Polypore genera Antella, Austeria, Butyrea, Citripora, Metuloidea and Trulla (Steccherinaceae, Polyporales)

Abstract

The polypore genera Antrodiella, Junghuhnia and Steccherinum have been proven polyphyletic, and are here partly revised based on phylogenetic and morphological evidence. The new genera Antella Miettinen, Austeria Miettinen, Butyrea Miettinen, Citripora Miettinen and Trulla Miettinen & Ryvarden are described, and the poroid-hydnoid Metuloidea re-introduced. Two new species are described: Citripora bannaensis Miettinen from subtropical China and Trulla meridae Miettinen & Ryvarden from the Venezuelan Andes. Twenty-one new combinations in the above-mentioned genera and Steccherinum are made. Lectotypes are selected for Physisporus flavicans P. Karst. (= Butyrea luteoalba) and P. variecolor P. Karst. (= Trechispora hymenocystis).

Full text

Ann. Bot. Fennici 53: 157–172 ISSN 0003-3847 (print) ISSN 1797-2442 (online)
Helsinki 13 May 2016 © Finnish Zoological and Botanical Publishing Board 2016
Polypore genera Antella, Austeria, Butyrea, Citripora,
Metuloidea and Trulla (Steccherinaceae, Polyporales)
Otto Miettinen1,* & Leif Ryvarden2
1) Finnish Museum of Natural History, P.O. Box 7, FI-00014 University of Helsinki, Finland
(*corresponding author’s e-mail: otto.miettinen@helsinki.)
2) Institute for Biological Sciences, University of Oslo, P.O. Box 1066 Blindern, N-0316 Oslo,
Norway
Received 17 Mar. 2016, nal version received 4 Apr. 2016, accepted 5 Apr. 2016
Miettinen O. & Ryvarden L. 2016: Polypore genera Antella, Austeria, Butyrea, Citripora, Metuloidea
and Trulla (Steccherinaceae, Polyporales). — Ann. Bot. Fennici 53: 157–172.
The polypore genera Antrodiella, Junghuhnia and Steccherinum have been proven
polyphyletic, and are here partly revised based on phylogenetic and morphological evi-
dence. The new genera Antella Miettinen, Austeria Miettinen, Butyrea Miettinen, Citri-
pora Miettinen and Trulla Miettinen & Ryvarden are described, and the poroid-hyd-
noid Metuloidea re-introduced. Two new species are described: Citripora bannaensis
Miettinen from subtropical China and Trulla meridae Miettinen & Ryvarden from the
Venezuelan Andes. Twenty-one new combinations in the above-mentioned genera and
Steccherinum are made. Lectotypes are selected for Physisporus avicans P. Karst.
(= Butyrea luteoalba) and P. variecolor P. Karst. (= Trechispora hymenocystis).
Introduction
The basidiomycete genera Antrodiella, Junghuh-
nia and Steccherinum have long been considered
related. Traditionally the name Antrodiella has
been used for polypore species with small and
small-pored basidiocarps, dimitic, clamped and
non-encrusted hyphae, and tiny spores. Jung-
huhnia has contained similar species but with
encrusted skeletocystidia. Steccherinum spp.
were considered similar to Junghuhnia but with
a hydnoid hymenophore.
Miettinen et al. (2012) produced a multi-lo-
cus phylogeny of the family Steccherinaceae,
which includes the three above-mentioned
genera. They showed clearly that none of the
previous concepts of those three genera fully
reect the evolutionary history, and suggested a
revised classication. They refrained from for-
mally describing any new taxa in their treatment.
This paper continues that work, introducing two
new species, ve new genera and new combina-
tions for the species previously included in the
genera Antrodiella, Junghuhnia and Steccheri-
num.
Material and methods
We relied on Miettinen et al. (2012) as a phylo-
genetic basis for the taxonomic conclusions of
this paper. To supplement, we compiled a mul-
tigene data set for the Antrodiella clade (genera
Antella, Antrodiella, Butyrea, and Metuloidea).
It included nrDNA ITS (24 seq.), nrDNA LSU
(17 seq.), mtDNA atp6 (6 seq.), and mtDNA

158 Miettinen & Ryvarden • ANN. BOT. FENNICI Vol. 53
Small Subunit, mtSSU (9 seq.). The sequences
derive from Vampola and Vlasák (2011), Miet-
tinen et al. (2012), Yuan (2013), and Kout et al.
(2014). In addition four ITS sequences with Gen-
Bank accession numbers KU926963–KU926966
were produced for this study with the methods
described in Miettinen et al. (2012).
Phylogenetic analyses were conducted with
MrBayes 3.2.2 (Ronquist et al. 2012). The data
set was partitioned into four subsets: ITS1 +
ITS2 (454 bp), 5.8S + LSU (912 bp), atp6 (685
bp) and mtSSU (559 bp). Partition models were
derived from BIC of JModelTest 2.1.6 (Darriba
et al. 2012): ITS1 + ITS2 (model HKY + G),
5.8S + LSU (model K80 + I), atp6 (model F81
+ I), mtSSU (model HKY + I). We conducted
three parallel runs of 8 chains each with 4 million
generations. Average standard deviation of split
frequencies went below 0.01 well before the 25%
burn-in. Also ML analysis with GTR + G settings
were done in RAxML 8.1.3 (Stamatakis 2014):
100 parallel runs to nd the best tree, and 1000
bootstrap replicas. Results are practically identi-
cal with MrBayes and are not repeated here.
In microscopic studies the basic mountant
was Cotton Blue (CB) made in lactic acid,
but also Melzer’s reagent (IKI), 5% KOH, and
Cresyl Blue (CRB) were used. In the micro-
scopic descriptions, the following abbreviations
are used: L = mean spore length, W = mean spore
width, Q = L/W ratio. Entry CB+ means cyano-
phily, CB– acyanophily; IKI– means the lack of
amyloid and dextrinoid reaction. While reporting
pore and spore measurements, the whole range is
given in parentheses; 90% range excluding 5%
extreme values from both ends of variation is
given without parentheses; in case the values are
identical, parentheses are omitted. For hyphal
width measurements, the 20% tails are in paren-
theses. For more details on microscopic method-
ology, see Miettinen et al. (2006).
Results
Our revised phylogeny of the Antrodiella clade
resolves four well-supported clades that support
the recognition of Antrodiella s. stricto, revised
Metuloidea, and the new genera Antella and
Butyrea (Fig. 1). It would appear that Antella
and Antrodiella are sister clades in this analysis,
but the signal from genetic loci is contradictory
— ITS–LSU data alone, for instance, would not
place the two as sister clades.
We see two possibilities for a phyloge-
ny-based genus classication within the Antro-
diella clade: either one big genus (which would
be called Metuloidea) or four genera as delin-
eated here. We have opted for the latter, mostly
because one genus would be morphologically
so heterogeneous that it would not be practical.
In contrast, the four genera can be dened mor-
phologically. This solution also minimizes new
combinations needed, since Metuloidea, which
formerly included only one species, is the oldest
genus name in the clade.
Taxonomy
Antella Miettinen, gen. nov.
MB 816250. — Type species: Antella niemelaei (Vampola
& Vlasák) Miettinen, comb. nova. — Basionym: Antrodiella
niemelaei Vampola & Vlasák, Czech Mycol. 63: 197. 2011.
MB 816251.
eTymology: Antella is a construct of letters from the
genus name Antrodiella.
Antella americana (Ryvarden & Gilb.) Miettinen, comb.
nova. — Basionym: Antrodiella americana Ryvarden &
Gilb., Mycotaxon 19: 138. 1984. MB 816252.
Antella chinensis (H.S. Yuan) Miettinen comb. nova. —
Basionym: Antrodiella chinensis H.S. Yuan, Mycol. Progr.
12: 439. 2013. MB 816253.
Resupinate, light-colored polypores with
middle- to large-sized pores (1–4 per mm) that
split to form irpicoid, yellowish hymenophore
when old. Hyphal system dimitic throughout,
clamps present, weakly to moderately cyanophi-
lous, skeletal hyphae yellowish in masses, rather
narrow (2–3.5 µm in diam) with a distinct lumen
of 1/3–1/6 of total hyphal width. Gloeocystidia
thin-walled, of variable sizes, well-differentiated
and easy to recognize. Spores ellipsoid, thin-
walled, small (below 4.5 ¥ 2.5 µm).
Ryvarden and Melo (2014) provide an excel-
lent microscopic drawing of A. niemelaei by
Tuomo Niemelä. For the photographs, see also
Yuan (2013) and Kout et al. (2014). The genus
resembles Butyrea with its rather cyanophilous
skeletal hyphae and gloeocystidia; differences

