Chloridium and some other dematiaceous hyphomycetes growing on decaying wood

W. Gams
Centraalbureau voor Schimmelcultures, Baarn
V. Holubova–Jechova
Botanical Institute, Czechosl. Academy of Sciences, Průhonice near Praha


Phialidic species described in Chloridium are revised, described from the natural substrate and/or cultures and connected with perfect states wherever possible. The correct name of the type species of the genus is Chloridium virescens (Pers. ex Pers.) comb. nov. (basionym Dematium virescens Pers.), and the connection with Chaetosphaeria vermicularioides (Sacc. & Roum.) comb. nov. (syn. Eriosphaeria vermicularioides Sacc. & Roum.) has been established. C. chlamydosporis (van Beyma) Hughes and C. caudigerum (Höhn.) Hughes are considered as varieties of C. virescens. These three taxa constitute the section Chloridium which is characterized by phialides with multiple conidiogenous loci active within the collarette. Section Gongromeriza (Preuss) comb. nov. has phialides with a single basipetal sequence of conidia formed within a collarette, in section Psilobotrys (Sacc.) comb. nov. the phialides are constricted at the tip to a narrow collarette and mostly proliferate sympodially. Sect. Gongromeriza contains nine species (incl. C. pachytrachelum, C. phaeosporum, C. matsushimae and C. preussii spp. nov.) of which four are connected with Chaetosphaeria perfect states. Sect. Psilobotrys contains two species, Chaetosphaeria innumera Berk. & Br. ex Tul. (with C. botryoideum (Corda) Hughes var. botryoideum and C. botryoideum var. minutum (Sacc.) comb. nov. as conidial states), and Chaetosphaeria f. fusispora sp. nov. (with C. cylindrosporum st. nov. as its conidial state).

Twenty–five other dematiaceous phialidic genera with simple or little branched (but not penicillate) conidiophores are keyed out and five considered in detail. Cylindrotrichum oligospermum (Corda) Bonord. is close to Chloridium sect. Psilobotrys, but has two–celled conidia formed on polyphialides and sterile setae; C. hennebertii sp. nov. has similar conidiophores, slightly shorter conidia and no sterile setae; C. zignoëllae (Höhn.) comb. nov., the presumptive conidial state of Chaetosphaeria abietis (Höhn.) comb. nov. (syn. Zignoëlla abietis Höhn.) and Cylindrotrichum clavatum sp. nov. do not proliferate sympodially and have no sterile setae. Exochalara gen. nov. is described to accomodate Chalara longissima Grove and Catenularia guadalcanalensis Matsushima with large, catenulate, one–celled conidia. Smaller species with usually discrete, thin–walled phialides and catenulate conidia are accomodated in Phialophora sect. Catenulatae sect. nov., viz. Lasiosphaeria hirsuta (Fr.) Ces. & De Not. with Phialophora lasiosphaeriae st. nov., P. clavispora, P. oxyspora, P. olivacea, P. phaeophora, P. brachyconia, P. hyalina and P. brevicollaris spp. nov. and P. rhodogena (Mangenot) comb. nov. (syn. Scopulariopsis rhodogena Mangenot). The very similar Chalara state of Ceratocystis autographa Bakshi and Chalara microchona sp. nov. as well as the conidial state of Cryptendoxyla hypophloia Malloch & Cain are included for comparison. Four species (one new) and three new varieties of Gonytrichum are keyed out: G. mirabile sp. nov., G. caesium var. subglobosum and var. chloridioides and G. chlamydosporium var. simile vars. nov. The perfect state of G. chlamydosporium is Chaetosphaeria chloroconia sp. nov. Verticillium cyclosporum (Grove) Mason & Hughes and V. tenuissimum Corda are accomodated in the new genus Phaeostalagmus and described from pure culture. Single new species are described in both Codinaea and Fusichalara. [p. 2]


1. Introduction

The present study was initiated as a revision of the genus Chloridium sensu stricto, i.e. those species with unbranched, dark conidiophores and integrated phialides; those with blastoconidia borne on sympodially proliferating conidiogenous cells will be revised by G. S. de Hoog in due course. A clear circumscription of the genus proved difficult and, after some hesitation, it was decided to maintain one large genus (Chloridium) with several sections instead of several genera with few species. During this study we were able to demonstrate perfect state connections in several species. While considerable effort was taken to find the appropriate epithets for perfect states now placed in Chaetosphaeria Tul., no attempt has been made to review the numerous published names in Chaetosphaeria and the similar Zignoëlla Sacc.
While material was collected in several countries by us independently for several years, Chloridium-like fungi proved to be particularly abundant in Czechoslovakia and the available information was then combined in this communication. It is thus probably not merely a chance that A. C. J. Corda (1809-1849), working in Bohemia, was one of the first to recognize and describe Chloridium species. No geographical limits were imposed to this study, but European material naturally predominated. All specimens consist of decaying twigs or wood, unless otherwise stated. Most of the PRM collections were gathered by V. H.-J., mainly from Czechoslovakia (the different countries are abbreviated: Boh. = Bohemia, Mor. = Moravia, Slov. = Slovakia). The recent CBS strains were isolated by W.G., unless otherwise indicated. Herbaria are abbreviated according to the 6th Edition of the Index Herbariorum (Holmgren and Keuken, 1974). G.L.H. refers to the personal herbarium of Prof. Dr. G. L. Hennebert, Louvain-la-Neuve.
Chloridium chlamydosporis is generally known as a soft rot fungus which occurs in advanced stages of decay on wood (Mangenot, 1952; further references in Domsch and Gams, 1970, and Omvik, 1970). The same probably holds true for all fungi described in this paper as can be judged from their occurrence, but studies on their wood-decomposing capacities are usually not available. Species of Chloridium seem to have little (C. virescens) or no (C. lignicola) polyphenoloxidase activity and are unable to decompose lignin, but they nevertheless commonly occur on strongly decayed wood, the lignin of which may already have been decomposed by other fungi. Gonytrichum caesium, however, shows considerable laccase activity (Holubová-Jechová, 1971).
On strongly decayed substrates a dense intermixture of various Hyphomycetes and other fungi often occurs and so particular care is required to obtain pure cultures. Perfect-imperfect relationships can be proven reliably only by growing ascospores and studying conidium formation in culture. The latter does not present any problems on the conventional agar media, such as 2% malt extract, oatmeal, cornmeal and [p. 3] cherry decoction agars which were generally used during this study. Cultures were incubated at room temperature, i.e. 20-22°C. The structure of the perithecia was studied in freezing-microtome sections.

  fig. 1

Fig. 1
Chloridium virescens var. virescens, CBS 919.73, three conidiophores and chlamydospores in pure culture (left), percurrent proliferation and conidial cirrhus on the natural substrate, IMI 179577 (right). x 1000


2. Morphology and conidium ontogeny in Chloridium virescens (Fig. 1, 2)

The type species of the genus Chloridium shows some unusual features amongst the phialidic Hyphomycetes which are described first as a basis for the subsequent discussion.
Conidiophores are simple and, in the first instance, of determinate length; before the phialide opens and conidia are formed, the conidiophore becomes repeatedly septate and the uppermost integrated cell, which becomes the phialide, is delimited rather late. The first conidium is formed in a typical phialidic manner, being liberated through the bursting of the primary phialide wall which becomes a shallow collarette. Subsequently formed conidia take a lateral position in relation to the first and there are generally two, three or more conidia attached to the phialide tip. Successive conidium formation leads first to the formation of slimy heads, and, in some strains, subsequently to typical cirrhi (i.e. columns consisting of five or more imbricate rows of conidia). The mechanism of conidium formation has been elucidated by Hammill (1972) using TEM of ultrathin sections (fig. 2). The conidia are blown out sequentially from new growing points in the phialide apex but within the collarette. [p. 4].

  fig. 2

Fig. 2.
Chloridium virescens var. chlamydosporum, conidiogenesis in TEM, schematically redrawn from Hammill (1972).

The mechanism of conidium formation is intermediate between phialidic and sympoduloconidiogenous’. In view of the great length of the cirrhi it is highly probable that after some time the same conidiogenous locus continues to produce conidia without any further elongation of the phialide tip, so that a true basipetal sequence occurs. Since the ensuing elongation of the tip is normally not pronounced and in view of the taxonomic implications (cf. below), we prefer not to exclude this kind of conidiogenesis from the phialidic type but to distinguish it as a special modification with mutiple conidiogenous loci. Exactly the same mode of conidiogenesis is present in Gonytrichum (Swart, 1959). A similar phenomenon in Codinaea setosa Kendrick & Hughes was no reason for Kendrick and Hughes (1968) to exclude this species from Codinaea; moreover, in other species of that genus the distinction between single and multiple conidiogenous loci is scarcely possible by light microscopy. A very conspicuous analogous case is conidium formation in Cacumisporium capitulatum (Corda) Hughes (Goos, 1969; Sutton, 1973).
After completion of a series of conidia, percurrent proliferation commonly occurs; a new wall layer is apposed throughout the previous phialides (Hammill, 1972). The proliferating part may be as long as the original phialide or shorter, and further septa are formed near or below the points of proliferation. The whole process of sporulation can thus be repeated several times, leading to distinctive geniculate conidiophores.


3. Historical survey and generic concepts

Since the first description by Link (1809) of Chloridium viride, the identity and delimitation of Chloridium has not been well known and [p. 5] species with very different types of conidiogenesis have been referred to it. Mainly as a result of Saccardo’s (1877) illustration of Chloridium minutum, reproduced by Lindau (1906), even species with denticulate, polyblastic conidiogenous cells were included in Chloridium. Only since Hughes (1958) reexamined the type specimens and declared Cirrhomyces Höhn. and Bisporomyces van Beyma as congeneric, has the generic concept been connected with phialidic fungi. Hughes recognized five species, viz. C. botryoideum (Corda) Hughes, C. caudigerum (Höhn.) Hughes, C. chlamydosporis (van Beyma) Hughes, C. minutum (Sacc.) Sacc. and C. viride Link. Booth (1957, 1958) described connections of Chaetosphaeria innumera Berk. & Br. ex Tul., C. myriocarpa (Fr.) C. Booth and C. bramleyi C. Booth with Catenularia-like conidial states now regarded as belonging to Chloridium.
Mangenot (1952) gave a full description of Bisporomyces chlamydosporis van Beyma, in which he recognized both the multiple conidium formation and the percurrent proliferation of the conidiophores. He kept Cirrhomyces caudigerus Höhn. apart because of its hyaline mycelium, formation of conidia in cirrhi and often intercalary chlamydospores. In 1953 he described a further species as B. lignicola. Tubaki, who (1958) first described as Chloridium minutum (Sacc.) Sacc. a polyblastic fungus now recognized as Acrodontium crateriforme (van Beyma) de Hoog, later corrected his generic interpretation (Tubaki, 1963) and gave a good description of Chloridium chlamydosporis. Cultural variability and physiology of this species were studied by Omvik (1970). The peculiar multiple conidium formation was further investigated by Swart (1959), who also found similar structures in Gonytrichum. Barron (1968) discussed C. chlamydosporis and the conidial state of Chaetosphaeria myriocarpa, while Ellis (1971) treated only three of the species recognized by Hughes (1958), viz. C. viride, C. caudigerum and C. chlamydosporis. C. caudigerum is also illustrated by Sivasithamparam (1975) and C. chlamydosporis by Matsushima (1975).
Problems of generic delimitation arise with the classification of some related species, such as C. minutum and the conidial state of Chaetosphaeria myriocarpa. The former, as well as C. botryoideum, has polyphialides developing by sympodial proliferation. Conidia are protruded in a single row through a narrow neck. There is no recent description, illustration or discussion of these structures.
The imperfect state of Chaetosphaeria myriocarpa was described by Mangenot (1952) and Booth (1957, 1958) in Catenularia. Hughes (1965) refused to include it in Catenularia because that genus can be sharply delimited by the presence of large, thick-walled conidia and capitate hyphae. In 1969 Hughes discovered a first description of this conidial state under the name Gongromeriza clavaeformis Preuss (1851) (which thus affords a valid epithet), but placed the fungus in Chloridium. On the other hand, M. B. Ellis (pers. commun.) did not include it in Chloridium because the conidia are formed in a single row within a rather deep collarette. [p. 6]
Conidium ontogeny is recognized as a principal criterion in generic delimitation, although with some restriction. We consider that multiple and single conidiogenous loci in the phialide can occur within one genus. The slight protrusion of the phialide tip is not regarded as significant even at the specific level in Chloridium virescens, where the length of elongation is rather variable. C. pachytrachelum and C. lignicola in which conidia form in a single row present a link to the polyphialidic C. botryoideum and to C. clavaeforme with a deeper collarette. Instead of resurrecting Psilobotrys Sacc. and Gongromeriza Preuss as genera, we prefer to relegate these to section rank. The main character for the delimitation of Chloridium is thus the unbranched, pluricellular, strongly pigmented conidiophore with an integrated phialide, and often percurrent or sometimes sympodial proliferation. This generic disposition is further corroborated by the discovery of several related perfect states (cf. below).

  fig. 3

Fig. 3
Schematic representation of the sections distinguished in Chloridium: a. sect. Chloridium, b. sect. Gongromeriza, c. sect. Psilobotrys.

Because of the intricate generic and specific relationships we decided to extend our study to some related genera as well, while many more are included in the generic key. Common features of these genera are the phialidic and dematiaceous character and the unbranched or little branched (not penicillate) dark conidiophores.
A similar, but hyaline species with mostly simple phialides was described by Mangenot (1952) as Scopulariopsis rhodogena. Since Hughes (1953) recognized that Scopulariopsis can be defined by annellophores, this fungus has not found an appropriate genus. In 1951 Hughes described the conidial state of Lasiosphaeria hirsuta (Fr.) Ces. & De Not. which is similar to S. rhodogena, but has a slightly pigmented mycelium and equally catenulate conidia with a truncate base. Hughes regarded it as belonging to Phialophora; this conclusion is further corroborated by our observation (Gams, 1973) that the closely related L. ovina (Fr.) Ces. & De Not. forms a similar conidial state but with capitulate conidia. The genus Phialophora as circumscribed by Schol-Schwarz (1970) is rather heterogeneous, but for the time being obviously an appropriate taxon in [p. 7] which to accomodate a range of species also with catenulate conidia. It is perhaps because of the lack of a convenient generic name, that a number of catenulate Phialophora species have never been formally described. Their delimitation from Chalara raises many problems. Until the publication of the monograph of Chalara and related genera by Nag Raj and Kendrick (1976) we regarded some species as belonging to Phialophora which we now place in Chalara (in particular Ch. microchona). Such species have been included in the Phialophora key here, in order to prevent misidentifications.
Whilst the integrated phialides in Chloridium and the discrete phialides in Phialophora have diagnostic value at the genus level, the presence or absence of several stalk cells in the conidiophore of Chalara (which decides whether a phialide is integrated or discrete), was not found to have any generic value (Nag Raj and Kendrick, 1976).


4. Perfect state connections

Chaetosphaeria Tul. was described by Tulasne and Tulasne (1863) as having smooth ascomata, whithout setae but sometimes beset with some conidiophores, pellucid ascospores, and being connected with a conidial state similar to Dematium virescens. Saccardo (1883), however, altered the generic concept to perithecia villosa vel glabrescentia . . . sporidia tota vel partim fuliginea’. Since then a host of unrelated species has been placed in Chaetosphaeria. Booth (1957, 1958) redescribed the type species, Chaet. innumera Berk. & Br., but could not observe the decisive features on the type specimen (i.e. the three-septate ascospores and conidiophores with polyphialides) and thus the identity of this fungus has remained somewhat doubtful. In his description of Chaetosphaeria myriocarpa, on the other hand, no doubts about the identity were left. Some other Chaetosphaeria species recognized by Booth have typical Catenularia, Menispora or Codinaea states and four-celled ascospores. Müller and von Arx (1962) followed Booth and characterized Chaetosphaeria as a genus of the Sphaeriaceae having dark, smooth-walled perithecia with a two-layered wall, thin-walled asci with a simple apical plate and two- or more-celled, hyaline ascospores. Dennis (1960, 1968), however, regarded Chaetosphaeria as having two-celled ascospores only, while the related genus Zignoëlla Sacc. would be reserved for phragmosporous species (e.g. Z. ovoidea (Fr.) Sacc.). Munk (1953, 1957), after studying Z. fallax (Sacc.) Sacc., placed the genus in the Diaporthaceae. This species, however, was not amongst the 27 original species of the genus (Saccardo, 1878) and cannot be regarded as its type species, nor was it designated as such by Munk. The only selection of a type species is apparently that of Clements and Shear (1931) who chose Z. pulviscula (Curr.) Sacc., a species with four-celled ascospores and a Menispora state, placed by Booth in Chaetosphaeria. Hence Zignoëlla [p. 8] becomes a synonym of Chaetosphaeria in the broader sense, while for the diaporthaceous fungi another generic name has to be found. Zignoëlla ovoidea (Fr.) Sacc., the first species mentioned by Saccardo (1883), is probably a Chaetosphaeria. The only specimen preserved in Fries’s herbarium (UPS) labeled ‘Sphaeria ovoidea v. minor, Upsaliae’, contains Lophiostoma quadrinucleatum Karst. var. triseptatum (Peck) Chesters & Bell; this fungus does not agree with the protologue, since the semi-immersed ascomata have a distinct cleft-like ostiolum; it therefore cannot be considered as type of the species. Other specimens preserved at UPS and PAD as Sphaeria and Zignoëlla ovoidea contain various Chaetosphaeria-like fungi without conidiophores.
Even if only hyaline-spored species with phialidic conidial states are recognized in Chaetosphaeria, the genus still remains large and somewhat heterogeneous. It now contains conidial states belonging to Chloridium, Catenularia, Zanclospora, Menispora, Codinaea, Stachybotrys and Sporoschisma (Kendrick and Hughes, 1968). The conidial states are particularly important in differentiating between species with hardly distinguishable perfect states.
The perfect state of Gonytrichum caesium Nees ex Leman was described in a separate genus, Melanopsammella, by von Höhnel (1919) because of the fragmenting ascospores; this genus was reduced to synonymy with Trichosphaerella Bomm., Rouss. & Sacc. by Müller and von Arx (1962). Since species of Trichosphaerella have very distinct, simple or branched black setae on the ascomata and Melanopsammella has not (except for some hyphae bearing conidiophores), and since the conidial states are different (for the conidial state of Trichosphaerella arecae (Sydow) E. Müller see Gams, 1971), these two genera should not be combined. Melanopsammella is very close to Chaetosphaeria and can best be regarded as a section of it which is distinct by ± early separation of the two-celled ascospores. This section then also accomodates the perfect state of Chloridium virescens. It is not surprising to find Gonytrichum spp. and Chloridium virescens having closely related perfect states, since both have the same kind of conidiogenesis with multiple conidiogenous loci.
It is of particular note that, as far as we know, the three sections distinguished in Chloridium on the basis of the imperfect states alone correspond to Chaetosphaeria sect. Melanopsammella, didymosporous Chaetosphaeria (syn. Lentomita Niessl) and phragmosporous Chaetosphaeria. An exception to this rule in Chaetosphaeria preussii W. Gams & Hol.-Jech. which has a conidial state of section Gongromeriza but fragmenting ascospores. This finding has led us to decide to not retain the generic distinction between Chaetosphaeria and Melanopsammella after all.
Although it is not general practice, most perfect and imperfect states are described and named separately in this paper for various reasons: The connection between both states has often not yet been proved by ascospore isolations and is based only on statistical probability. The perfect states in this group of the Sphaeriaceae are often less differentiated than the conidial states in which several varieties, species or even genera [p. 9] can be distinguished with almost identical perithecia and ascospores. It is therefore often not warranted to conclude that a given conidial state will produce a certain perfect state under adequate conditions on the natural substrate. In some cases species delimitation of the conidial states also requires further studies.


5. Key to the genera (cf. Fig. 4)


Conidiophores unbranched (although sometimes showing sympodial elongation), pluricellular, with integrated terminal (and sometimes intercalary) phialides



Conidiophores branched or consisting of a single phialide






Upper part of the conidiophore consisting of a series of intercalary phialides with lateral collarettes (cf. Sutton, 1973)

Zakatoshia Sutton


Terminal phialides only (sometimes sympodially proliferating)






Conidia formed within a long (at least the length of the conidium), tube–like collarette



Conidia formed within a shorter, widening or almost absent collarette






Conidia dark, phragmosporous; capitate hyphae present (cf. Nag Raj and Kendrick, 1976)

Sporoschisma Berk. & Br.


 Conidia pigmented or hyaline, unicellular or septate; capitate hyphae absent






Sterile setae present (cf. Nag Raj and Kendrick, 1976)

Chaetochalara Sutton & Piroz.


Sterile setae absent






Conidia fusiform (cf. Nag Raj and Kendrick, 1976)

Fusichalara Hughes & Nag Raj


Conidia ± cylindrical (cf. Nag Raj and Kendrick, 1976)

Chalara (Corda) Rabenh.





Conidia more than two–celled



Conidia one– or two–celled






Conidia cylindrical, arising within a flaring collarette (cf. Holubová–Jechová and Hennebert, 1972)

Sporoschismopsis Hol.–Jech. & Henneb.


Conidia elongate–ellipsoidal to allantoid, arising from a meristem protruding beyond the collarette (cf. Goos, 1969; Sutton, 1973)

Cacumisporium Preuss





Conidia cuneate, thick–walled, pigmented



Conidia mostly of other shapes, thin–walled, hyaline or slightly pigmented [p. 10]


 fig. 4

Fig. 4.
Survey of the genera mentioned in the key, schematic and not drawn to scale (genera roughly arranged according to similarity). [p. 11]

10a. .

Capitate hyphae present (cf. Hughes, 1965)

Catenularia Grove in Sacc.


Capitate hyphae absent (cf. Linder, 1933)

Haplochalara Linder





Conidia elongate, allantoid or lunate, with or without terminal setulae; phialides sympodially proliferating (cf. Kendrick and Hughes, 1968)

Codinaea Maire


 (if conidiophores synnematous around a dark seta Menisporopsis Hughes)



Conidia of other shapes, usually not curved






Conidia fusiform or ellipsoidal, adhering end–to–end in long chains; phialides proliferating only percurrently

Exochalara W. Gams & Hol.–Jech.


Conidia globose, ellipsoidal, cylindrical, clavate or slightly cuneate, forming globose heads (rarely chains), imbricate chains or cirrhi






Conidia one–celled

Chloridium Link ex Fr.


Conidia two–celled






Conidia obclavate; sympodial elongations long, repeatedly septate (cf. Matsushima, 1971)

Phialogeniculata Matsushima


Conidia cylindrical; sympodial elongations mostly present, short and generally without further septation; sterile setae sometimes present

Cylindrotrichum Bonord.





Conidiophores consisting of single or a cluster of phialides with little differentiated stalks; collarette usually well developed



Conidiophores distinct and pigmented, with a clearly differentiated stalk






Phialides densely packed on a basal stroma, sack–like, filled largely with a phragmosporous conidium released by apical rupture (cf. Nag Raj and Kendrick, 1976)

Ascoconidium Seaver


 Phialides arising from discrete vegetative hyphae, more slender






Collarette typically cylindrical, usually as long or longer than a conidium; conidia catenulate, cylindrical or clavate and elongate (cf. Nag Raj and Kendrick, 1976)

Chalara (Corda) Rabenh.


(if conidiophores form pustular sporodochia and conidia cuboid Bloxamia Berk. & Br.)



Collarette ± divergent or even reflexed, usually shorter; conidia usually shorter and of different shape






Conidia borne in heads (cf. Schol–Schwarz, 1970)

Phialophora Medlar (most species)


Conidia borne in chains, usually with truncate bases

Phialophora sect. Catenulatae W. Gams (





Conidiophores dichotomously branched, with integrated apical phialides (cf. Holubová–Jechová, 1972)

Craspedodidymum Hol.–Jech.


Conidiophores not dichotomously branched



[p. 12]



Phialides in appressed whorls arising from the central cells of the conidiophore stipes (cf. Ellis, 1971)

Zanclospora Hughes & Kendrick


Phialides at the end of ± verticillate branches






Phialides with dense sympodial proliferations (polyphialides) reminiscent of polyblastic conidiogenous cells



Phialides usually with a single apical opening (monophialides)






Conidiophores with branches in terminal whorls (cf. Sutton, 1973)

Selenosporella Arnaud ex MacGarvie


Conidiophores with short fertile branches in the lower part, upper part sterile (cf. Hughes, 1951; Barron, 1968)

Chaetopsis Grev.





Verticillate phialides arising from an irregularly contorted basal cell



Verticillate or solitary branches arising directly from the conidiophore stipe



Phialides with a wide opening and multiple conidiogenous loci; conidia globose to cylindrical

Gonytrichum C. G. & F. Nees ex Leman


(if conidia allantoid Codinaeopsis gonytrichoides (Shearer & Crane, 1971) Morgan–Jones)



Phialides with a narrow opening and a single conidiogenous locus; conidia narrowly cylindrical (cf. Barron, 1968; Ellis, 1971)

Chaetopsina Rambelli





Conidia allantoid or lunate, often with terminal setulae; phialide openings often recurved (cf. Hughes and Kendrick, 1963; Kendrick and Hughes, 1968)

Menispora Pers.


Conidia short–ellipsoidal; phialide openings straight






Phialides with a conspicuous, wide, dark collarette

Phaeostalagmus W. Gams


Phialides constricted at the tip, without a distinct collarette (cf. Hughes, 1951b)

Stachylidium Link ex Gray


[p. 13]



6. Chloridium Link ex Fr.

Chloridium Link – Mag. Ges. naturf. Freunde, Berl. 3: 13. 1809; ex Fr. – Syst. mycol. 1: XLVI. 1821.
Gongromeriza Preuss – Linnaea 24: 106. 1851.
Rhinotrichum Corda sect. Psilobotrys Sacc. – Michelia 1: 87. 1877 = Psilobotrys (Sacc.) Sacc. – Michelia 1: 538. 1879.
Cirrhomyces Höhn. – Annls mycol. 1: 529. 1903.
Bisporomyces van Beyma – Antonie van Leeuwenhoek 6: 276. 1940.
Monostachys Arnaud – Bull. trimest. Soc. mycol. Fr. 69: 278. 1953, nom. inval. (Art. 36). Sphaeromycetella Arnaud – Bull. trimest. Soc. mycol. Fr. 69: 279. 1953, nom. inval. (Art. 36).
Holotype species Chloridium viride Link ex Link = Chloridium virescens (Pers. ex Pers.) W. Gams & Hol.–Jech.
Ascigerous states in Chaetosphaeria Tul.

