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Fomitiporia tenuis and Fomitiporia aethiopica (Basidiomycetes, Hymenochaetales), two undescribed species from the Ethiopian highlands: taxonomy and phylogeny

     Mycothèque de l’Université catholique de Louvain (MUCL^,2, MBLA), Croix du Sud 3, B-1348 Louvainla-Neuve, Belgium

Adane Bitew

     School of Medical Laboratory Technology, University of Addis Ababa, P.O. Box 1176, Addis Ababa, Ethiopia

Gabriel Castillo

     Laboratoire d’Algologie, de Mycologie et de Systématique Expérimentale, Département de Biologie Végétale, (B22), Université de Liège, 4000 Liège, Belgium

	ABSTRACT

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Fomitiporia tenuis sp. nov. and Fomitiporia aethiopica sp. nov. from the Ethiopian highlands are described. Both are characterized by a resupinate habit, globose, dextrinoid basidiospores, cystidioles, and respectively scarcity or absence of hymenial setae. Fomitiporia pseudopunctata and F. robusta are reported also in the same area. The preliminary phylogenetic relationships of F. tenuis, F. aethiopica and F. pseudopunctata are inferred from parsimony analysis based on partial nuclear ribosomal large subunit (nrLSU). A preliminary key to the poroid Hymenochaetales with dextrinoid basidiospores (Fomitiporia, Phellinus s.l., Pseudoinonotus) is proposed.

Key words: Africa, Fomitiporia, taxonomy, Phellinus

	INTRODUCTION

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The taxonomy of the Phellinus Quél. and Inonotus P. Karst. complexes, and more generally of the Hymenochaetaceae Pat., has evolved considerably with the gathering of nonmorphological, especially biochemical (Fiasson 1983, Fiasson and Niemelä 1984) and DNA sequences data (Fischer 1996; Wagner and Fischer 2001, 2002). These data, complementary to morphological features, supported either the re-appraisal of previously defined generic entities but generally reduced to synonymy with Phellinus or Inonotus or the recognition of new, homogeneous alliances within these complexes. Most of these entities are now taxonomically recognized at generic level (see Fomitiporia Murrill, Fuscoporia Murrill, Phellopilus Niemelä et al, Pseudoinonotus T. Wagner & M. Fisch., etc., Fiasson and Niemelä 1984; Fischer 1996; Niemelä et al 2001; Wagner and Fischer 2001, 2002). However, because most of these genera are based on data gathered mainly from northern temperate taxa, their legitimacy (their "naturalness") still should be tested by inclusion of more tropical/subtropical taxa that represent the majority of species in these complexes (Fischer 1996, Wagner and Fischer 2001).

During a revision of some poroid Hymenochaetaceae from the Ethiopian highlands, two collections of two distinct Phellinus s.l. species were found that could not be identified satisfactorily as belonging to any of the existing taxa of the latter complex.

A resupinate habit, absence or scarcity of hymenial setae, presence of cystidioles and of hyaline, dextrinoid, cyanophilous, globose to subglobose basidiospores characterized both collections, with features that, within the Phellinus/Inonotus complex of genera, would place them in Fomitiporia (the Phellinus punctatus/robustus complex; Fischer 1996, Wagner and Fischer 2002).

To confirm the affinities of both taxa and to infer their relationships with other Fomitiporia or Phellinus s.l. species, preliminary phylogenetic analyses were carried out based on partial nuclear ribosomal large subunit (nrLSU) sequences data. The results of both the morphological and DNA studies supported the recognition of two distinct species and their placement in Fomitiporia. The two taxa are described below as Fomitiporia tenuis and Fomitiporia aethiopica.

Two other Fomitiporia species also are reported from the Ethiopian highlands: Fomitiporia robusta (P. Karst.) Fiasson & Niemelä and F. pseudopunctata (A. David et al) Fiasson.

A preliminary key to the poroid Hymenochaetales with dextrinoid basidiospores (Fomitiporia, Phellinus s.l., Pseudoinonotus) is proposed.

	MATERIALS AND METHODS

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Cultures and herbarium specimens are preserved at MUCL, LG (herbarium acronyms are from Holmgren et al 1990) and the mycological herbarium of the University of Addis Ababa, Ethiopia. Specimens were examined in Melzer’s reagent, lactic acid cotton blue (Kirk et al 2000), and KOH 4%. Colors are described according to Kornerup and Wanscher (1981). All microscopic measurements were done in Melzer’s reagent. In presenting the size range of the microscopic elements, 5% of the measurements were excluded from each end and are given in parentheses, when relevant. x = arithmetic mean, R = the ratio of length/width of basidiospores, and x_R = arithmetic mean of the ratio R.

