Antonie van Leeuwenhoek 84: 209 – 216, 2003.  2003 Kluwer Academic Publishers. Printed in the Netherlands. 209 A new species and three new combinations in Cyphellophora, with a note on the taxonomic affinities of the genus, and its relation to Kumbhamaya and Pseudomicrodochium 2 ´ C. Decock 1, *, G. Delgado-Rodrıguez , S. Buchet 3 and J.M. Seng 3,4 1w ` Mycotheque de l’ Universite´ catholique de Louvain ( MUCL, MBLA), Place Croix du Sud 3, B-1348 ´ y Sistematica ´ Louvain-la-Neuve, Belgium; 2 Instituto de Ecologıa , Carretera de Varona Km 3.5, Capdevila, Boyeros, A.P. 8029, 10800 C. Habana, Cuba; 3 BIOTRANSFER, rue Emile Zola 41, 93107 Montreuil, ´ Cedex, France; 4 Laboratoire de Phytopathologie Moleculaire , Universite´ de Paris-Sud, Centre d’ Orsay, ˆ . 630, 91405 Orsay Cedex ´ , France; Institut de Biotechnologie des Plantes ( IBP), Bat * Author for correspondence; (e-mail: decock@ mbla.ucl.ac.be) Received 26 March 2002; accepted in revised form 20 January 2003 Key words: Chaetothyriales, Herpotrichiellaceae, Taxonomy Abstract Cyphellophora guyanensis sp. nov., isolated from leaf litter in French Guyana, is described and illustrated. This fungus is characterized by ampulliform to flask-shaped phialides, with a conspicuous, funnel-shaped collarette, and nearly straight to falcate or slightly sigmoid, (22)3–6 septate conidia. Cyphellophora guyanensis is compared to the other Cyphellophora species, and the latter genus is compared to Kumbhamaya and Pseudomicrodochium. Kumbhamaya indica, P. fusarioides, and P. suttonii are transferred to Cyphellophora. The phylogenetic relationships of Cyphellophora with other genera of ascomycetes and related mitosporic fungi were inferred from the analysis of the partial nuclear ribosomal DNA SSU sequences data. The analysis supports a placement of Cyphellophora in Chaetothyriales, and probably in the Herpotrichiellaceae. Introduction The genus Cyphellophora De Vries (1962) was erected with C. laciniata de Vries as type species and was mainly characterized by having simple, intercalary, lateral phialides, mostly reduced to a distinct collarette, and sickle-shape, brown, 1–3 septate conidia, adhering in bundles. De Vries et al. (1986) broadened the generic description to include a species having well-developed, more or less swollen phialides, Cyphellophora pluriseptata de Vries et al. Both C. laciniata and C. pluriseptata were isolated from human skin (and nails for the latter), although without clinical diagnosis. Later, Walz and de Hoog (1987) added Cyphellophora vermispora Walz & de Hoog while Matw MUCL is a part of the Belgian Co-ordinated Collections of Micro-organisms, BCCM. sushima (1987) described Cyphellophora taiwanensis Matsush., both isolated from plant materials viz. in roots of Triticum aestivum and Hordeum vulgare and from rotten wood of Phyllostachis edulis, respectively. Cyphellophora vermispora was characterized by also having ampulliform to flask-shaped phialides with a conspicuous, funnel-shaped collarette, and vermiform, mostly curved, 30–55 mm long, 4(28) septate conidia while C. taiwainensis has intercalary, lateral, and sessile phialides reduced to a collarette and fusiform conidia, 16–35 3 1.5–2.0 mm, with (2–)3(26) septa. During a study of leaf litter microfungi from French Guyana, an interesting species of Cyphellophora, apparently not yet described, was isolated. The fungus may be compared to C. vermispora in its conidiogenous cells and long, septate conidia. However, it differs from the latter by having smaller (19–28 mm long), nearly straight to more commonly slightly falcate or 210 slightly sigmoid, (2–)3–6 septate conidia. The species is therefore described as C. guyanensis sp. nov. Cyphellophora is compared to two other genera of mitosporic fungi, Kumbhamaya M. Jacob & D.J. Bhat (Jacob and Bhat 2000) and