ANN. BOT. FENNICI Vol. 53 • Polypore genera Antella, Austeria, Butyrea, Citripora, Metuloidea and Trulla 159
include encrusted skeletocystidia and cylindrical
spore shape of Butyrea (Table 1).
Species of Antrodiella s. stricto are also quite
similar, but do not possess as well-differentiated
gloeocystidia. They have tougher consistency,
wood- or bone-like when cut with a razor blade
when dry, as opposed to board-like in Antella.
Pores are typically smaller, over 5 per millime-
tre. Skeletal hyphae in mature Antrodiella s.
stricto have usually an indistinct lumen, and they
are less cyanophilous.
Phylogenetic difference of Antella from Antr-
odiella s. stricto has been recognized earlier
(Johannesson et al. 2000, Miettinen et al. 2012,
Yuan 2013). Johannesson et al. (2000) pointed
to morphological differences between the core
Antrodiella and A. americana. Sequencing
results by Vampola and Vlasák (2011) indicate
that there may be two Antella species present in
North America.
Austeria Miettinen, gen. nov. (Fig. 2)
MB 816254. — Type species: Austeria citrea (Berk.) Miet-
tinen comb. nova. — Basionym: Polyporus citreus Berk., J.
Linn. Soc., Bot. 13: 162. 1873. MB 816255.
eTymology: Auster (Lat.), southern.
Polypore genus with one species: pileate
small basidiocarps with sulfur-yellow colors and
tiny pores (8–12 per mm). Hyphal structure dim-
itic throughout, clamps present, weakly cyano-
philous, skeletal hyphae narrower in trama (2.8–
3.5 µm in diam.), wider in context (3–5 µm),
with distinct lumen, 1/3–3/5 of the hyphal width,
generative hyphae thin- to thick-walled, in older
parts generative hyphae with short side branches
or twisted and swollen (2.8–4 µm). Intermediary
hyphae present particularly in context and differ-
ence between generative and skeletal hyphae not
always clear. Spores subcylindrical, thin-walled,
2.6–3.1 ¥ 1.5–1.9 µm.
Fig. 1. Phylogenetic Bayesian consensus tree of the Antrodiella clade based on nrDNA ITS and LSU, mtSSU, and
atp6. Numbers on nodes denote posterior probabilities (0–1) and bootstrap support values (50–100) derived from
maximum likelihood analysis.

160 Miettinen & Ryvarden • ANN. BOT. FENNICI Vol. 53
Table 1. Morphological comparison of selected polypore genera in Steccherinaceae.
Genus Basidiocarp Hyphal system Generative hyphae Skeletal hyphae Encrusted Gloeocystidia Spore shape Other
thick-walled
form color cystidia
Antella resupinate white to dimitic throughout thin-walled, narrow narrow, regular, – conspicuous, in ellipsoid CB+ stronger than
yellow lumen distinct tube bottoms Antrodiella
Antrodiella resupinate to white to dimitic throughout thin-walled, narrow narrow, regular, – absent, but rarely ellipsoid, bone hard when dry
s. stricto pileate yellow lumen mostly small, inconspicuous cylindrical
indistinct cystidia present
Austeria pileate sulfur- dimitic throughout in context slightly regular, lumen – – subcylindrical
yellow thick-walled, distinct
(partly) intermediary to
skeletals; partly
sinuous, swollen,
short-celled
Butyrea resupinate straw- dimitic throughout thin-walled, narrow narrow, regular, present conspicuous, cylindrical CB+ stronger than
colored lumen distinct large Antrodiella
Citripora pileate citric- indistinctly dimitic, in context wide, winding, branching – – ellipsoid
yellow upper context thick-walled in subiculum, with
nearly monomitic only slightly to
moderately thick walls
Flaviporus pileate white, dimitic to indistinctly thin- to slightly rather wide (often 4–5 µm), present or – ellipsoid thin- to thick-walled
reddish dimitic, trama thick-walled, lumen distinct; absent cystidia of tramal
±monomitic, very narrow intermediary hyphae origin bending into
dense to agglutinated hymenium
Junghuhnia resupinate whitish dimitic thin-walled, narrow narrow, regular, abundant, – ellipsoid CB(+) to CB–,
lumen rather distinct short, coarsely cystidia arranged
encrusted irregularly
Metuloidea pileate brown dimitic throughout, thin-walled, narrow regular, rather wide, indistinct – ellipsoid, sweet smell; also
approaching to slightly thick-walled lumen distinct, to well subcylindrical hydnoid species
trimitic and moderately wide branching commonly differentiated
Steccherinum resupinate to white, dimitic throughout thin-walled, narrow mostly narrow, present – ellipsoid, also hydnoid
pileate pinkish, regular, lumen cylindrical species
reddish distinct
brown
Trulla pileate to light trama dimitic, in context wide, regular, lumen – – cylindrical
semistipitate yellowish, context nearly thick-walled distinct curved
brownish monomitic