Genus of phialidic dematiaceous Hyphomycetes with erect, unbranched, pigmented conidiophores arising from submerged hyphae, and apical, integrated phialides. Phialides with a wide or narrow, deep or short collarette, in some species with multiple conidiogenous loci, budding conidia in succession within the collarette. Percurrent proliferation of conidiophores common, sympodial proliferation occurring in some species. Conidia arranged in slimy heads or gradually extending cirrhi, rarely in simple chains, but then mostly imbricate; one–celled, globose to ellipsoidal or cylindrical, hyaline or almost hyaline, smooth–walled, mostly containing one or two guttules. Chlamydospores present in some species, pigmented, globose or elongated, terminal, lateral or more rarely intercalary.
Three sections can easily be distinguished: Chloridium with multiple conidiogenous loci and pigmented chlamydospores, Gongromeriza with single conidiogenous loci and percurrent proliferation, and Psilobotrys with single conidiogenous loci and sympodial proliferation.

Key to the species


Proliferation of the conidiophores only percurrent; collarette over 2 µm diam



Proliferation mainly sympodial (besides rarely percurrent); collarette generally not exceeding 1.5 µm diam Sect. Psilobotrys






Multiple conidiogenous loci active within the collarette; conidia forming complex cirrhi or slimy heads; chlamydospores generally present Sect. Chloridium



(if chlamydospores absent and conidiophores arising from aerial hyphae or if conidia pigmented see Gonytrichum)



Single conidiogenous loci producing a basipetal sequence of conidia within the collarette; conidia forming slimy heads or sometimes chains; chlamydospores absent Sect. Gongromeriza



[p. 14]



Conidia always in heads, 3.0-5.0 x 1.7-2.7 length/width 1.7-2.2

C. virescens var. chlamydosporum


Conidia at least partly in cirrhi






Cirrhi yellow-green to herbage green; conidia 2.7-3.8 x 2.0-2.5 µm, length/width 1.2-1.5 (in culture up to 4.0 µm long)

C. virescens var. virescens


Cirrhi whitish; conidia 3.5-5.0 x 1.8-2.5 µm, length/width 1.7-2.2

C. virescens var. caudigerum





Conidiophores dark brown, also in the phialide; phialides tapering gradually towards the tip, with usually conspicuous, beaker-shaped or flaring collarettes



Conidiophores much less pigmented in the phialides than in the basal part; phialides ± constricted only below the tip, collarette usually less conspicuous






Conidia ellipsoidal, hyaline, 2.5-3.5 x 1.5-2.0 µm

C. preussii


Conidia cuneate, with truncate base and rounded tip






Conidia slightly pigmented, not more than two enclosed within the collarette

C. clavaeforme


Conidia hyaline, 3-5 included in a long collarette

Chloridium state of Chaetosphaeria bramleyi





Conidia cylindrical or clavate, length/width exceeding 3.0



Conidia shorter, ellipsoidal or reniform






Conidia cylindrical, with a slimy apical appendage, 15-19 x 3.6 5.3 µm

C. matsushimae


Conidia fusiform or clavate, without any appendage, 5-9 x 1.0-1.5 µm

C. codinaeoides





Conidia reniform, 3.5-5.5 x 2.0-3.0 µm

C. reniforme


Conidia ellipsoidal






Conidia pigmented, 4.0-5.0 x 1.7-2.2 µm

C. phaesosporum


Conidia hyaline






Phialides with a 2-3 µm deep beaker-shaped collarette which often includes two conidia at a time; conidia 3.5-5.0(-7.0) x 1.5-2.0 µm; colonies olivaceous-grey

C. lignicola


Phialides with a 3-8 µm deep collarette which includes only one conidium at a time, and a pronounced wall thickening below it; conidia 3.5-6.0 x 2.0-3.0 µm; colonies light brown

C. pachytrachelum





(from 1) Conidia cylindrical, 8.5-11.5 x 2.2-3.0 µm

C. cylindrosporum


Conidia ellipsoidal or if cylindrical, not exceeding 4.0 x 1.7 µm






Conidia narrow ellipsoidal to cylindrical, 3.0-4.0 x 1.0-1.7 µm

C. botryoideum var. botryoideum


Conidia ellipsoidal to subglobose, mostly obliquely apiculate at the base, 2.5-4.0 x 1.5-2.0 µm

C. botryoideum var. minutum


[p. 15]



Chloridium section Chloridium

Colonies rather slow-growing. Conidia formed on a non-constricted phialide tip within a shallow collarette with multiple conidiogenous loci, often in elongate cirrhi; percurrent proliferation commonly occurring. Pigmented chlamydospores present.
Type species Chloridium virescens (Pers. ex Pers.) W. Gams & Hol.-Jech.


1a. Chaetosphaeria vermicularioides (Sacc. & Roum.) W. Gams & Hol.-Jech. comb. nov. - Fig. 5 a-b.

Eriosphaeria vermicularioides Sacc. & Roum. - Revue mycol. 5: 235. 1883.

  fig. 5

Fig. 5
Chaetosphaeria vermicularioides, a. asci, spores and conidiophores of var. caudigerum from isotype specimen in L; b. CBS 675.74, immature and mature asci, spores and conidiophores of var. chlamydosporum. x 1000.

Perithecia superficial, scattered, broadly attached to the substrate, without subiculum, black, subglobose, ostiolar region slightly conical, 120-225(-270) µm diam; wall 13-20(-25) µm thick, composed of numerous layers of dark cells (textura epidermoidea). Asci cylindrical, thin-walled, 55-80 µm long, 3.5-4.5(-6.0) µm wide, with a thin apical plate. Ascospores arranged in a single row, hyaline, smooth or faintly roughened, [p. 16] ellipsoidal to slightly fusiform, with a thick, refringent and chromophilic septum in the middle, mostly breaking apart in a rather early stage within the ascus; part spores of almost equal size and shape, conical with rounded ends, 3,2-5.0 x 2.0-3.0 µm. Paraphyses regularly cylindrical, 1.0-1.5 µm wide. Conidial state Chloridium virescens.

Perithecia superficial, scattered, broadly attached to the substrate, without subiculum, black, subglobose, ostiolar region slightly conical, 120-225(-270) µm diam; wall 13-20(-25) µm thick, composed of numerous layers of dark cells (textura epidermoidea). Asci cylindrical, thin-walled, 55-80 µm long, 3.5-4.5(-6.0) µm wide, with a thin apical plate. Ascospores arranged in a single row, hyaline, smooth or faintly roughened, [p. 16] ellipsoidal to slightly fusiform, with a thick, refringent and chromophilic septum in the middle, mostly breaking apart in a rather early stage within the ascus; part spores of almost equal size and shape, conical with rounded ends, 3,2-5.0 x 2.0-3.0 µm. Paraphyses regularly cylindrical, 1.0-1.5 µm wide. Conidial state Chloridium virescens.

Material examined

Cultures from ascospores (and herbarium specimens)
CBS 675.74, ex Carya alba, Arboretum d’Amance nr. Nancy, France, 16 Sept. 1974.
CBS 676.74, ex deciduous wood, Schaelsberg nr. Valkenburg, Z.L., Netherlands, 22 July 1914.
CBS 667.75, ex deciduous wood, Le Roptai nr. Ave-et-Auffe, Belgium, Sept. 1975.

Herbarium specimens
Eriosphaeria vermicularioides Sacc. & Roum., isotype, Fungi Gall. exs. 2693, Malmedy, Belgium (L 9,0.236-665) (with conidia of var. caudigerum).
Sub ‘Eriosphaeria vermicularia (Nees) Sacc. on Robinia (?)’ (PAD).
Sub. ‘Chloridium ... an nova species’, ex Herb. Link (B, slide DAOM 40814).
PRM 793968, Fraxinus excelsior, Úpor Forest nr. Mĕlník, Boh., leg. Z. Pouzar, 6 Nov. 1971 (conidia of var. virescens and var. chlamydosporum).
PRM 793974, Quercus petraea, Vůznice valley nr. Nižbor, Boh., leg. Z. Pouzar, 7 Oct. 1951 (conidia of var. virescens).
PRM 793998, Fraxinus excelsior, Žebračka Forest nr. Přerov, Mor., 14 Sept. 1972 (conidia of var. virescens and var. chlamydosporum).
PRM 794033, Abies alba, Dúbrava Forest nr. Banská Bystrica, Slov., 20 Sept. 1971 (conidia of var. caudigerum).
PRM 794040, Fagus sylvatica, Kremnické pohorie Mts. nr. Banská Bystrica, Slov., 9 Aug. 1969 (conidia of var. virescens).
PRM 794063, Fagus sylvatica, Mt. Kámen nr. Nepomuk, Boh., 3 Oct. 1975 (conidia of var. chlamydosporum).
PRM 796735 and 296761, Popalus nigra, 796752, Fraxinus excelsior, 796762, Ulmus effusa, Úpor Forest nr. Mĕlník, Boh., 24 June 1976 (conidia of var. virescens and var. caudigerum).


The connection between Chaet. vermicularioides and C. virescens has been proven repeatedly by isolation of ascospores. The original material of CBS 676.74 and 667.75 had conidia arranged in cirrhi matching var. caudigerum and measuring mostly 3.0 x 2.0 µm. The original material of CBS 675.74 contained some typical conidiophores of var. chlamydosporum besides typical var. virescens; the cultures from ascospores matched the former variety. All three varieties of conidial states were represented in the PRM collections and no correlation could be found between slight differences in ascospore size and conidium shape and arrangement.
Chaet. vermicularioides is close to Chaet. inaequalis (Grove) W. Gams & Hol.-Jech., but the ascospores are slightly longer.
Eriosphaeria vermicularioides distributed by Petrak in Fl. Boh. Moray. exs. Lfg. 1, Nr. 2 (K) is a different fungus, probably belonging to Trichosphaeria. [p. 17]


1b. Chloridium virescens (Pers. ex Pers.) W. Gams & Hol.-Jech. comb. nov. var. virescens - Fig. 1, 6.

Dematium virescens Pers. - Tent. Dispos. meth. Fung. p. 41. 1797; Synops. meth. Fung. p. 698. 1801; ex Pers. - Mycol. Europ. 1: 14. 1822; Fries - Syst. mycol. 3: 363. 1832.
Chloridium viride Link - Mag. Ges. naturf. Freunde, Berl. 3: 13. 1809; ex Link - Linné Sp. Pl., Ed. 4, 6(1): 38. 1824 = Cirrhomyces viridis (Link ex Link) Höhn, apud Lindau - Rabenh. Krypt.-Fl. 1(9): 793. 1910 = Chloridium dispersum Nees - Syst. Pilze Schwämme, p. 66. 1816/17 (incl. of C. viride); ex Pers. - Mycol. Europ. 1: 15. 1822 = Dematium asserculorum Pers. - Mycol. Europ. 1: 15. 1822 (incl. of C. viride).

  fig. 6

Fig. 6
Chloridium virescens var. virescens, a. holotype of Mesobotrys flavovirens (FH), b. PRM 793974. x 1000. [p. 18]

Sporotrichum virescens Link - Mag. Ges. naturf. Freunde, Berl. 3: 13. 1809; ex Pers. -Mycol. Europ. 1: 14. 1822; Link - Linné Sp. Pl., Ed. 4, 6(1): 16. 1824 (fide Link, 1824).
Doratomyces viridis Corda - Weitenwebers Beitr. ges. Nat.-Heilwiss. p. 83. 1836.
Haplaria chlorina Ell. & Ev. - Bull. Torrey bot. Club 10: 97. 1883.
Mesobotrys flavovirens Höhn. - Sber. K. Akad. Wiss. Wien, Math.-nat. Kl. 111: 1048. 1902 (Fragm. Mykol. No. 58) = Cirrhomyces flavovirens (Höhn.) Höhn. apud Lindau - Rabenh. Krypt.-Fl. 1(8): 760. 1907.

On the natural substrate forming a dense layer of yellow-green to grass-green conidial cirrhi. Conidiophores relatively short, mostly 30-60(-over 100) µm long, but of variable length, 2.5-3.5 µm wide, tapering to 2.0-2.5 µm, with a very slight constriction below the collarette; on ageing repeatedly proliferating percurrently; collarette inconspicuous, 2.0-2.5 µm wide. Conidia short ellipsoidal, hyaline, 2.5-3.8 x 2.0-2.5 µm, length/ width ratio 1.2-1.5, mostly uniguttulate.
In culture attaining 15 mm diam in 14 days at 20-22°C, brownish grey, with a hyaline margin of variable width. Vegetative hyphae partly hyaline, partly pigmented and strongly encrusted, 2-4 µm wide. Conidiophores arising from the substrate mycelium (phalacrogenous), erect, 30-60 (-100) x 2.5-4.0 µm, several times septate, light brown, sometimes roughened above, repeatedly percurrent after three weeks or more. Phialides less pigmented than the lower conidiophore cells, hardly constricted below the inconspicuous collarette, 1.5-2.5 µm wide; the meristematic tip often protruding beyond the collarette and producing several conidia side by side from successively formed conidiogenous loci (for details of conidiogenesis see p. 3). Conidia mostly aggregated in slimy heads, in old cultures partly in cirrhi, ellipsoidal, smooth-walled, hyaline, 3.2-4.0 x 2.0-2.5 µm, length/width ratio 1.3-1.8, uni- or biguttulate. Chlamydospores in typical strains scarce, in terminal, lateral or also intercalary position, ovoid or ellipsoidal, solitary or in short chains, 4.5-6.0 x 2.5-4.5 µm, in other strains more numerous, ± globose, 4-5 µm diam.

General distribution

C. virescens (with its varieties) is very common on decaying twigs and wood of very many types in advanced stages of decay and has also been frequently isolated from soil (Domsch and Gams, 1970). The variety virescens is apparently rare in the Netherlands and Great Britain, but very common in Czechoslovakia and also in Belgium.

Material examined

Cultures (and herbarium specimens)
CBS 919.73 = IMI 179 577, Dinas Powys Wood, Powys, G.B., leg. J. P. Ellis, det. M. B Ellis, 3 Sept. 1973 (first record for G.B.).
CBS 982.73, Pijnenburg Forest nr. Baarn, Netherlands, 19 Aug. 1972.
CBS 224.74, ex Alnus glutinosa, nr. Kortenhoef, Netherlands, 3 Febr. 1974.
CBS 561.74, ex Quercus sp., Groeneveld nr. Baarn, Netherlands, Sept. 1974. [p. 19]
CBS 548.75, ex Quercus sp., nr. Leningrad, U.S.S.R., leg. G. S. de Hoog, Aug. 1975.
The specimens collected in the Netherlands did not show the typical green cirrhi but very short conidia.

Herbarium specimens
Chloridium virens, Rostock, Herb. Link (B), also slides IMI 110798 and DAOM 40815, lectotype for C. virens, neotype for C. virescens.
Doratomyces viridis Corda (PRM 155471, also slide DAOM 40986).
Haplaria chlorina Ell. & Ev., on rotten Magnolia, Newfield, July 1883 (DAOM 35346, slide ex NY).
Mesobotrys flavovirens Höhn., on decaying wood, Dammbachthal b. Purkersdorf, Wiener-Wald, 1 June 1902, holotype; second specimen on oak wood, Gelber Berg, Wiener-Wald, 29 May 1904, containing also var. chlamydosporum (FH).
DAOM 44805, Fagus sylvatica, Fontainebleau, France, leg. S. J. Hughes, July 1954.
DAOM 51685, Grib Skov, N-Sealand, Denmark, leg. J. Webster, 2 Oct. 1955.
DAOM 51695, Fagus sylvatica, Jaegersborg Dyrehave nr. Copenhagen, Denmark, leg. S. J. Hughes, 3 Oct. 1955.
G.L.H. 6858, Picea abies litter, Meerdael Bos nr. Heverlee, Belgium, leg. H. Maraite, 2 June 1964.
G.L.H. 8920 and 8984, Pteridium aquilinum, Meerdael Bos nr. Heverlee, Belgium, 16 May 1966.
Numerous specimens were examined from Czechoslovakia: PRM 793963, 793964, 793966, 793976, 793978, 793979, 793980, 793992, 793993, 793996, 793999, 794001, 794003, 794014, 794024, 794026, 794030, 794038, 794047, 794051, 794052, 794064, 794073, 794074, 794081, 794093, 794095, 794100, 794101, 796751, 796753, 796754, 796755. This variety occurs abundantly in Czechoslovakia throughout the year from riverside forests to the mountains; it was collected on rotten wood and bark on Acer sp., Alnus glutinosa, Carpinus betulus, Corylus avellana, Fagus sylvatica, Fraxinus angustifolia, Quercus cerris, Q. petraea, Q. robur, Q. rubra and Ulmus effusa. In some specimens listed conidial heads with longer, almost cylindrical conidia were present in patches adjacent to typical conidial cirrhi.


1c. Chloridium virescens (Pers. ex Pers.) W. Gams & Hol.-Jech. var. caudigerum (Höhn.) W. Gams & Hol.-Jech. comb. nov. - Fig. 5a, 7.

Cirrhomyces caudigerus Höhn. - Annls mycol. 1: 529. 1903 = Chloridium caudigerum (Höhn.) Hughes - Can. J. Bot. 36: 748. 1958.
? Stilbum capillamentosum Preuss - Linnaea 24: 132. 1851 = Stilbella capillamentosa (Preuss) Lindau - Rabenh. Krypt.-Fl. 1(9): 292. 1908.
Sphaeromycetella leucocephala Arnaud - Bull. trimest. Soc. mycol Fr. 69: 274. 1953, nom. inval. (Art. 36).

On the natural substrate forming whitish conidial cirrhi, conidia measuring 3.5-4.5 x 1.8-2.5 µm, length/width ratio 1.7-2.2.
In culture reaching 14-24 mm diam in 14 days at 20-22°C, hardly distinguishable from C. virescens var. chlamydosporum, except for the eventual formation of cirrhi in old cultures and the often predominating dark vegetative hyphae. Conidia ellipsoidal, sometimes slightly curved, uniguttulate, 3.0-5.0 x 1.6-2.7 µm, length/width ratio 1.6-2.2. Chlamydospores rather scarce, in terminal, lateral or also intercalary position, often elongate, sometimes in groups of two or three, 4-6(-8.5) x 3.0-5.0 µm. [p. 20]

  fig. 7

Fig. 7
Chloridium virescens var. caudigerum, a. from neotype specimen (FH), b. DAOM 51736. x 1000

  fig. 8

Fig. 8
Chloridium virescens var. chlamydosporum, CBS 526.73. x 1000.

Material examined

Cultures (partly with original specimens)
CBS 152.53, ex Acer sp., comm. F. Mangenot.
CBS 141.54 and 142.54, ex soil, Yangambi, Zaïre, comm. J. Meyer.
CBS 709.73A, 709.73B and 709.73C, ex wheat-field soil, Kiel-Kitzeberg, isol. W. Gams.
CBS 239.75C (strain 90), ex forest soil, Senart nr. Paris, RCP 40, comm. O. Reisinger.
CBS 260.75A, ex Quercus sp., Groeneveld nr. Baarn, Netherlands, 1974.
CBS 601.75B, nr. Neupont, Belg. Ardennes, Sept. 1975.
CBS 667.75, Le Roptai nr. Ave-et-Auffe, Belgium, Sept. 1975.
CBS 676.75, Pijnenburg Forest nr. Baarn, Netherlands, Aug. 1972.
CBS 263.76, Mt. Pidurutalagala, Ceylon (c. 2000 m alt.), leg. W. Gams, Jan. 1973.
CBS 421.76, ex Populus nigra, Úpor Forest nr. Mĕlník, Boh., V. Holubová-Jechová, 24 June 1976.
CBS 424.76 = MUCL 7830-A, ex Quercus robur, Parc d’Arenberg, Heverlee, Belgium, G. L. Hennebert, 29 Sept. 1965.

Herbarium specimens
Stilbum capillamentosum Preuss, on Alnus wood, Herb. Preuss (B), holotype.
Cirrhomyces caudigerus Höhn., three collections containing different fungi: Gelber Berg, Wiener Wald, on decaying Quercus wood, 29 May 1904, designated as neotype; Rekawinkel-Dürrwien, 28 July 1907 (also DAOM 54554), is var. chlamydosporum; on Fagus sylvatica, Georgenberg, Purkersdorf, 2 Aug. 1907, is C. botryoideum var. minutum (FH).
Sphaeromycetella leucocephala Arnaud, Parc de Versailles, two collections from 15 May 1946 and 16 May 1946, both with conidial cirrhi fitting this variety (PC).
IMI 50200, ex soil, Nottingham, R. H. Thornton, 1953.
IMI 55844, 55846, 55847, ex soil, Yangambi, Zaïre, J. Meyer, 1954.
IMI 68326, ex Woburn soil, E. A. Peterson.
DAOM 43966, South March, Ont., S. J. Hughes, 19 May 1954. [p. 21]
DAOM 51736, Picea abies, Ještĕd Mt. nr. Liberec, Boh., S. J. Hughes, 10 May 1955.
G.L.H. 6454, Quercus Sp, Egenhoven nr. Heverlee, Belgium, 24 Oct. 1964.
PRM 794043, Acer platanoides, Mt. Javořina, nr. Zborov, Slov., 9 Aug. 1972.
PRM 794048, Quercus petraea, North of Vel. Šariš, Slov., 8 Aug. 1972.
PRM 794065, Fagus sylvatica, Mt. Kámen nr. Nepomuk, Boh., 3 Oct. 1975.
PRM 794067, Picea abies, Mt. Sedlo nr. Sušice, Boh., 1 Oct. 1975.
PRM 794084, Quercus robur, Sirková Voda Forest between Brodské and Kopčany, Slov., 17 Sept. 1975.
PRM 794086, Quercus cerris, Mt. Krehora nr. Vel. Krtíš, Slov., 11 Aug. 1975.
PRM 796759, 796760, 796764, Populus nigra, 796763, Fraxinus excelsior, Úpor Forest nr. Mĕlník, Boh., 24 June 1976.
PRM 796765, Betula verrucosa, Lánská obora Forest nr. Rakovník, Boh., 12 June 1976.


1d. Chloridium virescens (Pers. ex Pers.) W. Gams & Hol.-Jech. var. chlamydosporum (van Beyma) W. Gams & Hol.-Jech. comb. nov. - Fig. 5b, 8.

Bisporomyces chlamydosporis van Beyma - Antonie van Leeuwenhoek 6: 277. 1940 = Chloridium chlamydosporis (van Beyma) Hughes - Can. J. Bot. 36: 748. 1958.
Sphaeromycetella ?macrospora Arnaud - Bull. trimest. Soc. mycol. Fr. 69: 281,. 1953, nom. inval. (Art. 36).

On the natural substrate forming only light-coloured conidial heads; conidia measuring 3.5-5.0(-6.0) x 1.7-2.7 µm, length/width ratio 1.7-2.2, uni- or biguttulate.
In culture reaching 12-25 mm diam in 14 days at 20-22°C; submerged mycelium mostly hyaline, producing abundant globose, brown chlamydospores, mostly in terminal and lateral positions, rarely intercalary, 4.5-5.5 x 4.0-5.0 µm. Conidiophores variable in density, 70- above 100 µm tall before proliferating, 3-7 times septate, 3.0-3.5 µm wide. Conidia never in cirrhi, 3.0-4.5(-5.5) x 2.0-2.7(-3.0) µm, length/width ratio 1.5-2.5, mostly uniguttulate. Some strains showing a more marked protrusion of the meristematic tip beyond the collarette in the course of conidium formation (Fig. 8) than others.

Material examined

Cultures (partly with original specimens)
CBS 114.41, type strain, ex soil, Baarn, Bakhuizen van den Brink, 1941.
CBS 142.61 = QM 1103, ex decaying wood, via G. Müller, 1961.
CBS 345.67, ex decaying wood, Höllbachg’spreng, Bayrischer Wald, F.R.G., July 1967.
CBS 581.73 (strain 26), comm. J.-E. Nylund, Stockholm.
CBS 916.73, ex volcanic ash soil, Chile, J. Grinbergs (526.73), 1973.
CBS 239.75B, ex forest soil, Sivrite nr. Nancy, France, O. Reisinger.
CBS 248.75, ex Cedrus atlantica, Plateau du Lubéron, France, Oct. 1975.
CBS 260.75A, ex Quercus sp., Baarn, Sept. 1974.
CBS 601.75A, Le Roptai nr. Ave-et-Auffe, CBS 601.75B, nr. Neupont,
CBS 621.75, Grande Tinémont near Han-sur-Lesse, Belg. Ardennes, Sept. 1975.
CBS 423.76, ex Betula verrucosa, Lánská obora Forest nr. Rakovník, Boh., V. Holubová-Jechová, 12 June 1976. [p.22]

Herbarium specimens
Material of Sphaeromycetella macrospora is not preserved (Nicot and Charpentié, 1971).
IMI 85335, ex organic soil, Guelph, Ont., Canada, G. L. Barron, 1960.
IMI 112913, ex deep litter, Boyle Crag Wood, T. F. Hering, 1965.
IMI 144475, ex soil under maize, Ont., Canada, G. C. Bhatt (UW 675), July 1967.
IMI 147291, ex Betala leaf litter, Meathorp Wood, J. C. Frankland, 14 March 1970.
IMI 149014, ex soil, Nottingham, R. H. Thornton, 1970.
DAOM 35345, leg. Morgan, Ohio (slide ex Herb. NY).
DAOM 40749, ex soil under maple-mixed hardwood, N-Wisconsin, M. P. Backus, 1956.
DAOM 60355, contaminant, Ottawa, Ont., Canada, W. B. Kendrick, 28 Nov. 1958.
DAOM 83101, on filter paper in moist chamber, Villa Hidalgo, Mexico, R. F. Cain (UT 36790), 6 March 1961.
DAOM 118919, ex roots of Caragana seedlings, Indian Head, Sask., Canada, M. I. Timonin, 1968.
G.L.H. 1613-F, Populus sp., Manotick, Ont., Canada, 4 Nov. 1960.
G.L.H. 6267, on twig with mosses, La Nfidi, Nsangi, Bas Congo, 26 Sept. 1964.
G.L.H. 6585, Picea abies root, Meerdael Bos nr. Heverlee, Belgium, H. Maraite, 24 Oct. 1964.
G.L.H. 8868, Juncus effusus, Meerdael Bos, 16 May 1966.
PRM 793969 and 793972, Quercus petraea and Fagus sylvatica, Týřovické skály nr. Rakovník, Boh., 28 Sept. 1967.
PRM 793973, Fraxinus excelsior, Úpořský potok valley nr. Skryje, Boh., 26 Apr. 1966.
PRM 793975, Salix sp., Mt. Studentý vrch nr. Stříbrná Skalice, Boh., Z. Pouzar, 29 May 1950.
PRM 793985, Sorbus aucuparia, Krušné hory Mts., Mt. Klínovec, Boh., 24 June 1969.
PRM 793994 and 794059, Quercus petraea, Hoja Forest nr. Znojmo, Mor., 29 Aug. 1971.
PRM 794049, Quercus petraea, Ruský Kazimír nr. Vranov, Slov., 14 Aug. 1972.
PRM 794068, Sorbus aucuparia, Orlické hory Mts., nr. Rokytnice, Boh., 15 Oct. 1975.
PRM 794069, Quercus robur, Mt. Kámen nr. Nepomuk, Boh., 3 Oct. 1975.
PRM 794071, Aesculus hippocastanum, Obora Hvĕzda Forest nr. Blatná, Boh., 30 Sept. 1975.
PRM 794104, Urtica dioica, Mt. Vlčí Vrch nr. Vel. Krtíš, Slov., 11 Aug. 1975.
PRM 796756, Populus nigra, and 796758, Fraxinus excelsior, Úpor Forest nr. Mélník, Boh., 24 June 1976.


