick, cream-colored to brownish.
hyPhAl structure di-trimitic; generative hyphae clamped, skeletal hyphae strongly
dextrinoid, distinctly cyanophilous (reactions more or less uniform in all parts of
basidiocarps).
coNtext: Hyphal structure dimitic; skeletal hyphae dominating, irregularly arranged
and occasionally branched, (2.4–)2.9–3.7(–3.9) µm in diam. (n = 20/1).
tuBes: Hyphal structure di-trimitic; trama irregular. Skeletal hyphae densely
interwoven, thick-walled (lumen usually distinct), (1.9–)2.0–3.3(–3.4) µm in diam.
(n = 40/2), occasionally branched (terminal branches 1–1.5 µm in diam.), some with
adventive septa or occasional inflations up to 5 µm in diam.; generative hyphae thin-
walled, 1.5–2.5(3) µm in diam.
BAsiDiosPores thick-walled (spore wall 1–1.5 µm thick), ellipsoid to broadly ellipsoid,
mostly regularly but sometimes obliquely truncate, (10.1–)10.2–12.8(–13.0) × (6.7–)
6.8–9.1(–9.3) µm, L = 11.17, W = 7.73, Q' = (1.2–)1.3–1.5(–1.6), Q = 1.44–1.46 (n
= 60/2), strongly dextrinoid, cyanophilous. Basidiospores detected on pileal surface
variably brownish, evidently absorbing a colored matter from underlying hyphae.
remArks: This species was first collected by L.V.Ljubarsky in Russian Far East and
reported as Fomitopsis ohiensis by Bondartsev (1953). That time this identification was
feasible due to lack of any striking morphological differences between T. ornata and
T. ohiensis. The only peculiar character of T. ornata is its dwarf-sized, conchate basidiocarps
(versus normally large and ungulate fruitbodies of T. ohiensis); however, these species are
certainly different according to DNA data (Figs 1, 2). Polyporus junctus was described by
Lloyd (1924: 1317) from Japan, and now it is considered as a synonym of T. ochroleuca
(Ryvarden 1990). Its picture published by Lloyd (1924, plate 311, fig. 3028) shows several
thick and light-colored caps fusing together – thus, this name would be hardly addressed
to T. ornata, having small-sized, solitary basidiocarps with brown upper surface. Identity
of Lloyd’s species versus the East Asian T. ochroleuca should be checked separately.
T. ornata is a temperate species, distributed in deciduous and mixed forests of Russian
Far East (Khabarovsk Reg. and Primorie) and North-East China but it seems to be
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rare. T. ornata grows on fallen logs and thick branches of frondose trees (Fraxinus
mandshurica, Maakia amurensis, Phellodendron amurense, Quercus mongolica) and
evidently avoids the man-induced ecosystems.
Truncospora tropicalis Vlasák & Spirin, sp. nov. Fig. 4
Pileus perennis, perpusillus, superficies cremea vel pallide ochracea, laevis, poris 4–5 per mm.
Systema hypharum dimiticum. Basidiosporae leniter crassitunicatae, ellipsoideae, truncatae, 9.3–12.2
× 6.2–9.1 µm, dextrinoideae, cyanophilae.
holotyPe: Puerto Rico. Rio Grande: El Yunque Nat. Forest, hardwood, XII.2011 J.Vlasák Jr.
1112/18J (H).
Mycobank no. MB808458
BAsiDiocArPs perennial, sessile, dimidiate, solitary, 10–20 mm long, 10–15 mm wide.
Upper surface first pale cream-colored, in mature basidiocarps pale ochraceous, matt,
indistinctly striate and zonate. Margin sharp but rather thick, even, fertile. Pore surface
first pale cream-colored, later pale ochraceous; pores angular, later elongated and partly
sinuous, in older basidiocarps showing a tendency to radial arrangement, 4–5(6) per
mm, dissepiments thin, entire (d/p = 0.3–0.8), pubescent under lens. Section: no distinct
pileal crust; context 1–2 mm thick, cream- to pale wood-colored; tubes one-layered,
1.5–5 mm thick, cream- to pale wood-colored.
hyPhAl structure di-trimitic; generative hyphae clamped, skeletal hyphae dextrinoid
(reaction moderate in context, weak to moderate in tube trama), weakly to moderately
cyanophilous.