ANN. BOT. FENNICI Vol. 53 • Polypore genera Antella, Austeria, Butyrea, Citripora, Metuloidea and Trulla 161
Buchanan and Ryvarden (2000a) report that
material previously determined as A. semisu-
pina represents in fact A. citrea. Their checklist
(Buchanan & Ryvarden 2000b) includes three
other species of Antrodiella (A. hunua, A. rata,
A. zonata), none of which belongs to Antrodiella
s. stricto. Thus Antrodiella in the strict sense has
not been reported from New Zealand, and we
have not seen any material from Australia either.
Butyrea Miettinen, gen. nov. (Fig. 3)
MB 816256. — Type species: Butyrea luteoalba (P. Karst.)
Miettinen comb. nova. — Basionym: Physisporus luteoalbus
P. Karst., Rev. Mycol. (Toulouse) 9: 10. 1887. MB 816257.
eTymology: Butyrum (Lat.), butter, a reference to the
gloeocystidia characteristic of the genus.
Butyrea japonica (Nuñez & Ryvarden) Miettinen &
Ryvarden comb. nova. — Basionym: Junghuhnia japonica
A photograph showing the characteristic col-
oration of the genus can be found in Gates and
Ratkowsky (2014). The closest relatives of A.
citrea are pileate hydnoid Mycorrhaphium spp.
The outer appearance and several microscopic
characters of A. citrea remind Antrodiella s.
stricto: small, tough basidiocarps, white, small
pores, dimitic hyphal structure, lack of cystidia,
and tiny spores. However, all species of Antro-
diella s. stricto have clearly delimited, always
dominant skeletal hyphae (mostly with indis-
tinct lumen) and thin-walled, indistinct gener-
ative hyphae. In contrast generative hyphae of
A. citrea are partly thick-walled, branching in a
characteristic manner, and in younger basidio-
carps dominating in context/subiculum. Inter-
mediary hyphae between skeletal and generative
hyphae occur frequently in Austeria, unlike in
Antrodiella s. stricto.
Fig. 2. Hyphal structures of Austeria citrea (Paulus 3946). — a: Tube trama. — b: Subiculum. — c: Hymenial cells.
— d: Spores of the holotype.

162 Miettinen & Ryvarden • ANN. BOT. FENNICI Vol. 53
Núñez & Ryvarden, Fungal Diversity 3: 112. 1999. MB
816258.
Polypores with annual, yellowish, resupinate
basidiocarps and rather small pores (4–8 per
mm). Hyphal system dimitic, with clamps, skel-
etal hyphae moderately cyanophilous, narrow
and sinuous, mostly < 2.5 µm in width, with a
distinct lumen 1/3–1/2 of the width. Two types
of cystidia present, thin-walled gloeocystidia in
hymenium and tube bottoms in particular, and
thick-walled, club-shaped encrusted cystidia
in trama (encrusted part 25–50 ¥ 10–18 µm).
Spores cylindrical, straight, thin-walled, 4–5 ¥
1.7–2.3 µm.
Butyrea is closely related to Antrodiella in
the strict sense as well as to Metuloidea (Fig. 1;
Miettinen et al. 2012). Species of Antrodiella do
not form cystidia. Metuloidea species do pro-
duce encrusted cystidia, but their pileate, brown-
ish basidiocarps with a distinct, sweet smell are
a clear separating character from Butyrea spp.
Morphologically the closest match are poroid
Steccherinum spp. with similar basidiocarps,
hyphal structure and encrusted skeletocystidia.
None of the species in Steccherinum species
possess gloeocystidia. With the exception of the
somewhat deviating species S. collabens, poroid
Steccherinum have ellipsoid spores and their
cyanophilous reaction is weaker.
Butyrea cystidia seem to develop encrus-
tation on top of a thin-walled appendix of the
thick-walled cystidial base (Fig. 3c). If cut when
young, it looks like crystal bits are sitting on a gel
or detached from any hyphal walls. When older,
they start to remind more typical, clearly thick-
walled, heavily encrusted Steccherinum cystidia.
Butyrea skeletocystidia are not as persistent as in
Steccherinum and are typically unevenly present
or possibly even absent from young collections.
Already Pouzar (1967) suggested that the
strong cyanophilic reaction is a basis of dividing
the genus Chaetoporus (corresponding to poroid
Steccherinum spp. of today in Pouzar’s concept)
in two sections, one of which, Cyanochaetopo-
rus with strong reaction included two species,
Butyrea luteoalba and Steccherinum collabens.
As noted above, the latter species has an excep-
tionally strong cyanophilic reaction among Stec-
cherinum and also deviating, curved cylindrical
Fig. 3. Butyrea luteoalba. — a: Hymenium with gloeocystidia and an encrusted skeletocystidium (Miettinen 7610).
— b: Gloeocystidia in a young basidiocarp (lectotype of Physisporus avicans. — Butyrea japonica (holotype). — c:
A gloeocystidium and a young, naked skeletocystidium. — d: Spores.
Butyrea luteoalba Butyrea japonica