Sequencing.— – DNA was extracted from freshly collected mycelium grown in liquid malt at 25 C in the dark. Extractions were carried out using the QIAGEN Dneasy plant Mini Kit (QIAGEN Inc.) and purified with Geneclean^® III kit (Q-Biogene), following the manufacturer’s recommendations. The primer pair LROR-LR6 (White et al 1990) was used to amplify the 5' end of the nr LSU DNA regions. Successful PCR reactions resulted in a single band observed on a 0.8% agarose gel, corresponding to approximately 1200 bps. Polymerase chains-reaction products were cleaned using the QIAquick^® PCR purification kit (250) (QIAGEN Inc.), following the manufacturer’s protocol. Sequencing reactions were performed using CEQ DTCS Quick Start Kit^® (Beckman Coulter), according to the manufacturer’s recommendations, with the primers LROR, LR3, LR3R, LR5 (biology.duke.edu/fungi/mycology/primers). Nucleotide sequences were determined with a CEQ 2000 XL capillary automated sequencer (Beckman Coulter). Initially, nucleotide sequences were automatically aligned with Clustal X for Macintosh (version 1.5b), then manually adjusted as necessary with the editor in PAUP* (version 4.0b10).

The final dataset comprised 28 sequences (25 taxa) and 895 characters, including gaps. Three small inserts were present in both Coltricia species (five and three nucleotides) and in the sequence of P. dryadeus (five nucleotides), and each were recoded as a single event. Phylogenetic analysis of the aligned sequences was performed using the maximum parsimony method of PAUP* 4.0b10 (Swofford 2002). Gaps were treated as fifth base. Optimal trees were identified using heuristic searches, and further evaluated by bootstrap analysis, retaining clades compatible with the 50% majority rules in the bootstrap consensus tree.

Sequences obtained from GenBank.— – Coltricia cinnamomea (Jacq.) Murrill AF311003 [GenBank] ; C. perennis (L.) Murrill AF311004 [GenBank] ; Fomes fomentarius (L.) J.J. Kickx AF261538 [GenBank] ; Fomitiporia hartigii (Allesch. & Schnabl.) Fiasson & Niemelä AF3110051; F. hippophaecola (H. Jahn) Fiasson & Niemelä AF311006 [GenBank] ; F. punctata (Fr.: P. Karst.) Murrill AF311007 [GenBank] ; F. robusta (P. Karst.) Fiasson & Niemelä AF311008 [GenBank] ; Fuscoporia contigua (Pers.) G. Cunn. AF311029 [GenBank] ; Fuscoporia torulosa (Pers.) T. Wagner & M. Fisch. AF311032 [GenBank] ; Fuscoporia ferruginosa (Schrad.) Murrill AF311032 [GenBank] ; Inocutis dryophila (Berk.) Fiasson & Niemelä AF311012 [GenBank] ; Inocutis tamaricis (Pat.) Fiasson & Niemelä AF311021 [GenBank] ; Inonotus hispidus (Bolton) P. Karst. AF311014 [GenBank] ; Inonotus obliquus (Fr.) Pilát; AF311017 [GenBank] ; Onnia triquetra (Pers.) Imazeki AF311024 [GenBank] ; Phellinus ignarius AF311033 [GenBank] ; Phellinus populicola Niemelä AF311038 [GenBank] ; Pseudoinonotus dryadeus (Pers. : Fr.) T. Wagner & M. Fisch. AF311011 [GenBank] .

New sequences generated at MUCL.— – Fomitiporia aethiopica Decock et al MUCL 44777 = AY 618204; F. hippophaeicola MUCL 31746 = AY 618207; F. mediterranea M. Fisch., MUCL 38514 = AY 618201; F. pseudopunctata (A. David et al) Fiasson, MUCL 44806 = AY 618202, MUCL 44799 = AY 618203; F. punctata MUCL 34101 = AY 618200; F. tenuis Decock et al MUCL 44802 = AY 618206; Fomitiporia sp., MUCL 45133 = AY 618205; Fuscoporia palmicola (Berk. & M.A. Curtis) Bondartseva & S. Herrera MUCL 44080 = AY 618209; Fuscoporia sp. MUCL 44180 = AY 618208.

	RESULTS

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A heuristic search with 1000 random additions produced three equally parsimonious trees, 691 steps in length (CI = 0.621, RI = 0.709). The topology of the trees was identical regarding the relative position of the different genera to some previous works (Wagner and Fischer 2001, 2002). One of the three EPTs is represented in FIG. 10.

View larger version (49K): [in this window] [in a new window]   FIGS. 8–10. Fomitiporia pseudopunctata. 8, 9. Cystidia respectively from MUCL 44799 and MUCL 44806; 10. Basidiospores, from MUCL 44799. Scale bar = 5 µm.

	TAXONOMY

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Fomitiporia tenuis Decock, Bitew, et Castillo, sp. nov. FIGS. 1–4

View larger version (53K): [in this window] [in a new window]   FIGS. 1–4. Fomitiporia tenuis, from the holotype. 1. Hymenial setae; 2. Basidium; 3. Cystidioles; 4. Basidiospores. Scale bar = 5 µm.

