Hydnoporia, an older name for Pseudochaete and Hymenochaetopsis, and typification of the genus Hymenochaete (Hymenochaetales, Basidiomycota).
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Abstract
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Hydnoporia, an older name for Pseudochaete and Hymenochaetopsis, and typification of the genus Hymenochaete (Hymenochaetales, Basidiomycota)
Abstract
The genus name Hydnoporia is reinstated to encompass the Hymenochaete tabacina group currently addressed to Pseudochaete (illegitimate genus) or Hymenochaetopsis. Identity of the type species of Hydnoporia, Sistotrema fuscescens (= Hydnoporia olivacea in current sense), is clarified, and a lectotype is selected. In total, 12 species are combined in Hydnoporia: H. corrugata, H. gigasetosa, H. lamellata, H. laricicola, H. latesetosa, H. lenta, H. rhododendri, H. rimosa, H. subrigidula, H. tabacina, H. tabacinoides, and H. yasudai. Hydnoporia diffissa is described as new. Analyses of all available ITS (94) and newly produced tef1 sequences (20) indicate that there are at least 20–27 species in the genus. Identity of the type species of Hymenochaete, H. rubiginosa, is clarified; the name is retained for the species so named in Europe while other species are present in North America and East Asia. Additionally, three new combinations in Hymenochaete are proposed: H. campylopora (= Cyclomyces fuscus), H. microcycla (= Cyclomyces tabacinus), and H. saepiaria.
INTRODUCTION
The genus Hymenochaete has traditionally included all brown-coloured corticioid fungi with simple-septate hyphae and setae, which are specialized sterile cells in the basidiocarp. After the study of Patouillard (1900) the concept of the genus remained stable until the application of DNA-based phylogenetics. Affinity of Hymenochaete to polypores with similar micromorphology and pigmentation also became well accepted, and both species of poroid and smooth hymenophores were included in the family Hymenochaetaceae early on (after Donk 1933).
The genus has received considerable attention from taxonomists, and Léger (1998) recognized no less than 132 species in the only published monograph of the genus. Yet before the DNA era, few questioned the unity of Hymenochaete, so similar are the species morphologically. For instance, in his extensive morphological analysis Parmasto (1995) did not find grounds for splitting Hymenochaete.
Wagner & Fischer (2002) were the first to use genetic data to show that Hymenochaete in the traditional sense was polyphyletic. They divided Hymenochaetaceae with smooth hymenophores to two separate lineages. The name Hymenochaete went with the larger clade, while they described a new genus for the other lineage, Pseudochaete. In their analysis Pseudochaete contained only one, albeit common species, Hymenochaete tabacina.
Since then more comprehensive order-level analysis have confirmed that the corticioid species within Hymenochaetaceae are divided in two clades, equal to well-established polypore genera such as Fomitiporia and Phellinus (Binder et al. 2005, Larsson et al. 2006). Recent studies have provided additional molecular data on Hymenochaete and demonstrated that 12 additional species belong to Pseudochaete (He & Dai 2012, He & Li 2013, Parmasto et al. 2014). Even though relationships between genera of the Hymenochaetaceae are to some degree unresolved, these studies have shown beyond doubt that Hymenochaete and Pseudochaete are separate clades that deserve generic status.
Unfortunately, the fungal genus name Pseudochaete T. Wagner & M. Fisch. turned out to be a homonym of Pseudochaete W. West & G.S. West 1903, a genus of green algae belonging to the family Chaetophoraceae. Consequently, Yang et al. (2016) suggested a new name (nomen novum) for the Pseudochaete group, Hymenochaetopsis S.H. He & Jiao Yang, making the appropriate combinations.
When describing a genus for Hymenochaete tabacina and its relatives, Wagner & Fischer (2002) and Yang et al. (2016) neglected the genus name Hydnoporia Murrill, which is typified by Sistotrema fuscescens Schwein., now considered a synonym of the common North American species Hydnochaete olivacea (Ryvarden 1982a). Parmasto et al. (2014), corroborated by Yang et al. (2016), showed that H. olivacea is a close, congeneric relative of H. tabacina. This makes Hydnoporia, fungal Pseudochaete and Hymenochaetopsis taxonomic synonyms.
As a genus described in 1907, Hydnoporia has priority over Hymenochaetopsis and Pseudochaete. We see no good reasons for conserving Pseudochaete for the fungal genus and even less so for Hymenochaetopsis. Conservation is recommended when a name change would affect names in wide and persistent use, e.g. species of great economic value or well-known and much studied parasites. Pseudochaete has been in use within kingdom Fungi for 17 yr only, and Hymenochaetopsis just for 3 yr. Neither does the genus contain enigmatic species, and thus arguments for conservation are weak. We therefore adopt the name Hydnoporia for the whole Hymenochaete tabacina clade, replacing Pseudochaete and Hymenochaetopsis.
In total, 14 species names are accepted in Hydnoporia after our revision, although true species number appears to be at least 20–27 judging from sequence data. The species range from resupinate species with smooth hymenophore to pileate and hydnoid (Fig. 1). Here we have studied the H. corrugata and H. tabacina species complexes more closely, resulting in the description of a new species, H. diffissa, and re-introduction of H. rimosa, an East Asian H. corrugata relative. Hymenochaete intricata is placed among the synonyms of Hydnoporia tabacina. In addition, the generic type of Hymenochaete, H. rubiginosa, is typified, and three other, closely related species are recognized in the H. rubiginosa group, although left unnamed.
Basidiocarp diversity of Hydnoporia. A. H. rimosa (RU-KHA, Spirin 10797). B. H. olivacea (US-MA, Miettinen 15.10.2011). C. H. tabacina (US-NC, Miettinen 19630). D. H. laricicola (RU-KHA, Spirin 3810). E. H. yasudai (RU-KHA, Spirin 10812).
MATERIALS AND METHODS
We studied type specimens and collections from herbaria H, O, UPS, NY, TNS, TAAM, BPI, and GB. Herbarium acronyms are given according to Thiers (2018). Studied collections with ITS sequences are marked with asterisk (*) throughout this paper. References to the International Code of Nomenclature for algae, fungi, and plants (ICN) all follow the Shenzhen code (Turland et al. 2018).
We measured all microscopic structures with Leica microscopes using Cotton Blue in lactic acid (CB, Merck 1275), with ×1250 magnification and phase contrast illumination. Additionally, we used 5 % KOH for microscopy to study tight hyphal structures. At least 30 basidiospores were measured per each specimen reported in Table 1. For presenting variation of hyphal width and basidiospores, the 20 % and 5 % extremes are given in parentheses, respectively (hyphal width variation is larger than spore size variation). Setal measurements are reported as for spores (Table 2). Sketches were made using a drawing tube with the exception of spores that were drawn with free hand after real measured spores. The sketches were then imported to CorelDRAW 2017 and redrawn to vector graphics. Spore statistics were produced in Microsoft Excel 2016.
Table 1.
Basidiospore measurements of the Hydnoporia specimens.
| Species / specimen | L’ | L | W’ | W | Q’ | Q | n |
|---|---|---|---|---|---|---|---|
| H. corrugata | (4.0)4.1–5.5(6.2) | 4.76 | (1.5)1.6–1.8(1.9) | 1.70 | (2.2)2.3–3.3(3.4) | 2.81 | 90 |
| Michelitsch 20 Mar. 1977 | (4.0)4.1–5.3(6.2) | 4.86 | (1.5)1.6–1.8(1.9) | 1.70 | (2.5)2.6–3.3(3.4) | 2.87 | 30 |
| Poelt 15 Sep. 1974 | (4.0)4.1–5.1(5.2) | 4.46 | (1.5)1.6–1.8(1.9) | 1.69 | (2.2)2.3–3.0(3.1) | 2.65 | 30 |
| Volobuev 1 Aug. 2013 | (4.3)4.4–5.5(5.8) | 4.97 | (1.5)1.6–1.8(1.9) | 1.72 | (2.3)2.4–3.2(3.4) | 2.90 | 30 |
| H. diffissa | (4.1)4.2–7.3(7.8) | 5.28 | 1.7–2.2 | 1.92 | (2.1)2.2–3.5(3.7) | 2.74 | 90 |
| Miettinen 17127.4 | (4.7)5.1–7.3(7.8) | 5.99 | 1.8–2.2 | 1.99 | (2.4)2.5–3.5(3.7) | 3.00 | 30 |
| Miettinen 19463 | (4.6)4.7–5.6(6.2) | 5.18 | 1.8–2.1(2.2) | 1.91 | (2.4)2.5–3.1 | 2.72 | 30 |
| Ryvarden 46500 | (4.1)4.2–5.2(5.6) | 4.67 | 1.7–2.0(2.1) | 1.87 | (2.1)2.2–2.8(3.0) | 2.50 | 30 |
| H. laricicola | (4.2)4.3–6.8(7.0) | 5.27 | (1.6)1.7–2.1(2.2) | 1.91 | (2.2)2.3–3.5(3.6) | 2.78 | 60 |
| Spirin 5400 | (4.2)4.3–5.6(5.8) | 4.91 | (1.6)1.7–2.1(2.2) | 1.93 | (2.2)2.3–2.8(2.9) | 2.56 | 30 |
| Ahti 50083 | (4.3)4.5–6.8(7.0) | 5.63 | (1.6)1.7–2.0(2.1) | 1.89 | (2.2)2.3–3.5(3.6) | 2.99 | 30 |
| H. rhododendri | (3.8)3.9–5.2(5.6) | 4.56 | (1.1)1.2–1.6(1.7) | 1.33 | (2.7)2.8–4.3(4.7) | 3.44 | 90 |
| Spirin 6450 | (3.8)3.9–5.2(5.6) | 4.51 | (1.1)1.2–1.5(1.6) | 1.29 | (2.7)3.1–4.1(4.3) | 3.51 | 30 |
| Spirin 6476 | (3.9)4.1–5.1(5.2) | 4.51 | (1.1)1.2–1.5(1.6) | 1.29 | (2.9)3.1–4.3(4.7) | 3.51 | 30 |
| TAAM 189353 | (3.8)4.0–5.2(5.3) | 4.65 | (1.1)1.2–1.6(1.7) | 1.42 | (2.7)2.8–4.0(4.1) | 3.31 | 30 |
| H. rimosa | (4.1)4.2–6.3(6.9) | 5.22 | (1.7)1.8–2.3(2.5) | 1.99 | (2.0)2.1–3.2(3.4) | 2.64 | 180 |
| Spirin 5060 | (4.2)4.3–5.7(6.0) | 4.94 | (1.7)1.8–2.0(2.1) | 1.84 | (2.3)2.4–3.0(3.2) | 2.69 | 30 |
| Spirin 5277 | (5.0)5.1–5.8(6.1) | 5.32 | (1.9)2.0–2.3(2.4) | 2.12 | (2.2)2.3–2.8(2.9) | 2.52 | 30 |
| Spirin 5617 | (4.1)4.2–5.9(6.1) | 4.89 | (1.8)1.9–2.3(2.4) | 1.99 | (2.0)2.1–2.9(3.0) | 2.47 | 30 |
| Spirin 5629 | (4.7)5.1–6.3(6.9) | 5.66 | (1.8)1.9–2.2(2.5) | 2.04 | (2.3)2.4–3.2(3.3) | 2.78 | 30 |
| Spirin 5678 | (4.1)4.2–5.3(5.5) | 4.82 | (1.7)1.8–2.0(2.1) | 1.88 | (2.2)2.3–2.9(3.0) | 2.58 | 30 |
| Spirin 6104 | (4.6)5.0–6.3(6.8) | 5.68 | (1.8)1.9–2.2(2.4) | 2.04 | (2.4)2.5–3.2(3.4) | 2.78 | 30 |
| H. tabacina | (3.9)4.1–6.2(6.4) | 4.93 | (1.4)1.5–2.1(2.2) | 1.81 | (2.0)2.1–3.6(3.8) | 2.74 | 420 |
| Kotiranta 20693 | (4.0)4.1–5.2(5.8) | 4.54 | (1.7)1.8–2.1(2.2) | 1.89 | (2.0)2.1–2.8(2.9) | 2.40 | 30 |
| Kotiranta 21006 | (4.3)4.5–6.1(6.3) | 5.20 | (1.7)1.8–2.0(2.1) | 1.89 | (2.2)2.3–2.9(3.3) | 2.75 | 30 |
| Kotiranta 25205 | (4.1)4.2–6.0(6.1) | 4.94 | 1.7–2.0 | 1.83 | (2.2)2.3–3.1(3.4) | 2.70 | 30 |
| Miettinen 17028.3 | (4.2)4.3–5.3(6.4) | 4.91 | (1.5)1.6–1.9(2.0) | 1.72 | (2.4)2.5–3.3(3.6) | 2.87 | 30 |
| Miettinen 18168 | (4.3)4.6–5.9(6.1) | 5.16 | (1.7)1.8–2.0(2.1) | 1.93 | (2.3)2.4–3.2(3.3) | 2.68 | 30 |
| Niemelä 2604 | (4.1)4.2–5.8(6.4) | 4.96 | (1.5)1.6–1.8(1.9) | 1.68 | (2.3)2.4–3.5(3.6) | 2.96 | 30 |
| Niemelä 3581 | (4.4)4.6–6.2(6.3) | 5.41 | (1.6)1.7–2.0(2.1) | 1.82 | (2.4)2.6–3.3(3.4) | 2.99 | 30 |
| Niemelä 8341 | (5.0)5.1–6.1(6.2) | 5.45 | (1.7)1.8–2.0(2.1) | 1.88 | (2.5)2.6–3.3(3.4) | 2.91 | 30 |
| Ryvarden 14421 | (4.1)4.2–5.7(6.0) | 4.74 | (1.5)1.6–1.8(1.9) | 1.69 | (2.2)2.3–3.4(3.6) | 2.82 | 30 |
| Spirin 5196 | (4.7)4.8–5.9(6.0) | 5.20 | 1.8–2.1(2.2) | 1.95 | (2.2)2.4–2.9(3.0) | 2.67 | 30 |
| Spirin 5496 | (3.9)4.0–5.2(6.2) | 4.37 | (1.4)1.5–1.8(1.9) | 1.70 | (2.1)2.2–3.4(3.6) | 2.59 | 30 |
| Spirin 6507 | (4.0)4.1–5.1(5.3) | 4.51 | (1.6)1.7–2.0(2.1) | 1.88 | (2.1)2.2–2.7(2.8) | 2.41 | 30 |
| Spirin 6517 | (4.2)4.3–5.6(5.7) | 4.76 | (1.5)1.6–2.0 | 1.79 | (2.2)2.3–3.2(3.3) | 2.67 | 30 |
| Spirin 6520 | (4.1)4.2–6.0(6.2) | 4.80 | (1.4)1.5–1.8 | 1.64 | 2.5–3.6(3.8) | 2.94 | 30 |
| H. yasudai | (4.3)4.4–6.1(6.4) | 5.28 | (1.9)2.0–2.3(2.4) | 2.09 | (1.9)2.2–2.9(3.0) | 2.53 | 30 |
| Spirin 6475 | (4.3)4.4–6.1(6.4) | 5.28 | (1.9)2.0–2.3(2.4) | 2.09 | (1.9)2.2–2.9(3.0) | 2.53 | 30 |
Table 2.