Pseudomicrodochium Sutton (Sutton 1975). The taxonomic affinities of Cyphellophora were also inferred by analysing the partial nuclear ribosomal DNA SSU (18S) sequences data. Material and methods Cultures were grown on Cornmeal agar (CMA) at 25 8C, with a 12 / 12h. incident light periodicity. Microscopic measurements were carried out in lactophenol cotton blue. In presenting the size range of several microscopic elements, 5 % of the measurements at each end of the range are given in parentheses, when relevant. In the text, the following abbreviations are used: (X ] 5 arithmetic mean. Colours are described according to Kornerup and Wanscher (1981). DNA was extracted from freshly collected mycelium grown on Petri dishes on OA, following a protocol of Lee et al. (1988) and purified with Geneclean  III kit (Q-Biogene), following the manufacturer’s recommendations. The primer pair NS1NS8 (biology.duke.edu / fungi / mycology / primers) were used to amplify the SSU (18S) regions of the nuclear ribosomal DNA. Successful PCR reactions resulted in a single band observed on a 0.8 % agarose gel, corresponding to approximately 1900 bp. All the PCR-products were cleaned using the QIAquick  PCR purification kit (250) (QUIAGEN Inc.), following the manufacturer’s recommendations. Sequencing reactions were performed using CEQ DTCS Quick Start Kit  (Beckman Coulter) following the manufacturer’s recommendations. Nucleotide sequences were determined with a CEQ 2000 XL capillary automated sequencer (Beckman Coulter). SSU (18S) DNA fragments were sequenced with primers NS1–NS8 (biology.duke.edu / fungi / mycology / primers). To begin with, nucleotide sequences were automatically aligned with Clustal X for Macintosh (version 1.5b) before being manually adjusted, when relevant, with Paup* (version 4.0b10). Phylogenetic analysis of the aligned sequences was performed using the maximum parsimony method of PAUP* version 4.0b10. The following sequences were imported from GenBank: Chaetothyriales: Capronia acutiseta AC232942, AF050262; C. pulcherrima AJ232944; C. mansonni X79318; Ceramothyrium linnaeae AF022715; Cladophialophora arxii AJ232948; C. devriesii AJ232947; Exophiala dermatidis X80702; E. castellani X78480; Glyphium elatum AF 346419, AF346420; Phialophora verrucosa AJ232945; Ramichloridium anceps AJ232951, AF050285; R. AJ232951; Rhinocladiella atrovirens AJ232950, AF050289; R. phaeophora AJ 232954. Eurotiales: Talaromyces flavus M83262; Monascus purpureus M83260; Eupenicillium javanicum U21298. Onygenales: Gymnoascus reessii AB015774; Gymnascella aurantiaca AB015772; Onygena equina U45442; Uncinocarpus reessii U29394; Pectinotrichum ilanense AB015783. Microascales: Microascus cirrosus M89994; Pseudallescheria boydii M89782; Ceratocystis fimbriata U43777. Hypocreales: Nectria cinnabarina AB 003949; Myrothecium verrucaria AJ302003. Ophiostomales: Ophiostoma piliferum AJ243295; Pesotum fragrans AB007656; Sporothrix schenckii M85053. Sordariales: Chaetomium elatum M83257; Sordaria fimicola X69851. Saccharomycetales: Saccharomyces cerevisiae V01335. The following sequences were obtained and are available at MUCL: Cyphellophora guyanensis MUCL 47737; Cyphellophora vermispora MUCL 43739 5 CBS 228.86; Cyphellophora laciniata MUCL 9569; Pseudomicrodochium fusarioides MUCL 44033; Pseudomicrodochium suttonii MUCL 44073; Pseudomicrodochium lauri MUCL 39134; Nectria pseudotrichia MUCL 39361. Taxonomy Cyphellophora guyanensis C. Decock & Delgado, sp. nov. ( Figure 1) Cultura in agaro ex zea farina: post 4 hebdomata coloniae 37 mm diam. sunt, tenues, firmae, centro chloro-olivaceae vel chloro-griseae, margine chlorogriseae vel griseae, reverso atrovirides vel fuscovirides ad centrum, zonatae et chloro-griseae ad marginem. Margo discreta, densa, appressa. Hyphae laeves et fere tenuitunicatae, 1 – 1.5 mm latae. 