Basidiome resupinate, effused, adnate, as much as 150 mm long x 30 mm wide, up to 2 mm thick, with a corky consistency when fresh, becoming harder on drying; margin slightly detached, yellowish (light yellow to yellowish orange, 3A(4–5) to 4A(5–6)), velutinous; pores surface brown, (mainly 6E7 (cognac) up to 6(E–F)8 (rust brown-brownish gray)), slightly glancing with light; pores small, round, 10–11/mm, (80–)85–115 µm diam (x = 96 µm diam); dissepiments entire, thin, 15–30(–60) µm thick (x = µm thick); subiculum reduced to a thin layer between the substrate and the tube layer, 100–250 µm thick, densely fibrous, brown (6E7, cognac); tube layer up to 2 mm thick, concolorous with the subiculum and pore surface, with a corky consistency and a fibrous texture; hyphal system dimitic in the context and the hymenophoral trama; hyphae subparallel in the hymenophoral trama; generative hyphae simple-septate, hyaline to pale yellowish, thin- to slightly thick-walled, sparingly branched 1.5–3 µm wide, difficult to find; skeletal hyphae identical in both the subiculum and the hymenophoral trama, yellowish brown, unbranched, thick-walled, nonseptate (or with occasional secondary septa), (1.8–)2.1–3.0 µm wide (x = 2.6 µm), occasionally locally swellings up to 4.5 µm in the subiculum and 2.3–3.2 µm (x = 2.6 µm) in the hymenophoral trama; hymenial setae rare, scattered, ventricose with the apex acute, straight, thick-walled, yellowish brown, directly protruding through the hymenium, 18–21 x 5.5–7.0 µm (x = 21 x 6.4 µm); basidia subglobose, with four sterigmata, 10–13 x 7.0–9.5 µm; basidiospores hyaline, slightly thick-walled, subglobose to globose, cyanophilous, and dextrinoid, 4.8–6.0 x 4.3–5.5 µm (x = 5.3 x 4.9 µm), R = 1.0–1.1(–1.2) (x_R = 1.1); cystidioles rare, fusiform to slightly ventricose ending in an elongated hyphoid apex, hyaline, the basal part thin- to (occasionally) slightly thick-walled, the hyphal-like apex thin-walled, 16–20 x 4.0–5.5 µm; chlamydospores absent; crystals commonly present in the hymenium, rhomboid, of variable size; substrate: living trunk of an unidentified angiosperm (liana); type of rot: white rot; cultural features: unknown; sexuality: unknown.

HOLOTY PE: ETHIOPIA, OROMIA REG. (former Arussi Prov.): Wondo Genet State Forest, 12 km southeast of Shashemene, on the lower part of a living trunk of an unidentified liana, close to the ground, 28 Jul 2003, C. Decock (ET-03/80), A. Bitew, and G. Castillo, MUCL 44802 (culture exholotype MUCL 44802).

Remarks. – The combination of a thin, resupinate basidiome, small pores, presence of rare, scattered hymenial setae, cystidioles with elongated, hyphoid apex, and small, on average smaller than 6 µm, globose, dextrinoid basidiospores, makes the species distinct within Fomitiporia and Phellinus s.l.

Morphologically F. tenuis is to be compared with F. bannaensis Y.C. Dai, described from southern China (Dai et al 2001). Both species have small pores, small basidiospores, and identical setae. The setae are rare in F. tenuis, while Dai et al (2001) note them as abundant in F. bannaensis. However the abundance of setae in Fomitiporia may be variable within a species (see for instance F. robusta, Ryvarden and Gilbertson 1994). Fomitiporia bannaensis differs from F. tenuis mainly by having a thicker basidiome (>1 cm thick), a paler pore surface (yellowish brown), slightly larger pores (8–10/mm), and subulate cystidioles, without an elongated hyphal-like apex (Dai et al 2001).

Within the Fomitiporia species with a resupinate basidiome reported from eastern Africa (Ryvarden and Johansen 1980, Ryvarden and Gilbertson 1994), Fomitiporia punctata (its sibling species, F. mediterranea, morphologically indistinguishable from F. punctata (Fischer 2002), has not been recorded from the area) and F. pseudopunctata form a morphologically homogeneous group that differs from F. tenuis by having usually thicker and cushion-shaped basidiomes, larger pores (6–8/mm), and larger basidiospores, on average longer than 7 µm (Fischer 2002, Ryvarden and Johansen 1980, Ryvarden and Gilbertson 1984, Larsen and Cobb-Poulle 1990). Furthermore F. punctata (and F. mediterranea) lack setae.