ANN. BOT. FENNICI Vol. 53 • Polypore genera Antella, Austeria, Butyrea, Citripora, Metuloidea and Trulla 163
spores, so Pouzar’s conclusion is understandable.
The two are, however, not closely related (Miet-
tinen et al. 2012), nor does S. collabens possess
gloeocystidia in any stage of development.
For now, we include two species in Butyrea:
B. japonica and B. luteoalba. Butyrea japonica
differs genetically from the North European B.
luteoalba (described from Finland) by 25 bp
(> 4%) in ITS. Nuñez and Ryvarden (1999)
separated the two morphologically by spore size
and presence of gloeocystidia in B. japonica.
The latter character does not separate the two
since both species bear gloeocystidia at least
when young (noted also by Vampola 2009). We
can conrm the spore size difference: W < 2 µm
in B. luteoalba and W > 2 µm in B. japonica.
The type of Junghuhnia japonica was collected
on Castanopsis sp. (Fagaceae) whereas Butyrea
luteoalba inhabits conifer wood.
Several epithets have been used for B. luteo-
alba in the past, among them Physisporus varie-
color P. Karst. 1881, which would be an older
name than P. luteoalbus P. Karst. 1887. Lowe
(1956) designated the lectotype for P. luteoalbus,
and it agrees well with the current concept of
Butyrea luteoalba. However, we have not been
able to nd lectotypication for P. variecolor¸
and therefore we select the following speci-
men as the lectotype (designated here): Finland.
Etelä-Häme, Tammela, Mustiala, on Alnus gluti-
nosa, Aug. 1880 P. A. Karsten s.n. (de Thümen:
Mycotheca universalis 1803, H 6034043). The
specimen is a typical member of Trechispora
hymenocystis.
We also studied the type of Physisporus
avicans P. Karst. 1986, which Lowe (1956)
considered, correctly, as a synonym of P. luteo-
albus. Lowe indicates that there is only one col-
lection, which can be considered to be the type,
but fails to clearly indicate which one, so we
designate here a lectotype for P. avicans: Fin-
land. Etelä-Häme, Tammela, Mustiala, Syrjä, on
Pinus, Aug.1895 P. A. Karsten s.n. (H 6004907).
Citripora Miettinen gen. nov. (Figs. 4 and 5)
MB 816259. — Type species: Citripora bannaensis Miettinen.
eTymology: Loosely refers to color of lemon (citrus,
Lat.).
Polypores with half-resupinate to pile-
ate, small-pored (5–11 per mm) basidiocarps
with lemon-yellow tints. Hyphal system dim-
itic, clamps present; lower tube trama appears
monomitic and subiculum dominated by gen-
erative hyphae or nearly monomitic. Skeletal
hyphae winding, somewhat irregular in outline,
3–4 µm in diameter, with a wide lumen, in
subiculum branched if present. Spores broadly
ellipsoid to subglobose, thin-walled, tiny (2.5 ¥
2 µm), with a shiny droplet inside.
The genus is easy to recognize by the com-
bination of color, hyphal structure with winding,
irregular skeletal hyphae with a wide lumen, and
tiny ellipsoid spores. It clearly belongs to Stec-
cherinaceae, but has no close relatives within
the family (Miettinen et al. 2012). Antrodiella s.
stricto includes one species with lemon-colored
basidiocarps, A. citrinella, but that species has a
typical Antrodiella hyphal structure: stiff, regular
skeletal hyphae with a narrow lumen dominating
throughout the basidiocarp. Citripora species
could be mixed with Tyromyces spp., but the
relatively thin basidiocarps, winding, irregular
skeletal hyphae and citric color should tell them
apart.
Citripora afrocitrina (Ipulet & Ryvarden)
Miettinen & Ryvarden, comb. nova
MB 816260. — Basionym: Antrodiella afrocitrina Ipulet &
Ryvarden, Syn. Fungorum 20: 88. 2005.
Ipulet and Ryvarden (2005) described Antro-
diella afrocitrina from Uganda. Their description
can be improved in some ways: the spores are not
globose but broad ellipsoid, the skeletal hyphae
in subiculum are often branched and the lumen of
the hyphae is wide, not “unbranched, thick-walled
to almost solid”, when studied in CB or Meltzer
(though KOH swells the walls inwards).
Citripora bannaensis Miettinen, sp. nova
(Figs. 4 and 5)
MB 816261. — Type: China. Yunnan: Xishuangbanna Bio-
sphere Reserve, 21.92°N, 101.19°E, decayed dicot log, 5
Aug. 2005 O. Miettinen 9999 (H 7008295 holotype, BJFC
isotype). ITS seq: JN710526.

164 Miettinen & Ryvarden • ANN. BOT. FENNICI Vol. 53
Fig. 5. Citripora bannaensis (holotype). — a: Context hyphae. — b: Tramal hyphae. — c: Hymenial cells. — Citri-
pora afrocitrina (Decock Ug08–33). — d: Subicular hyphae. — e: Spores of both species, drawn from holotypes.
Fig. 4. Citripora bannaen-
sis, fresh holotype.
Citripora afrocitrina
Citripora bannaensis