Nomenclature and typification: The epithet virescens Pers. was given preference over viride Link by Fries (1832) as being the oldest available name. Herb. L preserves a specimen (910.255-753) labeled ‘Dematium virescens Pers.’ in Herb. Persoon which contains an abundance of the Helicosporium state of Tubeufia cerea (Berk. & Curt.) C. Booth and no Chloridium at all (cf. also Hughes, 1958). Helicosporium vegetum Nees ex Pers. is very different from Chloridium and a confusion by Persoon could have occurred as a result of a macroscopic glance only. Persoon’s generic diagnosis clearly reads sporulis globosis aut ovalibus’; therefore this specimen can not possibly be the type of D. virescens and it must be concluded that Persoon’s original material has probably been lost. Since Persoon (1822) and Link (1824) both regarded each other’s species as identical, we accept Link’s specimen (the holotype of the genus) as neotype of the species, but the name employed must be that sanctioned by Fries (1832), viz. C. virescens. [p. 23]
There are few doubts about the identity of the older synonyms since the bright green conidial cirrhi of var. virescens are clearly recognizable in herbarium specimens even at the low power of the microscope. Mesobotrys flavovirens is also added as a synonym, since the ramification mentioned by von Höhnel (l.c.) (recognizable in the second specimen mentioned) appears to be a consequence of young hyphae growing along an old conidiophore and producing shorter conidiophores; this phenomenon in particularly common in Chloridium species of the section Psilobotrys.
Problems of species delimitation arise if the green colour and cirrhi are absent. The clearest distinction between Bisporomyces chlamydosporis and Cirrhomyces caudigerus was worked out by Mangenot (1952) and is summarized below:


Bisporomyces chlamydosporis

Cirrhomyces caudigerus





4.3-4.7 x 2.7 µm, in heads, biguttulate

3.0-4.0 x 2.0 µm, in cirrhi, uniguttulate


5-6 µm, solitary, terminal

4-7 x 4-5 µm, solitary or catenulate, intercalary or terminal.

Meyer (1959) regarded C. caudigerus, B. chlamydosporis and Sphaeromycetella leucocephala as synonymous. Ellis (1971) used only conidial size and arrangement to distinguish C. viride (2-3 x 2-2.5 µm), C. caudigerum (3-4 x 1.5-2 µm) and C. chlamydosporis (3.7-4.7 x 2-3 µm).
The pigmentation of the vegetative mycelium cannot be accepted as distinctive feature, since it is very variable in culture within one strain; sectors with dark and with hyaline vegetative mycelium arise, can be repeatedly subcultured, and remain distinct, and the amount of sporulation is also variable. In all strains examined some intercalary chlamydospores were found in addition to the predominating terminal ones. The extent of chlamydospore production is very variable, although generally more abundant and more commonly globose in var. chlamydosporum than in the other varieties, but this character proved too variable to be used as a distinctive criterion for all strains.
On the natural substrate separation of the three varieties can be achieved more easily than in culture. Isolates from representative var. virescens specimens yield more elongate conidia than are found on the natural substrate; these are mostly produced in heads and measure 3.2-4.0 x 2.0-2.5 µm. Therefore most strains studied only in culture were identified automatically by various authors as C. chlamydosporis. Strains received as or isolated recently as representatives of C. virescens, C. caudigerum and C. chlamydosporis, respectively, were also grown on sterile Acer twigs for six weeks, but this did not permit a clearcut separation between the three taxa.
It may be argued that on the natural substrate the formation of cirrhi [p. 24] is mainly age-dependent and always passes through a stage of conidial heads. But there remains the unusual fact, that typical green cirrhi are extremely rare in Great Britain and the Netherlands and very common in Czechoslovakia and Belgium. Therefore some generic differences or reactions on climatic differences are implicated.
The final argument for the treatment of these taxa within one species is that perfect states of all are indistinguishable.
For convenience the differences between the three varieties are tabulated here again:


var. virescens

var. caudigerum

var. chlamydosporum

On natural substrate


Conidial arrangement



Conidial shape


ellipsoidal cylindrical

length/width ratio



Conidial size

2.5-3.8 x 2.0-2.5

3.5-4.5 x 1.7-2.5

3.5-5.0 x 1.7-2.7

Conidial colour

bright green



In culture


Vegetative hyphae

predominantly pigmented

mostly hyaline


scanty, globose or often pyriform

abundant, mostly globose

Conidial arrangement

cirrhi in old cultures

only heads

Conidial shape



length/width ratio




Conidial size

3.2-4.0 x 2.0-2.5

3.0-5.0 x1.6-2.7

3.0-4.5(-5.5) x 2.0-2.7(-3.0)


Chloridium section Gongromeriza (Preuss)
W. Gams & Hol.-Jech. comb. nov.

Gongromeriza Preuss - Linnaea 24: 106. 1851.

Colonies very slow growing. Conidia produced from a single conidiogenous locus within a ± pronounced collarette. Percurrent proliferation commonly occurring. Chlamydospores absent.
Type species Chloridium clavaeforme (Preuss) W. Gams & Hol.-Jech.

2a. Chaetosphaeria lentomita W. Gams & Hol.-Jech. sp. nov. - Fig. 9.

Chaetosphaeria innumera Tul. sensu C. Booth - Mycol. Pap. 68: 2. 1957 [non Berk. & Br. ex Tul. (1863)]. [p. 25]

Perithecia subglobosa, nigra, levia, strato externo hyalino pruinosa, 150-165 µm diam, paries 15-22(-30) µm crassus. Asci tenuitunicati, cylindrici, 65-75 x 6.0-7.5 µm. Ascosporae oblique uniseriatae, fusiformes, utrinque rotundatae, hyalinae, asperulatae, bicellulares neque constrictae, 8-14 x 3.5-4.0 µm. Status conidialis Chloridium pachytrachelum W. Gams & Hol.-Jech.
Holotypus CBS 645.75 in ligno Fagi sylvaticae, prope Neupont in Belgio, Sept. 1975.

Perithecia broadly attached to the substrate, without subiculum, subglobose, ostiolar region slightly conical, black, smooth or slightly papillate but appearing slightly pruinose, 150-265 µm diam; wall 15-22(--30) µm thick, composed of rather wide (3-5 µm diam), very thick-walled, pigmented cells in textura epidermoidea or angularis, the outermost layer or at least some superficial cells hyaline. Asci thin-walled, cylindrical, with a thin apical plate, 65-75 x 6.0-7.5 µm. Ascopores obliquely uniseriate, fusiform with rounded ends, hyaline, finely roughened, at maturity bicellular, not constricted at the septum, each cell 1- of 2-guttulate, 8-14 x 3.5-4.0 µm. Paraphyses 1.2-1.5 µm wide, at the tip slightly swollen to 2.2 µm. Conidial state Chloridium pachytrachelum.

Material examined

CBS 645.75, ex holotype on Fagus sylvatica, nr. Neupont, Belg. Ardennes, Sept. 1975 (culture from ascospores).
DAOM 51738(b), Picea abies, Kundratice nr. Reichenberg (= Liberec), Boh., leg. S. J. Hughes, 11 May 1955.
Numerous specimens in the Chaetophaeria innumera folder at IMI on the following host plants in Great Britain: Acer pseudoplatanus: 19290; Alnus sp.: 69416; Betula sp.: 13405 (b); Fagus sylvatica: 19263, 19277, 32234, 178042; Fraxinus excelsior: 56772; Juglans regia: 54,15; Quercus spp.: 1684, 6925, 8122, 14474, 19247(k), 61210, 71013(a), 73450; Sambucus nigra: 14475; Ulmus sp.: 40660, 54114, 56771, 61205; indet. wood: 8118, 13221(b), 19287, 32235, 58458, 60254, 62966(a), 76684, 76686, 179572, 179573.
PRM 794016 and 794017, Fagus sylvatica, HrubýJeseník Mts., Bučina Forest, nr. Kouty n. Desnou, Mor., 4 Aug. 1971.
PRM 794023, Picea abies, Rychlebské hory Mts., valley of Bučínský potok nr. Javorník, Mor., 10 Aug. 1971.
PRM 796740, Fagus sylvatica, Malá Fatra Mts., in the valley of Sútovka, Slov., 15 July 1976.


Contrary to Chaetosphaeria innumera Berk. & Br. ex Tul., the ascospores are always two-celled from an early stage and remain so, although the spore size and shape of the two species agree completely. This is the reason for the confusion between the two species (cf. collections in IMI and Booth, 1957). Chaet. lentomita is also similar to Chaetosphaeria brevicollis (Niessl) E. Müll. (syn. Lentomita brevicollis Niessl), of which the ascospores are given as 8-13 x 5-8 µm (Müller and von Arx, 1962), but no material of this species could be traced at Herb. M. The generic name of this fungus is adopted for the epithet of the new species.
Two specimens (DAOM 51738(b) and PRM 794023) on Picea abies [p. 26] showed more coarsely warted ascospores and conidiophores with a shorter and more narrow collarette (1.0-1.5 µm diam) than is usual in this species.

  fig. 1

Fig. 9
Chaetosphaeria lentomita, PRM 794016, asci, a paraphysis and ascospores and conidiophores of Chloridium pachytrachelum, b. CBS 645.75, conidiophores from pure culture. x 1000.


2b. Chloridium pachytrachelum W. Gams & Hol.-Jech. st. nov. - Fig. 9.

Coloniae lentissime crescunt, dilute griseo-brunneae. Hyphae vegetativae hyalinae, 1.0-2.5 (-4.0) µm latae. Conidiophora erecta, 60-> 200 µm longa, 3.5-6.0 µm lata, in parte inferiore saepe cellulas breves, ± inflatas formantia, brunnea, crassitunicata, sub collari modice constricta et conspicue inspissata; proliferatio vulgo percurrens; collare caliculare, hyalinum, 3-5 µm altum. Conidia in capitulis mucidis aggregata, ellipsoidea, cylindrica vel fere reniformia, hyalina, levia, 3.5-7.5 x 2.0-3.5 µm. Chlamydosporae absunt.
Holotypus CBS 645.75, cultus ex ascosporis Chaetosphaeriae lentomitae e Fago sylvatica, prope Neupont in Belgio, Sept. 1975.

Colonies very slow growing, reaching 2-4 mm diam in 14 days at 20-22°C, light greyish-brown. Vegetative hyphae hyaline, 1.0-2.5(-4.0) µm wide or slightly monilioid and to 4.5 µm diam. Conidiophores erect, sometimes irregularly undulate, 60- over 200 µm tall, 3.5-6.0 µm wide, brown, wall 0.7-1.0 µm thick, several times septate, the lower part often [p. 27] with short and ± inflated cells, lighter towards the tip, slightly constricted below the collarette and with a pronounced wall-thickening after successive conidium formation; proliferation mostly percurrent, very rarely also sympodial; collarette beaker-shaped, almost hyaline, (3-)5-8 µm deep, 4-8 µm wide. Conidia arising singly and forming slimy heads, ellipsoidal, cylindrical or slightly reniform, hyaline, when dry brownish in mass, smooth-walled, (3.5-)5.0-7.5(-9.0) x (2.0-)2.5-3.5 µm, length/width ratio 1.5-2.5; in culture usually somewhat longer than on the natural substrate; uniguttulate. Chlamydospores absent.

Additional material examined

Living cultures
CBS 260.75B and 260.75C, and some further collections, ex Quercus petraea, Groeneveld nr. Baarn, Netherlands, Sept. 1974.
CBS 247.76, ex Fagus sylvatica, nr. Neuenheerse, Teutoburger Wald, F.R.G., Sept. 1972.
CBS 265.76, Pijnenburg forest near Baarn, Aug. 1972.

Herbarium specimens
Sub Chaetosphaeria innumera in Rabenh. Fungi cur. no. 925, nr. Lucknam, Pinus spec., C. E. Broome, Apr. 1865, and other coll. by Broome, 12 Apr. 1865 (K).
PRM 794021, Fagus sylvatica, Hrubý Jeseník Mts., valley of Hučivá Desná, Mor.. 6 Aug. 1971.
PRM 796748, Fagus sylvatica, between Mt. Klak and Mt. Revan nr. Nitran. Pravno, Slov., 14 July 1976.


3. Chloridium phaeosporum W. Gams & Hol.-Jech. sp. nov. - Fig. 10.

Conidiophora in strato velutino, fere brevia, 25-35 µm alta, 2.0-2.7 µm diametro, semel vel paucies septata, brunnea, sursum pallidiora et modice attenuata; collare 1.0-1.5 µm diametro. Conidia in capitulis aggregata ellipsoidea, basi truncata et obscuriora, pigmentata, levia, 4.0-5.0 x 1.7-2.2 µm.
Holotypus DAOM 40413, velut ‘Haplochalara ?’, in ligno putrido, Morgantown, in Virginia occidentali, U.S.A., H. L. Barnett, Majo 1953.

Conidiophores forming a velvety layer, relatively short, commonly 25-35 µm, with one or a few septa, 2.0-2.7 µm wide, brown, lighter towards the apex and slightly attenuate towards the very short, 1.0-1.5 µm wide collarette. Conidia aggregated in heads, ellipsoidal with a truncate base, distinctly pigmented, smooth-walled, with a darker basal hilum, 4.0-5.0 x 1.7-2.2 µm, length/width ratio 1.7-2.5.
Known only from the type collection.

It is possible that this species belongs to section Psilobotrys, since a few sympodial proliferations were also seen, but the conidiophores are unusually short and little constricted below the phialide, so that they do not fit into that section. [p. 28]

  fig. 10

Fig. 10
Chloridium phaeosporum, from holotype, x 1000.

  fig. 11

Fig. 11
Chloridium reniforme, habit sketch, conidiophores x 850, conidia x 1000 (with permission from Matsushima, 1975).

  fig. 12

Fig. 12
Chloridium matsushimae, habit sketch, conidiophores and conidia x 850 (with permission from Matsushima, 1975).
[p. 29]


4. Chloridium reniforme Matsushima - Fig. 11

Chloridium reniforme Matsushima - Icon. Microfung. a Mats. lect., Kobe, p. 28. 1975.

Description (after Matsushima): Conidiophores erect, straight, septate, thick-walled, dark brown below, more or less setiform, 100-180 µm tall, 4-5 µm wide at the base, 2.6-3.2 µm in the upper part and inflated to 3.6-4.4 µm near the tip, but strongly constricted at the hardly visible collarette. Conidia arising singly, more or less reniform, 3.5-5.5 x 2.0-3.0 µm, smooth-walled, hyaline, aggregated in a slimy head.
Holotypus (not seen): MFC-3183, on decaying branchlet of broadleaf tree, Kiyozumi Exp. Forest, Univ. of Tokyo, Chiba, Japan, Oct. 1967.

This species may also belong to Sect. Psilobotrys if sympodial proliferation occurs.


5. Chloridium matsushimae W. Gams & Hol.-Jech. sp. nov. - Fig. 12.

Chloridium sp. MFG-1640 Matsushima - Icon. Microfung. a Mats. lect., Kobe, p. 29. 1975.

In caule putrido: Coloniae effusae, sparsae, inconspicuae. Mycelium immersum. Conidiophora solitaria vel pauci-fasciculata, recta, erecta, simplicia, septata, 130-240 µm longa, prope basin 5-6 µm crassa, superne 4-5 µm crassa, sub apice angustata, apice infundibuliformiter collariata, frequenter per colla semel ad paucies elongascentia, basi atrofusca, apicem versus pallide fusca. Conidia singulatim successive in collo formata, cylindro-ellipsoidea, 15-19 x 3.6-5.3 µm, continua, levia, apice uniappendiculata, hyalina, conglutinata, in sicco pallide brunnea; appendices simplices gelatinosi.
Cultum in agaro potato-sucrose dicto: Coloniae tarde crescentes, coactae, pallide griseae, reverso fusco, steriles.
Holotypus: MFC-1640, in caule putrido Phyllostachydis edalis, Kyoto City, Kyoto, in Japonia, Feb. 1966.

Description from Matsushima: On the natural substrate conidiophores solitary or slightly fasciculate, erect, straight, septate, dark brown near the base, lighter in the upper part, 130-240 µm tall, near the base 5-6 µm wide, in the upper part 4-5 µm, tapering towards the tip, with a funnel-shaped collarette, frequently once or twice percurrently proliferating. Conidia aggregated in slimy droplets, arising singly in the collarette, cylindrical-ellipsoidal, 15-19 x 3.6-5.3 µm, length/width ratio 4.0-5.0, smooth-walled, hyaline, brownish when dry, with a slimy apical appendage. In culture on PSA slow-growing, felty, pale grey with dark reverse, remaining sterile.

C. matsushimae could not be examined for the possible occurrence of sympodial proliferation. It is unique in Chloridium in the size and the apical appendages of the conidia and may even deserve a generic [p. 30] distinction. The comparable species with cylindrical conidia (also of Cylindrotrichum) did not show any comparable structure even under phase contrast. Nevertheless C. matsushimae appears to be closer to Chloridium than to Codinaea, where both setulose and non-setulose species are known to occur.

6a. Chaetosphaeria myriocarpa (Fr.) C. Booth - Fig. 13a-b

Sphaeria myriocarpa Fr. - Syst. mycol. 2(2): 459. 1823 = Coniothyrium myriocarpum (Fr.) Sacc. - Syll. Fung. 3: 315. 1884 = Trichosphaeria myriocarpa (Fr.) Petr. & Syd. - Annls mycol. 22: 330. 1924 = Chaetosphaeria myriocarpa (Fr.) C. Booth - Mycol. Pap. 68: 5. 1957.
Rosellinia minima Fuckel & Nits., Symb. mycol. p. 149. 1870 = Wallrothiella minima (Fuckel & Nits.) Sacc. - Syll. Fung. 1: 455. 188z = Trichosphaeria minima (Fuckel & Nits.) Wint. - Rabenh. Krypt.-Fl. 1(2): 204. 1887 = Psilosphaeria minima (Fuckel & Nits.) Cooke - Grevillea 16: 50. 1887.
Sphaeria ostioloidea Cooke - Grevillea 4: 113. 1876 = Psilosphaeria ostioloidea (Cooke) Cooke - Grevillea 7: 84. 1879 = Zignoëlla ostioloidea (Cooke) Sacc. - Syll. Fung. 2: 104. 1883.
Synonyms fide Booth (1957).

  fig. 13

Fig. 13
Chaetosphaeria myriocarpa, a. ascus, spores and conidiophores of Chloridium clavaeforme, CBS 264.76, b. evanescent asci with spores of unequal maturity and septation, specimen Harkness in K; c. Chloridium clavaeforme in culture, CBS 619.75A, d. with catenulate conidia on the natural substrate, PRM 794045, and e. geniculate proliferations, PRM 794034. x 1000. [p. 31]

Detailed descriptions in Booth (1957, 1958). The connection with a phialidic conidial state was first proved in culture by Brefeld (1891).
Perithecia often densely aggregated, with or without a thin, black, basal pseudostroma. Perithecia black, brittle, subglobose, ostiole conical, smooth-walled, glossy, 110-155 µm diam. Wall 8-15 µm thick, consisting of several layers of strongly compressed, pigmented cells (textura epidermoidea). Asci thin-walled, narrow cylindrical, 45-55 x 4-6 µm. Ascospores mostly somewhat obliquely monostichous, chromophilic, cylindrical with rounded ends, one-septate at maturity, hyaline, smooth-walled, 5-7 x 1.5-2.0 µm (Booth: 5-7 x 2.5-3 pm). Paraphyses emerging between the asci, 1.5 µm wide, with a slightly curved pointed tip. Septation of the spores is sometimes delayed in several asci or in some regular groups of spores within an ascus (Fig. 13b), suggesting a genetic determination. Conidial state Chloridium clavaeforme.

Material examined

Selected herbarium specimens (conidial state generally also present)
Chaetosphaeria myriocarpa folder in K: Quercus, Brockley, 1845; Quercus, Langridge, Somers., 7 Apr. 1859; Quercus, Langley, Somers., Apr. 1868; Rubus, St. Catherines, Somers., May 1869, all coll. by C. E. Broome.
Chaetosphaeria innumera folder in K: Epping Forest, C. E. Broome, Feb. 1871; on Eutypa sp., Arboretum Kew, M. C. Cooke, Aug. 1880; on Acacia and Quercus, California, H. W. Harkness, ex Herb. M. C. Cooke; Carlisle, Cumberl. (Cumbria), Carlyle, Jan. 1884.
G.L.H. 1209, Foret de Lorgnes, Brabant, Belgium, 16 Aug. 1960.
G.L.H. 3247-A, Acer sp., Wieringermeer-Polder, Netherlands, 6 Apr. 1963.
G.L.H. 3265, coniferous wood, nr. Zeist, Netherlands, 7 Apr. 1963.
L 961.204-102, Quercus sp., Huis te Warmond, Netherlands, R. A. Maas Geesteranus, 19 Nov. 1961.
PRM 794050, Fagus sylvatica, nr. Rusky Kazimír, Slov., 14 Aug. 1972.
PRM 794099, Quercus Sp., Zudrok Hill nr. Nitra, Slov., 5 Aug. 1975.
Herb. CBS, decaying twigs, Le Roptai nr. Ave-et-Auffe, Belgium, Sept. 1975 (with isolate from ascospores).


6b. Chloridium clavaeforme (Preuss) W. Gams & Hol.-Jech. comb. nov. - Fig. 13a, c-e

Gongromeriza clavaeformis Preuss - Linnaea 24: 106. 1851 = Chloridium state of Chaetosphaeria myriocarpa, Hughes - Friesia 9: 62. 1969.
Catenularia heimii Mangenot - Revue gén. Bot. 59: 445. 1952 (Rech. méth. Champ. Bois Décomp. p. 27).

We abstain from correcting the epithet to claviforme as recommended in Art. 73G of the Code since clavus (nail) and clava (club) have a different meaning.

Colonies very slow-growing, reaching 4-10 mm diam in 14 days at 20-22°C, olivaceous black from the beginning, often glossy and iridescent at the margin; vegetative hyphae ± dark olivaceous, with strongly encrusted walls, 2-3 µm wide. Sporulating abundantly with simple conidiophores [p. 32] arising from the substrate. Conidiophores dark brown, 25-30 µm tall if consisting of simple phialides, but commonly 35-70(-100) µm tall and several times septate, 2.5-4.0 µm wide; subsequently many times percurrently proliferating; only slightly less pigmented in the upper part and often also darker at the collarette; tapering gradually or abruptly to 1.5-2.0 µm diam at the collerette; collarette flaring or vase- or funnel-shaped, 2.5-4.0 µm deep; by a renewed widening after proliferation assuming a characteristic geniculate appearance (Fig. 13e). Conidia typically short-cuneate or dacryoid with a truncate base and rounded tip, smooth-walled and slightly pigmented, mostly forming slimy heads, sometimes also regular chains, 2.3-3.0 x 1.6-2.5 µm; smaller and hyaline conidia sometimes occurring by submerged sporulation. Chlamydospores absent.

General distribution

The perfect and particularly the conidial state are extremely common on very many kinds of wood in all areas studied by the authors.

Selected material examined

Living cultures
CBS 141.53, type strain of Catenalaria heimii Mangenot, ex Fagus sylvatica, 1953.
CBS 116.57, comm. A. Rambelli, 1957 (= IFO 7527).
CBS 698.70 = PET 381, ex A1 horizon under Populus tremuloides, Ont., Canada, G. C. Bhatt.
CBS 878.70, ex old stroma of Diatrypaceae on Fagus sylvatica, Baarn, Netherlands, H. A. van der Aa, Sept. 1970.
CBS 241.75A, ex Cedrus atlantica, Plateau du Lubéron; CBS 241.75B, ex deciduous wood, Massif de Ste Baume nr. Marseille, France, Oct. 1974.
CBS 619.75A, nr. Neupont, 619.75E and 651.75, Grande Tinémont nr. Han-sur-Lesse, Belgium, Sept. 1975.
CBS 264.76, Pijnenburg Forest nr. Baarn, Aug. 1972.