Within the Phellinus s.l. species with dextrinoid basidiospores, Ph. spinescens Wright & Coelho (Coelho and Wright 1996), Ph. uncinatus Rajchenb. (Rajchenberg 1987) and Ph. bambusarum (Rick) M. Lars. (Larsen and Cobb-Poulle 1990, Rajchenberg 1987, Wright and Blumenfeld 1984) share with F. tenuis a resupinate basidiome, hymenial setae and globose basidiospores, on average smaller than 6 µm. They differ from the latter in that they have larger pores (respectively 4–6/mm, (4–)5–6/mm and 7–8/mm), grow on Bambusaceae and so far are known only in the neotropics. Phellinus uncinatus and Ph. spinescens also have different setae, apically hooked in the former, and with lateral spinulose processes in the latter.

Phellinus transversus Corner (Corner 1991) also has small pores (70–100 µm diam) and globose, slightly thick-walled, hyaline basidiospores of a size comparable to those of F. tenuis but differs in that it has thicker basidiomes, up to 20 mm thick, slightly curved setae and (presumably) nondextrinoid basidiospores. (Actually Corner [1991] did not mention a reaction of the basidiospores in Melzer’s reagent, what could be interpreted, although with caution, as a negative result.)

A specimen originating in Papua New Guinea (PAPUA NEW GUINEA, MADANG PROV.: 04 Oct. 1989, E. Quanten EQ 726 (GENT) and tentatively identified by Quanten (1997) to F. robusta (under the name Ph. robustus [P. Karst] Bourd. & Galz.) forma resupinatus Bourd. & Galz.) might be compared to F. tenuis. The Papua New Guinea specimen shared with F. tenuis a thin, resupinate basidiome (up to 2 mm thick), a brown pore surface ("leather to tan-colored" = 6E6, Kornerup and Wanscher 1981), very small pores (9–11/mm), ventricose setae (14–21 x 5.5–6.5 µm), and small (4.6–6.2 x 4.0–5.4 µm) dextrinoid basidiospores (Quanten 1997). Quanten (1997) did not mention cystidioles, but they might be difficult to find in dried material. These features deviate strongly from F. robusta, which has much larger pores (5–6/mm) and basidiospores on average longer than 7 µm (6–8.5 x 5.5–7.0 µm, Ryvarden & Gilbertson 1994) and in all probability, the PNG specimen does not belong there. The voucher specimen was not examined, but the characteristics provided (Quanten 1997) apparently agree with the main features of F. tenuis.

Fomitiporia aethiopica Decock, Adane, et Castillo, sp. nov. FIGS. 5–7

View larger version (20K): [in this window] [in a new window]   FIGS. 5–7. Fomitiporia aethiopica, from the holotype. 5. Cystidioles; 6. Basidium; 7. Basidiospores. Scale bar = 5 µm.

Basidiome resupinate, effused, adnate, seasonal (to perennial), up to 200 x 50 mm, up to 5–6 mm thick, of a corky consistency when fresh, hard corky when dried; margin narrow, up to 0.5 mm wide, white at the very outside, pale yellow up to concolorous to the pore surface, cottony or with stiff, erect hyphae; pores surface light, grayish orange (5B4), brownish orange (5C(4–5), golden blonde), to yellowish brown (5D4, dark blonde), glancing with light to grayish brown (6E4, brown); pores 5–6/mm, round to elongated on sloppy part, 125–140 µm diam (x = 135 µm); dissepiments thick, smooth, entire, 50–110 µm thick (x = 73 µm); subiculum 200–750 µm thick, brown, dark cinnamon brown, with a hard, corky consistency and a fibrous, dense texture; tube layer single, 2–5 mm deep, elongated on sloppy part, brown (6E[6–7], cocoa brown, cognac), contrasting with the much paler pore surface; hyphal system dimitic in the subiculum and the hymenophoral trama; hyphae in the subiculum densely packed, perpendicular to the substrate, somewhat aggregated in bundles; generative hyphae difficult to find, simple-septate, hyaline to pale yellowish, thin- to slightly thick-walled, slightly branched, 2.0–3.0 µm wide; skeletal hyphae identical in the subiculum and the hymenophoral trama, yellowish brown, unbranched, thick-walled, nonseptate (or with occasional secondary septa), 2.5–4.0(–4.3) µm wide (x = 3.3 µm) in the subiculum and 2.5–4.2 µm (x = 3.2 µm) in the hymenophoral trama; setae absent in all parts of the basidiome; basidia with four sterigmata, 13–18 x 8–9.5 µm; basidiospores globose to subglobose, slightly thick-walled, hyaline, cyanophilous and dextrinoid, 6.0–8.8 x 5.5–7.2(–7.5) µm, (x = 7.2 x 6.5 µm), R = 1.0–1.2(–1.3), (x_R = 1.1); cystidioles scattered, fusiform to slightly ventricose, hyaline, thin-walled, 17–20 x 3.5–6.0 µm, (x = 18–4.8 µm); crystals commonly present in the hymenial layer, rhomboid, of variable size; chlamydospores absent; substrate: dead trunk, angiosperm; type of rot: white rot; cultural features: unknown; sexuality: unknown.