ANN. BOT. FENNICI Vol. 53 • Polypore genera Antella, Austeria, Butyrea, Citripora, Metuloidea and Trulla 165
eTymology: Refers to the area of collecting, Xishuang-
banna in Yunnan, China.
Basidiocarp pileate, bright yellow from
above, yellowish cream-colored below, caps pro-
jecting 2–5 cm, 2–10 cm wide, 1–4 mm thick.
Consistency rather tough but exible when fresh,
drying light-weight and rigid, but rather easy to
break apart. Pores regular, thin-walled, rounded
angular, mouths with hyphal tufts but other-
wise smooth, 8–11(13) per millimetre, 1–3 mm.
Subiculum white, lighter than tube layer, 0.2–1
mm thick, in parts with an agglutinated thin layer
visible under the lens. Cap margin sharp.
Hyphal system indistinctly dimitic, most
clearly in tube trama, also intermediary hyphae
present, CB(+), IKI–, KOH–, CRB lilac; clamps
present, but simple septa not rare either. Con-
text varying from nearly monomitic (particularly
upper part) to dimitic, mostly dominated by
loosely interwoven, slightly thick-walled (walls
up to 1 µm) generative hyphae, (2.6)3.4–4.2(4.9)
µm in diameter; single skeletal hyphae present
even then and distinction to generative hyphae
not clear. Tramal hyphae interwoven, dominated
by interwoven, sinuous skeletals with only mod-
erately thickened walls (0.3–1 µm) and a very
wide, distinct lumen, (2.0)3.2–4.3(4.8) µm in
diameter; tramal generative hyphae thin-walled,
(2.0)2.5–3.5(5.5) µm in diameter. Oily substance
present in dissections. Cyanophilic reaction,
though not strong, is stronger than in Antrodiella
s. stricto.
Hymenium very regular, small-celled pal-
isade. Basidia clavate to barrel-shaped, 7–9 ¥
3.8–4.4 µm, with 4 sterigmata, 1.8–2.2 µm.
Basidiospores ellipsoid, thin-walled, with a
shiny droplet inside, 2.2–2.8(3.0) ¥ 1.7–2.0(2.1)
µm, L = 2.49 µm, W = 1.86 µm, Q´ = 1.2–
1.5(1.6), Q = 1.34, n = 30.
DisTriBuTion: China (southern Yunnan, foot-
hills of the Himalayas).
ecology: The type collection was found on
a decayed dicot log, 40 cm in diameter (decay
stage 4/5) in a subtropical seasonal forest (at
least Lauraceae, Fabaceae, Annonaceae, Sapin-
daceae).
The two Citripora species are very similar
under the microscope, and the best character
to separate them is basidiocarp morphology: in
C. afrocitrina basidiocarps are effused-reexed,
whereas they are pileate in C. bannaensis. Pores
of C. bannaensis are distinctly smaller, 8–11 per
millimetre vs. 5–7 in C. afrocitrina.
Metuloidea G. Cunn. (Fig. 6)
Bull. New Zealand Dept. Sci. Industr. Res., Pl. Dis. Div. 164:
250, 263. 1965. — Type species: Metuloidea tawa (G. Cunn.)
G. Cunn. 1965. — Basionym: Trametes tawa G. Cunn. 1948.
Metuloidea cinnamomea (Iturr. & Ryvarden) Miettinen
& Ryvarden, comb. nova. — Basionym: Antrodiella cinna-
momea Iturr. & Ryvarden, Syn. Fungorum 27: 78. 2010. MB
816262.
Metuloidea fragrans (A. David & Tortič) Miettinen,
comb. nova. — Basionym: Trametes fragrans A. David &
Tortič, Acta Bot. Croat. 38: 133. 1979. MB 816263.
Metuloidea murashkinskyi (Burt) Miettinen & Spirin,
comb. nova. — Basionym: Hydnum murashkinskyi Burt,
Ann. Missouri Bot. Garden 18: 477. 1931. MB 816264.
Metuloidea rhinocephala (Berk.) Miettinen, comb nova.
— Basionym: Polyporus rhinocephalus Berk. in Hooker,
Bot. Antarct. Voy., III, Fl. Tasman. 2: 253. 1859 [1860]. MB
816265.
Polypores and hydnoid fungi with pile-
ate, brownish basidiocarps with a sweet smell.
Hyphal system dimitic throughout, skeletal
hyphae yellowish-brownish, weakly to distinctly
cyanophilous, with an easily visible lumen and
frequent branching at least in context, relatively
wide, (2)3–5 µm in diameter. Encrusted hyphal
ends or differentiated, encrusted skeletocystidia
present in tube mouths. Spores ellipsoid to short
cylidrical, small (3–4 ¥ 2–2.8 µm), thin-walled.
Photographs showing the characteristic col-
oration of the genus can be found in Miettinen et
al. (2012) and Ryvarden and Melo (2014).
Branched skeletal hyphae are common in
Metuloidea, and in some species, e.g. context of
M. murashkinskyi, the branched skeletal hyphae
(binding hyphae), can be distinguished from
straight, unbranched, wider skeletal hyphae. The
difference between binding and skeletal types is
not as clear in other species, but hyphal system
in all Metuloidea spp. approaches trimitic. This
was also the conclusion of David and Tortič
(1979) for M. fragrans: “structura dimitica, ad
trimiticam vergens”.
Presence of cystidia in Metuloidea spp. is
somewhat ambiguous. They can be identied

166 Miettinen & Ryvarden • ANN. BOT. FENNICI Vol. 53
without hesitation in old basidiocarps of M.
murashkinskyi, but are hardly differentiated
in younger basidiocarps of M. fragrans and
M. rhinocephala, being just slightly widened
and lightly encrusted ends of skeletal hyphae
(Fig. 6). David and Tortič (1979) also provide a
good illustration of such a structure. It is a matter
of taste if one calls them poorly differentiated
cystidia or just encrusted hyphal ends.
The sweet smell so typical for M. fragrans
has not been reported for all species in the genus.
This is probably just an oversight; Spirin et
al. (2007) noted that Metuloidea murashkinskyi
has the distinct pleasant smell although no one
reported it before. Ryvarden and Iturriaga (2010)
do not comment on the smell of M. cinnamo-
mea, and the specimen does not smell anymore.
Recent herbarium specimens of M. rhinocephala
still have a strong smell.
Metuloidea resembles morphologically most
closely Steccherinum, which are generally not
brown-colored and have no sweet smell. Stec-
cherinum skeletal hyphae are typically narrower
with a narrower lumen, very rarely branched,
and their skeletocystidia are more distinct. The
closest relative phylogenetically, Butyrea spp.,
Fig. 6. Metuloidea cinnamomea (holotype). — a: Tramal hyphae, encrusted hyphae cystidia-like. — b: Hymenial
cells. — Metuloidea murashkinskyi (Spirin 2367). — c: Tramal skeletocystidia and subicular hyphae. — d: Spores
of Metuloidea spp. from holotypes, except M. murashkinskyi (Spirin 2367) and M. rhinocephala (Gates 13 Mar.
2007).
Metuloidea cinnamomea
Metuloidea cinnamomea
Metuloidea murashkinskyi
Metuloidea murashkinskyi
Metuloidea fragrans
Metuloidea rhinocephala
Metuloidea tawa

