The genus Cladosporium and similar dematiaceous ... - CBS - KNAW

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available online at www.studiesinmycology.org doi:10.3114/sim.2007.58.09 Studies in Mycology 58: 235–245. 2007. Taxonomy, nomenclature and phylogeny of three cladosporium-like hyphomycetes, Sorocybe resinae, Seifertia azaleae and the Hormoconis anamorph of Amorphotheca resinae K.A. Seifert, S.J. Hughes, H. Boulay and G. Louis-Seize Biodiversity (Mycology & Botany), Eastern Cereal and Oilseed Research Centre, Agriculture & Agri-Food Canada, Ottawa, Ontario K1A 0C6 Canada *Correspondence: Keith A. Seifert, seifertk@agr.gc.ca Abstract: Using morphological characters, cultural characters, large subunit and internal transcribed spacer rDNA (ITS) sequences, and provisions of the International Code of Botanical Nomenclature, this paper attempts to resolve the taxonomic and nomenclatural confusion surrounding three species of cladosporium-like hyphomycetes. The type specimen of Hormodendrum resinae, the basis for the use of the epithet resinae for the creosote fungus {either as Hormoconis resinae or Cladosporium resinae) represents the mononematous synanamorph of the synnematous, resinicolous fungus Sorocybe resinae. The phylogenetic relationships of the creosote fungus, which is the anamorph of Amorphotheca resinae, are with the family Myxotrichaceae, whereas S. resinae is related to Capronia (Chaetothyriales, Herpotrichiellaceae). Our data support the segregation of Pycnostysanus azaleae, the cause of bud blast of rhododendrons, in the recently described anamorph genus Seifertia, distinct from Sorocybe; this species is related to the Dothideomycetes but its exact phylogenetic placement is uncertain. To formally stabilize the name of the anamorph of the creosote fungus, conservation of Hormodendrum resinae with a new holotype should be considered. The paraphyly of the family Myxotrichaceae with the Amorphothecaceae suggested by ITS sequences should be confirmed with additional genes. Key words: Amorphothecaceae, Cladosporium resinae, creosote fungus, Hormoconis resinae, jet fuel fungus, kerosene fungus, Myxotrichaceae, Pycnostysanus, resinicolous fungi. INTRODUCTION The ascomycete Amorphotheca resinae Parbery (1969) grows in hydrocarbon-rich substrates such as jet fuel, cosmetics and wood preserved with creosote or coal tar. This fungus is widely known by the anamorph name Hormoconis resinae (Lindau) Arx & G.A. de Vries or its obligate synonym Cladosporium resinae (Lindau) G.A. de Vries. It produces lightly pigmented, warty conidiophores, and branched, acropetally developing chains of lightly pigmented ameroconidia lacking conspicuous scars (Fig. 1B–E). This species is known colloquially as the “creosote fungus”, the “kerosene fungus” or the “jet fuel fungus”; to avoid confusion caused by the many heterotypic names with the epithet “resinae”, in this paper we generally will use the oldest of these informal names, “creosote fungus”, when referring to A. resinae or its anamorph. This fungus grows in jet fuel contaminated with small amounts of water, and the mycelium clogs fuel lines and corrodes metal parts. Consequently, fuel tanks in airports are monitored for this fungus by private companies using various physiological or biochemical tests. Sorocybe resinae (Fr.) Fr. produces dark black colonies on conifer resin, comprising dark synnemata and an effuse mononematous synanamorph, both with cladosporium-like conidiogenous cells and conidia. Unlike the anamorph of the creosote fungus, the conidia of Sorocybe resinae are dark brown and the lateral walls are conspicuously thicker than the poles (Fig. 2D–G). Colonies with only the mononematous anamorph sometimes occur, and the mononematous anamorph can be sparse on colonies bearing synnemata. However, the conidia of the mononematous anamorph have identical pigmentation and lateral wall thickening to that of the synnematous anamorph. The mononematous anamorph rarely has been referred to by its own binomial name although, as we will show, there is a species epithet available. For the same reasons given above for Amorphotheca Parbery, generally we will refer to Sorocybe resinae herein as “the resin fungus”. Despite the micromorphological differences noted above, there is disagreement about whether the creosote fungus is conspecific with the mononematous synanamorph of the resin fungus (Parberry 1969). The name for the anamorph of the creosote fungus is based on Hormodendrum resinae Lindau (1906). Christensen et al. (1942) presented a study of a cladosporium-like fungus commonly isolated from wood impregnated with creosote and coal tar and applied Lindau’s name without examining its type. A later ecological study by Marsden (1954) employed the same name for the same fungus. An extra dimension was added to the confusion when de Vries (1952, using the name Cladosporium avellaneum G.A. de Vries) described four formae for the creosote fungus (differing in the colours of their conidia, the production of setae, or the total absence of conidia), each based on single conidium isolates made from one parent culture. De Vries (1955) and Parberry (1969) examined the holotype of Hormodendrum resinae and concluded that it represented the creosote fungus. Hughes (1958), prior to the description of Amorphotheca or Hormoconis Arx & G.A. de Vries, examined the same specimen and considered it to be the mononematous synanamorph of the resin fungus. If Hughes (1958) is correct, then neither the species Hormodendrum resinae, nor the genus that it typifies, Hormoconis, can represent the creosote fungus, as intended by Parberry (1969) or von Arx and de Vries (in von Arx 1973). In this paper, we present micromorphological, cultural and molecular evidence that the resin fungus is a different species from the creosote fungus. Combined with re-examination of the holotype of Hormodendrum resinae, this information is used to provide a revised taxonomy and nomenclature for these two species. A third cladosporium-like fungus, Seifertia azaleae, is also considered in our discussion of generic concepts. Historical review The history of the fungus now known as Sorocybe resinae began with Fries (1815), who described Racodium resinae Fr. as follows: “310. Racodium resinae, expansum molliusculum dense contextum nigrum, filis inaequalibus. In resina Pini Abietis in silvis Suecia passim. Habitu et loco natali distinctum. Fila divaricato-ramosa; alia rigidula apice capituli sera, sub miscrosc. Coremio Link similia, Demat. villosum Schleich. huic simile; sed sub microsc. fila maxime differunt.” 235