HOLOTY PE: ETHIOPIA, OROMIA REG. (former Shoa Prov.): Menagesha forest, mixed Podocarpus—Juniperus forest, ca. 8°57'N, 038°32'E, ca. 2470 m a.s.l., on a dead standing trunk of an unidentified angiosperm, 30 Jul 2003, C. Decock (ET-03/97), A. Bitew, and G. Castillo, MUCL 44777, LG-GC (culture ex-holotype 44777).

Remarks. – The morphology of F. aethiopica, both at macro- and microscopic level, first prompted us to relate it to F. punctata because both taxa share a thick basidiome, absence of setae and basidiospores on average longer than 7 µm (Ryvarden and Gilbertson 1994). Fomitiporia aethiopica differs mainly by having a paler, grayish orange, golden brown (dark blonde) pores surface, contrasting with the chocolate brown tube layer. The pore surface is darker, grayish brown and concolorous with the tube layers in F. punctata (Ryvarden and Johansen 1980, Ryvarden and Gilbertson 1994). Fomitiporia aethiopica also is related morphologically to F. pseudopunctata, which differs by the presence of setae, in addition to the feature already described above for F. punctata.

F. aethiopica also is phylogenetically distinct from F. punctata but closely related to F. pseudopunctata (FIG. 11), and both taxa formed a well supported clade (bootstrap value 100%).

View larger version (45K): [in this window] [in a new window]   FIGS. 11. One of the three equally parsimonious trees obtained from an heuristic search. Branches in bold = bootstrapping > 70% (see text for details).

	OTHER FOMITIPORIA SPECIES FROM THE ETHIOPIAN HIGHLANDS

TOP ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS TAXONOMY OTHER FOMITIPORIA SPECIES FROM... DISCUSSION PRELIMINARY KEY TO THE... LITERATURE CITED

 
Fomitiporia pseudopunctata (A. David et al) Fiasson, Karstenia 24:25, 1984. FIGS. 8–10

Phellinus pseudopunctatus A. David et al, Mycotaxon 14:171, 1982.

For a description of the species, see David et al (1982) or Ryvarden and Gilbertson (1994).

Specimens examined. – ETHIOPIA, SOUTHERN NATIONS NATIONALITIES PEOPLES ADMINISTRATIVE REGION (former Kefa Prov.), ca. 20 km from Jimma, on the road to Bongo, 7°03'N, 036°45'E, ca. 1750 m a.s.l., at the lower side of a dead hanging branch of an unidentified angiosperm, ombrophilous afro-mountainous forest, 20 Jul 2003, C. Decock, A. Bitew, and G. Castillo ET-03/19, MUCL 44799 (culture ex-MUCL 44799) ; OROMIA REGION (former Shoa Prov.): ca. Ginshi, 9°03'N, 038°28'E, on the underside of a dead branch still attached to a living unidentified angiosperm tree, 25 Jul 2003, C. Decock, A. Bitew, and G. Castillo, ET-03/71, MUCL 44806, LG-GC.

Remarks. – Fomitiporia pseudopunctata was first described from southern France (David et al 1982) and is widespread in the Mediterranean area (Ryvarden and Gilbertson 1994). Ryvarden and Gilbertson (1994) noted that F. pseudopunctata also occurred in Zimbabwe and presumed that the taxon probably was more widespread in eastern Africa. Niemelä and Mrema (2002) also reported from eastern Africa (Tanzania) F. sublaevigata (Cleland & Rodway) Y.C. Dai, an Australian taxon apparently very similar to F. pseudopunctata, from which it apparently differs by having slightly wider hymenial setae (Buchanan and Ryvarden 1993). The distinction between these two species remains uncertain so far and would necessitate a closer comparison of more specimens. However, assuming that two taxa are represented by these names, it would be more likely from a bio-geographical point of view to be F. pseudopunctata that occurs in eastern African countries instead of F. sublaevigata.

Two Fomitiporia collections from the Ethiopian highlands showed most characteristics of F. pseudopunctata as perennial (with up to 15 tube layers (in MUCL 44806), individual layers 1–3 mm each), thick (<55 mm thick), cushion-shaped basidiomes, with margin turning black in older parts (in MUCL 44806), grayish brown surface of the pores (6–)7–8 pores/mm, 125–165 µm diam (x = 142 µm), ventricose to subulate setae, 17–35 x 5–9 µm (x = 25 x 7.2 µm), and globose, thick-walled, slightly to strongly dextrinoid (the reaction might take time to develop), cyanophilous basidiospores, 6.5–8.0 x 6.2–7.5 µm (x = 7.3 x 6.9 µm). David et al (1982) reported the setae to be "numerous and widespread in the whole dissepiments". In our two collections, the setae, although not difficult to find, were rare and scattered.

Fomitiporia robusta (P. Karst.) Fiasson & Niemelä, Karstenia 24:24, 1984.