211 Phialides ampulliformes, ex hyphis nascentes, terminales vel laterales, solitariae vel aggregatae, 6.0 – 9.3(211.0) x 3.6 – 4.8 mm, (X ] 5 7.8 3 4.1 mm, hyalinae vel pallide olivaceo-brunneae, collariculo infundibuliform, hyalino vel pallide olivaceo-brunneo, 2 – 3 mm diam. Conidia hyalina, laevia, recta vel falcata vel sigmoidea, basi truncata, sursum rotundata, (2 –)3 – 6 septata, (18 –)19.7 – 28(229) x 1.5 – 2.0 mm. Colonies on Cornmeal agar slow growing, attaining a diameter of 37 mm after four weeks, avers thin, regular, azonated, with a dense cottony texture, Figure 1. Cyphellophora guyanensis. (A) Phialides. (B) Conidia. Scale bar 5 10 mm. 212 olivaceous green (30E(5–6), greyish green) in the centre discolouring progressively to pale greyish green to pale greyish near the margin. Reverse blackish green to dark green in the centre, progressively paler toward the margin, pale greenish to pale greyish, faintly radially striated. Margin regular, dense, and appressed. Hyphae thin- to slightly thick-walled, hyaline to pale olivaceous brown, smooth, septate, 1–1.5 mm diam. Conidiophore absent or rarely reduced to a short cell basal to the conidiogenous cells. Conidiogenesis enteroblastic, phialidic. Phialides discrete, ampulliform to flask-shaped, mostly arising at right angles from undifferentiated hyphae, occasionally terminal, solitary or in small clusters, sub-hyaline to pale olivaceous brown, thin-walled, 6.0– 9.3(211.0) x 3.6–4.8 mm, (X ] 5 7.8 3 4.1 mm, with a conspicuous collarette, funnel-shaped, sometimes darker than the body of the phialides, 2.0–3.0 mm diam. Conidia hyaline, smooth, nearly straight to more commonly falcate or slightly sigmoid, with slightly truncate basis and rounded apices, (2–)3–6 septate, (18.0–)19.7–28.0(229.0) x 1.5–2.0 mm, (X ] 5 24.1 3 1.8 mm), aggregating in bundles. Chlamydospores and sclerotia absent. Holotype: (dried culture on CMA, one month old). FRENCH GUYANA, Cayenne area, Matouri, ‘‘sen´ tier d’interpretation de la nature Lamirande’’, Isolated from a dead leaf of an undetermined angiosperm, Jan. 1996, C. Decock et M. Henry de Frahan, MUCL 43737. Partial nrDNA 18S sequences available at MUCL. Discussion Cyphellophora guyanensis is similar to C. vermispora in the general morphology of the phialides but the latter produces longer and slightly narrower (30–55 3 1.2–1.5 mm), vermiform, mostly curved conidia, having up to 8 septa. Cyphellophora guyanensis has smaller (19–28 mm) and slightly wider (1.5–2.0 mm), (straight), falcate to sigmoid conidia, with up to 6 septa. The phialides of C. vermispora are also narrower (2.0–3.5 mm) and occasionally reduced to a sessile collarette on hyphal cells, a feature absent in our species. Walz and de Hoog (1987) also noted the presence of sclerotia, which are absent in C. guyanensis. Both C. guyanensis and C. vermispora were isolated from plant material. Cyphellophora taiwanensis Matsush. may also be compared to C. guyanensis in conidial morphology. The former has fusiform conidia, 16–35 3 1.5–2.0 mm, with (2–)3(26) septa, but the presence of only intercalary phialides reduced to a sessile collarette separates clearly it from C. guyanensis. Within the genera of mitosporic fungi with (mostly) septate phialoconidia aggregating in bundles, Kumbhamaya M. Jacob & D. J. Bhat (Jacob and Bhat 2000) and Pseudomicrodochium Sutton (Sutton 1975) present some evident similitude with Cyphellophora. Kumbhamaya is typified by K. indica M. Jacob & D. J. Bhat (type specimen in PC!, permanent microscopic slides, the ex-type culture being not available), which was isolated as an endophyte from leaves of Carissa carandas (Jacob and Bhat 2000). The latter genus shares with Cyphellophora the flaskshaped, pale brown phialides with a conspicuous collarette, and septate, fusiform conidia. Kumbhamaya seems to differ from the