Setal measurements of selected Hydnoporia specimens.
| Species / specimen | L’ | L | W’ | W | Q’ | Q | n |
|---|---|---|---|---|---|---|---|
| H. corrugata | (49.8)52.1–86.0(110.9) | 66.8 | (6.9)7.2–14.1(17.8) | 9.8 | (4.5)4.7–9.6(9.7) | 7.00 | 40 |
| Poelt 15 Sep. 1974 | (49.8)52.3–86.0(110.9) | 66.9 | (6.9)7.2–14.1(17.8) | 10.1 | (4.8)5.1–8.3(8.6) | 6.77 | 20 |
| Volobuev 1 Aug. 2013 | (51.8)52.1–84.2(94.2) | 66.6 | (7.2)7.3–11.3(13.7) | 9.5 | (4.5)4.7–9.6(9.7) | 7.23 | 20 |
| H. diffissa | (46.8)50.7–73.0(90.2) | 60.6 | (6.2)6.8–11.4(12.9) | 9.1 | (4.7)5.0–8.8(9.2) | 6.82 | 60 |
| Miettinen 17127.4 | (46.8)50.7–70.2(72.0) | 58.2 | (6.2)7.0–9.9(11.6) | 8.4 | (5.1)5.2–8.8(9.2) | 7.08 | 20 |
| Miettinen 19463 | (49.8)52.7–70.3(79.8) | 60.2 | (6.4)6.8–11.1(11.5) | 8.8 | (4.7)5.0–8.1(8.6) | 6.94 | 20 |
| Ryvarden 46500 | (50.2)53.2–73.0(90.2) | 63.4 | (7.9)8.0–11.4(12.9) | 10.0 | (4.7)5.0–7.9(8.7) | 6.44 | 20 |
| H. laricicola | (30.8)31.2–68.8(83.6) | 43.9 | (5.3)5.6–12.0(12.1) | 8.1 | (3.8)3.9–8.2(8.9) | 5.41 | 40 |
| Spirin 5400 | (30.8)31.2–54.9(59.2) | 41.0 | (5.5)6.0–11.1(12.2) | 8.1 | (3.8)3.9–6.4(7.5) | 5.16 | 20 |
| Ahti 50083 | (31.8)32.3–68.8(83.6) | 46.8 | (5.3)5.6–12.0(12.1) | 8.1 | (3.8)4.1–8.2(8.9) | 5.86 | 20 |
| H. rhododendri | (53.2)56.1–95.3(98.6) | 71.2 | (6.2)6.3–17.2(18.7) | 10.5 | (4.2)4.7–11.5(13.3) | 7.19 | 60 |
| Spirin 6450 | (53.2)56.3–82.3(88.2) | 68.7 | (6.2)6.3–12.0(12.5) | 8.7 | (4.6)5.8–11.5(13.3) | 8.24 | 20 |
| Spirin 6476 | (54.3)56.1–95.3(98.6) | 74.0 | (8.2)9.6–15.6(17.8) | 11.7 | (4.2)4.8–8.7(8.8) | 6.48 | 20 |
| TAAM 189353 | (53.8)61.7–88.2(96.3) | 71.0 | (7.0)7.6–17.2(18.7) | 11.0 | (3.8)4.7–9.9(10.5) | 6.85 | 20 |
| H. rimosa | (50.2)53.3–130.5(144.3) | 74.6 | (6.6)6.8–18.0(18.2) | 10.3 | (4.4)5.1–12.1(12.7) | 7.41 | 100 |
| Spirin 5060 | (50.5)53.3–73.0(76.0) | 62.5 | (6.6)6.7–10.5(12.0) | 8.7 | (4.4)6.0–8.9(9.6) | 7.34 | 20 |
| Spirin 5277 | (59.7)62.1–130.5(144.3) | 96.8 | (8.2)8.8–18.0(18.2) | 13.1 | (5.0)5.3–10.7(12.7) | 7.65 | 20 |
| Spirin 5617 | (50.2)55.2–89.5(107.5) | 70.7 | (7.3)7.6–11.7(12.3) | 9.9 | (4.9)5.5–9.8(10.3) | 7.25 | 20 |
| Spirin 5629 | (59.0)59.2–99.8(126.2) | 75.1 | (8.2)8.3–15.2(15.6) | 10.6 | (4.5)5.1–12.1(12.5) | 7.31 | 20 |
| Spirin 5678 | (53.2)58.2–90.3(92.2) | 68.1 | (6.7)6.8–10.8(11.2) | 9.2 | (5.5)5.8–9.4(9.9) | 7.51 | 20 |
| H. tabacina | (51.2)54.3–119.6(131.3) | 78.6 | (6.8)7.0–17.4(18.8) | 11.4 | (4.1)4.4–11.0(12.5) | 7.02 | 380 |
| Kotiranta 20797 | (66.7)74.5–119.6(131.3) | 94.6 | (7.7)9.3–14.8(15.1) | 12.4 | (5.0)5.7–11.0(11.8) | 7.81 | 20 |
| Kotiranta 21006 | (56.4)58.3–95.6(104.5) | 78.9 | (7.8)8.2–13.7(14.4) | 10.6 | (5.6)6.1–8.2(8.8) | 7.51 | 20 |
| Miettinen 7428 | (56.2)62.6–89.3(95.2) | 77.6 | (9.5)10.3–13.7(15.6) | 12.2 | (4.8)4.9–8.0(9.0) | 6.42 | 20 |
| Kotiranta 20693 | (52.3)54.3–85.5(89.8) | 67.3 | (8.0)8.9–14.6(18.0) | 11.2 | (4.2)4.5–8.0(8.7) | 6.15 | 20 |
| Kotiranta 25205 | (51.2)60.5–83.1(112.0) | 71.5 | (6.8)7.0–14.0(14.4) | 10.5 | (5.0)5.7–11.0(11.8) | 7.81 | 20 |
| Kotiranta 25386a | (53.6)61.9–83.2(85.3) | 70.1 | (8.4)8.8–13.8(14.8) | 10.7 | (5.4)5.8–7.5(7.6) | 6.64 | 20 |
| Miettinen 17028.3 | (68.7)69.0–104.5(109.9) | 85.3 | (9.3)10.1–12.5(12.6) | 11.1 | (6.0)6.8–8.7(10.1) | 7.73 | 20 |
| Miettinen 18168 | (62.3)63.3–97.3(111.2) | 76.5 | (7.2)7.6–12.7(14.8) | 10.0 | (5.4)5.8–7.5(7.6) | 7.86 | 20 |
| Niemelä 2604 | (69.2)70.2–98.0(109.6) | 80.9 | (8.5)8.6–16.5(18.0) | 11.3 | (4.9)5.4–8.8(10.1) | 7.40 | 20 |
| Niemelä 3581 | (61.3)62.3–108.2(125.3) | 85.0 | (9.0)10.0–14.8(15.7) | 11.9 | (4.1)4.5–10.3(12.5) | 7.35 | 20 |
| Niemelä 8341 | (57.2)58.3–88.6(95.7) | 73.3 | (7.3)7.6–13.2(14.2) | 10.1 | (4.8)5.2–9.5(9.8) | 7.50 | 20 |
| Ryvarden 14421 | (73.1)74.2–91.2(101.0) | 82.5 | (9.2)9.8–13.7(14.2) | 11.3 | (5.7)5.8–8.5(9.3) | 7.42 | 20 |
| Spirin 5057 | (61.2)63.3–102.8(104.6) | 83.8 | (9.7)10.4–15.4(17.2) | 12.7 | (4.5)4.9–8.4(10.6) | 6.71 | 20 |
| Spirin 5196 | (49.0)55.7–79.0(83.2) | 64.7 | (6.0)8.3–12.0(13.2) | 9.8 | (5.0)5.1–8.1(8.5) | 5.86 | 20 |
| Spirin 5496 | (63.8)69.3–88.4(108.2) | 79.6 | (9.8)11.1–17.4(18.8) | 13.9 | (4.1)4.4–7.6(9.0) | 5.86 | 20 |
| Spirin 6066a | (57.3)58.8–85.2(94.3) | 71.2 | (7.3)8.7–12.8(14.0) | 10.4 | (4.8)5.4–8.6(9.2) | 7.01 | 20 |
| Spirin 6507 | (55.3)63.1–99.4(108.7) | 78.8 | (7.3)9.2–14.4(14.7) | 12.1 | (5.4)5.5–9.0(9.3) | 6.59 | 20 |
| Spirin 6517 | (69.2)71.2–100.1(104.8) | 85.0 | (8.5)10.1–13.8(14.5) | 11.8 | (6.2)6.4–8.5(8.7) | 7.24 | 20 |
| Spirin 6520 | (61.7)63.7–114.9(126.0) | 84.1 | (10.2)10.6–15.4(16.1) | 13.1 | (4.2)4.4–8.7(9.8) | 6.49 | 20 |
| H. yasudai | (54.8)61.4–103.7(106.0) | 84.0 | (8.0)10.3–16.6(17.1) | 12.0 | (4.0)4.1–9.2(10.3) | 7.07 | 40 |
| holotype | (55.2)61.4–96.5(97.6) | 85.6 | (9.3)10.3–16.6(17.1) | 12.4 | (4.0)4.1–9.2(10.3) | 6.89 | 20 |
| Spirin 6475 | (54.8)62.2–103.7(106.0) | 82.3 | (8.0)10.3–14.2(16.0) | 11.6 | (4.9)5.0–9.2(9.5) | 7.25 | 20 |
We sequenced nuclear ribosomal DNA internal transcribed spacer (ITS) from 25 samples and translation elongation factor 1-α (tef1) from 20 samples. Sequences were produced as described in Miettinen et al. (2018). The resulting sequences are available in INSDC (Table 3). In addition, all publicly available sequences of Hydnoporia and Hymenochaete rubiginosa complex were included in our analyses based on BLAST and name searches: GenBank - 67 sequences; UNITE - 2 (Nilsson et al. 2018); BOLD - 1 (Ratnasingham & Hebert 2007).