ANN. BOT. FENNICI Vol. 53 • Polypore genera Antella, Austeria, Butyrea, Citripora, Metuloidea and Trulla 167
have very different, thin effused basidiocarps
without unusual odor, and they produce gloeo-
cystidia and well differentiated skeletocystidia
unlike any of the poroid Metuloidea.
We include four species in Metuloidea for
now, but the number will rise in the future. It
is not fully clear whether M. tawa from New
Zealand is conspecic with M. rhinocephala
from Tasmania. Since spores are identical and
ITS sequence differences between two Tasma-
nian and one New Zealand specimen only 3 bp,
we consider them conspecic for now. Metu-
loidea rhinocephala closely resembles M. fra-
grans from Europe, but spore size (Table 2) and
genetic differences (10 bp in ITS) indicate that
the European and Austral species are separate.
Metuloidea is a somewhat exceptional genus
in containing both clearly hydnoid and poroid
species. Interestingly young developing hymeno-
phores of the hydnoid M. murashkinskyi look
like irregular pore surface, which only later turns
distinctly hydnoid (Spirin et al. 2007). It is likely
that we’re seeing a shift from a poroid to hyd-
noid hymenophore within the genus, since basal
species in the genus are polypores, and so are
the closest relatives of Metuloidea in the genera
Antrodiella and Butyrea. Miettinen et al. (2012)
stated that splitting the genus into monophyletic
hydnoid and poroid parts “seems possible”. Our
results contradict this, since the poroid M. cin-
namomea appears to be more closely related to
hydnoid than to other poroid Metuloidea species
(Fig. 1).
Key to the species of Metuloidea
1. Hymenophore distinctly hydnoid or irpicoid ..................
............................................................ M. murashkinskyi
1. Hymenophore poroid ................................................... 2
2. Spores subcylindrical, mostly < 2 µm wide ....................
.............................................................. M. cinnamomea
2. Spores ellipsoid, > 2 µm wide .................................... 3.
3. European species, W < 2.5 µm, Q < 1.45 ... M. fragrans
3. Southern Australia and New Zealand, W > 2.5 µm, Q >
1.45 ............................. M. rhinocephala (incl. M. tawa)
Table 2. Spore measurements of Austeria, Metuloidea and Trulla species. L = average length, W = average width,
Q´ = length/width of individual spores, Q = L/W. Metuloidea rhinocephala compound statistics are based on Tasma-
nian specimens only.
Species/specimen Length L Width W Q´ Q n
Austeria citrea 2.6–3.1(–3.5) 2.84 1.5–1.9(–2.0) 1.70 1.5–1.9 1.67 90/3
holotype 2.7–3.1(–3.3) 2.91 (1.6–)1.7–2.0 1.81 1.5–1.8 1.61 30
Coveny 52/1982 (2.6–)2.7–3.2(–3.5) 2.90 1.6–1.8(–1.9) 1.69 1.6–1.9 1.71 30
Ryvarden 37392 2.6–2.9 2.73 1.5–1.7 1.60 1.5–1.9 1.71 30
Metuloidea cinnamomea (2.7–)2.8–3.2(–4.2) 3.08 1.8–2.0(–2.1) 1.89 (1.4–)1.5–1.8(–2.0) 1.62 30
Metuloidea fragrans (2.8–)2.9–3.8(–4.0) 3.29 (2.0–)2.1–2.8(–3.0) 2.40 (1.1–)1.2–1.5(–1.8) 1.37 110/3
holotype (2.8–)2.9–3.7(–3.8) 3.18 (2.0–)2.1–2.8 2.38 (1.1–)1.2–1.4(–1.5) 1.34 70
Hausknecht 20 Sep. 1996 2.8–3.6 3.34 2.0–2.7 2.30 1.2–1.8 1.45 10
Spirin 2031 3.1–3.9(–4.0) 3.52 2.3–2.8(–3.0) 2.49 (1.2–)1.3–1.6 1.41 30
Metuloidea murashkinskyi (3.1–)3.2–4.3(–4.5) 3.59 1.8–2.1(–2.2) 1.96 (1.5–)1.6–2.1(–2.2) 1.84 61/2
Spirin 2388 3.2–4.2(–4.3) 3.61 1.8–2.1 1.94 (1.6–)1.7–2.2 1.86 30
Spirin 2367 3.1–4.4(–4.5) 3.57 1.8–2.2 1.98 (1.5–)1.6–2.1(–2.2) 1.81 31
Metuloidea rhinocephala (3.4–)3.5–4.5(–4.9) 3.80 (2.2–)2.3–2.8(–3.1) 2.55 (1.3–)1.4–1.7(–1.8) 1.49 70/2
Gates 13 Mar. 2007 3.4–4.2(–4.5) 3.79 (2.2–)2.3–2.8(–3.1) 2.59 (1.3–)1.4–1.6 1.46 30
Gates 24 May 2007 3.5–4.5(–4.9) 3.82 (2.2–)2.3–2.7(–2.8) 2.52 1.4–1.8 1.52 40
Ryvarden 39981 3.8–4.8 4.23 (2.4–)2.5–2.9(–3.0) 2.69 (1.4–)1.5–1.8 1.57 30
Trulla crustulina (3.4–)3.6–4.5(–4.6) 4.00 1.4–1.8 1.61 (2.1–)2.2–2.9 2.48 30
Trulla dentipora 4.3–5.2 4.70 (1.4–)1.5–1.8 1.59 (2.5–)2.6–3.3 2.96 30
Trulla duracina (4.2–)4.3–5.3(–5.4) 4.77 1.6–1.9(–2.0) 1.76 2.4–3.0(–3.2) 2.71 39
Trulla meridae (3.7–)3.8–4.4(–4.8) 4.09 (1.6–)1.8–2.0(–2.1) 1.89 (1.9–)2.0–2.4(–2.6) 2.16 63/2
Ryvarden 43589 3.7–4.3(–4.4) 4.03 (1.6–)1.8–2.0 1.87 1.9–2.3(–2.6) 2.16 31
Ryvarden 43619 (3.7–)3.8–4.4(–4.8) 4.15 1.8–2.1 1.91 2.0–2.4 2.17 32
Trulla ochrotinctella 4.0–4.7(–4.8) 4.25 1.4–1.7 1.55 2.5–3.0(–3.1) 2.73 31
Trulla polyporoides (4.3–)4.4–5.2 4.66 1.3–1.5 1.39 3.0–3.8(–4.0) 3.35 30