latter by having the phialides commonly with an elongated hyphal-like base (Jacob and Bhat 2000), a feature, however, also observed occasionally in C. pluriseptata (De Vries et al. 1986; Guarro et al. 1995). Therefore, we do not consider that the difference noted above is significant at genus level, and Kumbhamaya is here reduced to synonymy with Cyphellophora. The new combination Cyphellophora indica (M. Jacob & D.J. Bhat) C. Decock comb. nov. is proposed (basionym: K. indica M. Jacob & D.J. Bhat, Cryptogamie, Mycologie 21: 82, 2000). ˜ Pseudomicrodochium Sutton (Castaneda Ruiz et al. 1998; Sutton 1975) also shares with Cyphellophora the phialidic conidiogenesis, relatively similar conidiogenous cells, and, for many species, septate conidia. The genus was originally intended for two plant saprophytic species forming white to pale salmon colonies, with hyaline conidiogenous cells aggregating in sporodochia and hyaline conidia: P. aciculare Sutton, type species, and P. cylindricum Sutton (Sutton 1975). Several species with light or dark coloured colonies, associated either with plants or mammals, were later added to the genus (Ajello et ˜ al. 1980; Castaneda Ruiz and Kendrick 1991; de Hoog 1978; Kirk 1983; Matsushima 1995, 1996; Sutton et al. 1991), broadening the generic circumscription, and making the distinction between Cyphellophora and Pseudomicrodochium critical. In particular, most of the main morphological features of the Pseudomicrodochium species having dark colonies such as P. fusarioides Sutton et al. (Sutton et 213 al. 1991), P. suttonii Ajello et al. (Ajello et al. 1980), and P. triseptata Matsush. (Matsushima 1995) are identical to those in Cyphellophora, what would better place them in the latter genus. In P. fusarioides, for example, the conidiogenous cells are reduced to a pale brown, flared, sessile collarette, very similar to those observed in C. laciniata. However, the position of the Pseudomicrodochium species having light coloured colonies remains more problematic. More importantly, most latter species also differs from Cyphellophora by lacking the large, flared collarette found in Cyphellophora. The phylogenetic relationship of Cyphellophora with P. fusarioides, P. suttonii, and P. lauri, a species with light coloured colonies, and their relationships with other genera of mitosporic fungi and ascomycetes were inferred from a parsimony analysis based on partial 18S sequences data. Unfortunately, there was no culture available of the type species of Pseudomicrodochium (P. aciculare), impeding any sound phylogenetic inference of the relationships of the latter genus with Cyphellophora or other genera of mitosporic fungi or ascomycetes. Preliminary indications of the relationships of C. laciniata, C. vermispora, and C. guyanensis were obtained from a search in GenBank based on partial 18S sequence data. The search yielded exclusively members of the so-called ‘‘black yeasts’’ belonging to the Chaetothyriales Barr, e.g. the ascomycete genus Capronia Sacc. and the genera of mitosporic fungi Exophiala Carmich., and Rhinocladiella Nannf. A 18S sequences data matrix (39 taxa, 1738 characters) was constructed based on sequences either imported from GenBank or obtained at MUCL. A heuristic search with 100 random additional sequences yielded twelve most parsimonious trees, 1095 steps in length, CI 5 0.646, RI 5 0.838. The robustness of the branch was assessed by a bootstrap analysis (1000 bootstrap replication). One of the MPT is presented in Figure 2. The tree topology is similar to that of some trees published previously (e.g. Sterflinger et al. 1999). The other trees do not differ in the major clades nor in the position of Cyphellophora and Pseudomicrodochium. The three Cyphellophora species, together with P. fusarioides and P. suttonii, form a well-supported clade (bootstrap100 %). Cyphellophora laciniata, C. vermispora, P. fusarioides, and P. suttonii form a very well supported sub-clade (bootstrap 95%), while C. guyanensis appears somewhat isolated. These results support the conclusions based on the morphological data, indicating that these species form a very homogeneous group and are in fact congeneric. Consequently, two new combinations are proposed: Cyphellophora suttonii (Ajello et al.) C. Decock comb. nov. (basionym: Pseudomicrodochium suttonii Ajello et al., Mycotaxon 12: 133, 1980); Cyphellophora fusarioides (Campbell & Sutton) C. Decock comb. nov. (basionym: Pseudomicrodochium fusarioides Campbell & Sutton, Mycopathologia 114: 160, 1991). Cyphellophora suttonii and C. fusarioides have an identical 18S sequence, which confirms their very close relationship. The position of P. triseptata remains uncertain, and as the strain was not available for comparison, we will not propose any new combinations for the time being. Our preliminary molecular data also revealed that Cyphellophora belongs to the Chaetothyriales; more precisely the genus may belong to the Herpotrichiellaceae Munk (Figure 2). However, our data show that the strain of P. lauri (MUCL 39134) clusters neither with Cyphellophora nor within the Chaetothyriales but within the Hypocreales Lindau. Pseudomicrodochium lauri MUCL 39134 is thus not congeneric with P. fusarioides or P. suttonii, and Pseudomicrodochium can be considered as polyhyletic in its previous acceptation. Pseudomicrodochium lauri differs from C. fusarioides, C. suttonii, and Cyphellophora in general, by having light coloured, whitish colonies, hyaline conidiogenous cells lacking a large flared phialidic collarette, and hyaline, non-septate conidia. Pseudomicrodochium aciculare, P. cylindricum (Sutton 1975), P. antillanum ˜ ˜ Castaneda & Kendrick (Castaneda Ruiz and Kendrick ˜ ˜ 1991), and P. nectandrae Castaneda et al. (Castaneda Ruiz et al. 1998) also differ from Cyphellophora by the absence of the large, flared collarette at the apex of the phialides, in addition to have light coloured colonies. It is possible that Pseudomicrodochium could then be restricted to the light coloured species mentioned above, but the homogeneity (the monophily) of this group should be tested by molecular methods, which is hardly possible for the time being, due to the lack of living culture. The Herpotrichiellaceae contains many members of so-called ‘‘black yeasts’’. Detailed taxonomic and phylogenetic studies of this group were presented recently by Untereiner and Naveau (1999) and Haase et al. (1999), but it is beyond the scope of this paper to deal with these aspects in detail. However, it can be noted that the conidiogenous features of Cyphel- 214 Figure 2. One of the two twelve parsimonious tree resulting from an heuristic search based on a data set of 39 taxa and 1738 characters (including gaps) of the rDNA SSU. The heuristic search produced twelve most parsimonious trees (MPT), of 1095 steps in length (100 random additions sequences), CI 5 0.646, RI 5 0.838. Saccharomyces cerevisiae was used as outgroup. The branches in bold are those with a bootstrap value between 80–100 %. Cyphellophora and Pseudomicrodochium species are in bold. 215 lophora are also found in several members of the Herpotrichiellaceae, e.g. in several Capronia species, such as Capronia acutiseta Samuels or C. dactylotricha Untereiner et al., which both have flask-shaped phialides with a flared, pale brown collarette. However, they produce smaller, non-septate conidia (Untereiner 1995a, 1995b). Phialophora Medlar also has similar phialides, but small, non septate conidia. The ecological requirements of many members of the Herpotrichiellaceae are still poorly known (Haase et al. 1999). Many species are found in association with plant or fungal