Table 3.
Specimens of Hydnoporia and Hymenochaete sequenced for this study with GenBank accession numbers. Sequences marked with ¤ derived from cultures.
| Species | Collector, coll. no. (herbarium) | Country (province/state) | ITS | tef1 |
|---|---|---|---|---|
| Hydnoporia corrugata | Jon Klepsland 11.021 (O F-247869), neotype | Norway | MK514613 | MK552138 |
| Hydnoporia diffissa | Otto Miettinen 17127.4 (H) ¤ | USA (NY) | MK514598 | – |
| Otto Miettinen 19463 (H 7008917), holotype | USA (NC) | MK514611 | MK552136 | |
| Hydnoporia laricicola | Viacheslav Spirin 5400 (H) ¤ | Russia (KHA) | MK514606 | MK552132 |
| Hydnoporia olivacea | Otto Miettinen 16956.2 (H 7008916), epitype | USA (NY) | MK514610 | – |
| Otto Miettinen & Kelo Käppi 16044 (H 7005770), epitype | USA (MA) | MK514612 | MK552137 | |
| Hydnoporia rhododendri | N. Gerhold 3 Jun 2005 (H, ex TAA 189353) | Austria | MK514593 | – |
| Viacheslav Spirin 6476 (H) ¤ | Russia (PRI) | MK514599 | MK552127 | |
| Viacheslav Spirin 6450 (H) ¤ | Russia (PRI) | MK514603 | – | |
| Hydnoporia rimosa | Viacheslav Spirin 5277 (H) | Russia (KHA) | MK514592 | MK552122 |
| Viacheslav Spirin 5678 (H) | Russia (KHA) | MK514594 | MK552123 | |
| Viacheslav Spirin 6104 (H) | Russia (KHA) | MK514595 | MK552124 | |
| Hydnoporia tabacina | Heikki Kotiranta 20797 (H) | Russia (PER) | MK514591 | MK552121 |
| Heikki Kotiranta 25205 (H) | Russia (KK) | MK514596 | MK552125 | |
| Viacheslav Spirin 6066a (H) ¤ | Russia (NIZ) | MK514600 | MK552128 | |
| Otto Miettinen 17028.3 (H) ¤ | United States (NY) | MK514601 | MK552129 | |
| Otto Miettinen 22126 (H) | Finland | MK782755 | MK787232 | |
| Viacheslav Spirin 5196 (H) ¤ | Russia (KHA) | MK514602 | MK552130 | |
| Viacheslav Spirin 6582 (H) ¤ | Russia (KHA) | MK514604 | – | |
| Viacheslav Spirin 6566 (H) ¤ | Russia (KHA) | MK514605 | MK552131 | |
| Viacheslav Spirin 6520 (H) ¤ | Russia (KHA) | MK514607 | MK552133 | |
| Viacheslav Spirin 6507 (H) ¤ | Russia (KHA) | MK514608 | MK552134 | |
| A. M. Ainsworth & A. Lucas 17 Jan 2017 (K(M) 233332) ¤ | UK | MK514614 | MK890223 | |
| Hydnoporia yasudai | Viacheslav Spirin 5533 (H) ¤ | Russia (KHA) | MK514597 | MK552126 |
| Viacheslav Spirin 6475 (H) ¤ | Russia (PRI) | MK514609 | MK552135 |
We constructed three sequence datasets for analyses:
Hydnoporia ITS-dataset includes 94 sequences. Alignment length was 738 with 250 variable site patterns. We also analysed the same alignment after excluding fastest evolving sites (651 bp, 191 variable sites). Results were topologically identical and we report below results based on the longer alignment.
Hydnoporia tef1-dataset includes 20 sequences. After removing unalignable, ambiguous alignment positions, alignment length was 721 with 151 variable site patterns.
Hymenochaete rubiginosa ITS dataset includes 12 sequences. The closest relatives of H. rubiginosa s. str., determined by BLAST similarity (≥ 94 %) were included in the analyses. Alignment length was 660 with 71 variable site patterns.
PRANK online version at the European Bioinformatics Institute website was used for aligning sequences (Löytynoja & Goldman 2005, Cowley et al. 2017). Resulting alignments were refined, and alignment positions with unclear homology were excluded manually (tef1-dataset) using PhyDE v. 0.9971 (Müller et al. 2010).
Phylogenies were constructed with MrBayes v. 3.1.2 and 3.2.6 (Ronquist et al. 2012). Nucleotide substitution models were chosen with jModelTest v. 2.1.10 based on AIC (Darriba et al. 2012): GTR+I+G for the Hydnoporia ITS-dataset, GTR+G for the Hydnoporia tef1-dataset, and GTR for the Hymenochaete rubiginosa ITS-dataset. Analyses were run with eight chains in three parallel runs, temp = 0.1 for 5 M generations (10 M for Hydnoporia ITS-dataset) sampling a tree every 2 000 generations. All runs had converged to below 0.01 average standard deviation of split frequencies by the end of the run. A burn-in of 25 % was used before computing the consensus tree.
The alignments and phylograms have been deposited in TreeBASE (submission 24281). The resulting phylograms were edited in FigTree v. 1.4.3 (Rambaut 2014); since our focus was in species delimitation and in the absence of reliable rooting data, we used midpoint rooting in all cases.
Specimens examined (sequenced collections marked by asterisk):
Clavariachaete rubiginosa. Venezuela, Río Negro, Rio Mawarinuma, 3 Feb. 1985, Halling 4151 (NY).
Hydnoporia corrugata. Austria, Steiermark, Koralpe, branches, 15 Sep. 1974, Poelt (H), Fagus sylvatica, 20 Mar. 1977, Michelitsch (H). Norway, Hordaland, Bømlo, Corylus avellana, 24 Jul. 2011, Klepsland 11.021* (O F-247869, neotype). Russia, Kaluga Reg., Ulyanovo Dist., Kaluzhskie Zaseki Nat. Res., C. avellana, 1 Aug. 2013, Volobuev* (H ex LE).
Hydnoporia diffissa. USA, New Jersey, Gloucester Co., Newfield, hardwood, Jun. 1893, Ellis & Everhart (Fungi Columbiani #8) (H, NY), Quercus ilicifolia (branches), ‘October’ (no year indicated), Ellis (Rabenhorst-Winter Fungi Europaei #3136) (NY); New York, Essex Co., Arbutus Lake, dry branches, 22 Sep. 2013, Miettinen 17127.4* (H); North Carolina, Buncombe Co., Blue Ridge Assembly, Rhododendron maximum, 24 Sep. 2015, Miettinen 19463* (H); Ohio, Hocking Co., Old Man’s Cave St. Park, Fagus grandifolia, 15 Nov. 1975, Cooke 51769 (H, O); Pennsylvania, Butler Co., Saxonburg, 18 Sep. 1939, Sumstine 12125 (NY); Indiana Co., Marion Center, 27 Sep. 1949, Henry (NY); Tennessee, Great Smoky Nat. Park, dead hardwood, 14 Jul. 2004, Ryvarden 46500 (H, O).
Hydnoporia laricicola. Russia, Altai, Shebalino Dist., Cherga, Larix sibirica, 16 Aug. 1985, Alanko 52913 (H); Karelia, Pudozh Dist., Vodlozero Nat. Park, L. sibirica, 22 Jun. 1991, Ahti 50083 (H); Khabarovsk Reg., Khabarovsk Dist., Malyi Kukachan, Larix gmelinii, 19 Aug. 2012, Spirin 5400* (H); Solnechnyi Dist., Igdomi, L. gmelinii, 5 Aug. 2011, Spirin 3810 (H); Verkhnebureinskii Dist., Kyvyty, L. gmelinii, 17 Aug. 2014, Spirin 7457 (H); Dublikan, L. gmelinii, 23 Aug. 2014, Spirin 7954 (H).
Hydnoporia olivacea. USA, New Jersey, Gloucester Co., Newfield, hardwood, 1874 Ellis (NY 00742784, lectotype of Hymenochaete agglutinans), Nov. 1881, Ellis (NY ex C. Roumeguére, Fungi Selecti Exsiccati #4540, epitype of H. agglutinans); New York, New York: Essex Co., Huntington Wildlife Forest, Adjidaumo flow, Fagus grandifolia, 18 Sep. 2013, Miettinen 16956.2* (H 7008916, epitype of Thelephora episphaeria); North Carolina, Burke Co., Salem, Schweinitz (UPS F-173549) (lectotype of Sistotrema fuscescens), the same locality, Schweinitz (UPS F-173519) (lectotype of Thelephora episphaeria); Durham/Orange Co., Eno River State Park, dicot, 2 Nov. 1997, E. Larsson 90-97* (GB 0150286); Massachusetts, Worcester, Columbus Park, hardwood, 11 Apr. 2013, Miettinen 16044 & Käppi* (H 7005770, epitype of Sistotrema fuscescens).
Hydnoporia rhododendri. Austria, Tyrol, Kaunertaler Gletscherstrasse, Rhododendron ferrugineum, 3 Jun. 2005, Gerhold* (H ex TAAM 189353, isotype?); Russia, Buryatia, Barguzin Dist., Shumilikha, Rhododenron tomentosum, 28 Aug. 2003, Ryvarden 45907 (O); Primorie Reg., Krasnoarmeiskii Dist., Valinku, Rhododendron hypoleucum, 28 Aug. 2013, Spirin 6450* (H); 6452 (H); 29 Aug. 2013, Spirin 6476* (H). Sweden, Uppland, Roslagen, Calluna vulgaris (or Empetrum nigrum), 16 Jul. 1906, Starbäck (H). Switzerland, Graubünden, St. Moritz, R. ferrugineum, Aug. 1882, Winter (H ex Rabenhorst-Winter Fungi Europaei #2932, as Stereum tabacinum).
Hydnoporia rimosa. Japan, Tôhoku, Sendai, 24 Oct. 1920, Yasuda (TNS F-203210, lectotype). Russia, Khabarovsk Reg., Khabarovsk Dist., Malyi Niran, Corylus mandshurica, 8 Aug. 2012, Spirin 5060 (H); Ulika, C. mandshurica, 14 Aug. 2012, Spirin 5277* (H); Ulun, Ulmus japonica, Rhododendron dauricum, Syringa amurensis, 26 Aug. 2012, Spirin 5617, 5629, 5678* (H); Komsomolsk Dist., Boktor, C. mandshurica, 18 Aug. 2013, Spirin 6104* (H) (basidiocarps and sterile stage); Solnechnyi Dist., Igdomi, Quercus mongolica, 7 Aug. 2011, Spirin 3952 (H); Duschekia fruticosa, 2 Sep. 2016, Spirin 10797 (H); Sonakh, Q. mongolica, 15 Aug. 2014, Spirin 7294 (H).