Phellinus robustus (P. Karts.) Bourd. & Galz., Hym. Fr.: 616, 1927.

For a description of the species, see Gilbertson and Ryvarden (1987) and Ryvarden and Gilbertson (1994).

Specimen examined: ETHIOPIA, OROMIA REG. (former Shoa Prov.): ca. Ginchi, 9°03'N, 038°28'E, on a living trunk of an unidentified angiosperm, 25 Jul 2003, C. Decock, A. Bitew and G. Castillo, ET-03/70, MUCL 45131.

Remarks. – The collection was characterized mainly by a semicircular, applanate basidiome, 90 mm long and 110 mm wide, up to 50 mm thick at the base, down to 15 mm thick at the rounded margin. The pileus was dark brown to blackish, slightly rimose, in great part covered by mosses. The surface of the pores was brown (rust brown, 6E[7–8]), and the pores round, even, (4–)5(–6) pores/mm. The basidiospores are globose, subglobose, hyaline, cyanophilous, and dextrinoid, 6.5–7.5 x 6.2–7.0 µm (x = 7.1 x 6.7 µm). Setae were not observed. The specimen was growing on the trunk of a living tree of an unidentified angiosperm, ca. 2 m above ground. Attempts to obtain a culture of the species and DNA from the herbarium specimen were unsuccessful.

Fomitiporia robusta seems to be a variable species (Ryvarden and Johansen 1980) but its present concept might encompass a complex of taxa. For instance, the number of pores/mm, found in literature, range from 4 to 5 (Ahmad 1972), 5 to 6 (Ryvarden and Johansen 1980), 7 to 9 (Gilbertson and Ryvarden 1987, Ryvarden and Gilbertson 1994), approximately 9 (Quanten 1997) and 9 to 11 (Quanten 1997, in fa. resupinatus). Basidiospore range is 4.6–6.2 x 4.0–5.4 µm (Quanten 1997, in fa. resupinatus), 5.0–6.5 x 4.5–6 µm (Quanten 1997) to 6–8.5 x 5.5–7.0 µm (Ryvarden and Gilbertson 1994). More detailed morphological and molecular studies based on more collections from the presumed range of distribution would be necessary to ascertain variation within the species and any possible taxonomic signification of the variations observed in this species.

	DISCUSSION

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Fomitiporia first was established by Murrill (1907) with F. langloisii (Murrill) Murrill as type species, a name now considered as a taxonomic synonym of F. punctata (Ryvarden 1985). The status of the genus was debated for a long time but considered by most authors as a taxonomic synonym of Phellinus (Ryvarden 1991, Ryvarden and Gilbertson 1994, Ryvarden and Johansen 1980, Larsen and Cobb-Poulle 1990). However, considering the morphological, biochemical and molecular data available, the genus recently was reconsidered for a number of species known as the Ph. Punctatusrobustus complex (Fiasson and Niemelä 1984; Fischer 1996; Wagner and Fischer 2001, 2002).

Fischer (1996) and Wagner and Fischer (2002) then defined Fomitiporia as having perennial basidiomes, hyaline, ampulliform cystidioles, hymenial setae rare to absent, and globose, cyanophilous, dextrinoid basidiospores. The hyphal system was defined as intermediate between mono- and dimitic (Wagner and Fischer 2002). Wagner and Fischer (2002) also characterized the genus by a "brass"-colored context, a feature however difficult to handle and to ascertain in some species having a thin subiculum.

The main morphological features of F. aethiopica and F. tenuis justified the choice of the genus, as defined by Wagner and Fischer (2002). This placement also was supported by our preliminary phylogenetic analysis based on partial nuclear ribosomal DNA sequences data (FIG. 11).

Fomitiporia aethiopica phylogenetically appeared to be closely related to the two Ethiopian specimens of F. pseudopunctata, and both taxa belong to the (although nonsupported) F. punctate–F. robusta clade. Fomitiporia tenuis so far has an isolated position within the genus. An unidentified culture originating from Cuba (MUCL 45133; CUBA, PROV. GUANTA-NAMO: Municipio Baracoa, Quibijan-La Perrera, on a dead stump of "albaricoque", 6 Sep 2003, C. Decock, J. Ortiz, L. Del Castillo, K. Licea CU-03/35, culture MUCL 45133 and HAC [herbarium specimen at HAC]) also clustered within the Fomitiporia clade in a rather isolated position. The voucher specimen being not available for the time being, we cannot draw conclusions concerning the status and identity of this isolate.

For many Fomitiporia species, molecular data still are missing and DNA sequences are available for only about half of the known species (without including the Phellinus s.l. species with dextrinoid basidiospores, among which some most probably will prove to belong to Fomitiporia). The genus’ internal phylogenetic relationships thus remain largely unknown. However, the present concept of Fomitiporia (Wagner and Fischer 2002) was supported by our phylogenetic analysis including (sub) tropical taxa.