Herbarium specimens
Gongromeriza clavaeformis, holotype, on Alnus wood, Herb. Preuss No. 220 (B).
DAOM 44851, on dead wave’, Fallacofield, Ont., D. E. Wells, 5 Oct. 1954.
DAOM 44912, nr. Old Chelsea, P.Q., 21 Oct. 1954, S. J. Hughes.
G.L.H. , 597-B = DAOM 71552(B), G.L.H. 1677-C = DAOM 74611(B), Fagus grandifolia Gatineau Park, Québec, Canada, 31 Oct. 1960.
G.L.H. 2796-D, Lorith March, Ont., Canada, 14 Oct. 1961.
G.L.H.2944 and 3003, Fagus grandifolia, Stoddard Brook Road, Alleghany State Park, N.Y., 11 June 1961.
G.L.H. 3247-B and 3249-C, Acer Sp., Wieringermeer-Polder, Netherlands, 6 Apr. 1963. G.L.H. 326,-A, Fagus sylvatica, G.L.H. 3262-B and 3263, Pinus sylvestris, nr. Zeist, Netherlands, Apr. 1963.
G.L.H. 4357, on a grass, Richmond Park, Surrey, G.B., 16 June 1963.
PRM 793990, Fagus sylvatica, Jizerské hory Mts., nr. Liberec, Boh., 7 Aug. 1965.
PRM 794020, Sorbus aucuparia, Mt. Keprník, Hrubý Jeseník Mts., Mor., 11 Aug. 1971.
PRM 794027, Fagus sylvatica, Mt. Velká Javořina, White Karpathian Mts., Mor., 27 July 1970. [p. 33]
PRM 794034, Fagus sylvatica, Harmanecká dolina valley nr. Malý Stúrec, nr. Banská Bystrica, Slov., 10 Aug. 1969.
PRM 794036, Fagus sylvatica, Mt. Kalužná, Martinské hole Mts., nr. Martin, Slov., 17 Sept. 1971.
PRM 794041, Quercus robur, Mt. Jedlina nr. Zborov, Slov., 9 Aug. 1972.
PRM 794080, Carpinus betalus, nr. Žuhráčka nr. Levice, Slov., 6 Aug. 1975.
PRM 794089, Quercus robur, Dúbrava Forest nr. Záhorská Ves, Slov., 19 Sept. 1975.
PRM 794096, Fagus sylvatica, Kačín nr. Lamac, Slov., 16 Sept. 1975.


7. Chaetosphaeria bramleyi C. Booth - Fig. 14a

Chaetosphaeria bramleyi C. Booth - Naturalist, Hull 1958: 85. 1958.

With Chloridium state.

Booth (1958) described Chaet. bramleyi from a decorticated branch of Rosa sp. as differing from Chaet. myriocarpa by perithecia 140-220 µm diam, ascospores slightly longer, 7-10 x 2.0-2.5 µm, conidiophores 25-135 x 4 µm at the base and phialides 20-30 x 2-3 µm with a 7-12 µm deep collarette and catenulate, cuneate, hyaline conidia measuring 1.5-2.5 x 1.0-1.5 µm. The connection with the conidial state was established by cultures from ascospores.

  fig. 14

Fig. 14
a. Chaetosphaeria bramleyi, IMI 67848, asci, ascospores and conidiophores;
b. specimen C. E. Broome from Langridge (K), ascospores and Chalara-like conidiophores. x 1000. [p. 34]

Material examined

IMI 67848, holotypus, on Rosa sp., Whitecliffe Wood, Richmond, Yorksh., leg. W. G. Bramley, 1 Oct. 1956.
IMI 27368, Sambucus nigra, Penglais Woods, Aberystwyth, S. J. Hughes, 26 March 1948. Sub Sphaeria innumera Berk. & Br., Langridge, Somerset, C. E. Broome, March 1871 (K) (Fig. 14b).


Since no living culture was available we abstain from naming the conidial state separately.
In the type collection the collarette usually includes two or three conidia at a time; the conidia are completely hyaline, but otherwise identical with those of C. clavaeforme. The collection in K contains perithecia with ascospores measuring 6.5-8.5 x 2.0-2.2 µm and Chalara-like conidiophores with a c. 10 µm long collarette including five or more conidia at a time (Fig. 14b, cf. also Chalara brevispora Nag Raj & Kendrick). Chaet. bramleyi is undoubtedly most closely related to Chaet. myriocarpa.

8a. Chaetosphaeria preussii W. Gams & Hol.-Jech. sp. nov. - Fig. 15a

? Chaetosphaeriae myriocarpae simillima; perithecia globosa, atra, levia, splendentia, 140-220 µm diametro, paries 7-15(-18) µm crassus, textura epidermoidea. Asci cylindrici, 45-55 x 3.5-4.0 µm. Ascosporae oblique monostichae, cylindricae ad ellipsoideae, aequaliter bicellulares, hyalinae, fere leves, 5.0-7.0 x 1.7-2.0 µm, maturae diffractae. Status conidialis Chloridium preussii W. Gams & Hol.-Jech.
Holotypus PRM 796750, in ligno Alni glutinosae, prope Kolno ad meridiem urbis Augustów in Polonia, V.H.- J., 11 Sept. 1974.

Ascigerous state similar to Chaetosphaeria myriocarpa (Fr.) C. Booth. Perithecia arising from a dense mat of dark hyphae, globose, with a minute apical papilla, black, glossy, 140-220 diam, wall 7-15(-18) µm thick, composed of 6-8 layers of opaque, very thick-walled cells (textura epidermoidea). Asci cylindrical, 45-55 x 3.5-4.0 µm. Ascospores obliquely uniseriate, cylindrical to ellipsoidal, two-celled, with a thin median septum, hyaline, almost smooth-walled, 5.0-7.0 x 1.7-2.0 µm, breaking apart at maturity, sometimes already within the ascus. Paraphyses indistinct, evanescent, emerging beyond the asci, thread-like, 1.0-1.5 µm wide. Conidial state Chloridium preussii.

Additional material examined

PRM 794010, Fagus sylvatica, Hrubý Jeseník Mts., nr. Louná, Mor., 3 Aug. 1971.
IMI 57856, Fagus sylvatica, Denny Area, New Forest, G.B., A. H. S. Brown, 12 Sept. 1954. [p. 35]

  fig. 15

Fig. 15
Chaetosphaeria preusii, a. asci and ascospores from PRM 796750, b. conidiophores and conidia on the natural substrate, CBS 230.75, c. conidiophores and conidia from Acrostalagmus geniculatus, Herb. Preuss (B). x 1000.


The specimens examined contain numerous ascomata and conidiophores of C. preussii, but not a single C. clavaeforme. This is taken as circumstantial evidence for the connection between the perfect and imperfect states. In the mature material of the PRM specimens ascospores are often found breaking apart, sometimes while still in the ascus, whilst in the less mature IMI 57856 this was never observed. The separation occurs somewhat later and the septum in the young spores is less differentiated than in the other species of sect. Melanopsammella of Chaetosphaeria. The separating ascospores as well as the conidial state are the main distinctive features to separate this species from Chaet. myriocarpa.


8b. Chloridium preussii W. Gams & Hol.-Jech. st. nov. - Fig. 15b-c

? Acrostalagmus geniculatus Preuss - Linnaea 24: 125. 1851 = Acrostalagmus (?Stachylidium) geniculatus (Preuss) Sacc. - Syll. Fung. 4: 164. 1886 = Stachylidium geniculatum (Preuss) Lindau - Rabenh. Krypt.-Fl. 1(8): 740. 1906.

Coloniae lentissime crescunt, 14 diebus 8-10 mm diametro, olivaceo-griseae, reverso fusciore. Hyphae vegetativae hyalinae, 1.5-2.0 µm latae, vel pigmentatae et incrustatae, [p. 36] 2.5-4.0 µm latae. Conidiophora erecta, crassitunicata, fusca, sursum nonnumquam pallidiora, levia vel asperulata, 40-> 80 alta, 2.5-4.0 µm lata; phialides paulatim angustatae ad 1.0-1.5 µm, collari dilatato obscuriore terminatae; conidiophora saepe percurrenter, diametro latiore proliferant. Conidia in capitulis mucidis aggregata, hyalina, ellipsoidea, nonnumquam ad basim oblique apiculata, 2.5-3.5 x 1.5-2.0 µm.
Holotypus CBS 230.75, isolatus e ligno Taxi baccatae, Groeneveld prope Baarn in Neerlandia, H. A. van der Aa, 17 Dec. 1972.

Colonies very slow-growing, reaching 8-10 mm diam in 14 days at 20-22°C, olivaceous grey from the beginning, with a darker reverse. Vegetative hyphae hyaline, 1.5-2.0 µm wide or, more commonly, pigmented and strongly encrusted, 2.5-4.0 µm wide. Conidiophores erect, thick-walled (to 0.7 µm), dark brown, in the upper part sometimes lighter, smooth-walled or slightly roughened, 40- over 80 µm tall before proliferation, 2.5-4.0 µm, wide; phialides gradually tapering to 1.0-1.5 µm and expanding in a flaring, sometimes darker collarette; percurrent proliferation continues at a greater diameter, giving a characteristic geniculate appearance. Conidia forming slimy heads, hyaline, ellipsoidal, sometimes obliquely apiculate at the base, 2.5-3.5 x 1.5-2.0 µm, length/width ratio 1.4-2.0. Chlamydospores absent.

Additional material examined

Living cultures
CBS 257.75, ex Fraxinus excelsior, Elm nr. Schöppenstedt, F.R.G., Jan. 1975.
CBS 263.75, ex Quercus sp., Groeneveld, nr. Baarn, Netherlands, Sept. 1974.
CBS 262.76, Donoper Teich nr. Detmold, F.R.G., Sept. 1972.

Herbarium specimens
Acrostalagmus geniculatus Preuss in Herb. Preuss, labeled Fig. 1354, auf Erlenholz’. Another specimen in Herb. Preuss and Rabenhorst-Klotzsch’s Herb. 1278 (all on Alnus wood, Pinka nr. Hoyerswerda, 1849) did not allow an identification (B).
‘Hyalopus ater Corda (?)’, on decayed wood, Dr. Uppold Lane, Herb. W. B. Grove, 24 Apr. 1884 (K, ex BM).
G.L.H. 2330-B, Acer saccharum, Bell’s Corner, Ont., Canada, 31 July 1961.
G.L.H. 3481, Fraxinus excelsior, Caube Botton Shere, Surrey, 16 June 1963.
G.L.H. 3484-A and 3487, Dogkennel Green, Dorking, Surrey, June 1963.
PRM 793981, Quercus petraea, nr. Chomutov, Boh., 23 June 1969.
PRM 794004, Lanžhotský prales Forest nr. Břeclav, Mor., 28 July 1970.
PRM 794057, Quercus petraea, Bĕšický chochol Hill nr. Kadan, Boh., 5 July 1972.


Specimens of Acrostalagmus geniculatus contain several fungi which respond to the meagre original diagnosis as noted by Hughes (1958: 825) and the epithet was therefore considered a nomen confusum by him. An original specimen from Preuss’s herbarium, then not available to Hughes, clearly shows the above fungus, while Preuss’ specimens distributed in Rabenhorst-Klotzschii Herbarium No. 1278 which are equally eligible as type remain a source of confusion. [p. 37]
The gradually tapering, dark brown conidiophores distinguish this species from C. lignicola, while the completely hyaline, ellipsoidal conidia separate it from C. clavaeforme.


9. Chloridium lignicola (Mangenot) W. Gams & Hol.-Jech. comb. nov. - Fig. 16.

Bisporomyces lignicola Mangenot - Revue Mycol. 18: 136. 1953.

Colonies very slow-growing, reaching about 10 mm diam in 14 or more days at 20-22°C, olivaceous-grey in the centre with a broad lighter or hyaline margin. Vegetative hyphae hyaline, 0.8-2.0 µm wide. Conidiophores pigmented, 50- over 100 µm tall before proliferation; three or more septate, later becoming several hundred µm tall, 3-4 µm wide, olivaceous-brown, lighter towards the tip and strongly constricted below the collarette; collarette often darker than the phialide tip, cup-shaped, 2.0-3.0 µm deep. Conidia abstricted singly at the point of constriction at the base of the collarette, forming a slimy head; often with more than one conidium remaining within the collarette. Conidia oblong-ellipsoidal, slightly truncate at the base, sometimes slightly allantoid, hyaline, smooth-walled, 3.5-5.0(-7.0) x 1.5-2.0 µm length/width ratio 2.2-3.0, usually biguttulate. Chlamydospores absent.

Material examined

Living cultures
CBS 143.54, type strain, ex Fagus sylvatica, Vallée Grand Rû, Argonne, France, F. Mangenot, 1952.
CBS 171.76, nr. Neupont, Belg. Ardennes, Sept. 1975.

Herbarium specimens
G.L.H. 1528-B, Quercus sp., between Hull and Aylmer, Quebec, Canada, 15 Oct. 1960.
G.L.H. 2983-C, Fagus grandifolia, Stoddard Brook Rd., Alleghany St. Park, nr. Salamanca, N.Y., 11 June 1961.
G.L.H. 4420, Hotchkiss Hollow, Alleghany St. Park, N.Y., 10 June 1961.


Mangenot (1953) described and illustrated two conidia situated adjacent to each other within a single collarette and thus placed this fungus in Bisporomyces. The conidia are, however, abstricted singly at the narrow base of the collarette. This species is thus placed in section Gongromeriza here. The type strain has longer conidia (3.5-5.0 x 1.5-2.0 µm) than others assigned to the same species which measure 3.0-3.5 x 1.5-2.0 µm, while Mangenot reported them as 3.8-8.0 x 1.5-2.7 µm on the natural substrate. [p. 38]

  fig. 16

Fig. 16
Chloridium lignicola, a. type strain CBS 143.54, b. G.L.H. 4420 from the natural substrate, c. CBS 171.76 in culture. x 1000.

  fig. 17

Fig. 17
Chloridium codinaeoides, IMI 106158(e), x 1000. [p. 39]


10. Chloridium codinaeoides Pirozynski - Fig. 17

Chloridium codinaeoides Pirozynski - Mycol. Pap. 129: 8. 1972.

Full description in Pirozynski (1972). Forming isabelline, effused, velvety colonies on the natural substrate. Conidiophores formed singly or often gregariously, straight, bent or geniculate, monophialidic but with percurrent elongations, pale brown, lighter towards the tip, smooth and thick-walled; simple phialides 20-30 µm tall, up to 60 µm when repeatedly septate (before proliferation), 3.0-5.0 µm wide, with a conical basal cell, 9-12 µm diam. Phialides slightly tapering towards 1.5-2.0 µm at the collarette; collarette slightly widening or almost cylindrical, 3-5 µm deep. Conidia produced singly, but often several included within the collarette, narrowly fusiform or clavate with the upper end rounded, sometimes slightly allantoid, 5-9 x 1.0-1.5 µm (given as 4-9 x 0.5-1.0 µm by Pirozynski, 1972).

Material examined

IMI 106158(e), holotype and DAOM 133944 (isotype), on dead rachis of Elaeis guineensis, Kakombe, Kigoma, Tanzania, K. A. Pirozynski, 12 Jan. 1964.

C. codinaeoides occupies an intermediate position between Chloridium and Codinaea, but as polyphialides with sympodial proliferation have never been observed in it and the phialide opening is much wider than in Codinaea, it is retained in Chloridium here.


Chloridium section Psilobotrys (Sacc.) W. Gams & Hol.-Jech. comb. nov.

Rhinotrichum Corda sect. Psilobotrys Sacc. - Michelia 1: 87. 1877 = Psilobotrys (Sacc.) Sacc. - Michelia 1: 538. 1879.

Colonies very slow-growing. Conidiophores often produced in two layers, a darker and taller layer with hardly proliferating phialides, and a less pigmented and shorter layer with strongly sympodially proliferating phialides. Conidia produced from a single conidiogenous locus within a shallow collarette which is much narrower than the phialide; sympodial proliferation commonly and percurrent proliferation rarely occurring. Chlamydospores absent.
Holotype species Chloridium botryoideum (Corda) Hughes var. minutum (Sacc.) W. Gams & Hol.-Jech. [p. 40]

  fig. 18

Fig. 18
Chaetosphaeria innumera, a. ascus and ascospores from holotype specimen, b. ascus, ascospores and conidiophore of var. botryoideum, PRM 794005, c. the same PRM 793988. x 1000.

11a. Chaetosphaeria innumera Berk. & Br. ex Tul. - Fig. 18.

Sphaeria innumera Berk. & Br. apud Berk. - Outl. Br. Fungol. p. 395. 1860, nom. inval. (Art. 34) = Chaetosphaeria innumera Berk. & Br. ex Tul. - Sel. Fung. Carpol. 2: 252. 1863 = Lasiosphaeria innumera (Berk. & Br. ex Tul.) Stevenson - Mycol. Scot. p. 391. 1879 = Byssosphaeria innumera (Berk. & Br. ex Tul.) Cooke - Grevillea 15: 123. 1887.
Non Chaetosphaeria innumera sensu C. Booth (1957).

Perithecia mostly loosely scattered amongst conidiophores, black, glossy, brittle, subglobose, ostiolar region conical, 150-200 µm diam; wall 18-25 µm thick, composed of numerous layers of flattened pigmented cells (texture epidermoidea). Asci thin-walled with a thin apical plate, 65-90 x 7-9 µm. Ascospores ± distichous, fusiform with rounded ends, at maturity one- and finally three-septate, hyaline, thin- and smooth-walled, (10-)12-16 x 3.5-4.0(-4.5) µm, each cell guttulate with guttules often visible before septation. Conidial state Chloridium botryoideum.

General distribution

Very common on deciduous wood of various kinds, particularly Fagus sylvatica. [p. 41]

Material examined

Living cultures (and herbarium specimens)
CBS 600.75B, conidial isolate of var. minutum and 629.75, ascospore isolate, nr. Abbaye de St. Rémy, Belg. Ardennes, Sept. 1975.
CBS 259.76, ascospore isolate (var. botryoideum), Neuenheerse, Teutoburger Wald, F.R.G., Sept. 1972.

Herbarium specimens
Sphaeria innumera Berk. & Br., Somerset, leg. C. E. Broome, Dec. 1858 (PC, holotypus, the only material preserved by Tulasne).
Chaetosphaeria innumera folder in K: Sphaeria innumera Berk. & Br., Batheaston, Somerset, Dec. 1858 (two isotype collections); on rotten stumps, Banner Down, nr. Corsham, Wiltsh., Dec. 1858; on elm, Batheaston, with conidia of var. minutum, 30 March 1865; St. Catherines, with conidia of var. minutum, 30 March 1871, all coll. by C. E. Broome. Arboretum Kew, with conidia of var. minutum, M. C. Cooke, Aug. 1880.
IMI 4975(c), Sorbus aucuparia, Howldale Wood, Yorks., S. J. Hughes, Apr. 1946.
G.L.H. 6936-A, Acer campestre, Grijfersvijver, Egenhoven nr. Heverlee, Belgium, 24 June 1965 (with conidia of var. minutum).
PRM 793988, Fagus sylvatica, nr. Liberec, Boh., 6 Aug. 1965 (with conidia of var. botryoideum).
PRM 794005, 794006, Fagus sylvatica, Mt. Černava, Hostýnské vrchy Mts., Mor., 26 July 1970 (with conidia of var. botryoideum).
PRM 794009, 794109, 794110, Fagus sylvatica, Mt. Mravenečník, Hrubý Jeseník Mts., Mor., 3 Aug. 1971 (with conidia of var. botryoideum and var. minutum).
PRM 794018, Fagus sylvatica, Mt. Cervená hora, Hrubý Jeseník Mts., Mor., 11 Aug. 1971 (with conidia of both varieties).
PRM 794032, Fagus sylvatica, nr. Stratená, Slov., 26 June 1965 (with conidia of var. botryoideum).
PRM 794037, Fagus sylvatica, Martinské hole Mts., nr. Martin, Slov., 17 Sept. 1971 (with conidia of var. minutum).
PRM 794039, Fagus sylvatica, Badínský prales Forest, Kremnické pohorie Mts., Slov., 9 Aug. 1969 (with conidia of var. minutum).
PRM 794054, Fagus sylvatica, Mt. Veža in the Vihorlat Mts., nr. Remetské Hámre, 10 Aug. 1972 (with conidia of var. minutum).
PRM 794079, Acer pseudoplatanus, Litavica valley nr. Cerovo, nr. Vel. Krtíš, Slov., 12 Aug. 1975 (with conidia of var. minutum).
PRM 794102, Quercus petraea, Mt. Korpaš nr. Modrý Kameň, Slov., 13 Aug. 1975 (with conidia of both varieties).
PRM 794105, Cornus sanguinea, Kopáč Insula in river Danube nr. Podunajské Biskupice, Slov., 27 Aug. 1974 (with conidia of var. minutum).
PRM 794114, Fagus sylvatica, Mt. Bukový vrch, Cerchovské pohorie Mts., 13 Aug. 1972 (with conidia of var. botryoideum).
PRM 796742, Robinia pseudacacia, valley of Csurgo brook, west of Vel. Krtíš, Slov., 12 Aug. 1975 (with conidia of var. minutum).
PRM 796741, Fagus sylvatica, Bučina Forest nr. Kouty n. Desnou, Hrubý Jeseník Mts., Mor., 4 Aug. 1971 (with conidia of var. botryoideum).


The epithet innumera is sometimes ascribed to Tulasne (1863) only and at others to Berkeley and Broome; the latter course was adopted by the Tulasnes and appears to be justified, particularly since only British material has been kept as representing the holotype in PC. In the type [p. 42] material there is no conidial state, but clearly three-septate, thin-walled ascospores are recognizable in the specimen kept at Paris as well as the isotypes at Kew.

11b. Chloridium botryoideum (Corda) Hughes var. botryoideum - Fig. 18 b-c; 19

Fusidium botryoideum Corda - Icon. Fung. 1: 3. 1837 = Chloridium botryoideum (Corda) Hughes - Can. J. Bot. 36: 748. 1958.
Chloridium minutisporum Lindau - Rabenh. Krypt.-Fl. 1(8): 724. 1906.

  fig. 19

Fig. 19
Chloridium botryoideum var. botryoideum, a. DAOM 40962 from the holotype specimen, with a conidial cirrhus of C. virescens var. caudigerum; b. in culture, CBS 246.76 x 1000.

Conidiophores on the natural substrate typically forming two layers; (a) very long (to 150 or even 250 µm), 3-4(-5) µm wide, dark brown, thick-walled (to 1 µm), setiform, but generally fertile, rarely sympodially proliferating; and (b) short (20-40(-100) µm), 2-3 µm wide, light brown, repeatedly sympodially proliferating. Both types tapering to very narrow [p. 43] (1.0 µm), almost hyaline phialide tips with a very short collarette. Sympodial proliferations extending over long or short distances, percurrent proliferations occurring infrequently. Conidia forming whitish or yellowish slimy heads or imbricate simple chains, elongate-ellipsoidal to cylindrical, often obliquely apiculate at the base, (2.5-)3.0-4.5(-5.0) x 0.8-1.2 (-1.5) µm, length/width ratio 2.0-4.0.
In culture extremely slow-growing, reaching 4 mm diam in approx. three weeks at 20-22°C; colonies grey velvety in the centre, blackish and smooth at the margin; vegetative hyphae hyaline, 1.0-1.5 µm wide, to dark brown and to 3.5 µm wide. Conidia in dry heads, somewhat bigger than on the natural substrate, to 6.5 x 2.0 µm. Chlamydospores absent.

Material examined

Living cultures (and specimens)
CBS 246.76, ex ?Fagus sylvatica, Neuenheerse, Teutoburger Wald, F.R.G., Sept. 1972.
CBS 429.76, ex Fagus sylvatica, Fyledalen, Skåne, Sweden, 19 June 1976.

Herbarium specimens
Fusidium botryoideum Corda, holotype, in Trichomycetibus prope Reichenberg’ (= Liberec)), Boh., PRM 155500; slide W.G. 2507 contains C. virescens var. caudigerum; slide DAOM 40962 from the same specimen contains the fungus described here which agrees with the protologue, but also C. virescens var. caudigerum.
Chloridium minutisporum Lindau, holotype, Fagus sylvatica, Sachsenwald nr. Hamburg, leg. O. Jaap, 12 Apr. 1904 (B); Chloridium minutum Sacc., an unpublished variety by Bresadola, same locality and date (S).
DAOM 57921, Underhill, Vt., Canada, R. F. Cain, 20 June 1951.
DAOM 88980, Lake Opinicon, Leeds Co., Ontario, Canada, W. B. Kendrick, 1 July 1962. Numerous specimens were examined from Czechoslovakia: PRM 793971, 793983, 793984, 793991, 794002, 794013, 794015, 794019, 794022, 794029, 794042, 794044, 794055, 794056, 794060, 794061, 794076, 794091, 794092, 794097, 794112, 794113, 794115, 796743, 796744.
The species grows commonly on dead wood and bark of Fagus sylvatica mainly in mountain areas and in the late summer (August to September); it was found also on Alnus sp. and Sorbus aucuparia.


11c. Chloridium botryoideum (Corda) Hughes var. minutum (Sacc.) W. Gams & Hol.-Jech. comb. nov. - Fig. 20

Rhinotrichum minutum Sacc. - Michelia 1: 87. 1877; Fungi ital. autogr. del. Tab. 62. 1877 = Psilobotrys minuta (Sacc.) Sacc. - Michelia 1: 538. 1879 = Chloridium minutum (Sacc.) Sacc. - Syll. Fung. 4: 321. 1886.
Monostachys transversalis Arnaud - Bull. trimest. Soc. mycol. Fr. 69: 278. 1953, nom. inval. (Art. 36).

Very similar to var. botryoideum, but differing in the short ellipsoidal to subglobose conidia, mostly obliquely apiculate at the base, (2.0-)2.5-4.0 x 1.5-2.0 µm, and a length/width ratio of 1.4-2.0. Vegetative hyphae in some strains are monilioid and to 5 µm wide. [p. 44]

  fig. 20

Fig. 20
Chloridium botryoideum var. minutum, a. CBS 261.76B, b. CBS 213.74, c. catenulate, non-apiculate conidia of PRM 796745. x 1000.