168 Miettinen & Ryvarden • ANN. BOT. FENNICI Vol. 53
Steccherinum Gray
Nat. Arr. Brit. Pl. 1: 651. 1821.
Steccherinum autumnale (Spirin, Zmitr. & Malysheva)
Miettinen comb. nova. — Basionym: Junghuhnia autumnale
Spirin, Zmitrovitch & Malysheva, Ann. Bot. Fennici 44: 303.
2007. MB 816266.
Steccherinum mbriatellum (Peck) Miettinen, comb.
nova. — Basionym: Polyporus mbriatellus Peck, Rep.
(Annual) New York State Mus. Nat. Hist. 38: 91. 1885. MB
816267.
Steccherinum formosanum (T.T. Chang & W.N. Chou)
Miettinen, comb. nova. — Basionym: Antrodiella formosana
T.T. Chang & W.N. Chou, Mycol. Res. 102: 400. 1998. MB
816268.
Steccherinum semisupiniforme (Murrill) Miettinen,
comb. nova. — Basionym: Tyromyces semisupiniformis Mur-
rill, Bull. New York Bot. Garden 8: 148. 1912. MB 816269.
Steccherinum subnitidum (H.S. Yuan & Y.C. Dai) Miet-
tinen, comb. nova. — Basionym: Junghuhnia subnitida H.S.
Yuan & Y.C. Dai, Sydowia 60: 148. 2008. MB 816270.
Miettinen et al. (2012) concluded that Stec-
cherinum contains both polypores and hydnoid
fungi. Most species in their Steccherinum clade
are dimitic species with encrusted skeletocystidia
and clamped septa, but others are highly deviat-
ing such as monomitic, simple-septate species
of “Candelabrochaete” and “Odonticium”. For
now we are stuck with a wide concept of Stec-
cherinum, so below we provide a few “practical”
combinations based on phylogenetic results of
Miettinen et al. (2012) and Yuan and Dai (2008).
Trulla Miettinen & Ryvarden, gen. nov.
(Figs. 7 and 8)
MB 816271. — Type species: Trulla dentipora (Ryvarden
& Iturr.) Miettinen & Ryvarden, comb. nova. — Basionym:
Antrodiella dentipora Ryvarden & Iturr., Mycologia 95:
1066. 2003. MB 816272.
eTymology: Trulla (Lat.), ladle, refers to the spatulate
shape of the basidiocarps.
Trulla crustulina (Bres.) Miettinen, comb. nova. —
Basionym: Polyporus crustulinus Bres., Hedwigia 56: 293.
1915. MB 816273.
Trulla duracina (Pat.) Miettinen, comb. nova. — Basi-
onym: Leptoporus duracinus Pat., Bull. Soc. Mycol. France
18: 174. 1902. MB 816274.
Trulla meridae Miettinen & Ryvarden (see below).
Trulla ochrotinctella (Murrill) Miettinen, comb. nova.
— Basionym: Coriolus ochrotinctellus Murrill, North Ameri-
can Flora 9 (1): 22. 1907. MB 816275.
Trulla polyporoides (Ryvarden & Iturr.) Miettinen &
Ryvarden, comb. nova. — Basionym: Tyromyces polyporoi-
des Ryvarden & Iturr., Mycologia 95: 1075. 2003. MB
816276.
Basidiomes pileate or laterally stipitate, com-
monly spathulate, upper surface smooth, shiny,
usually light brownish with colored zones, light-
Fig. 7. Trulla meridae
(holotype and Ryvarden
43619 (the largest basidi-
ocarp) photographed from
herbarium specimens).
Scale = 2 cm.

ANN. BOT. FENNICI Vol. 53 • Polypore genera Antella, Austeria, Butyrea, Citripora, Metuloidea and Trulla 169
weight and hard when dry but easy to crack into
pieces. Context white to cream, mostly lighter
than upper surface and relatively thick. Pores
small and regular (5–12 per mm) in all except
one species (2–4 per mm). Hyphal structure
dimitic, CB+, generative hyphae with clamps.
Context monomitic or nearly so, dominated by
slightly thick-walled generative hyphae with fre-
quent clamps and branching (3–4.5 µm in diam-
eter). Trama dominated by interwoven skeletal
hyphae (2.5–3.8 µm) with a well-visible lumen
of 1/2 to 1/6 of hyphal width, generative hyphae
narrow (1.8–3 µm); some intermediary hyphae
between skeletal and generative hyphae occur.
Hymenial cells thin-walled and short, 7–10 µm
long. Spores thin-walled, cylindrical and curved,
with a tiny or non-existent apiculus, 3.5–5.5 ¥
1.3–2 µm.
Fig. 8. Trulla meridae (holotype). — a: Cap context. — b: Tube trama. — c: Hymenial cells. — d: Spores of Trulla
spp. from holo/lectotypes.
Trulla crustulina
Trulla dentipora
Trulla duracina
Trulla meridae
Trulla ochrotinctella
Trulla polyporoides

170 Miettinen & Ryvarden • ANN. BOT. FENNICI Vol. 53
This genus is characterized foremost by its
monomitic context contrasting to dimitic trama.
Curved cylindrical spores, cyanophilic skeletal
hyphae and slightly thick-walled, wide gener-
ative hyphae in context are further characters
typical to the genus. It differs from its closest
relative Nigroporus by its light-colored basidi-
omes and monomitic context (dimitic in Nigro-
porus spp.). Antrodiella spp. are dimitic through-
out, and hyphal structure in Flaviporus is much
tighter.
Most species of Trulla are tropical, although
some extend to south temperate areas in Asia
and North America. Here, we have included six
species, but it is likely that T. duracina complex
comprises more species. Morphological differ-
ences are small, and sorting out species taxon-
omy in this complex requires abundant fresh
collections and DNA comparisons. For instance
separation of T. ochrotinctella from T. duracina
deserves to be conrmed with a more in-depth
study.
The type of Tyromyces elmeri Murrill, for-
merly considered a synonym of Trulla durac-
ina (Ryvarden 1985), was checked, and being
monomitic or nearly so, it does not belong to
Trulla.
Key to the species of Trulla
1. Pores mostly 2–4 per mm, with sinuous mouths ............
..................................................................... T. dentipora
1. Pores smaller than 5 per mm, mouths regular ............. 2
2. Spores ≤ 1.5 µm wide, narrow cylindrical, Q > 3 ...........
................................................................ T. polyporoides
2. Most spores ≥ 1.5 µm wide and not as narrow, Q < 3 .... 3
3. Pores 5–9 per mm, spores 4–5 µm long ....................... 4
3. Pores 9–12 per mm, spores < 4.5 µm long .................. 5
4. Pores 5–7 per mm, spores mostly < 4.5 µm long and
around 1.5 µm wide ............................. T. ochrotinctella
4. Pores 8–9 per mm, spores mostly > 4.5 µm long and >
1.5 µm wide ................................................. T. duracina
5. Most spores > 1.8 µm wide, strongly curved, American
species ........................................................... T. meridae
5. Most spores < 1.8 µm wide, only moderately curved, old
world species .............................................. T. crustulina
Trulla meridae Miettinen & Ryvarden, sp.
nova (Figs. 7 and 8)
MB 816277. — Type: Venezuela. Mérida: Zona de la Car-
bonera, around Bosque San Eusebio, 8.6°N, 71.3°W, alt.
2100 m, on dead hardwood, 31 Jan. 2001 L. Ryvarden 43589
(holotype O 917470; isotype H). ITS seq: JN710513.
eTymology: Refers to Estado de Mérida, the type locality.
Basidiocarp pileate, spathulate, upper surface
light brown with weak color zones, smooth and
a little shiny, pores cream-colored with pink-
ish-greyish tints, caps projecting 2–6 cm, equally
wide, caps 2–6 mm thick, thinning towards the
margin. Consistency light and hard when dry,
easy to crack to pieces with force. Pores regular,
round to angular, rather thin-walled, 10–12(–13)
per millimetre, pore layer thickness 0.3–2 mm.
Subiculum cream-colored, lighter than pores and
upper surface, up to 15 mm at base, 0.5–5 mm
in caps. Margin acute in young caps, blunt when
older, with a light-colored 1 mm border zone at
lower surface.
Hyphal system dimitic, clamps present. Cap
context monomitic, dominated by slightly thick-
walled, mostly horizontally oriented generative
hyphae that branch frequently, (2.3)3.3–4.2(5.4)
µm in diameter; a few skeletal hyphae in lower
context, 3.2–4.2 µm. Tube trama dimitic, hyphae
interwoven, dominated by winding skeletal
hyphae with a distinct lumen, (2.5)2.8–3.6(4.2)
µm; generative hyphae (1.6)2.0–3.0(3.7) µm.
Hyphal pegs common, mostly composed of
short-celled, rather wide hyphae.
Hymenium. Basidia clavate or barrel-like,
8–10 ¥ 4–5 µm, with 4 sterigmata of 2 µm in
length. Cystidioles absent.
Basidiospores cylindrical, curved, thin-walled,
(3.7)3.8–4.4(4.8) ¥ (1.6)1.8–2.0(2.1) µm, L =
4.09 µm, W = 1.89 µm, Q´ = (1.9)2.0–2.4(2.6), Q
= 2.16, n = 63/2.
DisTriBuTion: Northern Andes in Venezuela.
ecology: The known collections derive from
hardwood logs at 2100 m.
Member of the Trulla duracina complex, T.
meridae differs from T. duracina by its smaller
pores and strongly curved, shorter spores
(Table 2). This species was treated as Antro-
diella duracina by Miettinen et al. (2012).
Specimens studied (substrate given when
known)
Antella chinensis. Russia. Khabarovsk Reg., on Betula,
Spirin 6612 (H 7024229), on Acer ukurunduense, Spirin