Hydnoporia tabacina. Canada, Québec Nordique, Poste-de-la-Baleine, Alnus crispa, 29 Jul. 1982, Niemelä 2569 (H); 10 Aug 1982, Niemelä 2720 (H); Salix, 31 Jul. 1982, Niemelä 2604 (H). China, Jilin Prov., Antu Co., Huang Song Pu, Acer, 28 Aug. 2005, Miettinen 10618.2 (H). Estonia, Jõgevamaa, Imukvere, Picea abies, 22 Sep. 1956, Parmasto (Mycotheca Estonica 1: 7, 1957) (H ex TAAM); Tartumaa, Kastre, P. abies, 20 Jun. 1930, Lepik (Fungi Estonici Exsiccati 1: 9, 1931) (H); Valgamaa, Paluperä, Käpa, Corylus avellana, 13 Sep. 2012, Kotiranta 25386a (H). Finland, Varsinais-Suomi, Tammisaari, Lökudden Nat. Res., Ribes alpinum, 1 Sep. 2004, Kotiranta 20693 (H); Uusimaa, Helsinki, Viikki, C. avellana, 18 Mar. 2014, Miettinen 18168* (H); Veräjämäki, P. padus, 7 Apr. 2019, Miettinen 22126* (H); Etelä-Häme: Lammi Biol. Station, Quercus robur, 17 Sep. 2013, Spirin 6723a (H); Untulanharju, P. abies, 17 Sep. 1986, Niemelä 3581 (H); Lapinjärvi, P. abies, 8–9 Sep. 2003, Miettinen 7428 (H); Luhanka, Sepänmäki, Juniperus communis, 21 Oct. 2004, Kotiranta 21006 (H); Pohjois-Savo, Enonkoski, Kolovesi Nat. Park, Salix caprea, 30 Sep. 2006, Niemelä 8341 (H). Japan, Hokkaido, Tomakomai, Iburi, Alnus, 20 Dec. 1904, Niidjima 706 (TNS F-200336, isotype of Hymenochaete intricata); Honshu, Kantô, Tochigi, Larix, 31 Oct. 1991, Ryvarden 30234 (O). Russia, Khabarovsk Reg., Khabarovsk Dist., Malyi Niran, Pinus koraiensis, 8 Aug. 2012, Spirin 5057 (H); Ulika, Actinidia kolomikta, 13 Aug. 2012, Spirin 5196* (H); Bolshoi Khekhtsir, Picea ajanensis, 2–3 Sep. 2013, Spirin 6517, 6582* (H); C. mandshurica, 2 Sep. 2013, Spirin 6520* (H); Acer ukurunduense, 3 Sep. 2013, Spirin 6566* (H); Levyi Ulun, R. dauricum, 22 Aug. 2012, Spirin 5496 (H); Solnechnyi Dist., Igdomi, Alnus hirsuta, 4 Aug. 2011, Spirin 3776, 3780 (H); Khakassia, Abakan Dist., Erkagi, Salix viminalis, 15 Aug. 2011, Kotiranta 25205* (H); Krasnoyarsk Reg., Krasnoyarsk, Stolby Nat. Res., Abies sibirica, 13 Aug. 1980, Parmasto (O ex TAAM 102904); Nizhny Novgorod Reg., Lukoyanov Dist., Sanki, C. avellana, 10 Aug. 2013, Spirin 6066a* (H); Razino, Pinus sylvestris, 8 Aug. 1998, Spirin (H ex LE 211323); Perm Reg., Krasnovisherskii Dist., Kvarkush, S. caprea, 6 Aug. 2005, Kotiranta 20797* (H); Primorie Reg., Krasnoarmeiiskii Dist., Valinku, A. ukurunduense, 27 Aug. 2013, Spirin 6409 (H); Picea koreana, 29 Aug. 2013, Spirin 6507* (H). Sweden, Skåne, Fries (UPS F-175737, lectotype of Thelephora tabacina); Uppland, Uppsala, Silva Nosten, P. abies, 16 May 1929, Lundell (H ex Fungi Suecici #487). UK, England, South Hampshire, New Forest, Salix, 17 Jan. 2017, Ainsworth & Lucas* (K(M) 233332). USA, Minnesota, Clearwater Co., Itasca, 16 Sep. 1977, Ryvarden 14421 (H, O); New York, Tully, Fagus, 10 Aug. 1963, Larsen 320 (H); New York, Essex Co., Catlin Lake, deciduous tree, 19 Sep. 2013, Miettinen 17028.3* (H); North Carolina, Swain Co., Great Smokey Mtns NP, Clingmans Dome, Abies fraseri, 1 Oct. 2015, Miettinen 19630 (H); dicot, Miettinen 19631 (H).
Hydnoporia yasudai. Japan, Tokyo, Minami-Tama Gun, Sainokami Goryochi, Abies firma (dry branches), 10 Oct. 1939, Imazeki (TNS F-207357); Shiga Pref., Kutsuki, Pinus, 3 Nov. 1991, Hattori (H ex O 14614). Russia, Khabarovsk Reg., Khabarovsk Dist., Levyi Ulun, Pinus pumila (dry branches), 23 Aug. 2012, Spirin 5533* (H); Solnechnyi Dist., Igdomi, P. pumila, 2 Sep. 2016, Spirin 10812 (H); Primorie Reg., Khasan Dist., Kedrovaya Pad’ Nat. Res., Abies holophylla, 22 Jul. 1985, Parmasto (H ex TAAM 106187); Krasnoarmeiskii Dist., Valinku, P. pumila (dry branches), 29 Aug. 2013, Spirin 6475* (H).
Hymenochaete allantospora. Mexico, Veracruz, Xalapa-Coatepec, Santa Inés, 29 Mar. 1990, Bandala (TAAM 171364, holotype).
Hymenochaete iodina. Brazil, Pernacumbo, Cabo de Santo Agostinho, Mata de Gurjaú, Jul. 2001, Gibertoni 365 (O F-917605); Caruaru, Paque Ecológico Municipal Vasconcelos Sobrinho, Oct. 2000, Cáceres & Lücking 396 (O F-917951).
Hymenochaete saepiaria. Australia, New South Wales, Sydney, no date, Clealand (Lloyd’s herb. #24124) (BPI US0325520, lectotype).
Hymenochaete rubiginosa. Russia, Nizhny Novgorod Reg., Lukoyanov Dist., Panzelka, Quercus robur, 6 Aug. 2014, Spirin 7134 (H). Sweden, Småland, Femsjö, Fries (UPS F-175700, lectotype of Thelephora rubiginosa). Switzerland, Basel-land, Liestal, Wildenstein, Quercus (?), 14 Jun. 2009, Miettinen 13667.2* (H). USA, Massachusetts, Worcester, Wachusett Mtn. State Reservation, Quercus, 28 Oct. 2014, Miettinen 19066* (H); Oklahoma, Payne Co., Carl Blackwell Lake, Quercus, 8 Nov. 1979, Cooke 56530 (H ex O F14563); Tennessee, Cocke Co., Cosbey Creek, Quercus (?), 2 Oct. 2015, Miettinen 19659.2 (H); Sevier Co., Great Smokey Mtns., Ramsey Cascade Trail, Quercus, 30 Sep. 2015, Miettinen 19593* (H).
RESULTS
Our phylogenetic analyses focused on species delimitation within the genus Hydnoporia (= Pseudochaete, Hymenochaetopsis). We included all publicly available Hydnoporia ITS sequences in the analysis, altogether 94. According to our interpretation, ITS variation indicates presence of at least 20 species, but possibly up to 27 species (Fig. 2). Two of the sequences may be contaminants (KC785573, KC785579), supposedly originating from soil in a barren Antarctic cave (Connell & Staudigel 2013), while representing common North American species (Hydnoporia diffissa, H. olivacea).
Phylogram of Hydnoporia nrDNA ITS sequences. Bayesian consensus tree with midpoint rooting. Numbers denote posterior probabilities. Two-letter codes after species names denote country, followed by the province of origin (ISO 3166). Asian specimens have been coloured blue, European green and North American red. Sequences with the accession number starting with “MK” have been produced for this study, the rest are from public sources. Asterisk (*) denotes specimens that may represent separate species within the annotated lineages.
Our estimate of the species number is based on combination of two approaches: extrapolating from sequence data of our own specimens, where morphological differences are documented, and comparing sequence differences within and between clades (Fig. 2). All except two of the clades we have annotated as species contain several ITS sequences. ITS differences vary, but generally intraspecific variation is 0–3 bp (< 0.5%), while sister species show differences above 5 bp (> 0.8%). However, the difference between H. tabacina and its morphologically and ecologically distinct sister species H. rhododendri is merely 2 bp (0.3 %).
In contrast, differences between tef1 sequences (primer pair 983-G) are much larger between species, for instance 14 bp (1.7 %) between the aforementioned H. tabacina vs. H. rhododendri case (Fig. 3). The tef1 dataset supports separation of Far East H. rhododendri from the rest of H. tabacina, as well as all other divisions visible between species in the ITS dataset. Hydnoporia yasudai, in particular, would deserve further study - according to ITS data at least three species are present in East Asia in this complex, but it is unclear which of them is the real H. yasudai.
Phylogram of Hydnoporia nrDNA tef1 sequences. Bayesian consensus tree with midpoint rooting. Numbers denote posterior probabilities. Two-letter codes after species names denote country, followed by the province of origin (ISO 3166). Asian specimens have been coloured blue, European green and North American red.
While we have indicated the presence of 20 or more species based on the ITS dataset (Fig. 2), we accept only 14 formally described Hydnoporia species in this paper. Clearly more research effort is needed to sort out species names and true diversity in Hydnoporia. The species diversity of Hydnoporia seems to be high in East Asia compared to other temperate areas, although skewed sampling likely explains this pattern partially.
Taxonomy
Hydnoporia Murrill, North Am. Fl. 9: 3. 1907.
Type species: Sistotrema fuscescens Schwein. [current name Hydnoporia olivacea (Schwein.) Teixeira].
Basidiocarps brown, resupinate to effused-reflexed with smooth to hydnoid hymenophore. Hyphal system dimitic with intermediary hyphae or setal hyphae, and simple septa. Setae very large (up to 140 × 18 μm), pointed, born deep in trama. Basidia narrow, tightly packed, narrowly clavate. Basidiospores curved cylindrical or allantoid, small- to middle-sized, 4–7 × 1–2.5 μm. Some species are known to produce compact mycelial mats infecting living branches.
Notes: Description and typification by Murrill (1907) leaves no doubt that he had in mind the species commonly known as Hydnochaete olivacea when describing the monotypic genus. Hydnoporia can be morphologically divided in two groups: effused-reflexed species (H. tabacina coll.) and resupinate (H. corrugata coll.). This division is not supported by our phylogeny (Fig. 2). Resupinate species are found in the H. corrugata - H. olivacea clade only, but pileate species are otherwise present in several clades separate from the H. tabacina complex.
When describing Hymenochaetopsis (i.e. Hydnoporia), Yang et al. (2016) made no serious attempt to define the genus morphologically in comparison to Hymenochaete. They listed one possible character to separate the genera: basidiospores, which are narrowly and shortly cylindrical with a maximum size of 7 × 2.5 μm in Hymenochaetopsis.
While spore size and shape are useful characters, they are not alone sufficient for genus identification. All Hydnoporia species have relatively small cylindrical, slightly curved basidiospores, but so do some Hymenochaete spp. For instance, Hymenochaete fuliginosa and Hydnoporia corrugata have very similar cylindrical basidiospores. Other members of Hymenochaete with small cylindrical spores include Hymenochaete nanospora (sequenced in Wagner & Fischer 2002, He & Dai 2012) and Hymenochaete pinnatifida (Wagner & Fischer 2002, Parmasto et al. 2014) to mention a few further examples. Morphological differences exist, however, and are discussed below under Hymenochaete.
We base our species concept in Hydnoporia on sequence data from two markers, indicating lack of gene flow. When defining species, sequence data are used in combination with geographic range, host specificity, basidiocarp morphology, and microscopic characters (setae and spore shape and size in particular). Species delimitation without DNA sequencing is possible with the combination of the above-mentioned traits (Table 4), but in many cases requires careful study under the microscope. As discussed under H. tabacina, it may be that morphologically cryptic species are found in the genus, but this needs to be confirmed in future studies.
Table 4.