Material examined

Living cultures (and specimens)
CBS 877.73 and 223.74 and some other isolates, ex Acer pseudoplatanus, Buren, Isle of Ameland, Netherlands, 28 Oct. 1973.
CBS 564.74, Schaelsberg nr. Valkenburg, Z.L., Netherlands, 23 July 1974.
CBS 685.74, ex Fagus sylvatica, Le Chitelet, Vosges, France, Sept. 1974.
CBS 247.75, ex Fraxinus excelsior, Elm nr. Schöppenstedt, F.R.G., Jan. 1975.
CBS 600.75A and 600.75B, 652.75, nr. Neupont, nr. Abbaye de St. Remy and Le Roptai, Belg. Ardennes, Sept. 1975.
CBS 669.75, ex Eucalyptus roots, Canberra, Australia, comm. G. C. Johnson, 1975.
CBS 261.76A, nr. Neuenheerse, CBS 261.76B, Remminghausener Berg, Teutoburger Wald, F.R.G., Sept. 1972.

Herbarium specimens
Rhinotrichum (Psilobotrys) minutum, in Quercu, Selva, Sept. 1874 (holotypus, PAD, also slide DAOM 41259).
Psilobotrys minuta Sacc. (Chaetopsis fusca var.), Populus fastigiata (slide DAOM 41258 ex PAD).
Monostachys transversalis, sur éclat de bois moisissant sur le sol, Parc de Versailles,’ 19 May 1926 (good material); same locality, 12 Jan. 1946, not recognizable (Herb. Arnaud in PC).
Sub Chaetosphaeria ‘immersa’ Tul. (misspelling of innumera), Rabenh. Fungi europ. 925, Lucknam, in Pini segmentis, C. E. Broome, Apr. 1865 (L).
DAOM 29263, Kingsmere, P.Q., D. E. Wells, 6 Oct. 1952.
DAOM 51735, Picea abies, Kundratice nr. Liberec, Boh., S. J. Hughes, 11 May 1955.
DAOM 112935, Kitchener, Ont., Canada, G. Morgan-Jones, 21 Sept. 1965. [p. 45]
G.L.H. 1577, Ridrelle Island, Ottawa River, Ottawa, Canada, 30 Oct. 1960.
G.L.H. 3001-B, Stoddard Brook Road, Alleghany St. Park, nr. Salamanca, N.Y., 11 June 1961.
G.L.H. 3262-A, Pinus sylvestris, nr. Zeist, Netherlands, 4 Apr. 1963.
G.L.H. 6430-A, Egenhoven, nr. Heverlee, Belgium, 15 Oct. 1964.
G.L.H. 6973-F, Heikendorf nr. Kiel, F.R.G., 27 July 1965.
G.L.H. 8629, Populus canadensis, Lovenjoel, Belgium, 26 March 1966.
Numerous specimens were examined from Czechoslovakia: PRM 793967, 793970, 793977, 793982, 793989, 793995, 793997, 794000, 794007, 794012, 794025, 794031, 794035, 794046, 794070, 794072, 794075, 794077, 794085, 795088, 794094, 794098, 794103, 796745, 796746, 796747. C. botryoideum var. minutum is very common in Czechoslovakia, from river-side forests up to mountainous areas (in summer from July to October) on dead wood and bark of Abies alba, Betula verrucosa, Carpinus betulus, Fagus sylvatica, Fraxinus excelsior, Populus alba, P. nigra, Quercus petraea, Q. robur, Robinia pseudacacia, and Sorbus aucuparia.


The two conidial varieties described here were both obtained from ascospore isolates of indistinguishable perfect states. The conidial shape is usually distinguishable both on the natural substrate and in culture, but on natural substrates both types often occur side by side; therefore we prefer to distinguish them only at the varietal level. Var. minutum is variable in different strains; most commonly the conidia are obliquely apiculate and tend to adhere in imbricate chains. In the holotype specimen and in PRM 796745 (Fig. 20c) the conidia are perfectly round and subglobose and adhere in regular chains as illustrated by Arnaud (1953) for Monostachys transversalis; on Arnaud’s specimen, however, both round and apiculate conidia occur.

12a. Chaetosphaeria fusispora W. Gams & Hol.-Jech. sp. nov. - Fig. 21a.

Perithecia atra, levia, splendentia, fere globosa, ostiolo conico praedita, stromate basilari carentia, 140-180 µm diametro, paries 12-18 µm crassus. Asci cylindrici ad fusiformes, tenuitunicati, 75-95 x 6-7 µm. Ascosporae in ternis vel quaternis seriebus dispositae, 30-40 (-52) x 2-3 µm, fusiformes, sursum rotundatae, plerumque triseptatae. Status conidialis Chloridium cylindrosporum W. Gams & Hol.-Jech.
Holotypus PRM 794008, in ramo putrido iacente Abietis albae, in Monte Mravenečník, Hrubý Jeseník Montes in Moravia, 3 Aug. 1971.

Perithecia black, smooth-walled, shining, subglobose, ostiole conical, covered with a few conidiophores, without a basal stroma, 140-180 µm diam; wall 12-18 µm thick, consisting of numerous layers of thick-walled, pigmented cells. Asci cylindrical to somewhat fusiform, thin-walled, with a thin apical plate 75-95 x 6-7 µm, with spores in three to four rows. Ascospores 30-40(-52) x 2-3 µm, fusiform with a somewhat rounded upper end, mostly 3-septate, each cell 1- or 2-guttulate. Paraphyses threadlike, 1.5 µm wide, soon evanescent. Conidial state Chloridium cylindrosporum. [p. 46]

  fig. 21

Fig. 21
Chaetosphaeria fusispora, a. ascus and ascospores and conidiophores of Chloridium cylindrosporum, PRM 794008; b. sympodially proliferating conidiophores of PRM 794111. x 1000.

Material examined

PRM 794008 (holotype), 794011, 794111, all on Abies alba, Mt. Mravenečník, Hrubý Jeseník Mts., Mor., 3 Aug. 1971.


Chaet. fusispora differs from the similar Chaet. abietis (Höhn.) W. Gams & Hol.-Jech. in the longer ascospores and non-septate conidia. Although not yet grown in culture, the regular association between perfect and imperfect states make the connection highly probable.


12b. Chloridium cylindrosporum W. Gams & Hol.-Jech. st. nov. - Fig. 21, 22.

Conidiophora in duobus stratis formata, breviora saepe sympodialiter proliferantia, conidiis albidis vel flavidis pulverulenta, altiora fusciora, crassitunicata, setiformia, ad 160 (-250) µm alta, 5-6 µm lata prope basin, ad octies septata, sursum ad 3.5-4.0 µm angustata, [p. 47] sub collari ad 1.5-2.0 µm constricta. Conidia in capitulis siccis stellaribus aggregata, cylindrica, utrinque attenuata et rotundata vel truncata, in medio nonnumquam modice angustata, hyalina (acervata dilute ochracea), levia, plerumque uniguttulata, 8.5-11.5(13.0) x 2.2-3.0 µm.
Holotypus PRM 794008, ad ramum putridum iacentem Abietis albae, in monte Mravenecník, Hrubý Jeseník Montes in Moravia, 3 Aug. 1971.

Conidiophores forming two layers, the lower with abundant whitish to yellowish sporulation, the upper with darker, thick-walled (to 1.5 µm), setiform conidiophores, to 160(-250) µm tall, 5-6 µm wide at the base, to 8-septate, tapering to 3.5-4.0 µm in the upper part and to 1.5-2.0 µm immediately below the short, almost hyaline collarette; short conidiophores sympodially or sometimes percurrently proliferating. Conidia in dry, star-like heads, cylindrical with tapering and rounded or truncate ends, centrally sometimes slightly constricted, hyaline (ochraceous in mass), smooth-walled, mostly uniguttulate, 8.5-1 1.5(-13)  2.2-3.0 µm, length/width ratio 3.0-5.0.
Known from the same specimens as Chaet. fusispora.

  fig. 22

Fig. 22
Chloridium cylindrosporum, long setiform and short, sympodially proliferating conidiophores, PRM 794111. x 1000. [p. 48]


Insufficiently known or excluded species
(only species with presumed or proved phialidic conidiogenesis)

Chloridium trichostylum (Corda) Sacc. - Syll. Fung. 4: 321. 1886 = Menispora trichostyla Corda - Icon. Fung. 2: 12. 1838.
A doubtful species (cf. Hughes and Kendrick, 1963). Type material in PRM contains very dark conidiophores with incomplete tips, and no connection with any kind of conidia could be established.

Chloridium laeense Matsushima - Microfungi Solomon Isl. Papua-N. Guinea p. 14. 1971.
Conidiophores produced in compact fascicles and all structures much larger than known in Chloridium. The phialides have a pronounced apical wall thickening and this fungus is to be classified in a separate genus by B.C. Sutton.


7. Cylindrotrichum Bonord

Cylindrotrichum Bonord. - Handb. allg. Mykol. p. 88. 1851.

Conidiophores simple, erect, pigmented, repeatedly septate; the apical integrated phialide bearing a succession of cylindrical, mostly two-celled, hyaline conidia, subsequently sympodially or percurrently proliferating. Sterile dark setae sometimes present.
Lectotype species Cylindrotrichum oligospermum (Corda) Bonord.

Of the four species originally placed in Cylindrotrichum by Bonorden, C. inflatum Bonord. (= Menispora cylindrosperma Corda) is a Menispora, two (C. album Bonord. and C. repens Bonord.) are not documented by herbarium material, and the last one, C. oligospermum (Corda) Bonord., was selected as lectotype for the genus by Hughes (1958).
We include here two species without sympodial proliferation and thus broaden the scope of the genus in a way comparable to Chloridium. The delimitation of the two genera then becomes somewhat schematic on the Saccardian criterion of conidial septation. The species included in this genus have in common the fact that they sporulate tardily or not at all in culture.
A fifth species has recently been described as C. triseptatum M. B. Ellis - More demat. Hyphom. p. 470. 1976 - which has percurrently proliferating conidiophores and 3-septate conidia, 14-19 x 3-5 µm.

Key to the species


Sterile setae present; conidiophores with sympodial proliferation (polyphialides; conidia cylindrical, two-celled, 11-16 x 2.0-2.5 µm, length/width ratio 5-7

C. oligospermum


Sterile setae absent; length/width ratio of conidia about 3



[p. 49]



Conidiophores sympodially proliferating; conidia ellipsoidal-cylindrical to somewhat clavate 7-13 x 2.0-3.0 µm, sometimes two- or three-septate

C. hennebertii


Conidiophores not sympodially proliferating






Conidia ellipsoidal-cylindrical, 6-12 x 2.0-3.5 µm

C. zignoëllae


Conidia cylindrical to slightly clavate, 8.5-13 x 3.0-4.0(-5.0) µm

C. clavatum


1. Cylindrotrichum oligospermum (Corda) Bonord. - Fig. 23.

Menispora oligosperma Corda - Icon. Fung. 2: 12. 1838 = Cylindrotrichum oligospermum (Corda) Bonord. - Handb. allg. Mykol. p. 88. 1851.
Acrothecium delicatulum Berk. & Br. - Ann. Mag. nat. Hist., Ser. 3, 15: 402. 1865 (Notices No. 1055) = Cordana delicatula (Berk. & Br.) O. Kuntze - Rev. Gen. Pl. 2: 850. 1891 (fide Hughes, 1958).

  fig. 23

Fig. 23
Cylindrotrichum oligospermum, a. PRM 794107, b. CBS 557.74, three young conidiophores from the natural substrate and one sympodially proliferating conidiophore and conidia from culture. x 1000. [p. 50]

Conidiophores forming a velvety layer, accompanied by dark brown sterile setae, to 450 µm tall and 6 µm wide at the base; conidiophores light brown, usually less than 75 µm tall (sometimes to more than 125 µm and 6-septate), with numerous lateral phialidic openings arising from sympodial elongation (up to 20 collarettes); upper part almost hyaline, collarettes c. 1 µm diam. Conidia cylindrical, with rounded ends, sometimes with a truncate base, mostly two-celled, hyaline, smooth-walled, 11-16 x 2.0-2.5 µm (Barron’s drawing 13.8-18.0 x 2.5-3.0 µm; Ellis: 12-22 x 2.5-3.0 µm), length/width ratio 5-7; each cell usually biguttulate.
In culture very slow-growing, reaching less than 10 mm diam after 14 days at 20-22°C, olivaceous-grey, poorly sporulating with conidia in irregular heads; after one month with typical setae, polyphialides and typical conidia. Chlamydospores absent.

Material examined

Living culture
CBS 557.74, ex Salix purpurea, Průhonice nr. Praha, W. Gams and V. Holubová-Jechová, 21 June 1974.

Herbarium specimens
Menispora oligosperma Corda, holotypus, on Betula alba, Berezina, Bob., PRM 515149.
PRM 794106, Cirsium oleraceum, Bílichovské údolí Valley nr. Slany, Boh., 27 Aug. 1973.
PRM 794107, Robinia pseudacacia, Mt. Baske nr. Omšenie, Slov., 4 June 1970.
PRM 794108, Robinia pseudacacia, nr. Marcelová, Slov., 18 Sept. 1975.
PRM 796766, Sorbus aucuparia, Velká Fatra Mts., Gadierská dolina valley, 13 July 1976.


Barron (1968) described this fungus correctly as having sterile setae and proliferating phialidic conidiophores, whereas Ellis (1971) emphasized an absence of setae. This disagreement may arise from a confusion with what we regard as a distinct species, C. hennebertii. The type specimen (PRM) has conspicuous setae, 200-300 x 3.7-5.5 µm, and conidia 12-15.5 x 2.0-2.5 µm.


2. Cylindrotrichum hennebertii W. Gams & Hol.-Jech. sp. nov. - Fig. 24

Conidiophora erecta, simplicia, 100-150 alta, in parte inferiore fusca, crassitunicata, 4.0-5.5 µm lata, sursum pallidiora, ad 2.5-3.5 µm gradatim angustata, sub collari ad 1.0-1.5 µm constricta, compluries sympodialiter proliferantia. Conidia in capitulis irregularibus aggregata, ellipsoideo-cylindrica vel modice clavata, basi apiculata, hyalina, 7-13 x 2.0-3.0 µm, 0-3-septata, plurumque 1-septata, cellulis uni- ad pluriguttulatis. Setae steriles et chlamydosporae absunt. [p. 51]
Holotypus G.L.H. 7544, in ramo emortuo Symphoricarpi albi, Heikendorf prope Kiel in Germania, G. L. Hennebert, 29 Jul. 1963.

  fig. 1

Fig. 24.
Cylindrotrichum hennebertii, a. G.L.H. 7544, in the centre two conidiophores from the natural substrate, left and right conidiophores and conidia from died agar cultures (36 days old); b. CBS 570.76, in culture (40 days old). x 1000.

On the natural substrate forming velvety colonies of erect conidiophores without sterile setae; conidiophores 100-150 µm tall, dark brown, thick-walled and 4.0-5.5 µm wide in the lower part, paler above and gradually tapering to 2.5-3.5 µm, and constricted at the shallow collarette to 1.0-1.5 µm, a few times sympodially proliferating. Conidia aggregated in irregular heads, ellipsoidal-cylindrical to slightly clavate, with an apiculate base, hyaline, of variable size, 7-13 x 2.0-3.0 µm, 0-3-septate, mostly 1-septate, each cell uni- to pluriguttulate.
In vitro very slow growing, whitish, sporulating moderately after three weeks or more.

Material examined

G.L.H. 7544 (holotype) and CBS 570.76 = MUCL 7544 (ex holotype), and G.L.H. 7549, on dead twigs of Symphoricarpus albus, Heikendorf nr. Kiel, F.R.G., 28/29 July 1963.


C. hennebertii differs from C. oligospermum by the absence of sterile setae, less frequently proliferating conidiophores and more irregular, somewhat shorter and broader conidia. [p. 52]

  fig. 1

Fig. 25
Cylindrotrichum zignoëllae, PRM 794087, x 1000.

  fig. 25

Fig. 26
Chaetosphaeria abietis, a. ascospores, conidiophore and conidia from isotype specimen, b. ascopores from PRM 796749. x 1000.

  fig. 1

Fig. 27
Cylindrotrichum clavatum, conidiophores and conidia from the natural substrate, CBS 428.76. x 1000. [p. 53]


3a. Chaetosphaeria abietis (Höhn.) W. Gams & Hol.-Jech. comb. nov. - Fig. 26

Zignoëlla abietis Höhn. apud Rehm - Annls mycol. 5: 469. 1907; Sber. K. Akad. Wiss. Wien, Math.-nat. Kl. 118: 332. 1909 (Fragm. No. 230).

Description (largely after v. Höhnel, 1909): Perithecia gregarious, superficial on a brown hyphal subiculum, globose, ostiole conical, black, brittle, shining, c. 180 µm diam; asci 80-90(-120) x 8-10 µm. Ascospores tristichous, fusiform with slightly rounded ends, somewhat curved, hyaline, smooth-walled, indistinctly three-septate, 27-33 x 3.0-3.5 µm (orig. descr. 26-32 x 3-4 µm); each cell with a lipid droplet. Conidial state Cylindrotrichum zignoëllae.

Material examined

Zignoëlla, abietis, isotypus, Rehm Ascom. exs. No. 1740, on Abies alba, Dürrwien, Wiener Wald, July 1907 (BPI).
PRM 796749, Picea abies, Svidový potok valley, Lower Tatra, Slov., 13 Aug. 1969.


3b. Cylindrotrichum zignoëllae (Höhn.) W. Gams & Hol.-Jech. comb. nov. - Fig. 25, 26a

Acrotheca zignoëllae Höhn. - Sber. K. Akad. Wiss. Wien, Math.-nat. Kl. 118: 332. 1909 (Fragm. No. 230).

Conidiophores on the natural substrate simple, dark brown, to 230 µm tall, 3.5-6.0 µm wide at the base, the upper part 3.0-4.0 µm wide and tapering to 1.0-1.5 µm below the short, widening collarette, sometimes percurrently but not sympodially proliferating. Conidia ellipsoidal-cylindrical, base truncate, one- or two-celled, sometimes slightly pigmented, 6-12 x 2.0-3.5 µm (orig. descr.: 10-13 x 2-3 µm), each cell uniguttulate. Sterile setae absent.

Additional material examined

PRM 794082, Salix sp., nr. Tekovský Hrádok nr. Levice, Slov., 5 Aug. 1975.
PRM 794087 and 797096, Quercus robur, Súr Forest nr. Bratislava, Slov., 24 Aug. 1974.
PRM 797095, Fagus sylvatica, between Mt. Revan and Mt. Klak nr. Nitran. Pravno, Slov., 14 July 1976.


In the few specimens so far studied a sympodial proliferation of the phialides was never observed in spite of the narrow phialide tip. This is apparently the major difference from C. hennebertii. The connection with the perfect state is so far only probable because of the joint occurrence [p. 54] of both states on the type specimen. In PRM 796749 (Fig. 26b) the conidia are constantly one-celled and, except for the absent sympodial proliferation, very similar to Chloridium cylindrosporum.


4. Cylindrotrichum clavatum W. Gams sp. nov. - Fig. 27

Conidiophora 110-180 µm alta, fusca, sursum pallidiora, levia, prope basin 5-6 µm lata, raro percurrenter nec sympodialiter proliferantia; phialides 25-45 µm longae, 4.0-4.5 µm latae, ad 1.5-2.0 µm angustatae. Conidia in capitulis albis connexa, cylindrica ad modice clavata, basi distincte apiculata, uno vel raro duobus septis divisa, 8.5-13(-15) x 3.0-4.0 (-5.0) µm. Setae steriles absunt. Chlamydosporae absunt.
Holotypus CBS .128.76, in ligno putrido Ulmi scabrae, Örups Almskog, Skåne, Sweden, leg. W. Gams et B. Söderström, 19 June 1976.

Conidiophores on the natural substrate 110-180 µm long, dark brown, lighter towards the tip, smooth-walled, 5-6 µm wide at the base; phialides 25-45 µm long, 4.0-4.5 µm wide, tapering in the uppermost part to 1.5-2.0 µm, rarely percurrently proliferating. Conidia accumulating in white heads, cylindrical to slightly clavate, distinctly apiculate at the base, with a septum at or somewhat above the centre, rarely with two septa; each cell uni- or pluriguttulate, 8.5-13(-15) x 3.0-4.0 (-5.0) µm. Sterile setae absent.
In vitro reaching 1.5-2.0 cm diam in 3 weeks, cream, moist with some darker grey spots, remaining almost sterile. Vegetative hyphae mostly hyaline, 1.0-2.5 µm wide, sometimes light brown and densely branching, 2.5-4.0 µm wide. Chlamydospores absent.

Additional material examined

Sub Hyalopus ater Corda, on decayed wood, Sutton, W. B. Grove, 25 Oct. 1884 (K ex BM). PRM 796767, indet. wood, Velká Fatra Mts., Gadierská dolina valley, 13 July 1976.


C. clavatum is close to C. zignoëllae, but differs by more clavate and larger conidia.


8. Codinaea Maire

Codinaea Maire - Publnes Inst. Bot., Barcelona 3: 15. 1937
Menisporella Agnihothrudu - Proc. Indian Acad. Sci. 56(B): 98. 1962

For diagnosis and key to the species cf. Hughes and Kendrick, 1968

  fig. 28

Fig. 28
Codinaea clavulata, PRM 794083. x 1000

Codinaea clavulata Hol.-Jech. sp. nov. - Fig. 28

Coloniae velutinae, fuscae; conidiophora erecta, fuscissima, sursum pallidiora, 130-340 µm alta, ad 15-septata; nonnumquam rupta percurrunt; ad 8 µm lata prope basin, in media 3.5-4.5 µm, ad 1.0-1.5 µm sursum angustata; collare breve; incrementa sympodialia brevia. Conidia in capitulis albidis aggregata, clavata et curvata, latissima in parte superiore, deorsum, acutata et minime truncata, sursum rotundata, hyalina, levia,, 8-10 x 1.7-2.5 µm. Holotypus PRM 794083, in ramo putrido Carpini betuli, ad septemtrionem montis Vlčí vrch prope Čelovce, in Slovakia, 11 Aug. 1975.

Conidiophores forming a dense velvety layer on wood; very dark brown but becoming lighter in the uppermost part, 130-340 µm tall and to 15-septate; sometimes percurrently proliferating after rupture; to 8 µm wide at the base, 3.5-4.5 µm wide centrally, tapering to 1.0-1.5 µm with a very shallow collarette; short sympodial elongations occur. Conidia aggregated in pale heads, clavate and curved, widest above the centre, base tapering and hardly truncate, tip rounded, hyaline, smooth-walled, 8-10 x 1.7-2.5 µm. Chlamydospores were observed on the natural substrate, brown, 9-10.5 x 6.5-7.0 µm.
Only known from the type collection.


This species is similar to both Chloridium codinaeoides Piroz. from which it differs in the very narrow phialide openings and sympodial proliferation, and the Codinaea state of Chaetosphaeria callimorpha (Mont.) Sacc., from which it is separated by its forming only one layer of tall conidiophores with prominent clustered phialide openings and shorter conidia (10-14.5 x 2.0-2.9 µm in Chaet. callimorpha). [p. 56]


9. Exochalara W. Gams & Hol.-Jech. gen. nov.

Conidiophora erecta, simplicia, pigmentata, identidem septata, collari singulo apicali minuto praedita, catenas basipetales phialoconidiorum hyalinorum, simplicium formant; saepe percurrenter proliferant.
Species typica Exochalara longissima (Grove) W. Gams & Hol.-Jech.

Conidiophores erect, solitary, pigmented, repeatedly septate; phialides bearing a single apical small collarette, forming basipetal chains of hyaline, one-celled conidia; often percurrently proliferating.
Chalara longissima Grove was excluded from Chalara by Nag Raj and Kendrick (1976) because of the relatively short, obconical collarette and the absence of a deep-seated conidiogenous locus; these authors showed that its inclusion in the subgenus Euchalara by v. Höhnel (1902: 1050/64) was incorrect since it differs essentially from the type species Chalara fusidioides (Corda) Rabenh. Its classification in Chloridium (possibly as a fourth section) is also not recommendable, since the regular chains of fusiform conidia are not comparable to any other species in this genus. Catenularia guadalcanalensis Matsushima does not fall within the circumscription of this genus given by Hughes (1965); the conidia are not thick-walled or cuneate and capitate hyphae are absent. Although its dimensions are much larger, this species may be likened to Chalara longissima.

Key to the species


Conidia fusiform or clavate and with truncate ends, 6.5-10.5 x 2.5-3.5 µm

E. longissima


Conidia elongate-ellipsoidal, 16-20 x 6.5-9.5 µm

E. guadalcanalensis


1. Exochalara longissima (Grove) W. Gams & Hol.-Jech. comb. nov. - Fig. 29

Chalara longissima Grove - J. Bot., Lond. 1885: 12.
Chalara sp. Reisinger - Contribution à l’étude ultrastructurale de l’appareil sporifère chez quelques hyphomycètes à paroi mélanisée. These Nancy, 1972, p. 8.
Catenularia piceae M. B. Ellis - More demat. Hyphom. p. 443. 1976.

Detailed description in Nag Raj and Kendrick (1976: 177).

Colonies very slow growing, reaching 2 mm diam in 14 days at 20-22°C, blackish-brown. Vegetative hyphae hyaline and 1.5-2.0 µm wide, or slightly pigmented, 2.0-3.5 µm, partly monilioid and to 5 µm wide. Sporulation poor but improved on cherry decoction agar. Conidiophores erect, simple, 55- over 100 µm tall before proliferation, 4.0-6.5 µm wide, dark brown, lighter above; phialides gradually tapering to 1.5-2.0 µm below the collarette; repeatedly percurrently proliferating to more than 300 µm (very rarely also sympodially proliferating). Collarette funnelshaped, [p. 57] 2.5-3.0 µm deep, widening to 2.5-3.0 µm diam. Conidia cohering end-to-end in long chains, fusiform or slightly clavate, usually widest above the centre, base slightly truncate, tip rounded or acuminate and truncate, hyaline (in old material slightly pigmented), smooth-walled, 6.0-10.5 x 2.5-3.5(-4.5) µm, length/width ratio 2.0-3.0, one- to several-guttulate. Chlamydospores absent.


fig. 29

Fig. 29
Exochalara longissima, a. dried culture O. Reisinger; b. G.L.H. 3255, c. PRM 793986, both from the natural substrate, x 1000

Material examined

CBS 980.73, ex decaying wood, Pijnenburg forest nr. Baarn, 19 Aug. 1972.
Chalara sp., No. 5, dried culture, isolated as coloniser of filter paper exposed in soil in the Vosges, France, comm. O. Reisinger (Herb. CBS).
On Picea abies(?), wood, Láznĕ Kundratice, Boh., leg. W. Gams and V. Holubová-Jechová, 20 June 1974 (Herb. CBS).
G.L.H. 3255, Larix leptolepis, Baarn, 7 Apr. 1963.
PRM 793986, Fagus sylvatica, nr. Černý potok river, Jizerské hory Mts., Boh., 1 Aug. 1965. [p. 58]
PRM 794062, Picea abies, Mt. Královský kámen nr. Kašperské Hory, Boh., 1 Oct. 1975.
PRM 794078, Betula verrucosa, Kacín nr. Lama, Slov., 16 Sept. 1975.
IMI 167413, holotype of Catenularia piceae M. B. Ellis, Picea sp., Hartland Forest, N. Devon, G.B., M. B. Ellis, 22 May 1972.