coNtext: Hyphal structure dimitic; skeletal hyphae dominating, irregularly arranged
and occasionally branched, (3.1–)3.3–4.9(–5.1) µm in diam. (n = 40/2).
tuBes: Hyphal structure di-trimitic; trama irregular. Skeletal hyphae loosely interwoven,
often flexuous, with thickened walls (wall not exceeding 1 µm thick), lumen wide, a
few deeply arranged skeletals distinctly thick-walled and with a capillary lumen, (2.8–)
2.9–4.2(–4.3) µm in diam. (n = 40/2), regularly branched (terminal branches 1–1.5 µm
in diam.), some with adventive septa; generative hyphae thin-walled, 3–4 µm in diam.
BAsiDiosPores slightly thick-walled (spore wall 0.5–1 µm thick), ellipsoid to broadly
ellipsoid, a few spores almost subglobose, often regularly truncate, (9.2–)9.3–12.2
(–12.4) × (6.1–)6.2–9.1(–9.2) µm, L = 10.75, W = 7.40, Q' = (1.1–)1.3–1.7(–1.8),
Q = 1.43–1.50 (n = 60/2), weakly to moderately dextrinoid, unevenly cyanophilous
(reaction mostly weak, more or less visible in thickest parts of the spore wall only).
remArks: T. tropicalis, T. mexicana and T. floridana are pale-colored and dwarf-sized
species displaying weak morphological differences from each other. T. tropicalis is
distinguishable due to larger, thin-walled pores, more or less radially oriented in mature
basidiocarps, and relatively wide and loosely arranged tramal skeletals. In addition, spores
of T. tropicalis have thinnest walls if compared with the rest of species in this group, quickly
degrading and thus showing an uneven coloration in Cotton Blue.
T. tropicalis is so far found in South Florida (the Florida Keys), Hispaniola (Dominican
Republic) and Puerto Rico; it may be widely distributed in the eastern part of the
Caribbean.
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Species key for the T. ohiensis – T. ochroleuca group
1 Tramal skeletals 4–6 µm wide ........................................................................ T. ochroleuca coll.
1' Tramal skeletals up to 4 µm wide ...............................................................................................2
2 Pores 3–4 per mm, spores regularly reaching 14 µm long ......................................................... 3
2' Pores 4–7 per mm, spores reaching 14 µm long in exceptional cases ........................................4
3 Mature basidiocarps with thick, black crust. Arizona (USA) .................................... T. arizonica
3' Mature basidiocarps with ochraceous or brownish upper surface; no distinct crust. Macarone-
sia and the Iberian Peninsula ......................................................................................T. atlantica
4 Pores 4–5 per mm, in mature basidiocarps distinctly elongated and with a tendency to radial
arrangement. Dissepiments rather thin (d/p = 0.3–0.6). Spore wall up 1 µm thick, only partly
and weakly cyanophilous. South Florida, Hispaniola, Puerto-Rico ..........................T. tropicalis
4' Pores 5–7 per mm; no tendency to radial arrangement, dissepiments normally thick. Spore wall
1 µm thick or more, distinctly cyanophilous ..............................................................................5
5 Temperate species ....................................................................................................................... 6
5' Subtropical or tropical species (the Caribbean and the Gulf of Mexico) ...................................7
6 Basidiocarps ungulate, 3–10 mm thick. American North-East ...................................T. ohiensis
6' Basidiocarps conchate, 2–4 mm thick. East Asia .......................................................... T. ornata
7 Context equally thick to or thicker than tube layer. Florida ...................................... T. floridana
7' Context distinctly thinner than tube layer. Western and southern coasts of the Gulf of Mexico
...................................................................................................................................T. mexicana
Specimens examined
T. arizonica. USA. Arizona: Cochise Co., Portal Area, Chiricahua Mts., Quercus arizonica, 6.IX.2012
Vlasák 1209/69 (H – holotype, JV) (GenBank KJ410696, KJ410716); Santa Cruz Co., Coronado
Nat. Forest, Madera Canyon, Quercus hypoleucoides, 5.IX.1988 Gilbertson 16372 (CFMR),
Q. arizonica, 2.IX.2012 Vlasák 1209/21a (JV, H) (GenBank KJ410695, KJ410715), VII.2013 J.
Vlasák Jr. 1307/11b (JV, H) (GenBank KJ410697, KJ410717).