	PRELIMINARY KEY TO THE POROID HYMENOCHAETACEAE WITH DEXTRINOID BASIDIOSPORES (FOMITIPORIA, PHELLINUS S.L., PSEUDOINONOTUS)

TOP ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS TAXONOMY OTHER FOMITIPORIA SPECIES FROM... DISCUSSION PRELIMINARY KEY TO THE... LITERATURE CITED

 
Fifteen species so far have been accepted in Fomitiporia. They are: Fomitiporia aethiopica (see above), F. bannaensis (Dai et al 2001), F. erecta (A. David et al) Fiasson (David et al 1982, Ryvarden and Gilbertson 1994), F. hartigii (Ryvarden and Gilbertson 1994), F. hippophaeicola ( Jahn 1976, Ryvarden and Gilbertson 1994), F. mediterranea (Fischer 2002), F. pseudopunctata (David et al 1982, Ryvarden and Gilbertson 1994), F. punctata (Ryvarden and Gilbertson 1994), F. pusilla (Lloyd) Y.C. Dai (Ryvarden 1989, Núñez and Ryvarden 2000), F. robusta (Ryvarden and Gilbertson 1994), F. sonorae (Gilbn.) Y.C. Dai (Dai et al 2001, Gilbertson and Ryvarden 1987), F. sublaevigata (Cleland & Rodway) Y.C. Dai (Buchanan and Ryvarden 1993, Dai et al 2001), F. tenuis (see above), F. texana (Murrill) I. Nuss (Gilbertson and Ryvarden 1987) and F. tibetica Y.C. Dai & M. Zang (Dai and Zang 2002).

In addition, seven species of Phellinus s.l. have dextrinoid basidiospores and some might be affined to Fomitiporia. They are: Phellinus apiahynus (Speg.) Rajchenb. & Wright (Rajchenberg and Wright 1987, Wright and Blumenfeld 1984), Ph. bambusarum (Larsen and Cobb-Poulle 1990), Ph. juniperinus Bernicchia & Curreli, Ph. rosmarini Bernicchia (Bernicchia 1990; Pieri and Rivoire 2000), Ph. spinescens (Coelho and Wright 1996), Ph. tabaquilio (Urcelay et al 2000), and Ph. uncinatus (Rajchenberg 1987).

Many species of Fomitiporia form morphological "complexes" that are difficult to distinguish and to key (c.f. F. punctata and F. mediterranea (Fischer 2002), F. erecta and Ph. juniperinus (Pieri and Rivoire 2000) or F. pseudopunctata and F. sublaevigata (Buchanan and Ryvarden 1993, Dai et al 2001). Some species can be distinguished by their host (viz. F. mediterranea/F. punctata) or by their geographical distribution, but for many the hosts and geographical range are known incompletely. We hereby propose a key for Fomitiporia and Phellinus s.l. species with dextrinoid basidiospores.