Morphological comparison of northern temperate Hydnoporia species with a smooth hymenophore.
| Species | Substrates / distribution | Macroscopic characters | Subhymenial / tramal setae | Basidiospores |
|---|---|---|---|---|
| H. corrugata | Dead angiosperm branches; Europe | Resupinate, sterile stage absent; hymenial surface grey to pale brown, margin slightly lighter or concolorous | Sharp- to blunt-pointed, 52–86 × 7–14, L=66.8, W=9.8, Q=7.00 | Cylindrical, 4.1–5.5 × 1.6–1.8, L=4.76, W=1.70, Q=2.65–2.90 |
| H. diffissa | Dead or living angiosperm branches; North America (Northeast) | Resupinate, sterile stage sometimes present; hymenial surface grey to pale brown, margin slightly darker or concolorous | Sharp- to blunt-pointed, 51–73 × 7–11.5, L=75.8, W=11.0, Q=7.10 | Cylindrical, 4.2–7.3 × 1.7–2.2, L=5.28, W=1.92, Q=2.50–3.00 |
| H. laricicola | Larix spp. (bark and branches); northern Eurasia | Effused-reflexed; hymenial surface dark grey to chocolate-brown, margin concolorous | Blunt-pointed, 31–69 × 6–12, L=43.9, W=8.1, Q=5.41 | Cylindrical, 4.3–6.8 × 1.7–2.1, L=5.27, W=1.91, Q=2.56–2.99 |
| H. rhododendri | Ericaceae; boreal Eurasia | Effused-reflexed; hymenial surface pale brown, margin slightly lighter or concolorous | Rather sharp-pointed, 56–95 × 6–17, L=71.2, W=10.5, Q=7.19 | Allantoid, 3.9–5.2 × 1.2–1.6, L=4.56, W=1.33, Q=3.31–3.51 |
| H. rimosa | Dead or living angiosperm branches; temperate East Asia | Resupinate, sterile stage sometimes present; hymenial surface grey to pale brown, margin slightly lighter or concolorous | Rather blunt-pointed, 53–131 × 7–18, L=74.6, W=10.3, Q=7.41 | Cylindrical, 4.2–6.3 × 1.8–2.3, L=5.22, W=1.99, Q=2.47–2.78 |
| H. tabacina | Angiosperms, rarely gymnosperms; holarctic | Effused-reflexed; hymenial surface greyish brown to brown, margin slightly lighter or concolorous | Rather sharp-pointed, 54–120 × 7–17, L=78.6, W=11.4, Q=7.02 | Cylindrical, 4.1–6.2 × 1.5–2.1, L=4.93, W=1.81, Q=2.40–2.99 |
| H. yasudai | Pinus spp. (dead branches), rarely other gymnosperms; boreal/temperate East Asia | Effused-reflexed; hymenial surface bright ferruginous-brown, margin slightly lighter or concolorous | Sharp-pointed, 61–104 × 10–17, L=84.0, W=12.0, Q=7.07 | Cylindrical, 4.4–6.1 × 2.0–2.3, L=5.28, W=2.09, Q=2.53 |
Eleven generic names have been used for hymenochaete-like fungi in addition to Hydnoporia. Eight of them belong to the Hymenochaete clade (Cerrenella, Cyclomyces, Cycloporellus, Dichochaete, Hydnochaete, Hymenochaete, Hymenochaetella, Stiptochaete), and two to the Hydnoporia clade (Pseudochaete, Hymenochaetopsis). One genus, Clavariachaete, has not been sequenced. Below we summarize information for these genera.
Cerrenella Murrill, Bull. Torrey Bot. Cl. 32: 361. 1905.
Type species: Irpex tabacinus Berk. & Curt. (current name Hymenochaete odontoides S.H. He & Y.C. Dai)
He & Dai (2012) transferred the type species to Hymenochaete technically, based on sequences of Chinese material while the type originates from South Carolina. Authors of their nomen novum were incorrectly ascribed in the original paper (reference to Berk. & M.A. Curtis ex Fr. should not be used), but such a technical error does not affect validity of the new name (ICN Art. 41.6). Based on the description of Ryvarden (1982a) and our experience it is clear that this transfer was justified.
Clavariachaete Corner, Annals of Botany Memoirs 1: 689. 1950.
Type species: Clavariachaete rubiginosa (Berk. & M.A. Curtis ex Cooke) Corner.
When Corner (1950) described Clavariachaete, he mentioned confusingly that the genus is based on the bogus Clavariachaete McGinty, typified by C. peckoltii, while at the same time specifying C. rubiginosum as the generic type (page 689). The McGinty names by Lloyd are generally considered invalid, a filament of Lloyd’s sense of humour (Stevenson & Cash 1936). Thus, the valid description is that of Corner, and the valid generic type is C. rubiginosa, as generally acknowledged.
Clavariachaete reminds morphologically quite closely Hymenochaete and does not fit Hydnoporia: regular, middle-sized setae, unambiguously monomitic hyphal structure and broad ellipsoid spores (Parmasto 2010). We are confident that Clavariachaete is not congeneric with Hydnoporia, but whether the genus is a taxonomic synonym of Hymenochaete or a well-defined genus of its own, remains to be seen.
Parmasto (2010) described a new family, Clavariachaetaceae, based on this genus. Whereas the genus Clavariachaete may stand time, we do not think that the family is justified. Clavariachaete fits very well in the Hymenochaetaceae.
Cyclomyces Fr., Linnea 5: 512. 1830.
Type species: Cyclomyces fuscus Fr. [current name Hymenochaete campylopora (Mont.) Spirin & Miettinen].
All the species sequenced thus far, including the type species, belong to the Hymenochaete clade with the exception of C. lamellatus (Wagner & Fischer 2002, He & Dai 2012, Parmasto et al. 2014). While transferring C. fuscus to Hymenochaete, Wagner & Fisher (2002) changed its name to Hymenochaete cyclolamellata T. Wagner & M. Fisch, to avoid homonymy with Hymenochaete fusca P. Karst. However, they overlooked Polyporus campyloporus Mont., considered a synonym of C. fuscus (Ryvarden 1982b), which can serve as a basionym if C. fuscus is unavailable. Cyclomyces tabacinus presents a similar case: the epithet is not available in Hymenochaete and it was replaced with Hymenochaete porioides nom. nov. by Wagner & Fischer (2002). They overlooked that Polyporus microcyclus Zipp. ex Lév. is available as a taxonomic synonym (Ryvarden 1981), and its epithet is not previously occupied in Hymenochaete. Hence new combinations:
Hymenochaete campylopora (Mont.) Spirin & Miettinen, comb. nov. MB830586.
Basionym: Polyporus campyloporus Mont., Ann. Sci. Nat., Bot., sér. 4, 1: 132. 1854.
Synonyms: Cyclomyces fuscus Kunze ex Fr., Hymenochaete cyclolamellata T. Wagner & M. Fisch.
Hymenochaete microcycla (Zipp. ex Lév.) Spirin & Miettinen, comb. nov. MB830587.
Basionym: Polyporus microcyclus Zipp. ex Lév., Ann. Sci. Nat., Bot., sér. 3, 2: 188. 1844.
Synonyms: Cyclomyces tabacinus (Mont.) Pat., Hymenochaete porioides T. Wagner & M. Fisch.
Cycloporellus Murr., Bull. Torrey Bot. Cl. 34: 468. 1907.
Type species: Polyporus iodinus Mont. [syn. Cycloporellus iodinus (Mont.) Murrill, Cyclomyces iodinus (Mont.) Pat., current name Hymenochaete iodina (Mont.) Baltazar & Gibertoni].
Before the revision of Hymenochaete, Cycloporellus was considered a taxonomic synonym of Cyclomyces (Ryvarden 1991). The type species H. iodina is morphologically very close to sequenced species of other poroid Hymenochaete such as H. campylopora (= Cyclomyces fuscus) and H. microcycla (= Cyclomyces tabacinus) (Gomes-Silva et al. 2012). It is clear that H. iodina belongs to the Hymenochaete clade and Cycloporellus should be considered a taxonomic synonym of Hymenochaete for now.
Dichochaete Parmasto, Folia Crypt. Estonica 37: 56. 2000.
Type species: Thelephora setosa Sw. [syn. Dichochaete setosa (Sw.) Parmasto, current name Hymenochaete resupinata (Sw.) Parmasto].
Parmasto (2000) described this genus for two Hymenochaete species (the type H. setosa and H. ceratophora) based on dichohyphidia. Dichohyphidia are found and even more conspicuous in other members of Hymenochaete such as H. cruenta (Léger 1998). The type species of Dichochaete (under the name Hymenochaete resupinata) as well as H. cruenta have been shown to belong to Hymenochaete by molecular means (Parmasto et al. 2014).
Hydnochaete Bres, Hedwigia 35: 287. 1896.
Type species: Hydnochaete badia Bres. [syn. Hydnochaete peroxydata (Berk. ex Cooke) Dennis, current name Hymenochaete peroxydata (Berk. ex Cooke) Baltazar et al.].
The type species has been sequenced and belongs to Hymenochaete (Baltazar et al. 2014). Also H. asetosa, H. duportii, H. japonica, H. paucisetigera, H. resupinata and H. tabacina (=Hymenochaete odontoides) belong to Hymenochaete (Wagner & Fischer 2002, He & Dai 2012, Parmasto et al. 2014) but Hydnochaete tabacinoides has been shown to belong to Pseudochaete (= Hydnoporia in current sense; He & Dai 2012). Based on the type material of Hydnochaete saepiaria we have studied, the species belongs to Hymenochaete.
Hymenochaete saepiaria (Lloyd) Spirin & Miettinen, comb. nov. MB830588.
Basionym: Irpex saepiarius Lloyd, Mycol. Writings 5(48): 682. 1917.
Hymenochaete Lév., Ann. Sci. Nat., Bot., sér. 3, 5: 150. 1846.
Type species: Helvella rubiginosa Dicks. [current name Hymeno-chaete rubiginosa (Dicks.) Lév.]
The name is conserved against Cyclomyces Kunze ex Fr and Hymenochaeta P. Beauv. ex T. Lestib. (Angiospermae) (ICN Appendix III).
Parmasto et al. (2014) demonstrated that the type species of Hymenochaete and Hydnoporia (as Pseudochaete olivacea) belong to distinct clades separated by several polypore genera phylogenetically. Since Wagner & Fischer (2002) Hymenochaete has been expanded considerably to include also hydnoid and poroid taxa that belong to a well-supported clade in DNA-based analyses, while excluding species of the Hydnoporia clade. We discuss Hymenochaete here in that wide sense.
Hydnoporia and Hymenochaete are morphologically very similar. Differences are listed in Table 5. As can be expected in the case of such a large and variable genus as Hymenochaete, not all characters work for each single species. However, combination of characters (Table 5) does work, and with careful observations, it is possible to separate Hydnoporia and Hymenochaete morphologically.
Table 5.
Comparsion of general morphological differences between Hydnoporia and Hymenochaete.
| Characters | Hymenochaete | Hydnoporia |
|---|---|---|
| Hyphal system | Monomitic or dimitic with true skeletal hyphae | Resupinate species: monomitic with sclerified basal hyphae; effused-reflexed species: dimitic with less distinct differences between hyphal types, often with setal hyphae |
| Setae | Rather stable in length and width, originating mostly from subhymenium or lower trama | Variable in size and mostly very wide, originating at different levels of trama |
| Subhymenium | Rather loosely arranged, often short and inflated cells | Densely arranged and only slightly inflated cells |
| Basidia | Suburniform with a clear medial constriction | Narrowly clavate or sinuous, not constricted |
| Spores | Broadly ellipsoid to cylindrical, rarely allantoid | Cylindrical to allantoid and often glued together |
| Mycelial mats infecting adjacent substrate | Absent | Present in some if not all species (Stenlid & Holmer 1991, Parmasto 2001, our observations) |
The typification of the generic type, H. rubiginosa, requires clarification. Hymenochaete rubiginosa has been conserved as the generic type of Hymenochaete (ICN Appendix III). However, the original English material of Helvella rubiginosa Dicks. which could be used for the species typification does not exist (Parmasto 2001). Job (1990) selected Lloyd’s collection from Ohio as a neotype of H. rubiginosa, but we disagree with his decision. According to our analysis of ITS sequences, the current concept of H. rubiginosa covers probably at least four species: European (i.e. H. rubiginosa s. str.), two unnamed sibling species from North America, and one from East Asia (Fig. 4). The type of H. rubiginosa should have been designated from European material.
Phylogram of Hymenochaete rubiginosa and closely related species. Bayesian consensus tree based on nrDNA ITS sequences with midpoint rooting. Numbers denote posterior probabilities. Two-letter codes after species names denote country, followed by the province of origin (ISO 3166). Asian specimens have been coloured blue, European green and North American red. Sequences with the accession number starting with “MK” have been produced for this study, the rest are from public sources.