Seifert et al. Fig. 1. Amorphotheca resinae, colony characters and anamorph micromorphology. A. 10-d-old colony on PDA. B, D–E. Micromorphology of conidiophores, showing acropetal conidial chains, ramoconidia, and conidia. C. Conidia. DAOM 170427; for C, E see scale bar in D. The comparison with Coremium Link indicates the probability of a synnematous fungus, and an authentic specimen of Fries’ fungus, which as the only known authentic material we interpret as the holotype, is preserved in Link’s herbarium (see below). It represents the synnematous form of the resin fungus 1 . 1 Persoon (1822) described a form of R. resinae “β piceum”. Hughes (1968) examined the holotype of this form, and it represents the mycelium of the ascomycete Strigopodia resinae (Sacc. & Bres.) S.J. Hughes. This taxon is thus not relevant to the three species that are the focus of this paper. Fries (1832) later transferred his species to Sporocybe Fr. (1825), a genus then used for relatively conspicuous dark hyphomycetes with dry spores (Mason & Ellis 1953). The 1832 description explicitly stated... “capitulo rotundato inaequali, sporidiis seriatis, stipite aequali simplici.” The use of “capitulo” and “stipite” imply what would now be recognised as a synnematous fungus. Fries (1832) further characterised the habit of the fungus as “habitu stipitum Calicii,” a further comparison to a group of black, stipitate lichenized fungi classified in Calicium Pers., which under a hand lens look similar to a dark synnematous fungus. Fries (1849) next described the genus Sorocybe Fr. for this fungus, as follows: 236
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Studies in Mycology 58 (2007) The g
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Studies in Mycology The Studies in
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CONTENTS P.W. Crous, U. Braun and J
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lectotype for the genus by Clements
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Schubert K (2005a). Morphotaxonomic
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Crous et al. Table 1. Isolates for
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Crous et al. Table 1. (Continued).
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Crous et al. 100 10 changes 65 100
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Crous et al. Treatment of phylogene
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Crous et al. Teratosphaeria bellula
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Crous et al. 6. Conidiophores short
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Crous et al. Habit plant pathogenic
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Crous et al. Fig. 7. Catenulostroma
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Crous et al. system, consisting of
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Crous et al. Fig. 9. Penidiella col
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Crous et al. Fig. 12. Penidiella ri
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Crous et al. conidiophores, about 1
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Crous et al. have conidiomata rangi
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Crous et al. Schizothyriaceae clade
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Crous et al. Fig. 21. Stigmidium sc
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Crous et al. Gams W, Verkley GJM, C
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Crous et al. Table 1. Isolates for
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Crous et al. Fig. 3. Rachicladospor
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Crous et al. Fig. 4. Toxicocladospo
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Crous et al. Fig. 5. Verrucocladosp
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Crous et al. Fig. 7. Stenella aragu
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Crous et al. Helotiales, incertae s
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Crous et al. Fig. 10. Ochrocladospo
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Crous et al. 7. Conidiophores unbra
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Crous et al. Crous PW, Kang JC, Bra
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Arzanlou et al. To date 26 species
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Arzanlou et al. Table 1. (Continued
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Arzanlou et al. 10 changes Athelia
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Arzanlou et al. Athelia epiphylla A
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Arzanlou et al. Ramichloridium musa
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Arzanlou et al. Rhinocladiella mack
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Arzanlou et al. even further, thoug
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available online at www.studiesinmy
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Dichocladosporium gen. nov. 10 chan
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Dichocladosporium gen. nov. Fig. 3.
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Dichocladosporium gen. nov. to intr
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Dichocladosporium gen. nov. Czechos
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Cladosporium herbarum species compl
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Cladosporium sphaerospermum species
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Page 260 and 261: Ramichloridium epichloës, 60, 89 P
Page 262: Trimmatostroma salicis, 3 t , 5, 6
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