1b Basidiome principally resupinate 2 1b Basidiome pileate 12     2a Hymenial setae absent; basidiome usually thick and cushion-shaped 3     2b Hymenial setae present 5 3b Surface of the pores pale, grayish orange, golden blonde, contrasting with the chocolate brown tube layer; so far, only known from Ethiopia F. aethiopica 3a Surface of the pores darker, brown, grayish brown to golden brown, concolorous with the tubes 4     4a Basidiomes strictly resupinate; pores 5–6/mm; basidiospores 6.5–8.5 x 5.5–7.0 µm; cosmopolitan (?) (Europe, North America, Temperate Asia (If the host is Vitis vinifera [or Olea europea] , see also F. mediterranea, a taxon that morphologically is undistinguishable from (Fischer 2002) F. punctata).     4b Basidiomes resupinate to pseudopileate, pileus then arising by marginal accumulation of tube layers; pores 4–6/mm; basidiospores 6.0–7.5 x 4.8–6.5 µm; on living Polylepis australis (Rosaceae); South America, at altitude Ph. tabaquilio 5a Setae apically hooked; South America; growing on bamboo Ph. uncinatus 5b Setae straight 6     6a Setae with lateral, spinulose processes; pores 4–6/mm; South America; growing on bamboo Ph. spinescens     6b Setae without lateral, spinulose processes 7 7a Basidiospores on average smaller than 6 µm 8 7b Basidiospores on average longer than 6 µm 11     8a Setae up to 55 µm long, ventricose with an elongated (hyphal-like) apical portion; resupinate to slightly reflexed basidiomes; Southern USA and Mexico (Valenzuela and Chacón-Jiménez 1991) F. sonarae     8b Setae shorter than 25 µm, without elongated apical portion; tropical species 9 9a Pores 7-8/mm; South America taxon; growing on bamboo; basidiospores 5-6 x 5 µm in diam; setae 16-20 X 5-7 µm Ph. bambusarum 9b Pores 8-1 I/mm; paleotropical species; growing on other woody hosts 10     10a Pores 8–10/mm; pore surface yellowish brown; basidiome to 12 mm thick; cystidioles subulate; setae abundant; basidiospores 4.2-5.2 x 3.8-4.8 µm; so far known from southern China F. bannaensis     10b Pores 10-1 I/mm; pore surface rust brown; basidiomes to 2 mm thick; cystidioles with an elongated hyphoid apex; setae very rare; basidiospores 4.8-6.0 x 4.3-5.5 µm; so far known from Ethiopia only F. tenuis 11a Setae 15-28 x 7-10 µm; pores 6-8/mm; basidiospores 6.5–7.5 x 5.5–7.0 µm; Mediterranean Europe, east Africa F. pseudopunctata 11b Setae 21-30 x 4.5-8.5 µm; pores 5-7/mm; basidiospores 6.5–7.0 x 5.0–6.0 µm; Australia F. sublaevigata (See Buchanan and Ryvarden [1993] for a discussion of F. pseudopunctata and F. sublaevigata)     12a Hymenial setae absent 13     12b Hymenial setae present 18 13a On gymnosperm; pileate to effuse reflexed; temperate species F. hartigii 13b On angiosperm; temperate or tropical 14     14a Basidiospores on average smaller than 6 µm 15 14b Basidiospores on average longer than 6 µm 16     15a Basidiome pileate, sessile; pores 7–8/mm; basidiospores 5.0-6.0 x 4.5-5. 0(-5.7) µm; pileus not rimose; South America Ph. apiahynus     15b Basidiome pendant and conical, dorsally attached; pores 8-9/mm; warm temperate Asia. F. pusilla 16a On Hippophae; Europe and east to central Asia F. hippophaeicola 16b On other hosts 17     17a True pileus applanate-ungulate, brown, blackish, rimose with age; basidiospores 6.5-8.5 x 5.5–7.5 µm; cosmopolitan F. robusta     17b Pseudo-pileus formed by accumulation of tube layers fusing at their margin, not rimose with age; basidiospores 6.0-7.5 x 4.8-6.5 µm; on living Polylepis australis (Rosaceae); South America, at altitude Ph. tabaquilio 18a Setal hyphae (variably) present; hymenial setae 28-58 x 4.5-8.0 µm; basidiospores 6.0-7.5 x 5.5-6.5 µm; mainly on Juniperus, Mediterranean Ph. juniperinus/F. erecta 18b Setal hyphae absent 19     19a Basidiome annual, seasonal; setae abundant, straight to hooked 20     19b Basidiome perennial; setae never hooked 21 20a Setae straight to hooked, ventricose; on angiosperm (mainly Quercus); Europe, North America Ps. dryadeus 20b Setae always hooked, triquetrous; on gymnosperm (Abies, Picea); China F. tibetica     21a Basidiospores on average smaller than 6 µm 22     21b Basidiospores on average larger than 6 µm 23 22a Setae and cystidioles with (hyphal-like) slender apical portion; basidiomes mainly effused, with reflexed margin reflexed; southern USA and Mexico Ph. sonorae 22b Setae subulate to ventricose, cystidioles fusiform, basidiomes pileate; on Rosmarinus and Rhamnus; Mediterranean Europe Ph. rosmarini     23a Basidiome tuberculiform, fan-shaped to semiclavate; setae 20-40 x 5.5-6.0 µm; basidiospores 6.5–7.5 x 5.5–6.5 µm; mainly on Quercus, Mediterranean F. erecta     23b Basidiome regular in shape, typically applanate to ungulate 24 24a Setae absent to common, subulate to ventricose, without apical elongation, 18-37 x 5-8 µm; basidiospores 6.5-8.5 x 5.5-7.5 µm; cosmopolitan F. robusta 24b Setae with a slender apical portion, up to 60 µm long; basidiospores 7.0–9.0 x 6.5—9.0 µm; southern USA F. texana

	ACKNOWLEDGMENTS

 
The authors warmly thank the University of Addis Ababa, Ethiopia, for the facilities offered during two collecting trips to the Ethiopian highlands in Nov 2002 and Jul 2003. Professor C. Evrard (BOTA) also is thanked for his help in the Latin diagnoses and Dr Yu Cheng Dai for providing useful literature. Cony Decock gratefully acknowledges the financial support from the Belgian Federal Science Policy Office (contract BCCM C2/10/007 and C3/10/003) and, together with G. Castillo, from the "Fonds de la Recherche Fondamentale Collective" (FRFC, Contract 2.4551.99). The authors also thank Mrs S. Huret for her help in obtaining DNA sequences and the directors of MUCL and MBLA for providing facilities and continuous encouragement.

	FOOTNOTES

 
Accepted for publication May 31, 2004.

² MUCL is a part of the Belgian Coordinated Collections of Micro-organisms, BCCM^TM.

¹ Corresponding author. E-mail: decock{at}mbla.ucl.ac.be

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