Crous et al. Habit plant pathogenic, leaf-spotting or saxicolous-saprobic, occasionally isolated from opportunistic human mycoses. Mycelium internal <strong>and</strong> external; hyphae dark brown, septate, branched. Conidiomata in vivo vary from acervuli to sporodochia or fascicles of conidiophores arising from well-developed or reduced, pseudoparenchymatal stromata. Setae <strong>and</strong> hyphopodia absent. Conidiophores arising from hyphae or stromata, solitary, fasciculate to sporodochial, in biotrophic, plant pathogenic species emerging through stomata, little differentiated, semimacronematous, branched or not, continuous to septate, brown, smooth to verruculose. Conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, holoblastic-thalloblastic, meristematic, unilocal, delimitation of conidium by a single septum with retrogressive delimitation of next conidium giving an unconnected chain of conidia, brown, smooth to verruculose, conidiogenous scars (conidiogenous loci) inconspicuous, truncate, neither thickened nor darkened. Conidia solitary or usually forming simple to branched basipetal chains of transversely to muriformly eu- or distoseptate, 1– to multiseptate, brown, smooth, verruculose to verrucose conidia, conidial secession schizolytic. Type species: Catenulostroma protearum (Crous & M.E. Palm) Crous & U. Braun, comb. nov. Description: Crous & Palm (1999), Crous et al. (2004a; figs 364– 365). Notes: Catenulostroma contains several plant pathogenic species previously placed in Trimmatostroma, a morphologically <strong>similar</strong> but, based on its type species, phylogenetically distinct <strong>genus</strong> belonging to Helotiales (Fig. 1). Trimmatostroma s. str. is well-distinguished from most Catenulostroma species by being saprobic, living on twigs <strong>and</strong> branches of woody plants, or occasionally isolated from leaf litter, i.e., they are not associated with leaf spots. <strong>The</strong> conidiomata of Trimmatostroma species are subepidermal, acervular-sporodochial with a well defined wall of textura angularis, little differentiated, micronematous conidiophores giving rise to long chains of conidia that disarticulate at the surface to form a greyblack to brown powdery mass. <strong>The</strong> generic affinity of other species assigned to Trimmatostroma, e.g. those having a lichenicolous habit, is unresolved. Trimmatostroma abietis Butin & Pehl (Butin et al. 1996) clusters together with the plant pathogenic Catenulostroma species, but differs from these species in having a more complex ecology. Trimmatostroma abietis is usually foliicolous on living or necrotic conifer needles on which characteristic acervuli to sporodochia with densely arranged, fasciculate fertile hyphae are formed, comparable to the fasciculate conidiomata of the plant pathogenic species of Catenulostroma (Butin et al. 1996: 205, fig. 1). Although not discussed by Butin et al. (1996), T. abietis needs to be compared to T. abietina Doherty, which was orginally described from Abies balsamea needles collected in Guelph, Canada (Doherty 1900). Morphologically the two species appear to be synonymous, except for reference to muriformly septate conidia, which is a feature not seen in vivo in the type of T. abietis. Furthermore, as this is clearly a species complex, this matter can only be resolved once fresh Canadian material has been collected to serve as epitype for T. abietina. Isolates from stone, agreeing with T. abietis in cultural, morphological <strong>and</strong> physiological characteristics, have frequently been found (Wollenzien et al. 1995, Butin et al. 1996, Gorbushina et al. 1996, Kogej et al. 2006, Krumbein et al. 1996). Furthermore, isolates from humans (ex skin lesions <strong>and</strong> ex chronic osteomycelitis of human patients) <strong>and</strong> Ilex leaves are known (Butin et al. 1996). De Hoog et al. (1999) included strains of T. abietis from stone, man <strong>and</strong> Ilex leaves in molecular sequence analyses <strong>and</strong> demonstrated their genetical identity based on 5.8S rDNA <strong>and</strong> ITS2 data, but strains from conifer needles were not included. Furthermore, we consider T. abietis, as presently defined, to represent a species complex, with Dutch isolates from Pinus again appearing distinct from German Abies isolates, suggesting that different conifer genera could harbour different Catenulostroma species. Isolates from stone form stromatic, durable microcolonies, which are able to grow under extreme xerophilic environmental conditions. Cultural growth resembles that of other meristematic black yeasts (Butin et al. 1996, Kogej et al. 2006). Another fungus isolated from stone in Germany is in vitro morphologically close to C. abietis, but differs in forming conidia with oblique septa. Furthermore, a human pathogenic isolate from Africa clusters together