T. atlantica. Portugal. Estremadura: Mata do Bombarral, Arbutus unedo, 27.X.1978 Melo & Cardoso
469 (H ex LISU). Madeira: Funchal, Botanical Garden, vineyard support wood, 6.I.1974 Väisälä
22 (H). Spain. Canary Islands: Tenerife, Anaga, Erica arborea, 21.XII.2005 Alanko 128700 (H
– holotype, JV) (GenBank KJ410700, KJ410720), Myrica faya, 30.XI.2013 Kout 1311/4K (JV)
(GenBank KJ410699, KJ410719).
T. floridana. USA. Florida: Gainesville, Upper Sugarfoot Prairie, Persea borbonia (decorticated
stump), 14.II.1936 West (NY); Key Largo, John Pennekamp St. Pk., hardwood, 29.VIII.2010
Vlasák 1008/78 (H – holotype, JV) (GenBank KJ410704, KJ410723); Miami, Matheson Hammock,
hardwood, 30.VIII.2010 Vlasák 1008/85 (JV) (GenBank KJ410705, KJ410724), 1008/86 (JV, H)
(GenBank KJ410706, KJ410725).
T. mexicana. Cuba. Pinar del Río Prov.: Herradura, dead wood in dense thickets, 7–12.III.1905
Earle & Murrill 183 (NY). Mexico. Veracruz: Zamora Park, Casuarina equisetifolia (fallen branch),
1.X.2006 Kout 0610/U (H – holotype, JV, KBI) (GenBank KJ410707, KJ410726). USA. Texas:
Cameron Co., Brownsville, Resaca de la Palma St. Park, fallen hardwood branch, IX.2011 Vlasák
1109/83 (JV, H) (GenBank KJ410708, KJ410709, KJ410727).
T. ochroleuca coll. Belize. Cockscomb Basin, Antilope Trail, 30.X.2006 Kout JV0610/7B-Kout (JV,
H, KBI) (GenBank KJ410698, KJ410718). Costa Rica. Heredia Prov.: Uvita, hardwood fencepost,
13.VII.1963 Lowe 13123 (NY, CFMR). Jamaica. Castleton Bot. Garden, 14–15.XII.1908 Murrill
126 (NY). Singapore. Nee Soon, Botanical Garden, angiosperm branch, 15.III.2011 Tran &
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Skornickova (H ex SING 2011-131) (var. brevispora – Corner 1989). Tanzania. Kilimandjaro Reg.:
Kilimanjaro, Mandara, E. arborea, 27.I.1973 Niemelä 1178 (H). Tanga Reg.: Lushoto, Mazumbai
Forest Reserve, dead stem, 8.XII.1989 Niemelä 5268 (H). Iringa Reg.: Mufindi, Ludoga, Acacia
mearnsii, 4.II.1993 Niemelä 5647 (H).
T. ohiensis. Canada. Ontario: Middlesex Co., London, dead branch and timber wood, XII.1890
Dearness (NY); York Co., Nashville, Fagus grandifolia, 25.IX.1957 Cain (NY). USA. Florida:
Alachua Co., Gainesville, hardwood, 2.IX.1952 Rhoads (CFMR), Carya sp., 13.IX.1952 Rhoads
(CFMR). Georgia: Coweta Co., Coweta Watershed, Robinia pseudoacacia, 20.IX.1965 Ross
215 (CFMR). Illinois: Piatt Co., Cerro Gordo, rotten wood, 6.VII.1915 Overholts 3210 (NY).
Indiana: Putnam Co., Greencastle, Quercus sp. (rails and logs), XI.1892 Underwood (NY).
Iowa: Fayette, Quercus sp. (fence posts), III.1908 Wilson (NY); Winneshiek Co., Decorah,
X.1882 Holway 303 (NY). Kansas: Riley Co., Manhattan, dead wood, 24.IX.1951 Rogerson
(NY). Michigan: Washtenaw Co., Ann Arbor, dead tree, 20.XI.1893 Pieters (NY). New York:
Onondaga Co., La Fayette, hardwood log, 11.X.1961 Gilbertson 3034 (NY); Wayne Co., Clyde,
IX.1887 Cook (NY). North Carolina: Buncombe Co., Ashville, fence post, 22.XII.1920 Bache-
Wiij (NY); Transylvania Co., Pink Bed Valley, 13–23.VII.1908 Murrill 440 & House (NY).