The holotype specimen IMI 17047 on rotting wood, Birmingham, England, was examined by Nag Raj and Kendrick (1976).
The strain CBS 980.73 grows extremely slowly and sporulates rather poorly. A dried culture obtained from O. Reisinger showed abundant sporulation and probably had had a faster radial growth.


2. Exochalara guadalcanalensis (Matsushima) W. Gams & Hol.-Jech. comb. nov. - Fig. 30

Catenularia guadalcanalensis Matsushima - Microf. Solomon Isl., Papua-N. Guinea, Kobe, p. 10. 1971.


fig. 30

Fig. 30
Exochalara guadalcanalensis, two conidiophores, x 500, conidiophore tips and conidia x 1000, CBS 346.76.

Colonies rather fast growing, reaching 3-4 cm diam in 10 days at 20-22°C, dark brown with a wide hyaline margin, with lighter grey aerial mycelium; reverse olivaceous black; sporulation abundant. Vegetative hyphae hyaline or pigmented, mostly 2-4 µm wide. Conidiophores arising from dark, swollen, rhizoid-like cells in the substratum, dark brown, lighter at the tip, smooth-walled, slightly roughened near the tip, 150 over 400 µm tall, and to 14 times septate, 6.0-7.5 µm wide at the base, 3.0-4.5 µm at the phialide tip. Collarette little differentiated, 1-2 µm deep, darker than the wall below. Conidia in regular chains, ellipsoidal with a truncate base, hyaline, smooth-walled, containing numerous lipid droplets, 16-20 x 6.5-9.5 µm (orig. descr.: 16-30 x 8-10 µm). Chlamydospores absent. Conidia germinating in any direction and sometimes becoming two-celled.

Material examined

CBS 346.76 = MFC-2875, ex holotype, collected on leaf of Musa sp., Honiora, Guadalcanal Island, T. Matsushima, 3 Jan. 1970.


10. Phialophora Medlar

Phialophora Medlar - Mycologia 7: 202. 1915.

For monographic treatment see Schol-Schwarz (1970).

The genus Phialophora can briefly be characterized as having pigmented or hyaline colonies with weakly pigmented or hyaline, thin-walled, basitonously branched conidiophores or simple phialides. Phialides discrete, typically flask-shaped, with a distinct, ± cup-shaped collarette. Conidia one-celled, hyaline or slightly pigmented.
Type species Phialophora verrucosa Medlar.

The recognition of a new section Catenulatae automatically establishes a section PHIALOPHORA typified by P. verrucosa Medlar. This is not the place for a full discussion of all sections to be distinguished, but for the sake of homogeneity it is desirable to reserve the section Phialophora for species with pigmented colonies, flask-shaped phialides formed in branched conidiophores, with distinct collarette and ellipsoidal conidia. Section Phialophora then comprises mainly P. fastigiata, P. malorum, P. verrucosa, P. mustea, P. lagerbergii and P. repens.

Phialophora section Catenulatae W. Gams sect. nov.

Coloniae pigmentatae vel hyalinae; conidia plus minusve cuneata vel dacryoidea, basi truncata, catenulata.
Species typica status imperfectus Lasiosphaeriae hirsutae (Fr.) Ces. & De Not. [p. 60]

Colonies pigmented or hyaline; conidia ± cuneate or dacryoid with a truncate base, regularly arranged in chains.


The delimitation of this new section from sect. Phialophora is not sharp. Lasiosphaeria ovina (Fr.) Ces. & De Not. has dacryoid conidia which are generally aggregated in slimy heads. The hardly pigmented colonies and dacryoid conidia exclude this state from sect. Phialophora as characterized above but there are no doubts about its inclusion in the genus Phialophora. The natural affinity between L. ovina and L. hirsuta is a strong argument for the incorporation of the conidial state of the latter in Phialophora, as already proposed by Hughes (1951a). In strains close to P. cyclaminis van Beyma conidial chains also occur sometimes, but conidial heads are more common and typical of this group; therefore they are not considered here.
The conidial state of Meliolina mollis (Berk. & Br.) Höhn. (Pirozynski, 1974) is similar to Phialophora sect. Catenulatae, but the conidiophores are very darkly pigmented, thick-walled, and dichotomously branched. Attempts to culture this fungus have failed (V. Agnihothrudu, pers. comm.).
In Chloridium the species most strongly reminiscent of Phialophora sect. Catenulatae are C. clavaeforme and C. lignicola. In these the thick-walled, unbranched but septate conidiophores are a good criterion for their generic separation.
The delimitation of sect. Catenulatae from Chalara is particularly difficult and the proper placement of some species is likely to be satisfactorily determined only after the finding of a perfect state or by chemical techniques. For example the Chalara state of Ceratocystis autographa Bakshi might be referred to this section were the Ceratocystis state not known. Consequently the new species Ch. microchona is accomodated in Chalara here because of the almost perfect agreement in many characters with the former species and in spite of the short funnel-shaped collarette.
Some distinctive features characterizing the two genera are tabulated below:



Phialophora sect. Catenulatae


phialides integrated in a septate conidiophore or arising solitarily from a vegetative hypha

phialides discrete in a branched conidiophore or arising solitarily from a vegetative hypha


at least the length of one conidium, ± cylindrical, or slightly divergent

usually shorter than one conidium, funnel-shaped or flared


strictly cylindrical, clavate or slightly ellipsoidal, length/width > 3

cuneate, clavate, dacryoid or ellipsoidal with a truncate base, length/width < 2.5.

[p. 61]

In order to prevent misidentifications, some species accepted by Nag Raj and Kendrick (1976) in Chalara are also treated in the present key; the new species Chalara microchona, described below has also been included.

Key to the species


Colonies brown, grey or olivaceous



Colonies hyaline or pink






Conidia with gradually tapering (or sharply pointed) tips



Conidia with truncate tips






Conidia clavate with gradually tapering tips, 5-6 x 1.5-2.0 µm; colonies ochraceous brown

P. clavispora


Conidia sharply pointed, 5.0-7.5 x 1.0-1.7 µm; colonies olivaceous grey

P. oxyspora





Conidia short-clavate, slightly pigmented

P. olivacea


Conidia hyaline






Phialides arising from often fasciculate, dark vegetative hyphae; often sympodially proliferating in old cultures; conidia elongate-clavate



Vegetative hyphae not fasciculate; conidia dacryoid or cuneate






Conidia formed within a 6-10 µm long, cylindrical to obconical collarette which is darkened at the constricted base

Chalara state of Ceratocystis autographa


Conidia formed within a 1.0-2.5 µm deep, funnel-shaped collarette

Chalara microchona





Phialides hyaline, constricted below the collarette; conidia 3.0-4.5 x 2.0-2.7 µm

P. lasiosphaeriae


Phialides pigmented, gradually tapering towards the collarette; conidia 2.0-3.0 x 1.5-2.0 µm

P. phaeophora





(from 1) Colonies pink or red; conidia 2.5-4.0 x 1.5-2.5 µm

P. rhodogena


Colonies white or whitish






Conidia formed within a cylindrical collarette, cylindrical or slightly clavate with a rounded tip, 3.5-8.0 x 1.5-2.2 µm

Chalara state of Cryptendoxyla hypophloia


Collarette and conidia shorter






Collarette inconspicuous, rarely 2 µm deep; conidia 3.0-4.5 x 1.2-1.7 µm

P. brevicollaris


Collarette conspicuous, commonly 2 µm deep






Conidia scarcely longer than wide, 2.0-3.0 x 1.7-2.0 µm

P. brachyconia


Conidia about twice as long as wide, short-clavate, 3.0-4.0 x 1.5-2.0 µm

P. hyalina

[p. 62]


fig. 31

Fig. 31
Phialophora lasiosphaeriae, conidiophores and conidia, CBS 955.72, x 1000


fig. 32

Fig. 32
Phialophora clavispora, CBS 129.74, x 1000

1a. Lasiosphaeria hirsuta (Fr.) Ces. & De Not.

Lasiophaeria hirsuta (Fr.) Ces. & De Not. - Comment. Soc. crittog. ital. 1: 229. 1863.


1b. Phialophora lasiosphaeriae W. Gams st. nov. - Fig. 31

Coloniae lente crescunt, ad 6 mm diametro post 14 dies, olivaceo-nigrae vel brunneo-nigrae, velutinae, margine quasi nigra. Hyphae vegetativae ad 2.5-3.5 µm latae, saepe pigmentatae et crassitunicatae. Phialides plerumque simplices, nonnumquam conidiophora bicellularia, deorsum pigmentata et crassitunicata, 15-40 x 3.0-3.5 µm, sub collari constricta, nonnumquam percurrenter vel sympodialiter proliferantia; collare 2.5-4.0 µm altum, modice divergens. Conidia plerumque catenulata, guttuliformia ad clavata, basi truncata, hyalina, levia, 3.0-4.5 x 2.0-2.7 µm. Chlamydosporae absunt.
Holotypus CBS 955.72, isolatus ex ascosporis Lasiosphaeriae hirsutae prope Detmold-Heiligenkirchen in Germania, Sept. 1972.

Description also in Hughes (1951: 30) and Gams (1973) as "Phialophora state".

Colonies slow-growing, reaching 6 mm diam in 14 days at 20-22°C, olivaceous-black or brownish-black, velvety, with an almost black margin. Vegetative hyphae to 2.5-3.5 µm wide, often pigmented and thick-walled. Conidiophores mostly consisting of simple phialides, sometimes with an additional septum near the base, pigmented in the lower part, 15-40 x 3.0-3.5 µm, constricted below the collarette, sometimes percurrently or sympodially proliferating. Collarette 2.5-4.0 µm deep, cylindrical with slightly flaring margin. Conidia mostly forming chains, dacryoid to clavate with a truncate base, hyaline, smooth-walled, 3.0-4.5  2.0-2.7 µm, length/ width ratio 1.4-2.0. Chlamydospores absent.

Additional material examined

Isolate from ascospores, H. A. van der Aa, No. 1, Gerolstein, Eifel, F.R.G., 13 Sept. 1971.


The related Lasiosphaeria ovina (Fr.) Ces. & De Not. (ascospore isolates CBS 725.69, 958.72, 596.75), illustrated by Gams (1973), has less pigmented, ochraceous colonies, shorter phialides, 8-15  2.0-2.7 µm and guttuliform conidia, 2.5-4.0 x 2.0-2.5 µm, forming slimy heads.


2. Phialophora clavispora W. Gams sp. nov. - Fig. 32, 33.

? Sphaeromyces clavisporus Arnaud - Bull. trimest. Soc. mycol. Fr. 69: 279. 1953, nom. inval. (Art. 36).

Coloniae lente crescunt, ochraceo-brunneae, pigmento brunneo diffundente. Hyphae vegetativae hyalinae vel dilute brunneae, 1.0-2.5 µm latae. Plerumque phialides simplices, nonnumquam conidiophora bicellularia, dilute brunnea, 23-38 x 2.5-3.0 µm, sub collari modice angustata; collare paulum divergens, 2.5-3.5 µm altum; proliferatio sympodialis in coloniis vetustis. Conidia catenulata, nonnumquam capitulata, clavata, basi quasi acuta, apice rotundata vel acutata, hyalina, levia, 5.0-6.0 x 1.5-2.0 µm. Chlamydosporae absunt. Holotypus CBS 637.73, isolatus e ligno putrido, La Heutte in Montibus Jurae in Helvetia, W. Gams, 1973.

Colonies reaching 5 mm diam in 16 days at 20-22°C, ochraceous brown, the agar becoming discoloured brown. Vegetative hyphae hyaline or light brown, sometimes monilioid, 1.0-2.5 µm wide. Conidiophores mostly phalacrogenous with simple phialides, but often with an additional septum in the lower part, light brown, 23-38 x 2.5-3.0 µm, only slightly tapered below the collarette to 1.0-1.5 µm; collarette slightly divergent, 2.5-3.5 µm deep; sympodial proliferation (schizophialides) occurring in old cultures. Conidia in chains or sometimes heads, clavate with an almost pointed base and rounded or almost pointed tip, hyaline (in very old material slightly pigmented), smooth-walled, 5.0-6.0 x 1.5-2.0 µm, length/width ratio 2.2-3.6. Chlamydospores absent. [p. 64]


fig. 33

Fig. 33
Phialophora clavispora, CBS 637.73 x 1000


fig. 34

Fig. 34
Phialophora oxyspora, CBS 698.73, x 1000

Additional material examined

CBS 129.74, Waterleidingsduinen nr. Vogelenzang, Netherlands, Nov. 1969.
CBS 271.75, ex Picea abies, Láznĕ Kundratice, Boh., 1974.


P. clavispora is similar to Arnaud’s drawing of Sphaeromyces clavisporus of which the conidia were reported to be 12-14 x 3-4 µm and aggregated in heads. On Arnaud’s specimen (Parc de Versailles, 7 Apr. 1948, PC) this fungus could not be found and so its identity remains somewhat doubtful.


3. Phialophora oxyspora W. Gams sp. nov. - Fig. 34

Coloniae modice lente crescunt, olivaceo-griseae, pulverulentae, tomentosae. Hyphae vegetativae modice pigmentatae, 1.2-2.0 µm latae. Phialides plerumque simplices, raro conidiophora bicellularia adsunt; phialides brunneae, 20-30 x 2.0-2.5 µm, vix angustatae; collare plerumque paulo obscurius, 1.5-2.0 µm altum, vix divergens. Conidia in catenis longis aggregata, fusiformia, basi late truncata, sursum acutata, raro rotundata, dilute pigmentata, levia, 5.0-7.5 x 1.0-1.7 µm. Chlamydosporae absunt.
Holotypus CBS 698.73, isolatus ex aphidibus in Clerodendro monahassa, in horto botanico Peradeniya in insula Zeylania, Jan. 1973. [p. 65]

Colonies reaching 13-18 mm diam in 14 days at 20-22°C; olivaceous grey, powdery and tufted. Vegetative hyphae slightly pigmented, 1.2-2.0 µm wide. Conidiophores mostly formed as simple phialides, rarely with an additional septum, brown, 20-30 x 2.0-2.5 µm, hardly tapered towards the collarette; collarette usually slightly darker, 1.5-2.0 µm deep, hardly divergent. Conidia forming long chains, fusiform, with a broadly truncate base and a long, pointed or rarely rounded tip, slightly pigmented, smooth-walled, 5.0-7.5 x 1.0-1.7 µm, length/width ratio 3.0-5.0. Chlamydospores absent.
Known only from the type strain.


4. Phialophora olivacea W. Gams sp. nov. - Fig. 35

Coloniae fere lente crescunt, olivaceo-nigrae, tomentosae, reverso nigro. Hyphae vegetativae omnino dilute pigmentatae, leves, 0.7-2.5 µm latae. Sporulatio plectonematogena; phialides plerumque simplices, raro conidiophora bicellularia, modice pigmentata, collare versus fusciora, 12-33 x 1.5-2.5 µm, sub collari modice angustata; collare 1-2 µm altum, divergens. Conidia in longis catenis aggregata, clavata vel ellipsoidea, basi truncata, dilute pigmentata, hilum basilare obscurius, levia, 3.0-5.5 x 1.3-2.2 µm. Chlamydosporae absunt. Holotypus CBS 122.74, isolatus e tapeto, Kiel-Kitzeberg in Germania, W. Gams, 1966.

Colonies reaching 13-22 mm diam in 14 days at 20-22°C, olivaceous black, tufted, reverse black. Vegetative hyphae uniformly slightly pigmented, smooth-walled, 0.7-2.5 µm wide. Sporulation plectonematogenous. Conidiophores mostly simple phialides, rarely with an additional septum, slightly pigmented, darkest at the collarette, 12-33 x 1.5-2.5 µm, slightly constricted below the collarette; collarette 1-2 µm deep, slightly divergent. Sympodial proliferation observed rarely. Conidia forming long chains, clavate to ellipsoidal with a truncate base, slightly but distinctly pigmented, the basal hilum somewhat darker, smooth-walled, 3.0-5.5 x 1.3-2.2 µm, length/width ratio 2.2-3.0. Chlamydospores absent.

Additional material examined

CBS 435.69, ex stalactites, Lehman Caves, Eastern Nevada, comm. F. W. Went (No. 103d), 1969.
CBS 123.74, ex dead petioles of Chamaerops humilis, Cantonspark, Baarn, Netherlands, 1968.
CBS 124.74, ex Inonotus obliquus, Segeberger Forst, Kr. Segeberg, F.R.G., Nov. 1965.

5. Phialophora phaeophora W. Gams sp. nov. - Fig. 36

Coloniae lente crescunt, post 14 dies ad 7 mm diametro, griseae in medio, margine hyalinae, deinde olivaceo-brunneae, reverso griseo. Hyphae vegetativae hyalinae ad fuscae, 1.5-3.0 µm latae, fere crassitunicatae, leves vel verrucosae, nonnumquam monilioideae, ad 5.5 µm latae. [p. 66]


fig. 35

Fig. 35
Phialophora olivacea, a. CBS 122.74, b. CBS 123.74, x 1000.


fig. 36

Fig. 36
Phialophora phaeophora, CBS 699.74, x 1000.


fig. 37

Fig. 37
Phialophora rhodogena, a. CBS 886.73, b. H. A. 4378, c. CBS 127.74, x 1000. [p. 67]

Phialides plerumque simplices, raro conidiophora composita e cellula basilari et duabus phialidibus adsunt; phialides brunneae, collare versus fusciores, 10-20 x 2.0-2.5 µm, paulatim angustatae ad 1.0-1.5 µm; collare caliculare, 1.0-2.0 µm altum. Conidia plerumque catenulata, nonnumquam, capitulata dacryoidea, basi truncata, hyalina, levia, 2.0-3.0 x 1.5-2.0 µm. Chlamydosporae absunt.
Holotypus CBS 699.74, isolatus e ramo emortuo Fraxini excelsioris, Berenkuil prope Rheni Trajectum (Utrecht) in Neerlandia, W. Gams, Aug. 1974.

Colonies reaching 7 mm diam in 14 days at 20-22°C, grey in the centre and with a hyaline margin, later becoming olivaceous brown, reverse grey. Vegetative hyphae hyaline to dark brown, 1.5-3.0 µm wide, rather thick-walled, smooth-walled or warted, sometimes monilioid and to 5.5 µm wide. Phialides mostly simple, rarely two on a single stalk cell, brown, with a darker zone at and below the collarette, 10-20 x 2.0-2.5 µm, gradually tapering in the upper half to 1.0-1.5 µm wide. Collarette cup-shaped, 1.0-2.0 µm deep. Conidia mostly in chains, sometimes in heads, dacryoid with a truncate base, hyaline, smooth-walled, 2.0-3.0 x 1.5-2.0 µm, length/width ratio 1.2-1.5. Chlamydospores absent.

Additional material examined

CBS 260.76, ex Fagus sylvatica, Beller Holz, Teutoburger Wald, F.R.G., Sept. 1972.
DAOM 4552 (a), ex yellow-brown sapwood of Picea glauca, Victoria, B.G., Canada, D. E. Wells, 2 Oct. 1953.


P. phaeophora recalls Chloridium clavaeforme but the conidiophores consist usually of discrete phialides and are much shorter with the conidia more regularly catenulate.


6. Phialophora rhodogena (Mangenot) W. Gams comb. nov. - Fig. 37

Scopulariopsis rhodogena Mangenot - Revue gén. Bot. 59: 442. 1952 (Rech. méthod. Champ. Bois en Décomp., Paris, p. 25).

Colonies slow-growing, reaching 6-8 mm diam in 14 days at 20-22°C, at first white, then pale pink to intensely red, particularly in the centre and from the reverse, the pigment diffusing into the agar, with age becoming partly grey in the centre. Vegetative hyphae hyaline, rather thick-walled, 1.5-3.0 µm wide, some cells inflated to 5-6 µm diam (almost like chlamydospores). Sporulating structures arising mostly from submerged hyphae (phalacrogenous); conidiophores mostly simple phialides, sometimes with an additional septum in the lower part; phialides often terminal on creeping vegetative hyphae; mostly 15-30 x 2.0-3.0 µm, hyaline, smooth-walled or irregularly encrusted; collarette 2.0-3.0 µm deep, usually slightly divergent; sometimes showing percurrent proliferation. Conidia [p. 68] forming long chains or sometimes slimy heads, cuneate to dacryoid, with a truncate base, 2.5-4.0(-5.5) x 1.5-2.5 µm, length/width ratio 1.2-2.0 (-3.0). Chlamydospores absent.

General distribution

A very common saprophyte, particularly on decaying Piptoporus betulinus on which it is nearly always present. A strong antibiosis against a contaminating basidiomycete was observed in culture.

Material examined

CBS 335.53, type strain, ex decaying wood of Betula sp., France, comm. F. Mangenot, 1953.
CBS 490.67, ex Piptoporus betulinus, Aukrug, Kr. Rendsburg, F.R.G., Oct. 1965.
CBS 886.73, ex Piptoporus betulinus, Leusden, Netherlands, Sept. 1973.
CBS 127.74, ex Piptoporus betulinus, Segeberger Forst, Kr. Segeberg, F.R.G., 26 Apr. 1965.
CBS 303.74, ex stained bark of Fagus sylvatica, Gooilust nr. Hilversum, Netherlands, March 1974.
CBS 687.74, ex Quercus pubescens, Murs, Vaucluse, France, Oct. 1974.
CBS 231.75, ex Tilia sp., Hamritejn nr. Liberec, Boh., June 1974.
H.A. 4378, ex Ascotremella faginea on Quercus sp., Boschoord nr. Zorgvlied, nr. Diever (Dr.), Netherlands, leg. J. Geesink, 24 Nov. 1974 (Herb. CBS).


7. Phialophora brachyconia W. Gams sp. nov. - Fig. 38

Coloniae lente crescunt, albae, sed aurantiacae in extracto Pruni cerasi acido. Hyphae vegetativae hyalinae, 1.0-2.5 µm latae, nonnumquam ad 3.5 µm inflatae, saepe fasciculatae. Phialides simplices vel conidiophora composita ramosa, complures phialides ferentia, hyalina, crassitunicata; phialides 12-30 x 2.0-2.5 µm, sursum vix angustatae, fere crassitunicatae et sub collari intus inspissatae; collare cylindricum vel caliculare, 1.5-2.0 µm altum. Conidia plerumque in longis catenis aggregata, dacryoidea, basi truncata, 2.0-3.0 x 1.5-2.0 µm. Chlamydosporae absunt.
Holotypus CBS 700.73, isolatus e ligno putrido, Irrel prope Augustam Trevirorum (Trier) in Germania, Maj. 1973.

Colonies very slow-growing, reaching 8-16 mm diam in 14 days at 20-22°C, usually pure white but developing an intense orange-ochre pigmentation on cherry decoction agar (pH c. 4.5). Vegetative hyphae hyaline, < 1.0-2.5 µm wide, sometimes inflated to 3.5 µm, often fasciculate. Conidiophores consisting of simple phialides, sometimes with an additional septum in the lower part, or a few times septate and branched, hyaline, thick-walled. Phialides often formed at the end of prostrate (or fasciculate) vegetative hyphae (telophialides); 12-30 x 2.0-2.5 µm, hardly tapered towards the collarette, rather thick-walled, with a pronounced wall-thickening below the collarette; collarette cylindrical or cup-sphaped, 1.5-2.0 µm deep. Conidia generally forming long chains, dacryoid with a truncate base, 2.0-3.0 x 1.5-2.0 µm, length/width ratio 1.1-1.5. Chlamydospores absent. [p. 69]


fig. 38

Fig. 38
Phialophora brachyconia, a. CBS 700.73, b. CBS 630.75, x 1000.

Additional material examined

CBS 304.74, ex stained bark of Fagus sylvatica, Gooilust nr. Hilversum, Netherlands, March 1974.
CBS 630.75, Grande Tinémont nr. Han-sur-Lesse, Belgium, Sept. 1975.
CBS 647.75, Robbenoordbos, Wieringermeer-Polder, Netherlands, Oct. 1975.


The repeated isolation of P. brachyconia and Chloridium clavaeforme together might suggest that the former might be an albino mutant of the latter. Although the conidial shapes agree very closely, however, there are consistent differences in the conidiophores, these being much shorter and more frequently branched in P. brachyconia. [p. 70]


8. Phialophora hyalina W. Gams sp. nov. - Fig. 39

Coloniae lentissime crescunt, post 14 dies 9-12 mm diametro, albae lanosae ad tomentosae, deinde griseolae in medio. Hyphae vegetativae hyalinae, 1.5-4.5 µm latae, plus minusve monilioideae. Vulgo phialides simplices, sed etiam conidiophora ramosa adsunt; phialides cylindricae, 17-30 x 1.5-2.0 µm vel inflatae ad 3-5 µm; collare 1-2 µm altum, plus minusve divergens. Conidia catenulata vel capitulata, dacryoidea, breviter clavata vel ellipsoidea, basi truncata, 2.5-4.0 x 1.5-2.5 µm. Chlamydosporae absunt.
Holotypus CBS 130.74, isolatus e solo agresti, Kiel-Kitzeberg in Germania, 1965.

Colonies very slow-growing, reaching 9-12 mm diam in 14 days at 20-22°C, white, cottony, tufted, later becoming grey in the centre. Vegetative hyphae hyaline, 1.5-4.5 µm wide, ± monilioid. Conidiophores often simple phialides but also repeatedly basitonously branched. Phialides variable, either cylindrical, 17-30 x 1.5-2.0 µm, or inflated to 3-5 µm and constricted below the collarette; collarette 1.0-2.0 µm deep, ± flaring. Conidia forming chains or slimy heads, dacryoid, shortly clavate or ellipsoidal with a truncate base, 2.5-4.0  1.5-2.5 µm, length/width ratio 1.6-2.2. Chlamydospores absent.