ANN. BOT. FENNICI Vol. 53 • Polypore genera Antella, Austeria, Butyrea, Citripora, Metuloidea and Trulla 171
4095 (H 7028495). — Antella niemelaei. Finland. Uusimaa,
on Prunus padus + Hymenochaete tabacina, Niemelä 3223
(H, isotype). Russia. Khabarovsk Reg., on Acer ukurundu-
ense + Hymenochaete intricata, Spirin 6565 (H 7024114).
— Austeria citrea. Australia. New South Wales, Dr Bukler
(K(M) 147875, holotype); Coveny 52/1982 (O, H). Tasma-
nia, Gates 542 (O, H 18656). New Zealand. Northland,
Paulus 3946 (PDD 92144), Ryvarden 37392 (O, H). Auck-
land, Buchanan 89/044 (PDD 58375). — Butyrea japonica.
Japan. Kagoshima pref., on Castanopsis, Nuñez 1065 (O-F
450326, holotype). — Butyrea luteoalba. Finland. Uusimaa,
on Picea abies, Miettinen 14004 (H 6012624). Etelä-Häme,
on Picea abies, Miettinen 7610 (H); on Pinus sylvestris,
VIII.1895 Karsten (H 6004907, lectotype of Physisporus a-
vicans); on Pinus sylvestris, 13.X.1889 Karsten (H 6048899,
lectotype). — Citripora afrocitrina. Uganda. Kabarole,
Decock Ug08-33 (H). Kamwenge, 10.XII.2001 Ipulet (O,
holotype). — Citripora bannaensis. China. Yunnan (see
holotype). — Metuloidea cinnamomea. Venezuela. Mérida,
on hardwood log, Ryvarden 43626 (O, holotype). — Met-
uloidea fragrans. Austria. Steiermark, 20.IX.1996 Haus-
knecht (O, H). Russia. Nizhnyi Novgorod, on Corylus
avellana, Spirin 2031 (H). Slovenia. On Corylus avellana,
10.IX.1978 Tortić (CNF, holotype). — Metuloidea murash-
kinskyi. Russia. Tomsk, on Betula verrucosa, 1.IX.1928
Murashkinsky (PRM 156151, isotype). Nizhnyi Novgorod,
on Quercus robur, Spirin 2367, 2388 (H). — Metuloidea rhi-
nocephala. Australia. Tasmania (K(M) 147645, lectotype);
13.III., 24.V.2007 Gates (H). New Zealand. Wellington,
Ryvarden 40179 (O, H). Southland, X.1946 Rawlings (PDD
4822, holotype of Metuloidea tawa); Ryvarden 39981 (O, H).
— Steccherinum semisupiniforme. China. Yunnan, Miettinen
10026.1 (H). Mexico. Jalapa, 12–20.XII.1909 Murrill (NY
776468, type). — Trulla crustulina. Philippines. Luzon,
VII.1909 McGregor (S F5553, lectotype). — Trulla denti-
pora. Venezuela. Aragua, alt. 1000–1200 m, on hardwood,
Ryvarden 40822 (O 11156213, isotype). — Trulla duracina.
Guadeloupe. On Cecropia peltata (FH 292715, holotype).
— Trulla meridae. Venezuela. Mérida, on dead hardwood,
Ryvarden 43619 (O); see holotype. — Trulla ochrotinctella.
United States. Mississippi, on Quercus, 10.IX.1904 Earle
(NY 704975, 704976, lectotype of Coriolus ochrotinctel-
lus). — Trulla polyporoides. Venezuela. Bolívar, 24.II.2000
Iturriaga 7273/Ryvarden 42373 (O, isotype). — Tyromyces
elmeri. Philippines. Luzon, Elmer 6945 (NY 776451, lec-
totype).
Acknowledgements
We are indebted to Genevieve Gates (Hobart), Cony Decock
(Louvain), and Viacheslav Spirin (Helsinki) for providing us
valuable specimens. Yu-Cheng Dai and his students (Beijing)
are warmly acknowledged for arranging eld trips for the
authors in China, resulting in collecting of a new species
described here. Ellen Larsson (Gothenburg) and Karl-Henrik
Larsson (Oslo) were instrumental in providing sequences for
the reference data set.
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