with other Catenulostroma species. <strong>The</strong> habit <strong>and</strong> origin of this human pathogenic fungus in nature <strong>and</strong> its potential morphology on “natural” substrates, which typically deviates strongly from the growth in vitro, are still unknown. However, C. abietis, usually growing as a foliicolous <strong>and</strong> saxicolous fungus, has already shown the potential ability of Catenulostroma species to cause opportunistic human infections. Key to Catenulostroma species 1. Conidia formed in basipetal chains, smooth, 4-celled, consisting of two basal cells with truncate lateral sides, each giving rise to a secondary globose apical cell, that can extend <strong>and</strong> develop additional septa, appearing as two lateral arms ................. C. excentricum 1. Conidia variable in shape, but without two basal cells giving rise to two lateral arms ............................................................................... 2 2. Conidia smooth or almost so, at most very faintly rough-walled; usually foliicolous on conifer needles or saxicolous, forming stromatic, xerophilic durable microcolonies on stone, occasionally causing opportunistic human infections ............................................................ 3 2. Conidia distinctly verruculose to verrucose; plant pathogenic, forming leaf spots ..................................................................................... 5 3. Conidia (8–)20–35(–60) × 4–5(–7) µm, 1–10-septate .................................................................................... C. chromoblastomycosum 3. Conidia much shorter, 8–20 µm long, 0–5-septate .................................................................................................................................... 4 4. Conidia 0–5 times transversely septate, mostly two-celled; usually foliicolous on conifer needles or saxicolous ....................... C. abietis 4. Conidia 2–4 times transversely septate <strong>and</strong> often with 1–2 oblique septa; isolated from stone ........................................ C. germanicum 5. Conidia rather broad, usually wider than 10 µm ........................................................................................................................................ 6 14
Phylogenetic lineages in the Capnodiales Fig. 6. Catenulostroma chromoblastomycosum (type material). A. Sporodochium on pine needle in vitro. B–H. Chains of disarticulating conidia. Scale bars: A = 350, B, E, G, H = 10 µm. 5. Conidia narrower, width below 10 µm ....................................................................................................................................................... 7 6. Conidia distoseptate, rather long, (12–)25–35(–45) × (7–)10–15(–25) µm; conidiomata large, up to 250 µm diam, on Protea anceps ............................................................................................................................................................................................... C. protearum 6. Conidia euseptate, shorter, (9–)16–20(–36) × (10–)14–18(–27) µm; sporodochia 90–100 × 40–80 µm; on Protea gr<strong>and</strong>iceps ................................................................................................................................................................................................. C. elginense 7. Conidia 1- to multiseptate, (10–)15–17(–23) × (5–)6.5–7(–9) µm; on various Proteaceae .................................................. C. macowanii 7. Conidia in vivo predominantly 1-septate, (8–)13–15(–21) × (3.5–)5.5–6(–8) µm; on Protea cynaroides ......................................................................................................................................... C. microsporum (Teratosphaeria microspora) Catenulostroma abietis (Butin & Pehl) Crous & U. Braun, comb. nov. MycoBank MB504504. Basionym: Trimmatostroma abietis Butin & Pehl, Antonie van Leeuwenhoek 69: 204. 1996. Notes: Catenulostroma abietis needs to be compared to Trimmatostroma abietina Doherty (Abies balsamea needles Canada), which is either an older name for this species, or a closely related taxon. Presently T. abietina is not known from culture, <strong>and</strong> needs to be recollected. Catenulostroma chromoblastomycosum Crous & U. Braun, sp. nov. MycoBank MB504505. Fig. 6. Etymology: Named after the disease symptoms observed due to opportunistic human infection. Differt a C. abieti et C. germanico conidiis longioribus, (8–)20–35(–60) × 4–5(–7) µm, 1–10-septatis. Description based on cultures sporulating on WA supplemented with sterile pine needles. Mycelium consisting of branched, septate, smooth to finely verruculose, medium to dark brown, thick-walled, 3–4 µm wide hyphae. Conidiomata brown, superficial, www.studiesinmycology.org 15
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