For this reason, we suggest to typify H. rubiginosa with an authentic specimen from the Fries herbarium in Uppsala: Sweden. Småland: Femsjö, Fries (UPS F-175700, lectotype of Helvella rubiginosa Dicks. designated here, MBT386587). This specimen was with high likelihood collected prior to 1821 when the sanctioning work Systema Mycologicum was published (Fries 1821) and is suitable as a lectotype (ICN Art. 9.3). Our reasoning is as follows: Fries used the name Stereum rubiginosum for this species in Fries (1818: 274), and this is how Fries has labelled the lectotype (his handwriting is visible). In Systema Mycologicum (Fries 1821) he switched to the name Thelephora rubiginosa for this species. Thus, the lectotype predates the sanctioning and is the only authentic specimen of this species in Fries’s herbarium.
The four relatives of H. rubiginosa in North America and East Asia need other names in the future. For the time being they can be labelled H. rubiginosa coll. One of them is labelled H. ochromarginata (He & Dai 2012), but because H. ochromarginata was described from Africa, its applicability to Chinese material should be critically assessed.
Hymenochaetella P. Karst., Bidrag till Kännedom av Finlands Natur och Folk 48: 428. 1889.
Type species: Hymenochaetella arida P. Karst, chosen by Cooke (1953) [current name Hymenochaete cinnamomea (Pers.) Bres.].
The type species belongs to Hymenochaete, as shown by Wagner & Fischer (2002) and confirmed by subsequent authors.
Hymenochaetopsis S.H. He & Jiao Yang, Mycol. Prog. 15: 2. 2016.
Type species: Auricularia tabacina Sowerby [syn. Hymenochaetopsis tabacina (Sowerby) S.H. He & Jiao Yang, current name Hydnoporia tabacina (Sowerby) Spirin et al.].
Nomen novum for Pseudochaete, a taxonomic synonym of Hydnoporia.
Pseudochaete T. Wagner & M. Fischer, Mycol. Prog. 1: 100. 2002.
Type species: Auricularia tabacina Sowerby [syn. Pseudochaete tabacina (Sowerby) T. Wagner & M. Fisch., current name Hydnoporia tabacina (Sowerby) Spirin et al.].
Illegitimate name due to ICN Art. 53.1 and Pseudochaete W. West & G.S. West 1903. A taxonomic synonym of Hydnoporia as the close relation between type species was shown by Parmasto et al. (2014) and confirmed here.
Stipitochaete Ryvarden, Trans. British Mycol. Soc. 85: 537, 1985.
Type species: Stereum damicorne Link. [syn. Stipitochaete damicornis (Link) Ryvarden, current name Hymenochaete damicornis (Link) Lév.].
Ryvarden (1985) described this genus for two species of Hymenochaete with stipitate basidiocarps. Wagner & Fischer (2002) showed that the type species belongs to Hymenochaete.
Accepted species
Hydnoporia corrugata (Fr. : Fr) K.H. Larss. & Spirin, comb. nov. MycoBank MB830589. Figs 5, ,66.
Spores of Hydnoporia spp.
Setae of Hydnoporia spp.
Basionym: Thelephora corrugata Fr., Obs. Mycol. 1: 134. 1815.
Typus: Norway, Hordaland, Bømlo, Corylus avellana, 59.7002°: 5.2645°, 24 Jul. 2011, Klepsland 11.021 (O F-247869) (neotype designated here, MBT386588).
Notes: According to our data, H. corrugata is a European species (described from Sweden by Fries 1815). It usually inhabits dead, still attached or recently fallen branches of deciduous trees, especially Betula spp., Carpinus betulus and Corylus avellana.
Hydnoporia corrugata differs from its closest relatives, North American H. diffissa and East Asian H. rimosa, by having distinctly narrower basidiospores (see Tables 1, ,4).4). Chinese and Korean sequences labelled as ‘H. corrugata’ most probably belong to a different, closely related species (Fig. 2).
Hydnoporia diffissa Spirin & Miettinen, sp. nov. MycoBank MB830590. Figs 5, ,66.
Typus: USA, North Carolina, Buncombe Co., Blue Ridge Assembly, 35.58356° : -82.34159° standing dead Rhododendron maximum, 24 Sep. 2015, Miettinen 19463 (holotype H 7008917).
Etymology: Diffissum, derived from diffindo (Lat., v.), named after the cracking hymenophore.
Basidiocarps perennial, effused, leathery, 0.1–0.5 mm thick, covering several cm. Margin sharply delimited and firmly attached to the substrate, first concolorous with hymenium, later ferruginous brown (in mature basidiocarps normally darker than hymenial surface). Hymenial surface even or indistinctly tuberculate, first grey to greyish ochraceous, later pale chocolate brown, in mature basidiocarps strongly rimose. Sterile patches and films bright ferruginous brown, rather small-sized, compact, associated with basidiocarps or growing solitarily. Section: Subiculum poorly visible, dark brown to almost black, about 20 μm thick; trama cream coloured to pale ochraceous, 30–50 μm thick; hymenium / subhymenium grey to pale brown, 30–40 μm thick, with dark coloured, mostly embedded setae. Hyphal system monomitic though appearing dimitic; hyphae with simple septa. Setal hyphae present in sterile films and in basal parts of senescent basidiocarps, sharp- to blunt-pointed, very thick-walled, lateral or pleural, brown, up to 170 × 12 μm. Skeletal-like hyphae often present in subiculum and lowermost parts of trama, brownish to dark-brown, occasionally branched and septate, 2–3.5 μm in diam. Generative hyphae mostly with thickened walls, in sterile films yellowish to brownish, 2–4.5 μm in diam, irregularly arranged to subparallel; in subicular layer and upper trama pale to dark brown, clearly thick-walled, more or less horizontal and parallel to the substrate, in lower trama and subhymenium hyaline to pale brown, mostly vertically arranged (2.1–)2.2–3.3(–3.6) μm diam (n = 20/1). Tramal / hymenial setae sharp- to blunt-pointed, normally with distinctly widened basal part, dark ferruginous brown, 47–73(–90) × 6.2–12.9 μm, L = 61 μm, W = 9.1 μm, Q = 6.82, sometimes pleural (originating from thick-walled, dark brown hyphae in upper trama) and with an apical crystalline cape, embedded or projecting up to 15–50 μm above hymenial cells, associated with simple hyaline hyphidia 1–2.5 μm in diam. Basidia narrowly clavate, 4-spored, 12.2–16.2(–17.2) × (2.8–)3.0–4.1 μm (n = 20/2). Basidiospores broadly cylindrical to cylindrical, straight to slightly curved, (4.1–)4.2–7.3(–7.8) × 1.7–2.2 μm, L = 5.28 μm, W = 1.92 μm, Q = 2.74.
Distribution and ecology: Hard angiosperm wood; our material is from the Eastern United States, but ITS sequences indicate it is present also in Colombia and Peru (Fig. 2).
Notes: Here we describe this species to encompass ‘H. corrugata’ from the American Northeast. Four older names from North America had been listed among synonyms of H. corrugata but none of them, in our opinion, is applicable to H. diffissa. Three of them, Thelephora episphaeria, Hymenochaete agglutinans and H. ambiens are treated under H. olivacea (see below) while the identity of H. insularis is vague. Nevertheless, the white tomentose margin mentioned in the protologue (Berkeley 1873) and by Burt (1918), who studied the type material of H. insularis, precludes H. diffissa. Morphologically, H. diffissa is almost indistinguishable from the East Asian H. rimosa, except for the more variable setae in the latter species (Table 4).
Hydnoporia gigasetosa (Parmasto) Miettinen & K.H. Larss., comb. nov. MycoBank MB830591.
Basionym: Hymenochaete gigasetosa Parmasto, Folia Cryptog Estonica 40: 41. 2003.
Notes: Yang et al. (2016) reported this Indian species as new to China and concluded based on sequences that it belongs to Hymenochaetopsis (i.e. Hydnoporia). Their description agrees with the original description of Parmasto (2003). Large setae, allantoid basidiospores and the shape of basidia all confirm that H. gigasetosa belongs to Hydnoporia.
Hydnoporia lamellata (Y.C. Dai & Niemelä) Miettinen & K.H. Larss., comb. nov. MycoBank MB830592.
Basionym: Cyclomyces lamellatus Y.C. Dai & Niemelä, Ann. Bot. Fenn. 40: 384. 2003.
Notes: Hydnoporia lamellata is an East Asian species morphologically and phylogenetically very close to H. tabacinoides. Both species share all important features with other Hydnoporia species, and DNA data confirm their placement to the latter genus. It would be useful to compare the type of H. tabacinoides with sequenced material to confirm that the names H. lamellata and H. tabacinoides belong to separate species. Descriptions by Ryvarden (1982a) of H. tabacinoides and Dai et al. (2003) of H. lamellata let us believe that spore and pore size separate the two, but the description by Dai & Niemelä (2006) of H. tabacinoides muddles the picture. ITS data nevertheless indicates that there are two species in East Asia in this complex.
Hydnoporia laricicola (S.H. He & Jiao Yang) Spirin & Miettinen, comb. nov. MycoBank MB830593. Figs 5, ,66.
Basionym: Hymenochaetopsis laricicola S.H. He & Jiao Yang, Mycol. Prog. 15: 3. 2016.
Basidiocarps perennial, sessile or effused-reflexed, leathery to corky, imbricate, 1–3 cm long, 0.3–0.6 mm thick, resupinate part up to 6 cm wide. Pileal surface uneven, tomentose, matt, grey, with narrow, distinct, darker zones, in older basidiocarps ferruginous brown. Pileal margin sharp, more or less even, a bit incurved in dried specimens, margin of resupinate parts sharply delimited and partly detaching from substrate, ferruginous brown. Hymenial surface even or indistinctly tuberculate, first dark grey, later chocolate-coloured, in mature basidiocarps strongly rimose. Section: Tomentum rather loose, grey, 50–70 μm thick; cortex well visible, dark brown to almost black, about 20 μm thick; trama more or less uniformly greyish brown, 40–60 μm thick; hymenium / subhymenium grey to rusty brown, 30–50 μm thick, with dark coloured, mostly embedded setae. Perennial basidiocarps distinctly stratified (2–4 layers divided by dark lines). Hyphal system dimitic; hyphae with simple septa. Setal hyphae abundant, originating deeply in trama, blunt-pointed, very thick-walled, lateral or pleural, mostly vertically arranged, dark ferruginous brown, 90–300 × 4–12 μm. Skeletal hyphae rather poorly differentiated, located in the uppermost parts of basidiocarps, dark-brown, mostly unbranched and not septate, 2–4 μm in diam. Generative hyphae mostly with thickened walls, yellowish to brownish, 2.8–4.8 μm in diam (n = 20/1), vertically and tightly arranged in tomentum, more or less horizontal and parallel in cortex and upper tramal layer, mostly vertically arranger in lower trama; subhymenium distinct, 20–30 μm thick, hyphae pale brown, with thickened walls, vertically and tightly arranged, 2.4–4.1 μm in diam (n = 10/1). Tramal / hymenial setae blunt-pointed, normally with widened basal part, dark ferruginous brown, 31–69(–84) × 5.3–12.2 μm, L = 44 μm, W = 8.1 μm, Q = 5.41, rarely pleural or bifurcate, some with scarce encrustation, embedded or projecting up to 15–20 μm above hymenial cells, usually associated with simple hyaline hyphidia 1–2 μm in diam. Hymenial hyphidia present, abundant, hyaline to brownish, with thickened walls and a few slight constrictions (moniliform), 1–2.5 μm diam at the apical part, embedded or projecting up to 10 μm. Basidia narrowly clavate, 4-spored, 15.0–22.3 × 3.1–4.1 μm (n = 20/2). Basidiospores cylindrical, straight to slightly curved, (4.2–)4.3–6.8(–7.0) × (1.6–)1.7–2.1(–2.2) μm, L = 5.27 μm, W = 1.91 μm, Q = 2.78.
Distribution and ecology: Hydnoporia laricicola grows on both living and fallen trees of larch (Larix gmelinii and L. sibirica) in taiga forests of Eurasia. We have not seen material from Europe outside of Russia.
Notes: Its most striking microscopic characters are abundant setal hyphae and blunt-pointed, very short tramal and hymenial setae. Hydnoporia tabacina can also occur on coniferous hosts, including Larix, but it differs in having much larger setae (Table 2).
Hydnoporia latesetosa (S.H. He & Hai J. Li) Miettinen & K.H. Larss., comb. nov. MycoBank MB830594.