Material examined

CBS 130.74 (type strain), 177.74 and some other isolates from wheat field soils, Kiel-Kitzeberg and Kiel, F.R.G., 1965.


fig. 39

Fig. 39
Phialophora hyalina, CBS 130.74, x 1000. [p. 71]


fig. 40

Fig. 40
Phialophora brevicollaris, CBS 126.74, x 1000.


P. hyalina is close to the Chalara state of Cryptendoxyla hypophloia, but separable by the usually flaring collarette and shorter, clavate or ellipsoidal conidia.


9. Phialophora brevicollaris W. Gams sp. nov. - Fig. 40

Coloniae lentissime crescunt, post 14 dies 6 mm diametro, albae, deinde griseolae. Sporulatio phalacrogena ad synnematogena. Conidiophora plerumque compluries baritone ramosa, prope basin ad 3.5 µm lata, phialides discretas ferentia; phialides hyalinae, fere crassitunicatae, quasi cylindricae, 15-27 x 2.0-3.0 µm, paulatim angustatae ad 1.0-1.5 µm; nonnumquam sympodialiter proliferantes; collare inconspicuum, raro ad 2.0 µm altum. Conidia in longis catenis aggregata, ellipsoidea vel clavata, basi truncata, hyalina, levia 3.5-4.5 x 1.0-1.5 µm. Chlamydosporae absunt.
Holotypus CBS 126.74, isolatus e Phellino sp., Arbersee-Wand in Sylva Bavarica in Germania, W. Gams, 1966.

Colonies very slow growing, reaching 6 mm diam in 14 days at 20-22°C, white at first but later becoming light grey. Sporulation phalacrogenous [p. 72] to synnematogenous. Vegetative hyphae hyaline, smooth-walled, rather thick-walled, (1.5-)2.0-3.5 µm wide. Conidiophores usually a few times basitonously branched, at the base to 3.5 µm wide, with discrete phialides; phialides hyaline, rather thick-walled, almost cylindrical, 15-27 x 2.0-3.0 µm, gradually tapering to 1.0-1.5 µm at the tip; some-times sympodially proliferating; collarette hardly perceptible, rarely 2.0 µm deep. Conidia forming long chains, ellipsoidal to clavate, with a truncate and slightly inspissate base, hyaline, smooth-walled, 3.5-4.5 x 1.0-1.5 µm, length/width ratio 2.0-3.0. Chlamydospores absent.
Known only from the type strain.


10. Chalara state of Ceratocystis autographa Bakshi - Fig. 41

Ceratocystis autographa Bakshi - Ann. Bot., Lond., N.S. 15: 58. 1951. Chalara state described in Nag Raj and Kendrick (1976: 140).

Colonies reaching 12-15 mm diam in 10 days at 20-22°C, grey-brown, tomentose; aerial mycelium olivaceous brown, funiculose, consisting of warted pigmented hyphae, to 2.5 µm wide, giving rise to mostly simple phialides. Phialides in orthotropic or plagiotropic position, slightly darker than the subtending prostrate hyphae, flask-shaped, differentiated into a venter and a cylindrical to obconical collarette, pigmentation darker at the constriction, venter 7-12 x 3.0-3.5 µm, constricted to 1.0-1.5 µm wide above, collarette 5-7(-10) x 1.5-2.2 µm. Conidia catenulate, clavate with truncated ends, hyaline, smooth-walled, 3.5-4.0 x 1.0-1.5 µm; percurrent proliferation sometimes observed. Chlamydospores absent, but monilioid hyphae occurring.

Material examined

IMI 20162, holotype, ex galleries of Dryocoetes autographus and Hylurgops palliatus infesting Larix leptolepis (slide and dried cultures).
CBS 670.75, ex decaying needles of Juniperus communis lying on the ground, Lheedorp (Dr.), Netherlands, Nov. 1975.


This species is close to Chalara constricta Nag Raj & Kendrick, which is distinguished by its more complex phialophores which are up to 4-septate, and conidia to 8 µm long. Ch. constricta (CBS 248.76) has a stronger tendency to sympodial proliferation of the phialides, but this was also observed in C. autographa by Nag Raj and Kendrick. Perithecia are reported to be easily obtained in culture after mating of compatible strains (Bakshi, l.c.) and this feature should allow a reliable identification thus making a separate naming of the conidial state superfluous. [p. 73]


11. Chalara microchona W. Gams sp. nov. - Fig. 42

Coloniae lente crescunt, olivaceo-griseae ad nigrae, tomentosae, reverso nigro. Hyphae vegetativae pigmentatae, plus minusve incrustatae, ad 2.5 µm latae; conidiophora plerumque ex hyphis fasciculatis fuscis oriuntur, praecipue phialides simplices, rarius conidiophora bi- vel triseptata; phialides pigmentatae in parte inferiore, 18-35 x 2.5-4.0 µm, sub collarí modice angustatae ad 1.0-1.5 µm; collare 1.0-2.5 µm altum, infundibuliforme, ad 2.0-2.5 µm divergens; proliferatio sympodialis frequens. Conidia in longis catenis connexa, clavata, basi truncata, sursum rotundata, hyalina, levia, 3.5-5.0 x 1.3-2.5 µm. Chlamydosporae absunt.
Holotypus CBS 175.74, isolatus e cortice Pini sylvestris, in silva Elspeeter Bos’ dicta in Neerlandia, Feb. 1974.

Colonies very slow-growing, reaching 7-9 mm diam in 14 days at 20-22°C, olivaceous-grey to blackish, tufted, reverse black. Vegetative hyphae pigmented, ± encrusted, to 2.5 µm wide. Conidiophores often arising from dark fascicles of vegetative hyphae, either as simple phialides or with 1-2 additional septa. Phialides pigmented mainly in the lower part, 18-35 x 2.5-4.0 µm, tapered slightly below the collarette to 1.0-1.5 µm, collarette 1.0-2.5 µm deep, funnel-shaped, at the margin 2.0-2.5 µm wide; on ageing often sympodially proliferating. Conidia cohering in long chains, clavate, with a truncate base, hyaline, smooth-walled, 3.5-5.0 x 1.3-2.5 µm, length/width ratio 2.0-3.0. Chlamydospores absent.


fig. 41

Fig. 41
Ceratocystis autographa, Chalara state, a. dried culture IMI 20,62, b. living culture CBS 670.75 (with monilioid hypha). X 1000


fig. 42

Fig. 42
Chalara microchona, CBS 175.74, x 1000 [p. 74]


fig. 43

Fig. 43
Chalara cf. microchona, CBS 125.74, a. conidiophores with conidia, b. chlamydospores, x 1000


fig. 44

Fig. 44
Cryptendoxyla hypophloia, CBS 509.70 (14 days old). x 1000

Additional material examined

Living cultures
CBS 964.72, Pijnenburg Forest nr. Baarn, Netherlands, 19 Aug. 1972.
CBS 867.73, on dead twigs together with Inermisia fusispora, Maria Waldrast nr. Matrei am Brenner, Austria, May 1973.
CBS 889.73, ex Pinus sylvestris, Leusder Bossen, Netherlands, July 1973.
CBS 118.74, sent by Institut f. Holzkunde, München, F.R.G., 1966.
CBS 119.74, ex Pinus sylvestris bark, Kampina Heide, Netherlands, 1969.
CBS 120.74, ex roots of Picea abies, Denmark, D. S. Malla, 1973.
CBS 121.74, ex Fomitopsis pinicola, nr. Innsbruck, Austria, 22 Dec. 1965.

Herbarium specimens
DAOM 29256, Abies balsamea, Deer Lake, Newfoundland, D. R. Redmond, 25 Aug. 1952.
DAOM 33715, Picea glauca, Saskatoon, Sask., C. G. Riley.
DAOM 33778, root of Picea glauca, Saskatoon, Sask., 29 Oct. 1951.
DAOM 33981, root of Picea rubens, Fredericton, N.B., D. R. Redmond, 1951.
DAOM 45523, Larix occidentalis, Victoria, B.C., Craig, 1953.
IMI 114067, ex forest soil, GB., M. Hodkinson, 1965.


This is the least typical species of Chalara so far known because of its funnel-shaped, short collarette. It has been placed in this genus rather than in Phialophora because of the strong similarity in colony habit with conidial Ceratocystis autographa. The available living strains were mated in all possible combination, but failed to produce perithecia. [p. 75]
A further strain, CBS 125.74, isolated by T. Grinbergs from volcanic ash soil in Chile, is close to Ch. microchona but differs in often having one- to three-septate conidiophores, slightly longer collarettes (to 4 µm deep), and producing dark chlamydospores which often form complex balls (Fig. 43). This strain is a further link between Ch. microchona and more typical Chalara species.


12. Chalara state of Cryptendoxyla hypophloia Malloch & Cain - Fig. 44

Cryptendoxyla hypophloia Malloch & Cain - Can. J. Bot. 48: 1816. 1970.
Chalara state also described by Nag Raj and Kendrick (1976: 142).

Colonies rather slow-growing, reaching 11-20 mm diam in 14 days at 20-22°C; aerial mycelium white, cottony, somewhat tufted, reverse ochraceous-grey to fawn; vegetative hyphae hyaline but becoming partly light brown with age, mostly 1-2 µm wide. Phialides rather scarce, simple or with an additional septum in the lower part, cylindrical, 15-20 x 1.5-2.0 µm (orig. descr.: 17-50 x 2.0-3.0 µm; Nag Raj and Kendrick: 13-28 x 2.0-2.5 µm). Conidia catenulate or in slimy heads, cylindrical, slightly clavate with a truncate base, mostly 4.0-4.5  1.5-2.0 µm (orig. descr.: 3.0-7.0 x 1.5-2.5 µm; Nag Raj and Kendrick: 3.5-11 x 1.5-2.5 µm, length/width ratio 3.3). Chlamydospores absent. Numerous young ascomata develop after three weeks from spiral initials.

Material examined

CBS 508.70 = TRTC 45320, type culture, ex wood under bark of Acer sp., 13 Sept. 1967; and CBS 509.70 = TRTC 45682, ex Betula sp., 1 Nov. 1969, both South of Dorset, Muskoka Distr., Ontario, Canada, comm. R. F. Cain.


This is the only small hyaline Chalara species mentioned by Nag Raj and Kendrick (1976), but there are, however, some other probably distinct isolates from soil now known (CBS 972.72, 128.74A-C) which have slightly taller phialides (22-35 x 3-4 µm), a deeper collarette (5-6 µm long), conidia 4.5-5.0 x 1.5-2.2 µm, and no ascomata; these would key out as C. hypophloia in Nag Raj and Kendrick (1976).
Since C. hypophloia produces ascomata in culture, there is no need to give the conidial state a separate name as misidentifications can be excluded. [p. 76]


11. Fusichalara Hughes & Nag Raj

Fusichalara Hughes & Nag Raj - N.Z. Jl Bot. II: 662. 1973.

A diagnosis and key to the three original species is in Nag Raj and Kendrick (1976: 144).

Fusichalara minuta Hol.-Jech. sp. nov. - Fig. 45

Coloniae effusae, fuscae, pilosae. Conidiophora sparsa vel bina ad quaterna aggregata, brunnea ad atrobrunnea, erecta, nonnumquam flexuosa, ± cylindrica, levia, pariete 1.0-1.8 µm crasso, 110-280 µm longa, prope basin 5-7 µm, in medio 4-5 µm, sursum 3.0-3.5 µm, lata, 3-7-septata, saepe percurrenter proliferantia; phialides integratae, cylindricae, 20-35 µm longue. Conidia breviter catenulata, quasi hyalina, fusiformia ad clavata, plerumque bicellularia, utrinque, acutata sed sursum truncatula, (7-)8-12 x 1.5-1.8(-2.0) µm; conidia primaria sursum rotundata, ad 2.0-2.5 µm lata.
Holotypus: PRM 795927, in codice putrido Quercus petraeae, in silva Lánská obora dicta, inter Lány et Ploskov in Bohemia, V. Holubová-Jechová, 12 Jun. 1976.

Colonies on the natural substrate effuse, dark brown, hairy. Conidiophores scattered or in groups of 2-4, simple, brown to dark brown, erect, straight or slightly flexuose, ± cylindrical, smooth-walled, wall 1.0-1.8 µm thick, 110-280 µm tall, 5-7 µm wide at the base, 4-5 µm in the central part and 3.0-3.5 µm below the frilled apex, usually 3- to 7-septate, on ageing percurrently proliferating; phialides integrated, 20-35 µm long, regularly cylindrical, with a transparent but considerable internal wall thickening near the tip or at some distance below, through which the conidia are pierced out. Conidia in short chains, fusiform, almost hyaline, mostly 2-celled, with a truncate apical and narrowly conical basal end, (7-)8-12 x 1.5-1.8(-2.0) µm; the primary conidia clavate with a rounded apex and 2.0-2.5 µm wide.
In culture reaching 10 mm diam in 21 days at 20-22°C, brown with a broad white margin, without aerial mycelium, moderately sporulating; vegetative hyphae hyaline, monilioid, 1.5-2.5 µm wide in the narrow, 3-6 µm in the inflated portions. Conidiophores arising from submerged hyphae often in a terminal position.

Culture examined

CBS 571.76, ex holotype PRM 795927, on decaying trunk of Quercus petraea, Lánská obora Forest between Lány and Ploskov, Boh., isol. W. Gams, July 1976. [p. 77]


fig. 45

Fig. 45
Fusichalara minuta, a. from natural substrate PRM 795927, b. in culture CBS 571.76 (21 days old, inflated conidia to the right 28 days old). x 1000.


12. Gonytrichum C. G. & F. Nees ex Leman

Gonytrichum C. G. & F. Nees - Nova Acta Acad. Leop.-Carol. Nat. Cur. 9: 244. 1818; ex Leman - Dict. Sci. nat., Paris 19: 209. 1821.
Mesobotrys Sacc. - Michelia 2: 27. 1880.
? Piminella Arnaud - Bull. trimest. Soc. mycol. Fr. 69: 281. 1953, nom. inval. (Art. 36).

Conidiophore stipes simple or branched, brown, thick-walled, bearing one or several short curved lateral branches (collar hyphae) below the septa or at the branching nodes, from which clusters of phialides arise appearing verticillate. Phialides usually less pigmented than the conidiophores stipes, flask-shaped, tapering towards a very shallow collarette, rarely percurrently proliferating. Conidia arising from multiple conidiogenous loci, aggregated in slimy heads or cirrhi, one-celled, ellipsoidal to cylindrical, hyaline or pigmented. Simple conidiophores with terminal [p. 78] sporulation as in Chloridium occur in all species, particularly in degenerate (very old) cultures and on the natural substrate under suboptimal developmental conditions.

Species of Gonytrichum have been studied by Hughes (1951b), Swart (1959) and Barron and Bhatt (1967), whose work is summarized by Kirilenko (1972). Here we describe two new varieties of G. caesium, a new perfect state in Chaetosphaeria with G. chlamydosporium as its conidial state and a variety of it, and a new species, G. mirabile.
We mention here also Codinaea gonytrichodes Shearer & Crane (1971; also Matsushima, 1975) which is intermediate between Gonytrichum and Codinaea and has recently been transferred to a separate genus as Codinaeopsis gonytrichoides (Shearer & Crane) Morgan-Jones - Mycotaxon 4: 167. 1976. The conidiophores match Gonytrichum macrocladum, but the conidia are allantoid, one-celled and setulose and measure 10-13(-16.5) x 1.2-2.3 µm.

Key to the species and varieties


Conidiophore stipes much branched



Conidiophore stipes unbranched






Phialides clustered on collar hyphae at branching nodes



Collar hyphae mostly absent, simple phialides arising from the straight part of the conidiophore stipes

G. mirabile





Phialides in dense clusters (of at least 5 phialides), usually not exceeding 10 µm long, much paler than the conidiophore stipe; solitary conidiophores rare; conidia 2.5-3.0(-3.5)  1.0-1.5 µm

G. caesium var. caesium


Phialides usually in more irregular groups of 3-6, ± pigmented (although less than the conidiophore stipe); solitary conidiophores commonly present; conidia in range 3-5  1.5-2.5 µm






Conidia 3.0-3.3 x 2.0-2.5 µm

G. caesium var. subglobosum


Conidia 3.0-5.0 x 1.5-2.0 µm

G. caesium var. chloridioides





Conidiophores usually bearing clusters of sterile setae above; conidia olivaceous, short-ellipsoidal, 3.5-4.5 x 2.2-2.5(-3.5) µm; chlamydospores rare in culture, mostly one-celled

G. macrocladum


Conidiophores usually without sterile setae; conidia hyaline or very pale, cylindrical or ellipsoidal; chlamydospores often abundant in culture, many-celled






Conidia yellow-green or pale green in mass, cylindrical with rounded ends, 2.8-4.0 x 1.2-2.0 µm

G. chlamydosporium var. chlamydosporium


Conidia white in mass, ellipsoidal, 4-5 x 2.0-2.5 µm

G. chlamydosporium var. simile


[p. 79]


1a. Chaetosphaeria inaequalis (Grove) W. Gams & Hol.-Jech. comb. nov. - Fig. 46a, b.

Eriosphaeria inaequalis Grove - J. Bot., Lond. 24: 132. 1886 = Lasiosphaeria inaequalis (Grove) Massee - Grevillea 16: 37. 1887 = Melanopsammella inaequalis (Grove) Höhn - Annls mycol. 17: 121. 1919 = Trichosphaerella inaequalis (Grove) E. Mull. apud Muller & v. Arx - Beitr. Krypt.-Fl. Schweiz 11(2): 575. 1962.


fig. 46

Fig. 46
Chaetosphaeria inaequalis, a. asci, ascospores and conidiophore of Gonytrichum caesium var. caesium, from holotype specimen; b. PRM 796684, asci and ascospores, x 1000, section through perithecium, x 300; C. PRM 796688, conidiophore with ramification, x 1000 [p. 80]

Detailed descriptions in von Höhnel (1919) and Müller and von Arx (1962: 575).
Perithecia broadly attached to the substratum, black, globose, ostiole slightly raised, smooth-walled, except for a few investing vegetative hyphae and conidiophores, 150-225 µm diam; wall 12-20(-25) µm thick, consisting of numerous layers of dark, thick-walled cells, textura epidermoidea. Asci cylindrical, thin-walled, 40-70 x 4-5 µm, 8-spored. Ascospores obliquely monostichous, fusiform, when young with a thick, refringent, chromophilic septum, hyaline, roughened, the cells separating at an early stage within the ascus, part spores 2.5-3.5(-4.5) x 2.0-2.5 µm, the upper slightly shorter. Paraphyses indistinct. Conidial state Gonytrichum caesium.

Material examined

Eriosphaeria inaequalis Grove, holotype, on dead sticks, Barnt Green, Warwickshire, U.K., 29 Aug. 1885 (K).
PRM 796676, Quercus cerris, foot of hill Rataj, south of Ipel. Ulany, Slov., 10 Aug. 1975.
PRM 796677, Populus sp. and 796678, Salix sp., north-west of Tekovský Hrádok nr. Levice, Slov., 5 Aug. 1975.
PRM 796689, Fagus sylvatica, Forest Diana near Tachov, Boh., 28 Oct. 1970.
PRM 796683 and 796691, Carpinus betulus, Rocky hill Týřovické skály nr. Rakovník, Boh., 26 Apr. 1966 and 28 Sept. 1967.
PRM 796682, Populus nigra, Úpor Forest nr. Mĕlník, Boh., 21 Oct. 1971.
PRM 796684, Carpinus betulus, Kódské polesí Forest near Beroun, Boh., 24 Sept. 1967. PRM 796686, Alnus glutinosa, Hvĕzda Forest nr. Blatná, Boh., 30 Sept. 1975.
PRM 796685, 796687, Abies alba, Dúbrava Forest near Ponická Huta, Slov., 20 Sept. 1971. PRM 796688, 796709, 796714, Populus nigra, 796712, Ulmus effusa, Úpor Forest nr. Mĕlník, Boh., 24 June 1976.
All specimens with conidia of var. caesium, only PRM 796685 and 796687 with var. chloridioides.


1b. Gonytrichum caesium C.G. & F. Nees ex Leman var. caesium - Fig. 46a, c

Gonytrichum caesium C. G. & F. Nees - Nova Acta Acad. Leop.-Carol. Nat. Cur. 9: 244. 1818; ex Leman - Dict. Sci. nat., Paris 19: 194. 1821 = Myxotrichum caesium (C. G. & F. Nees ex Leman) Fr. - Syst. mycol. 3: 348. 1832.
Conoplea cinerea Pers. - Mycol. eur. 1: 12. 1822 = Dematium cinereum (Pers.) Sacc. Fung. 10: 590. 1892.
Gonytrichum fuscum Corda - Icon. Fung. 1: II. 1837.
Gonytrichum erectum Preuss - Linnaea 24: 110. 1851.
? Gonytrichum rubrum Pat. apud Pat. & Lagerh. - Bull. Soc. mycol. Fr. 7: 181. 1891. Synonymy after Hughes (1951b).

Detailed description in Hughes (1951b).
Colonies reaching 8-20 mm diam in 14 days at 20-22°, light grey, in fresh isolates abundantly sporulating, like on the natural substrate. Vegetative hyphae ± hyaline, 2-3 µm wide. Conidiophores on the natural substrate forming a velvety layer, often white and powdery due to the [p. 81] conidia. Conidiophore stipes dark brown, thick-walled, 2.5-3.0 µm wide, repeatedly ± dichotomously branched; phialides arising in dense clusters from groups of short, swollen collar hyphae, subhyaline to light brown, soon collapsing, awl- or flask-shaped, the larger ones cylindrical with tapering tip, 6-10(-20) x 2.0-3.0 µm, tapering to 0.6-1.2 µm, collarette shallow and hardly widening. Conidia arising from multiple conidiogenous loci, forming whitish heads or cirrhi, cylindrical with a slightly apiculate base, hyaline, 1-guttulate, (2.0-)2.5-3.0(-3.5) x 1.o-1.5 µm. Simple Chloridium-like conidophores occur uncommonly on the natural substrate and more frequently in culture, with the conidiogenous meristem usually protruding beyond the collarette, 2.0-2.5 µm wide. Chlamydospores absent.

Additional material examined

Living cultures
CBS 230.74, ex dead twigs of Acer pseudoplatanus, nr. Buren, Isle of Ameland, Netherlands, 28 Oct. 1973.
CBS 696.74 ex PRM 796691, ex Carpinus betulus, Týřovické skály nr. Rakovník Boh., V. Holubová-Jechová Apr. 1966.

Herbarium specimens
Gonytrichum erectum Preuss, Herb. Preuss No. 363 and 364; 364 labeled auf Himbeerstengel’ is considered as holotypus (No. 362 represents var. subglobosum).
J. Poelt No. 988, on a garden plank, Pöching Kr. Starnberg, F.R.G., 5 Sept. 1965 (B). Numerous specimens were examined from Czechoslovakia: PRM 796679, 796690, 796692, 796693, 796694, 796695, 796696, 796680, 796681, 796697, 796698, 796699, 796700, 796701, 796702, 796703, 796704, 796705, 796706, 796707, 796708, 796710, 796711, 796713, 796715, 796716. The species is common in Czechoslovakia on dead wood and bark of Abies alba, Alnus glutinosa, Betula verrucosa, Carpinus betulus, Evonymus sp., Fagus sylvatica, Fraxinus excelsior, Pinus sylvestris, Populus nigra, Quercus cerris, Qu. petraea, Robinia pseudacacia, Salix sp., and Ulmus species.


1c. Gonytrichum caesium C. G. & F. Nees ex Leman var. subglobosum W. Gams & Hol.- Jech. var. nov. - Fig. 47d.

Differt a var. caesium phialidibus longioribus (10-30 µm), pallide brunneis, cylindricis, apice angustatis, ternis ad senis acervatis, et conidiis subglobosis, 3.0-3.3 x 2.0-2.5 µm.
Holotypus PRM 796727, ad ramum putridum deiectum Tiliae cordatae, in horto publico ad castellum Krásný Dvůr dictum haud procul a Podbořany in Bohemia, 3 Jul. 1972.

Phialides arising from the branching nodes in groups of 3-6, pale brown, darker than in var. caesium, 10-30 µm long, sometimes integrated in septate lateral branches to 50 µm long and 1- or 2-septate. Conidia forming whitish cirrhi, subglobose, hyaline, or very pale pigmented, smooth-walled, 3.0-3.3(-3.8) x 2.0-2.5 µm, length/width ratio 1.2-1.5. Chloridium-like conidiophores also present on the type specimen. [p. 82]

Material examined

The type specimen PRM 796727 contains also Chaetosphaeria inaequalis and C. caesium var. caesium.
Sub Gonytrichum erectum Preuss, Herb. Preuss No. 362 (B).


fig. 47

Fig. 47
Gonytrichum caesium var. chloridioides, a. PRM 796720, x 1000, b. CBS 129.72, branching conidiophore in culture, x 500, detail and conidia x 1000, c. CBS 239.75A, Chloridium-like conidiophores, x 1000, d. Chloridium caesium var. subglobosum, PRM 796727, x 1000.


1d. Gonytrichum caesium C. G. & F. Nees ex Leman var. chloridioides W. Gams & Hol.-Jech. var. nov. - Fig. 47a-c

? Piminella castaneae Arnaud - Bull. trimest. Soc. mycol. Fr. 69: 281. 1953, nom. inval. (Art. 36).

Differt a var. caesium phialidibus longioribus (10-20 µm), pallide brunneis, cylindricis, apice angustatis, ternis ad senis acervatis, et conidiis maioribus cylindricis, 3.0-5.0 x 1.5-2.0 µm. Conidiophora Chloridii similia vulgo adsunt.
Holotypus PRM 796720, ad ramum putridum Quercus petraeae in silvis inter vicos Holasice et Veverská Bítýška haud procul a Brno in Moravia, 20 Aug. 1974.