materials (Haase et al. 2000) but, interestingly, a large number of mitosporic species have been associated with human ‘‘pathogenesis’’ (Guarro et al. 1995; Haase et al. 1999; Sterflinger et al. 1999). Several Cyphellophora species have also been associated with mammals, mostly humans, potential ‘‘pathogenesis’’: C. laciniata, C. pluriseptata, and C. suttonii have been isolated so far exclusively from human (skin, nail, and pulmonary fluids) while C. fusarioides was isolated from a dog ear lesion. On the other hands, C. guyanensis, C. indica, C. taiwainensis, and C. vermispora have been isolated so far exclusively from plant materials. Cyphellophora vermispora and C. indica were isolated as plant endophytes (Jacob and Bhat 2000; Walz and de Hoog 1987). Bills and Polishook (1994) also reported an endophytic Cyphellophora species tentatively identified as C. cf. taiwainensis from leaves of Heliconia mariae from Costa Rica. However, there are very few records of Cyphellophora species, either from mammals or from plant materials, most species being known from one or very few specimens, and their ecological requirements are still largely unknown. Acknowledgements ` de l’AgThe authors sincerely thank the ‘‘Ministere ˆ riculture et de la Peche’’, Paris, for having granted them the necessary importation permits. The authors also thank Prof. C. Evrard (BOTA, UCL) for revising the Latin description, Dr. J. Stalpers (CBS) and F. Symoens (BCCM / IHEM) for having provided the cultures of C. laciniata and P. suttonii, and of P. fusarioides, respectively. Part of this research was done when G. Delgado Rodriguez came to MUCL on a working visit, funded by a grant from UNESCOIUMS-MIRCEN-SGM, to whom we are grateful. Gregorio Delgado Rodriguez also acknowledges the facilities and financial support provided by the Cuban Ministry of Science, Technology, and Environment. Cony Decock gratefully acknowledges the financial supports received from the Belgian Federal Office for Scientific, Technical, and Cultural affairs (OSTC, contracts BCCM / MUCL 94-98 / 10 / 003 and BCCM / MUCL C2 / 10 / 007), and thanks the directors of the BCCM / MUCL cultures collection for providing facilities and continual encouragement. Marie Henry ´ de Frahan, Stephanie Huret, and Mouna Derfoufi are also thanked for their help in isolating and purifying the culture when we were working in French Guyana and at MUCL, and in obtaining the various DNA sequences. Keys to the species of Cyphellophora. 1. 19. 3. 39 5. 59. 7. 79. Phialides intercalary, mainly reduced to a sessile collarette Phialides well developed, cylindrical, flask-shaped, sessile or with an elongated base 2.Conidia 1–3 septate 29.Conidia up to 6 septate Conidia up to 2.5 mm wide (11–20 3 2–2.5 mm), 1(22) septate Conidia up to 5 mm wide (11–25 3 2–5 mm), 1–3 septate 4. Phialides short to long and cylindrical; conidia 1–1.2 mm wide, 2–3 septate 49 Phialides well developed, flask-shaped, sessile or with an elongated base Conidia mainly straight, on average smaller than 20 mm, 1–5 septate Conidia straight to more commonly falcate, curved, or sigmoid, on average longer than 20 mm 6. Conidia (1–)3 septate, on average wider than 3 mm, 25–40 3 3.5–5.5 mm; phialides commonly with an elongated base 69. Conidia 2–8 septate, on average narrower than 3 mm; phialides without elongated base Conidia vermiform, mostly curved, mostly 4–8 septate, 30–55 3 1.2–1.5 mm. Conidia (straight), falcate or slightly sigmoid, (2–)3–6 septate, (18–)19.7–28(229) x 1.5–2.0 mm 2 4 3 C. C. C. C. 5 C. 6 C. taiwanensis fusarioides laciniata suttonii pluriseptata indica 7 C. vermispora C. guyanensis 216 References Ajello L., Padhye A.A. and Payne M. 1980. Phaeohyphomycosis in a dog caused by Pseudomicrodochium suttonii sp. nov. Mycotaxon 12: 131–136. 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