Basionym: Pseudochaete latesetosa S.H. He & Hai J. Li, Mycol. Prog. 12: 333. 2013.
Notes: This species was recently described from tropical China (He & Li 2013). Large setae of varying size, setal hyphae and narrow basidiospores, as well as DNA data confirm the transfer of this species to Hydnoporia.
Hydnoporia lenta (G.A. Escobar ex Léger) Spirin & Miettinen, comb. nov. MycoBank MB830595.
Basionym: Hymenochaete lenta G.A. Escobar ex Léger, Cryptog. Mycol. 11: 302. 1990.
Notes: Hydnoporia lenta was described from Costa Rica (Léger 1990), and we have not seen the type. However, the morphological description by Léger (1990, 1998) is sufficient to conclude that the species should be included in Hydnoporia.
Hydnoporia olivacea (Schwein.: Fr.) Teixeira, Rev. Brasil. Bot. 9: 43. 1986.
Basionym: Sistotrema olivaceum Schwein., Schr. Naturf. Ges. Leipzig 1: 101. 1822. — Hydnum olivaceum (Schwein.: Fr.) Fr., Elench. Fung. 1: 134. 1828.
Synonyms: Sistotrema fuscescens Schwein., Schr. Naturf. Ges. Leipzig 1: 102. 1822. Lectotype: USA, North Carolina, Burke Co., Salem, Schweinitz (UPS F-173549, designated here, MBT386589). Epitype: USA, Massachusetts, Worcester, Columbus Park, 42.2574° : -71.83458°, hardwood, 11 Apr. 2013, Miettinen 16044 & Käppi (H 7005770, designated here, MBT386590).
Irpex cinnamomeus Fr., Epicrisis: 524. 1838 (fide Ryvarden 1982a).
Thelephora episphaeria Fr., Elenchus Fung. 1: 225. 1828. Lectotype: USA, North Carolina, Burke Co., Salem, Schweinitz (UPS F-173519, designated here, MBT386591). Epitype: USA, New York, Essex Co., Huntington Wildlife Forest, Adjidaumo flow, 42.2574° : -71.83458°, Fagus grandifolia, 18 Sep. 2013, Miettinen 16956.2 (H 7008916, designated here, MBT386592).
Hymenochaete agglutinans Ellis, Bull. Torrey Bot. Club 5: 46. 1874. Lectotype: USA, New Jersey, Gloucester Co., Newfield, hardwood, 1874 Ellis (NY 00742784, designated here, MBT387111). Epitype: The same locality, Nov. 1881 Ellis (NY ex C. Roumeguére, Fungi selecti exsiccati #4540, designated here, MBT386593).
Sistotrema fuscescens and H. olivaceum were described in the same paper. This very common North American fungus is conspicuous and easy to identify among Hydnoporia with its irpicoid hymenophore. Material we have seen is uniform, and the following discussion focuses on nomenclature. However, it should be said that ITS sequences show some variation, more than is typical for most other Hydnoporia spp. (Fig. 2). Our own sampling is limited, so we refrain making any conclusions based on this.
Banker (1914) chose to synonymize S. fuscescens with S. olivaceum, and as he has been widely followed by later authors, the latter name should be used for this species. Ryvarden (1982a) designated a lectotype for S. olivaceum (Schweinitz 540, PH). Parmasto et al. (2014) studied the lectotype as well as an isolectotype in UPS (Parmasto 2001). There are two specimens, possibly parts of the same collection, with the same Schweinitz number (540), but filed under different herbarium numbers. This makes it necessary to amend Ryvarden’s lectotypification. Lectotype for Sistotrema olivaceum Schwein., designated here: USA, Pennsylvania, Salem, PH00062632 (MBT386594).
Since S. fuscescens is the type species of Hydnoporia, we need to secure the identity of the species through typification. Banker (1914) stated a type probably does not exist, and mentioned only one specimen from the Schweinitz collection, ‘580–7 Syn. Fung. I. cinnamoneus Epic. 19. Irpex fuscescens Schw. Beth.’, which represents typical H. olivacea. That specimen is not available as a lectotype, because it was evidently collected in Bethlehem, Pennsylvania, where Schweinitz moved in 1821 (Rogers 1977), and not in North Carolina, the type locality (Schweinitz 1822). Schweinitz’s herbarium in PH is fully available online under plants.jstor.org, and it does not contain any other specimens under the name S. fuscescens. However, an authentic specimen of S. fuscescens from locus classicus exists in Fries’s herbarium in Uppsala, and therefore we designate it as the lectotype here. After considering the intraspecific ITS variation and the fact that the type specimen is sterile, we also designate a modern, sequenced epitype.
Thelephora episphaeria was described by Fries (1828) based on a Schweinitz collection from North Carolina. Parmasto (2001) selected another Schweinitz’s collection from Pennsylvania to serve as a type (Bethlehem, on Diatrype, Schweinitz 723, PH 00077588). In our opinion, this choice cannot be supported because Fries must be considered the single author of this species and material seen by him should be selected for typification. Since only one specimen exists in Fries’s herbarium, collected by Schweinitz, it should be designated as the lectotype of T. episphaeria (indicated above). In older literature, T. episphaeria was considered very close or probably identical to H. corrugata, although with some reservations (Bondartseva & Parmasto 1986, Parmasto 2001). After studying the lectotype of H. episphaeria, we cannot agree with this opinion - both setal and spore measurements certainly point towards H. olivacea. The latter species is able to produce rather thin and completely smooth basidiocarps. In this condition, it can be mistaken for other Hydnoporia species. Here we designate a recent, sequenced specimen of H. olivacea with a smooth hymenium as an epitype for T. episphaeria.
Another synonym of H. olivacea introduced here is Hymenochaete agglutinans. This species was described from New Jersey (Ellis 1874) and it represents sterile films on branches of angiosperm trees and shrubs (Graves 1914, Burt 1918, Parmasto 2001). Reeves & Welden (1967) placed H. agglutinans in the synonyms of H. corrugata although their opinion on connection of sterile mats with the latter species was merely a matter of belief. We studied the original specimen of H. agglutinans from NY (designated above as the lectotype) and cannot agree with Reeves & Welden, because the NY specimen belongs to a truly dimitic species with wide setae. Moreover, there is another specimen of H. agglutinans collected by Ellis in or near the type locality in 1881 (distributed in C. Roumeguére’s Fungi selecti exsiccati, #4540), which represents a basidiocarp developed on the surface of a sterile patch. Morphologically, the sterile stage of this collection is identical to the H. agglutinans lectotype, composed of very thick-walled, almost unbranched and dark-coloured skeletal hyphae and sharp-pointed setal hyphae, while the basidiocarp is undoubtedly a young specimen of H. olivacea with irregularly poroid to irpicoid hymenophore. Therefore, we consider H. agglutinans a synonym of H. olivacea. Burt (1918) stated that the type material of H. ambiens, also described from New Jersey, represents the same sterile stage as H. agglutinans.
Hydnoporia rhododendri (Corfixen & Parmasto) Spirin & Miettinen, comb. nov. MycoBank MB830596. Figs 5, ,66.
Basionym: Hymenochaete rhododendri Corfixen & Parmasto, Karstenia 57: 65. 2017.
Basidiocarps annual or perennial, effused-reflexed, leathery, first cupulate or umbonate, 0.5–1 cm in diam, 0.2–0.4 mm thick, later fusing together and up to 10 cm in longest dimension; reflexed part 1.5–4 mm wide. Pileal surface uneven, finely tomentose, matt, first indistinctly zonate and ferruginous brown, later darkening to blackish brown. Pileal margin sharp, more or less even, a bit incurved in dried specimens, margin of resupinate parts sharply delimited and partly detaching from the substrate, first pale cream coloured to yellowish, then concolorous with hymenium. Hymenial surface even or indistinctly tuberculate, pale brown, in older basidiocarps often rimose, with greyish hues. Section: Tomentum loose, brown, 20–50 μm thick; cortex poorly visible, dark brown, 10–30 μm thick; trama duplex, upper layer loose, straw-yellow to pale ochraceous, 60–80 μm thick, lower layer dense, dark brown, 40–80 μm thick; hymenium / subhymenium pale cream to pale ochraceous, 40–60 μm thick, with dark coloured embedded or projecting setae. Hyphal system dimitic; hyphae with simple septa. Setal hyphae originating deeply in upper tramal layer, sharp-pointed, very thick-walled, lateral or pleural, some distinctly bent, a few vertically arranged and penetrating hymenial layer, dark ferruginous brown, 90–240(–350) × 5.2–9.4 μm (n = 11/2). Skeletal hyphae rather poorly differentiated, located in the uppermost parts of basidiocarps, dark-brown, mostly unbranched and not septate, 2–3.5 μm in diam. Generative hyphae mostly with thickened walls, yellowish to brownish, 2.3–4.0 μm in diam (n = 40/1), vertically and tightly arranged in tomentum, more or less horizontal and parallel in cortex and upper tramal layer, irregularly interwoven to subparallel in lower tramal layer; subhymenium distinct, 20–40 μm thick, hyphae hyaline, with thin or slightly thickened walls, vertically and tightly arranged, 2.1–3.1 μm in diam (n = 10/1). Tramal / hymenial setae mostly sharp-pointed, normally with widened basal part, dark ferruginous brown, 53–99 × 6.2–18.7 μm, L = 71 μm, W = 10.5 μm, Q = 7.19, rarely pleural or bifurcate, some with scarce encrustation, embedded or projecting up to 15–40 μm above hymenial cells, usually covered by semi-translucent sheaths of hyphidia. Basidia narrowly clavate, 4-spored, 11.9–19.8(–22.0) × 2.4–3.2(–3.6) μm (n = 40/3). Basidiospores allantoid, slightly to distinctly curved, (3.8–)3.9–5.2(–5.6) × (1.1–)1.2–1.6(–1.7) μm, L = 4.56 μm, W = 1.33 μm, Q = 3.44.
Distribution and ecology: Grows on branches of Ericaceae. Widely distributed although uncommon in mountains of temperate Eurasia.
Notes: Hydnoporia rhododendri is phylogenetically a sister species of H. tabacina, but should be easily identifiable by its narrow and rather short spores and its host. The Central American H. lenta has similar spores but it differs in having distinctly shorter (45–65 × 8.5–16 μm), blunt setae with wide lumen (Léger 1990). The only European specimen (TAAM 189353) sequenced for this study shows 9 bp difference in ITS region from two East Asian collections, and no tef1 sequence is available for it. Because of negligible morphological differences between European and Asian specimens, we currently consider them to belong to the same species.
Hydnoporia rimosa (Lloyd) Spirin & Miettinen, comb. nov. MycoBank MB830597. Figs 5, ,66.
Basionym: Hymenochaete rimosa Lloyd, Bot. Mag. Tokyo 44: 92. 1930.
Typus: Japan, Tôhoku, Sendai, 24 Oct. 1920, Yasuda (TNS F-203210, lectotype designated here, MBT387146, studied).
Basidiocarps perennial, effused, leathery, 0.1–0.2(–0.3) mm thick, covering several cm. Margin sharply delimited and firmly attached to the substrate, first whitish, later ferruginous brown (in mature basidiocarps normally darker than hymenial surface). Hymenial surface even or indistinctly tuberculate, first grey to greyish ochraceous, later pale chocolate brown, in mature basidiocarps strongly rimose. Sterile patches and films bright ferruginous brown, up to 7 cm in longest dimension, rather firm, associated with basidiocarps or growing solitary. Section: Subiculum poorly visible, first pale coloured, in perennial basidiocarps dark brown to almost black, about 20 μm thick; trama cream coloured to pale ochraceous, 30–50 μm thick; hymenium / subhymenium grey to pale brown, 30–40 μm thick, with dark coloured, mostly embedded setae. Perennial basidiocarps stratified (2–3 layers divided by dark lines). Hyphal system monomitic; hyphae with simple septa. Setal hyphae present in sterile films, blunt-pointed, very thick-walled, lateral or pleural, brown, 75–205 × 6–9.5 μm. Skeletal-like hyphae often abundant in subiculum and lowermost parts of trama, brownish to dark-brown, occasionally branched and septate, 2–4.5 μm in diam. Generative hyphae mostly with thickened walls, in sterile films yellowish to brownish, 3–5.5 μm in diam, irregularly arranged to subparallel; in subicular layer and upper trama pale to dark brown, clearly thick-walled, more or less horizontal and parallel to the substrate, in lower trama and subhymenium hyaline to pale brown, mostly vertically arranged (2.1–)2.3–3.6(–4.0) μm in diam (n = 20/2). Tramal/hymenial setae rather blunt-pointed, normally with distinctly widened basal part, dark ferruginous brown, 50–144 × 6.6–18.2 μm (n = 100/5), L = 75 μm, W = 10.3 μm, Q = 7.41, often pleural and with apical crystalline cape, embedded or projecting up to 15–60 μm above hymenial cells, associated with simple hyaline hyphidia 1.5–3.5 μm in diam. Basidia narrowly clavate, 4-spored, (11.1–)11.3–18.5(–20.3) × (3.2–)3.5–4.3 μm (n = 20/3). Basidiospores cylindrical, straight to slightly curved, (4.1–)4.2–6.3(–6.9) × (1.8–)1.9–2.3(–2.5) μm, L = 5.22 μm, W = 1.99 μm, Q = 2.64.