North Dakota: Cass Co., Fargo, Quercus macrocarpa, IX.1921 Stevens & Brenckle (NY).
Ohio: Butler Co., Ohio, F. grandifolia, 26.VII.1911 Overholts 215 (NY); Erie Co., Milan,
26.VII.1974 Jones 74-741 (NY); Lorain Co., Henrietta Township, Maclura pomifera (living
tree), 12.X.1949 Bigelow 359 (NY), Vermillion River, III.1951 Bigelow 361 (NY); Preble Co.,
Fairhaven, dead wood, 23.VI.1939 Overholts (NY); Warren Co., Waynesville (NY – isotype,
see above). Oklahoma: Payne Co., Ripley Bluffs, Quercus sp., 16.VIII.1979 Rogerson (NY).
Pennsylvania: Bucks Co., Point Pleasant, Quercus sp., 21.VIII.2003 Vlasák 0108/116 (JV),
16.IX.2003 Vlasák 0309/114 (JV, H) (GenBank KJ410694); Montgomery Co., Schwenksville,
R. pseudoacacia, 31.VIII.2012 Vlasák 1208/21 (JV, H), (GenBank KJ410692, KJ410713).
Tennessee: Great Smoky Mts., Cove Hardwood Nat. Trail, R. pseudoacacia, 6.IX.2005 Vlasák
0509/64 (JV, H) (GenBank KJ410693, KJ410714). Virginia: Clarke Co., White Post, 16.IX.1910
(anonymous collector) (NY); Giles Co., Mountain Lake, Castanea dentata, 8–14.VII.1909 Murrill
370 (NY). West Virginia: Mason Co., Apple Grove, deciduous tree, 12.I.1935 Gould 10 (NY);
Monongalia Co., Morgantown, 26.X.1907 Hartley 57 (NY).
T. ornata. China. Jilin: Huadian Co., Dongxing, Maakia amurensis, 17.X.1993 Dai 1644 (H)
(GenBank KJ410691). Russia. Khabarovsk Reg.: Khabarovsk Dist., Bol’shoi Khekhtsir Nat. Res.,
Fraxinus mandshurica, 5.IX.2013 Spirin 6672 (H – holotype, JV) (GenBank KJ410690, KJ410712).
Primorie Reg.: Ussuri Dist., Ussuri Nat. Res., hardwood, 27.VIII.1964 Smekalkina (VLA M-14234),
hardwood, 19.VII.1964 Nazarova (VLA M-14236, 14237), 7.VIII.1964 Nazarova (VLA M-14238);
Khasan Dist., Kedrovaya Pad’ Nat, Res., Quercus sp., 5.IX.1958 Ryffa (VLA M-14235).
T. tropicalis. Dominican Republic. La Vega Prov.: Ebano Verde Res., hardwood, 29.V.1997
Ryvarden 40273 (CFMR DR-763 ex O). Puerto Rico. Rio Grande: El Yunque Nat. Forest, hardwood,
XII.2011 J.Vlasák Jr. 1112/18J (H – holotype, JV) (GenBank KJ410701, KJ410721).USA. Florida:
Long Key St. Pk., hardwood, VIII.2010 Vlasák 1008/45 (JV, H) (GenBank KJ410702, KJ410703,
KJ410722).
Truncospora sp. Kenya. Coast Prov.: Kwale, Makadara, 9.I.1970 Ryvarden 5042 (H ex O 15806).
Acknowledgements
We are very indebted to Dr. B.Thiers, E.D.Bloch (NY) and B.Ortiz-Santana (CFMR) for sending
collections of T. ohiensis coll. from North America. Nadezhda Bukharova (Vladivostok, Russia)
provided us with valuable specimens and data of T. ornata. Teuvo Ahti (Botanical Museum,
University of Helsinki) helped us with Latin diagnoses. The research of J.Vlasák was funded by
institutional support RVO: 60077344, and that one of J.Kout is supported by the project EXLIZ –
CZ.1.07/2.3.00/30.0013 (co-financed by the European Social Fund and the government of Czech
Republic).
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