Colonies reaching 21 mm diam in 14 days at 20-22°C, grey-brown with a broad hyaline margin. Vegetative hyphae hyaline and 1.0-2.5 µm wide, or brown and to 4.0 µm or more wide. Conidiophore stipes forming a dark brown felt on the natural substrate, dark brown, repeatedly branched, with 2-3 equal branches arising at the nodes and often short collar hyphae giving rise to a succession of 3-6 phialides. Internodes 40-60 µm long, repeatedly septate. Phialides brown, cylindrical, with a strongly tapering tip, 10-20 x 2.5-3.0 µm, tapering to 1.0-1.5 µm wide; collarette inconspicuous. Conidia formed on a protruding meristematic tip from multiple conidiogenous loci, accumulating in slimy whitish heads or cirrhi, hyaline, ellipsoidal, usually 2-guttulate, 3.0-5.0 x 1.5-2.0 µm, length/width ratio 1.7-2.5. Chlamydospores absent.
After several transfers cultures often forming only simple, Chloridium-like conidiophores often arising from pigmented aerial hyphae with a wider collarette (2.0-2.5 µm) and a protruding conidiogenous meristem beset with numerous conidia. Chloridium-like conidiophores usually also present on the natural substrate.

Material examined

Living cultures
CBS 129.72, ex humus layer of forest soil, Meerdinkbos nr. Winterswijk, Netherlands, Nov. 1971, B. Söderström Nov. 1971.
CBS 239.75A, ex decomposing petiole of Pteridium aquilinum under Quercus-Corylus coppice on slate soil, Roudsea Wood, Cumbria, G.B., J. C. Frankland via O. Reisinger.

Herbarium specimens
G.L.H. 6951, Quercus robur, Baarnse Bos., Netherlands, 20 July 1965.
PRM 794066, Tilia cordata, Obora Hvĕzda forest nr. Sedlice, Boh., 30 Sept. 1975.
PRM 796717, Carpinus betulus, Dřevíč Forest nr. Beroun, Bob., 17 Oct. 1965.
PRM 796718, Fagus sylvatica, Kákov Mt., south-west of Nepomuk, Bob., 3 Oct. 1975.
PRM 796719, Quercus petraea, Kódské polesí Forest nr. Beroun, Boh., 24 Sept. 1967.
PRM 796721, Quercus petraea, Prešovské vrchy Mts., nr. Ruská Nová Ves, Slov., 16 Aug. 1972.
PRM 796722, Fagus sylvatica, valley of Bučínský potok nr. Jeseník, Rychlebské hory Mts., Mor., 10 Aug. 1971.
PRM 796723, Fagus sylvatica, Mt. Javořina, White Karpathian Mts., Mor., 27 July 1970. [p. 84]
PRM 796724, Betula verrucosa, Mt. Sedlo, České Středohoří Mts., Boh., 8 July 1970.
PRM 796725, Fagus sylvatica, valley of Hučivá Desna, Hrubý Jeseník, Mts., Mor., 6 Aug. 1971.
PRM 796726, Quercus petraea, Lánská obora Forest nr. Rakovník, Boh., 12 June 1976.


Apart from the synonyms listed by Hughes (1951b) the synonymy of G. erectum Preuss could be ascertained from the study of type material. G. rubrum Pat., which is reported to have reddish colonies due to conidia fulventia’ 3-4 x 2.5 µm, however, remains doubtful, since no material is preserved in either FH or PC.
The connection between Chaetosphaeria inaequalis and Gonytrichum caesium var. caesium was established by means of ascospore cultures by E.W. Mason (Hughes, 1951b). The variability in Chaet. inaequalis and its conidial states is comparable to that of Chaet. vermicularioides. Intermediates occurring between the three conidial groups here distinguished make their determination difficult (such specimens have mostly been listed under var. caesium). The connection with indistinguishable perfect states is very probable for var. caesium and var. chloridioides. In culture the differences between these varieties can be very marked; while the former produces normal branched conidiophores, the latter can completely lose this capacity and appear Chloridium-like, the only difference being the origin of the conidiophores from often pigmented aerial hyphae and the absence of chlamydospores. In Chloridium virescens and its varieties the conidiophores always arise from the substrate hyphae.
No material of Piminella castaneae is preserved in PC (Nicot and Charpentié, 1971). Arnaud’s original notes and the published drawing strongly suggest the solitary Chloridium-like conidiophores of var. chloridioides arising from pigmented aerial hyphae.
From the distribution in Czechoslovakia it appears that the varieties caesium and chloridioides are ecologically different: Var. caesium occurs mainly in warmer lowland river-side forests or in hilly areas, while var. chloridioides prefers higher altitudes from hilly regions to mountains and areas of temperature inversion in stream valleys. Forms intermediate between the two varieties occur mainly in hilly areas.


2. Gonytrichum mirabile Hol.-Jech. sp. nov. - Fig. 48

Coloniae effusae, pulvillosae, griseo-brunneae. Conidiophora brunnea, crassitunicata, septata, 3.0-3.5 µm lata, irregulariter ramosa et anastomosantia, internodia 20-80 µm et longiora, apices steriles. Phialides plerumque e partibus rectis conidiophororum ipsae vel raro e cellulis collaribus oriuntur, binae ad ternae, dilute brunneae, 5-11 µm longae, basi inflatae et curvatae, 3-5 µm latae, ad collare 1.0-1.5 µm latum angustatae. Conidia in cirrhis albidis connexa, hyalina, ellipsoidea, levia, 3.3-4.0 x 1.7-2.2 µm.
Holotypus PRM 795928, ad ramum putridum Fraxini excelsioris, in silva Úpor dicta prope Mĕlník in Bohemia, 24 Jun. 1976. [p. 85]

Colonies in vitro reaching 18-22 mm diam in 10 days at 20-22°C, olivaceous-grey, velvety, with olivaceous-black margin and reverse, abundantly sporulating like on the natural substrate. Colonies on the natural substrate effuse, pulvinate, grey-brown. Conidiophore stipes rather light brown, thick-walled, septate, 3.0-3.5 µm wide, irregularly branched and anastomosing, intricately entangled and very difficult to tease apart; lateral branches often bent around the stipe at their origin; internodes 20-80 µm long and repeatedly septate; conidiophore tips sterile, blunt, slightly less pigmented. Phialides arising from the straight portions of the conidiophore stipes, mostly directly below septa, with a curved swollen base, or rarely from a collar hypha delimited from the phialide by a septum; phialides mostly in groups of 2-3, pale brown, much lighter than the conidiophore stipes, 5-11 µm long, inflated at the base to 3-5 µm and tapering to the 1.0-1.5 µm wide collarette. Conidia accumulating in whitish cirrhi, hyaline, ellipsoidal, smooth-walled, 1-guttulate, 3.3-4.0 x 1.7-2.2 µm. Chlamydospores absent.

Culture examined

CBS 408.76, ex holotype PRM 795928, ex decaying log of Fraxinus excelsior, Úpor Forest nr. Melnik, Boh., V. Holubová-Jechová 24 June 1976.


G. mirabile is unique in Gonytrichum because it generally lacks collar hyphae, but the phialides take over their basally curved appearance and surround the conidiophores stalks like pairs of claws, one of which is connected to the stalk.


fig. 48

Fig. 48
Gonytrichum mirabile, a. from the natural substrate, PRM 795928, b. from culture, CBS 408.76, x 1000. [p. 86]


3a. Chaetosphaeria chloroconia W. Gams & Hol.-Jech. sp. nov. - Fig. 49b

Perithecia globosa, atra, levia, 120-200(-250) µm diametro, paries 8-15 µm crassus, 1-3 stratis cellularum hyalinarum circumdatus. Asci 45-70 x 3.5-4.0 µm, tenuitunicati. Ascosporae fusiformes, bicellulares, in ascis dilabentes, leves, cellulis divisis 3.0-4.0 x 2.2-2.7 µm. Status conidialis Gonytrichum chlamydosporium.
Holotypus PRM 796736, ad ramum putridum deiectum Populi sp., in silva madida sept.-occid. a pago Tekový Hrádok merid.-occid. ab oppido Levice, in Slovakia, 5 Aug. 1975.

Perithecia broadly attached to the substratum, globose, ostiole slightly raised, black, smooth-walled, 120-200(-250) µm diam; wall 8-15 µm thick, consisting of several layers of dark cells and 1-3 superficial layers of hyaline cells. Asci cylindrical, thin-walled, 45-70 x 3.5-4.0 µm. Ascospores monostichous, fusiform, smooth-walled, 2-celled, when young with a thick, chromophilic septum, cells separating at maturity within the ascus (but somewhat less easily than in Chaet. inaequalis), part spores 3.0-4.0 x 2.2-2.7 µm, of almost equal shape and size. Paraphyses emerging beyond the asci, thread-like, c. 1.5 µm wide. Conidial state Gonytrichum chlamydosporium.

Additional material examined

PRM 796737, Angelica sylvestris, Skařina Forest nr. Mikulčice, Mor., 28 Aug. 1971.
PRM 796739, Fraxinus angustifolia, river-side virgin forest Cahnov nr. Lanžhot, Mor., 14 Oct. 1971.


3b. Gonytrichum chlamydosporium Barron & Bhatt var. chlamydosporium - Fig. 49a.

Gonytrichum chlamydosporium Barron & Bhatt - Mycopath. Mycol. appl. 32: 126. 1967.
? Gonytrichum gilvum Rabenh. - Krypt.-Fl. Deutschl. 1: 105. 1844.

Colonies on the natural substrate yellow-green or pale green due to the mass of conidia. Conidiophores similar to those of G. macrocladum, erect, dark brown, but usually without lateral sterile setae (where present these are up to 60 µm long), 100-300 µm tall, 4.0-5.0 µm wide at the base, with a conicial short basal cell, 2.5-3.5 µm wide in the middle and 1.0-1.5 µm in the paler upper part; fertile lower part c. 50-150 µm long, bearing 3-6 whorls of 2-4 phialides each; collar hyphae regularly present, irregularly curved, darker than the phialides. Phialides slender flask-shaped, pale brown, 5-15(-20) x 2.5-3.0(-3.5) µm, tapering to 1.5(-2.0) µm at the collarette. Conidia accumulating in bright green cirrhi, hyaline to slightly pigmented, cylindrical with rounded ends, (2.8-)3.0-4.0 x 1.2-2.0 µm (orig. descr.: 2.0-4.5 x 1.0-2.0 µm). Unbranched conidiophores with terminal sporulation and identical conidia as described above occur in [p. 87] patches besides the normal conidiophores. Chlamydospores have been described by Barron and Bhatt (1967) from cultures as being composed of several cells, brown, thick-walled, 20-30 x 8-12 µm.

All specimens of Chaet. chloroconia examined were associated with this conidial state.


fig. 49

Fig. 49
a. Gonytrichum chlamydosporium var. chlamydosporium, conidiophores and conidia of PRM 796736, b. ascus and ascospores of Chaetosphaeria chloroconia from the same specimen, c. Gonytrichum chlamydosporium var. simile, branching conidiophore PRM 796730 and d. Chloridium-like conidiophores PRM 796738. x 1000. [p. 88]

Additional material examined

PRM 796731, Corylus avellana, Kásovický revír Forest nr. Oparany, Boh., 2 June 1965.
PRM 796732, Populus nigra, Úpor Forest nr. Mĕlník, Boh., 24 June 1976.


3c. Gonytrichum chlamydosporium Barron & Bhatt. var. simile W. Gams & Hol.-Jech. var. nov. - Fig. 49c, d.

Differt a var. chlamydosporium conidiis hyalinis in cirrhis albidis connexis, ellipsoideis, vel modice asymmetricis, (3.0-)4.0-5.0 x 2.0-2.5 µm.
Holotypus PRM 796730, ad truncum putridum Fagi sylvaticae, Montes Hrubý Jeseník, in silva Bučina dicta in declivibus supra casam venatoriam Františkova myslivna, apud Kouty n. Desnou in Moravia, 4 Aug. 1971.

Similar to var. chlamydosporium, conidiophore stipes 280-400 µm tall and 4.0-5.0 µm wide at the base, 2.5-4.0 µm centrally, tapering into a sterile seta 100-200 µm long and 1.0-1.5 µm wide, bearing 4-8 whorls each of 2-6 phialides which arise from curved collar hyphae. Phialides pale brown, 10-20(-30) µm long, 2-3(-3.5) µm wide, tapering to 1.0-1.5 µm at the collarette. Conidia accumulating in whitish cirrhi, hyaline, slightly pigmented when dry, ellipsoidal or slightly asymmetrically curved, (3.0-)4.0-5.0 x 2.0-2.5 µm. Unbranched conidiophores with terminal sporulation and identical conidia as described above occur in patches besides the normal ones.

Additional material examined

PRM 647192, Fagus sylvatica, Mt. Simonka nr. Zlatá Baňa, Slov., 19 July 1964.
PRM 796728, Acer pseudoplatanus, nr. Hadinec, Orlické hory Mts., Boh., 20 July 1969.
PRM 796738, Fagus sylvatica, Bučina Forest nr. Kouty n. Desnou, Hrubý Jesenik Mts., Mor., 4 Aug. 1971.
PRM 796757, Fagus sylvatica, Dranec Forest nr. Nižní Komárnik Slov., 22 July 1964.
PRM 796729, Fagus sylvatica, Mt. Magura nr. Zborov, Nízké Beskydy Mts., Slov., 12 Aug. 1972.


The very different conidial states corroborate and justify the distinction of the similar perfect states of G. caesium and G. chlamydosporium. The main differences between Chaet. inaequalis and Chaet. chloroconia are in the hyaline outermost wall layer of the perithecia in the latter and in the ornamentation of the ascospores in the former species, but on the specimen PRM 796738 of var. simile scarce perithecia with finely verrucose ascospores were observed.
The most remarkable feature of the conidial state is the yellow-green appearance of the conidial cirrhi reminiscent of those of Chloridium virescens var. virescens. This colour was not mentioned by Barron and Bhatt (1967) in their cultures, but otherwise their diagnosis accords well [p. 89] with our observations. As in the case of G. caesium (cf. p. 84) we do not regard the differences in conidial size and shape between these two varieties as sufficient for their segregation as separate species.
No material of G. gilvum is preserved at Herb. B; this species is described as having globose conidia which might apply to var. simile. Gonytrichum luteo-viride Torrend (Broteria, Ser. Bot., 12: 68. 1914; Fungi sel. exs. 283) which might suggest G. chlamydosporium because of the colour, was found to represent probably an Oidiodendron species (S).


fig. 50

Fig. 50
Gonytrichum macrocladum, collar hyphae with phialides and conidia, from the natural substrate, PRM 796733, x 1000


4. Gonytrichum macrocladum (Sacc.) Hughes - Fig. 50.

Chaetopsis macroclada Sacc. - Michelia 1: 79. 1877 = Mesobotrys macroclada (Sacc.) Sacc. - Michelia 2: 27. 1880 = Gonytrichum macrocladum (Sacc.) Hughes - Trans. Br. mycol. Soc. 34: 565. 1951.
Chaetopsis fusca Corda - Icon. Fung. 1: 18. 1880 = Mesobotrys fusca (Corda) Sacc. - Michelia 2: 27. 1880.
Gonytrichum macrocladum (Sacc.) Hughes var. terricola Gambogi - Nuovo G. bot. ital. 103: 33. 1969

Detailed descriptions are to be found in Hughes (1951b: 565) and Swart (1959).
Colonies rather slow-growing, reaching 18 mm diam in 10-14 days at 20-22°C, grey-brown at first but becoming grey-olivaceous to dull green with age, velvety, with a broad hyaline margin, abundantly sporulating like on the natural substrate. Conidiophores crowded, arising singly, consisting of a straight erect stipe, 350-400 µm long, dark brown, septate, 6-8 µm wide at the base, 4.5-5.0 µm centrally, 2.0-2.5 above the ramifications, ending in a paler sterile subulate seta or a sporulating phialide; in the lower half beset with 4-8 whorls of phialides below each septum at 20-25 µm intervals. Phialides arising from irregularly curved, pigmented collar hyphae in groups of 1-6, slender flask-shaped, pale brown, 10-20 x 3.0-4.0 tapering to 1.5-2.0 µm at the collarette; occasionally percurrently proliferating. Conidia accumulating in dark cirrhi, olivaceous, ellipsoidal to oval, or one side more strongly curved than the other, [p. 90] slightly apiculate at the base, 1-guttulate, 3.5-4.5 x 2.2-2.5 µm. In the upper part of the conidiophores one to three whorls of 3-4 sterile lateral pale brown to subhyaline setae, up to 100 µm long, often present. Scarce chlamydospores were observed by several authors in fresh isolates. Simple Chloridium-like conidiophores may be predominant in some strains, but still produce olivaceous conidial cirrhi.

Material examined

Living cultures
CBS 201.55, ex soil, Ancaster, Ont., Canada, comm. R. G. Atkinson, 1955.
CBS 253.59 = DAOM 57699, ex soil on Pisum roots, isol. Rene Grete, det. S. J. Hughes.
CBS 195.60, ex soil, Frankenwald Exp. Stn nr. Johannesburg, S.-Africa, comm. H. J. Swart.
CBS 875.68 = ATCC 16306 and some other strains ex soil under wheat fields, Kiel-Kitzeberg, F.R.G., W. Gams, 1964.
CBS 861.73, ex volcanic ash soil (trumao) in Chile, comm. J. Grinbergs as 502.73.
CBS 352.76, ex underground parts of sugar cane, Taiwan, comm. T. Watanabe as G. chlamydosporium (X 61-353).
CBS 419.76, ex PRM 796734, ex Populus nigra, Úpor Forest nr. Mĕlník, Boh., 24 June 1976.

Herbarium specimens
PRM 796733, Quercus robur, forest north-east of Mudronovo nr. Komárno Slov., 18 Sept. 1975.
PRM 796734 and 796735, Populus nigra, Úpor Forest nr. Mĕlník, Boh., 24 June 1976.


13. Phaeostalagmus W. Gams gen. nov.

Genus Hyphomycetum dematiaceorum. Coloniae fuscae, tardissime crescentes. Conidiophora fusca, erecta, mesotone vel acrotone, semel vel compluries verticillata; phialides lageniformes, fuscae, collari divergente terminatae; conidia ovalia, simplicia, hyalina.
Species typica Stachylidium cyclosporum Grove.

Colonies darkly pigmented and very slow growing. Conidiophore stipes dark brown, erect, with mesotonous or acrotonous branching, giving rise either to whorls of phialides or a series of branches which eventually terminate in verticillate phialides; phialides flask-shaped, pigmented, with a pronounced flaring collarette; conidia one-celled, ellipsoidal, hyaline.

Two characteristic dematiaceous species hitherto placed in Verticillium (Hughes, 1951a; Ellis, 1971) deserve generic distinction. Verticillium Nees ex Wallr. is not yet clearly delimited, but it contains white or pigmented species which sometimes form differentiated, brown conidiophores [p. 91] (e.g. V. albo-atrum Reinke & Berth., V. theobromae (Turc.) Mason & Hughes), but these species still have a mucedinaceous character in their hyaline, spreading vegetative hyphae. The two dematiaceous species, on the other hand, have darkly pigmented, very slow-growing vegetative hyphae and even darker conidiophores. The phialides have a distinct, dark, flaring collarette which distinguishes them from the otherwise similar Stachylidium.

Key to the species


Conidiophores with a very dark main axis bearing several whorls of phialides, usually without further ramification; conidia 3.0-4.0 1.2-1.8 µm

P. tenuissimus


Conidiophore axis bearing several branches which are repeatedly verticillately branched again; conidia 1.5-3.0 x 1.0-2.0 µm

P. cyclosporus


1. Phaeostalagmus cyclosporus (Grove) W. Gams comb. nov. - Fig. 51.

Stachylidium cyclosporum Grove - J. Bot., Lond. 22: 199. 1884 = Verticillium cyclosporum (Grove) Mason & Hughes apud Hughes - Mycol. Pap. 45: 19. 1951.
Stachylidium ramosum Arnaud - Bull. trimest. Soc. mycol. Fr. 69: 281. 1953, nom. inval. (Art. 36).

Detailed description in Hughes (1951a: 19).
Colonies in vitro reaching 2-6 mm diam after 14 days at 20-22°C, grey to olivaceous black, later often iridescent, reverse light or dark grey; vegetative hyphae hyaline, 0.7-1.5 (-3.0) µm wide, to dark brown in the colony centre, often encrusted, partly aggregated in stroma-like crusts. Young cultures often forming simple phialides or short, atypical, Phialophora-like conidiophores; normal branched conidiophores appearing after 2-3 weeks, but less complex than on the natural substrate, where they are to 160 µm or more tall, with a dark brown, thick-walled stipe, 2.5-3.0 µm wide, and repeatedly verticillate above. Phialides almost hyaline, their supporting cells with intermediate pigmentation, all elements ± con-stricted at the base. Phialides flask-shaped, with a long narrow neck and darker flaring collarette, 0.5-1.0 µm deep, often finely warted, 6-12 x 1.7-2.5 µm, tip 0.7-1.0 µm wide. Conidia aggregating in slimy heads at the apex of the phialides, ellipsoidal, ovoidal to cylindrical (variable between strains), hyaline, 1.5-3.0 x 1.0-2.0 µm (in culture usually slightly bigger than on the natural substrate). Sometimes also sporulating within the agar with lighter conidiophores and subglobose conidia, 1.5-2.2 µm diam. Chlamydospores absent. [p. 92]

Material examined

Living cultures
CBS 221.65 = OAC 10290, ex soil, Ontario, Canada, G. L. Barron, 1964.
CBS 663.70, ex decaying wood, Schovenhorst nr. Putten, Netherlands, J. A. Stalpers, 4 June 1970.
CBS 241.74, ex Pinus sylvestris, Meervelder Bos nr. Uddel, Netherlands, Jan. 1974.
CBS 242.74, ex Alnus glutinosa, nr. Kortenhoef, Netherlands, Feb. 1974.
CBS 312.75, ex Corylus avellana, Park Broekhuizen nr. Leersum, Netherlands, March 1975.

Herbarium specimens
Verticillium cyclosporum folder in K as listed by Hughes (1951a); the holotype collection made at King’s Heath (S.)’ is not preserved in K; the second collection made by Grove near Bear Lane, Smethwick, near Three Shire Oaks on the border of Worcester, Stafford and Warwick, 25 Apr. 1885, is rather meagre; a rich specimen on a dead trunk, Shustoke, W. B. Grove, 15 May 1886, is designated as neotype. - In addition: on wood, Queen’s Cottage, Kew Gardens, S.M.W.; Betula ? sp., Staverton Thicks, Suffolk, A. H. S. Brown, 24 May 1956; Acer pseudoplatanus, Isle of Bossa, R. W. G. Dennis 20 Apr. 1961; Ulmus sp., Morlwood, Gloucestershire, R. W. G. Dennis, 9 Aug. 1962; Betula sp., Isle of Mull, R. W. G. Dennis, 4 Aug. 1968; Alnus sp., Burpham, W. Sussex, D. A. Reid, 20 Feb. 1971; Ulex sp., Warden’s Hut, West Sussex, D. A. Reid, 16 Feb. 1974; Ulex europaeus, Isle of Lewis, R. W. G. Dennis, 7 May 1974; Castanea sativa, Erddey Park, Denbighshire, J. T. Palmer (329489).


fig. 51

Fig. 51
Phaeostalagmus cyclosporus, a. CBS 242.74 and b. CBS 663.70, from the natural substrate, c. CBS 663.70, in 10 day-old culture. x 1000


fig. 52

Fig. 52
Phaeostalagmus tenuissimus, CBS 917.73, on the natural substrate. x 1000. [p. 93]

No holotype material of V. cyclosporum is in existence, but the secondary collections made by Grove all are identical with the species here described and the specimen from 15 May 1886 is designated here as neotype. Mason and Hughes in Hughes (1951a) placed the species in Verticillium with some hesitation and were obviously aware of the heterogeneity of the genus. They suggested that the name cyclosporum meant the conidial heads and not the single conidia.


2. Phaeostalagmus tenuissimus (Corda) W. Gams comb. nov. - Fig. 52.

Verticillium tenuissimum Corda - Icon. Fung. 1: 20. 1837.
Verticillium apicale Berk. & Br. - Ann. Mag. nat. Hist., Ser. 2, 7: 101. 1851 (Notices No. 531) = Verticicladium apicale (Berk. & Br.) Sacc. - Syll. Fung.4: 328. ,886.
? Stachylidium extorre Sacc. - Michelia 1: 84. 1877.
Synonyms after Hughes (1951a).

Detailed description in Hughes (1951a).
Colonies in vitro reaching 4 mm diam in 14 days at 20-22°C, light grey-brown, reverse light grey. Vegetative hyphae hyaline, 0.8-1.5 µm wide, brown in the centre, almost stromatic, to 2.5 µm or more wide; slow to sporulate; after 14 days mostly with simple phialides only. Fully developed conidiophores on the natural substrate dark brown, erect, generally to 200 µm long, 4-6 µm wide near the base, with numerous septa, bearing whorls of phialides in the upper part below the septa. Phialides (sub)hyaline, flask-shaped, 10-13 x 3-4 µm, the long, narrow tip 0.7-1.2 µm wide, with a slightly darker, flaring collarette, 0.5-1.0 µm deep. Conidia aggregating in slimy heads, cylindrical with rounded ends, (sub)hyaline, (2.5-)3.0-4.0 x 1.2-1.8 µm. Chlamydospores absent.

Material examined

CBS 917.73, ex decaying wood, National Park Hoge Veluwe, Netherlands, Oct. 1973.
PRM 515139, holotype, on rotten acorns of Quercus, Tuchomerice near Prague.


The simple conidiophores bearing only simple whorls of phialides are the most important feature distinguishing P. tenuissimus from P. cyclosporus. The conidia are longer and more regularly cylindrical than in the former species. [p. 94]



We thank all the individuals and curators of herbaria mentioned in the text for their kind collaboration, particularly Prof. Dr. G. L. Hennnebert, Louvain-la-Neuve, who contributed many specimens of his personal collection for this study and suggested several improvements of the manuscript. We are indepted to Dr. M. B. Ellis for his views on generic delimitation of the imperfect and Dr. J. A. von Arx for suggesting the genus concept of the perfect states. Dr. D. L. Hawksworth kindly provided the necessary linguistic corrections; Miss M. Nieuwstad is acknowledged for some technical assistance.


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