Distribution and ecology: Occurs on both living and on recently dead branches of angiosperms in East Asia.
Notes: Hydnoporia rimosa is an East Asian relative of H. corrugata and H. diffissa. It seems to be a pathogen, starting to develop abundant sterile mycelium while the host species is still alive and continuing to grow after the host dies. We have no Japanese sequenced material, but morphologically the type fits well with sequenced specimens from Russian Far East. However, our application of the name H. rimosa should be revised against material from Honshu. Sequence data shows that there is a fourth species in this complex in China and Korea (Fig. 2).
Hydnoporia rimosa was was described in Ito (1930a, as Hymenochaete r.). Ito directly cited Lloyd’s description of the species and the correct species author is Lloyd. Ito also provided a list of specimens, adding to Lloyd’s description. Both components are part of the protologue (see ICN Art. 6.13 footnote), and therefore the specimens Ito listed are to be considered syntypes, which have priority over other original material when selecting the lectotype (Art. 9.12). This is relevant because Stevenson & Cash (1936) chose a lectotype for H. rimosa, from Lloyd’s herbarium. That specimen may have been part of the original material, but not a syntype since it was not cited in the protologue. Thus our lectotypification supersedes that of Stevenson & Cash (Art. 9.19).
Hydnoporia subrigidula (S.H. He & Hai J. Li) Miettinen & K.H. Larss., comb. nov. MycoBank MB830598.
Basionym: Pseudochaete subrigidula S.H. He & Hai J. Li, Mycol. Prog. 12: 335. 2013.
Notes: The species was introduced as a close relative of Hymenochaete rigidula from China (He & Li 2013). However, identity of the latter species is still unclear (see discussion under Excluded species). Nevertheless, the morphological description as well as available DNA sequences confirm that P. subrigidula is a member of Hydnoporia.
Hydnoporia tabacina (Sowerby) Spirin, Miettinen & K.H. Larss., comb. nov. MycoBank MB830599.
Basionym: Auricularia tabacina Sowerby, Coloured Fig. English Fungi 1: 14. 1797.—Thelephora tabacina (Sowerby) Fr., Systema Mycologicum 1: 437. 1821.
Typus: Sweden, Skåne, Fries (UPS F-175737, lectotype designated here, MBT386595, studied).
Synonym: Stereum intricatum Lloyd, Mycol. Writings 7(67): 1157. 1922. Typus: Japan, Hokkaido, Tomakomai, Iburi, Alnus, 20 Dec. 1904, Niidjima 706 [TNS F-200336, lectotype by Ito (1930b), studied].
Notes: We consider H. tabacina a Holarctic species in this paper. There is, however, some minor variation in ITS (Fig. 2) and clearly two separate tef1 groups (Fig. 2). European specimens from Nizhny Novgorod, Russia, England and Finland show a divergent tef1 sequence of 11 bp (1.3 %) from the rest of the material, though the easternmost European specimen from Perm (Kotiranta 25205) falls within the Asian-North American clade. The five European ITS sequences (excluding the Perm specimen) show a 1 bp difference against the rest as well, in line with the tef1 division. We see no differences between these genotypes in morphology or host species range. The species inhabits branches and more rarely logs of many angiosperm trees and shrubs but sometimes occurs also on conifers (especially on Picea).
Clearly, the H. tabacina complex deserves to be studied further to determine if more than one species is involved. Answering this question is beyond the scope of this paper, and will require more extensive sampling particularly from North America and Europe, with sequences of several fast-evolving markers. If the tef1 genotypes would turn out to represent different species, then the European genotype should be called H. tabacina. There would be several American and Asian old names to choose from for the Asian-North American species as discussed below.
Hydnoporia tabacina was described many times. Its oldest names include Helvella nicotiana Bolton 1792 (England, Syringa), Auricularia tabacina Sowerby 1797 (England, angiosperms), Thelephora avellana Fr. 1821 (Sweden, Corylus), T. cerasi Pers. 1822 (France, Cerasus), T. juratensis Pers. 1822 (France, Fagus), T. crocata Fr. 1828 (South Europe, angiosperms), and T. imbricatula Schwein. 1832 (USA). Fries (1821: 437) sanctioned Sowerby’s A. tabacina, and we select here a lectotype from Sweden in accordance with ICN Art. 9.3 for fixing its concept.
The choice of lectotype requires justification. Fries treated the species in the sanctioning work Systema Mycologicum (Fries 1821: 437), where he mentions that he has seen fresh material (“v. v.”) and that the main host is Corylus. The specimen we selected as a lectotype was collected from Skåne by Fries and labelled Stereum tabacinum. Its host is beyond doubt Corylus avellana. As usual, no collecting date is provided. He himself mentions material of H. tabacina from Skåne (under the name Stereum ferrugineum) in Observationes 2 (Fries 1818: 274). Strid (1995: 44) points out that Observationes were “largely based on material that he gathered between 1813 and 1815”. Later Fries (1857) wrote that he received several collections from Skåne in 1819, which prompted him to describe several species in Systema Mycologicum. Whichever the case, all indications are that the specimen from Corylus from Skåne was collected before the sanctioning and was used as a source of the sanctioning description.
Hymenochaete konradii from the Czech Republic was described as having “dendrophyses” in hymenium (Léger 1985), and it is the only difference from H. tabacina. We observed those branched hymenial cells in several typical specimens of the latter species, and this character is certainly not sufficient for species delimitation in this group. Thus, H. konradii might be a synonym of H. tabacina, but this needs to be confirmed. Bresadola (1920) added Daedalea lirellosa Pers. to the synonyms of H. tabacina, but his conclusion seems to be wrong; we will discuss this question in a forthcoming paper.
Hymenochaete intricata was described from Japan (Ito 1930b). This species was considered as widely distributed on angiosperm substrates in Northeast China and Russian Far East, while H. tabacina was supposedly absent from this area (Bondartseva & Parmasto 1986, Dai 2010). There are small differences in spore and setal sizes between East Asian and other Eurasian material of H. tabacina (Tables 1, ,2).2). However, our DNA data (ITS and tef1) do not support separation of East Asian specimens, i.e. H. intricata, from other Asian and North American specimens of H. tabacina.
Hydnoporia tabacinoides (Yasuda) Miettinen & K.H. Larss., comb. nov. MycoBank MB830601.
Basionym: Irpex tabacinoides Yasuda, Bot. Mag. Tokyo 33: 189. 1919.
Notes: Results by He & Dai (2012) indicate that this species belongs to Hydnoporia. Ryvarden (1982a) provided a modern description of the species based on Japanese material (as Hydnochaete tabacinoides). He considered it similar to Hydnoporia olivacea. The description by Dai & Niemelä (2006) does not fully agree with Ryvarden’s, for instance the spore size is different. While it appears that sequences published as H. tabacinoides from China represent a separate species, they should be checked against authentic material to ascertain their identity. In any case, all morphological descriptions indicate that H. tabacinoides is a member of Hydnoporia.
Hydnoporia yasudai (Imazeki) Spirin & Miettinen, comb. nov. MycoBank MB830602.
Basionym: Hymenochaete yasudai Imazeki, Bull. Tokyo Sci. Mus. 2: 6. 1940.
Typus: Japan, Tokyo, Minami-Tama Gun, Sainokami Goryochi, Abies firma (dry branches), 10 Oct. 1939, Imazeki (holotype TNS F-207357, studied).
Notes: Hydnoporia yasudai is an East Asian species inhabiting dry branches and bark of Pinus spp., rarely other coniferous trees. It was described from Japan (Imazeki 1940), and found also in China (Dai 2010) and Russian Far East (Bondartseva & Parmasto 1986). Basidiocarps of H. yasudai are usually sterile; however, the species is easily identifiable due to peculiar moniliform, thick-walled hyphidia and its host preferences. Basidiospores of H. yasudai are similar to other Hydnoporia species, cylindrical and rather small, 4.4–6.1 × 2–2.3 μm, not ellipsoid and large as described in some recent manuals (Bondartseva & Parmasto 1986, Dai 2010).
ITS data indicates that H. yasudai is a complex containing three to six species (Fig. 2). Our collections from Russia derive from Pinus pumila and the type from Abies firma in Japan. The type is in bad condition and contains very few spores. It is otherwise similar to the Russian specimens we have sequenced except that the Russian specimens are bright-coloured whereas the type is quite dull, dark brown. For now, the Russian specimens are best called H. yasudai, but clearly Japanese, sequenced material is needed to settle nomenclature in the complex.
Excluded species
Hymenochaete allantospora Parmasto, Folia Crypt. Estonica 37: 58. 2000.
Notes: Allantoid basidiospores of this species are the only morphological feature, which might imply Hydnoporia. They are, however, much longer than in any Hydnoporia spp. treated above, ca. 8–12 × 2.2–2.9 μm. Large, suburniform basidia (17–27 × 4–5 μm), well-differentiated skeletal hyphae and regularly shaped and rather rare setae indicate that H. allantospora is a member of Hymenochaete s. str.
Hymenochaete rigidula Berk. & M.A. Curtis, J. Linn. Soc., Bot. 10: 334. 1869.
Notes: This species was described from Cuba (Berkeley & Curtis 1869), and no sequenced material is available from the Caribbean. Based on the description by Parmasto (2001) we are on the opinion that H. rigidula s. str. probably belongs to Hymenochaete. He & Dai (2012) reported a Chinese species under this name. Their “H. rigidula”, also present in Korea, is a separate species from other Hydnoporia (Fig. 2). They did not study the type, and we think it is unlikely that their H. rigidula belongs to the species described from the Caribbean
Hymenochaete sallei Berk. & M.A. Curtis, J. Linn. Soc., Bot. 10: 333. 1869.
Notes: A Chinese sequence (FJ481043) is deposited in the INSDC database under the name H. sallei. This sequence from Jiangsu or Anhui represents, if it can be trusted, a third species in the Hydnoporia tabacinoides complex (Fig. 2). Hymenochaete sallei is, however, a typical member of Hymenochaete (Parmasto et al. 2014) and considered a synonym of H. rheicolor by Parmasto (2001). The name has apparently been misapplied to the sequence in question.
ACKNOWLEDGEMENTS
Our thanks to curators of the herbaria UPS, NY, TNS, TAAM, BPI, and GB. Martyn Ainsworth (Kew) provided us a valuable specimen of Hydnoporia tabacina. Alexander Sennikov (Helsinki) provided invaluable advice in nomenclatural questions.
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Article citations
Umbellaceae fam. nov. (Hymenochaetales, Basidiomycota) for Umbellus sinensis gen. et sp. nov. and Three New Combinations.
J Fungi (Basel), 10(1):22, 28 Dec 2023
Cited by: 0 articles | PMID: 38248932 | PMCID: PMC10817349
Notes of Five Wood-Decaying Fungi from Juwangsan National Park in Korea.
Mycobiology, 52(1):30-41, 24 Jan 2024
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Two new species of Hymenochaetaceae from tropical Asia and America.
Front Cell Infect Microbiol, 12:1100044, 19 Jan 2023
Cited by: 0 articles | PMID: 36741979 | PMCID: PMC9892452
Data
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BioStudies: supplemental material and supporting data
Nucleotide Sequences (3)
- (1 citation) ENA - FJ481043
- (1 citation) ENA - KC785573
- (1 citation) ENA - KC785579
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