Studies in Mycology, No. 6

10 June 1974

 

Paecilomyces and some allied Hyphomycetes

 

 

R. A. SAMSON

Centraalbureau voor Schimmelcultures, Baarn

 

 

Summary

 

The genus Paecilomyces (type species: P. variotii Bainier) is monographed and re­defined. The delimitation of the genus and its relationship to other genera are discussed; the perfect states belong to Byssochlamys Westl., Talaromyces C. R. Benjamin and Thermoascus Miehe. A key to the described genera and some relatives is given. The mor­phological characters of Paecilomyces are described; a subdivision into two sections is proposed: In the Sect. Paecilomyces some mesophilic to thermophilic species are ac­comodated with yellow-brown to brown colonies; several species are accompanied by a perfect state. Mesophilic species with white or other bright colonies are accomodated in the new Sect. Isarioidea; among them are several entomogenous taxa. 31 species are keyed out, described and illustrated. The descriptions are mainly based on their morpho­logical characters in pure culture. Paecilomyces aerugineus, P. ghanensis, P. ramosus, P. coleopterorum, P. reniformis and P. niphetodes are described as new species. New com­binations for 9 species, originally described in Isaria, Spicaria, Verticillium, Penicillium and Cylindrodendrum are proposed. The genus Mariannaea Arnaud is re-introduced and validated. Penicillium elegans Corda is transferred to this genus; a new variety punicea of this species and a new species M. camptospora are described and illustrated. The genus Nomuraea Maublanc is retained for two entomogenous species, Spicaria rileyi (Farlow) Charles and Isaria atypicola Yasuda. Paecilomyces berolinensis Olįh & Ackermann and P. herbarum Brown & Smith are accomodated in the genus Septofusidium W. Gams. The concept of this genus is briefly discussed. The new genus Gabarnaudia Samson & W. Gams is proposed to accomodate Oospora betae Delacr. and the conidial state of Sphaeronae­mella fimicola Marchal. Gabarnaudia tholispora Arnaud ex Samson & W. Gams and G. humicola Samson & W. Gams (ascigerous state: Sphaeronaemella humicola Samson & W. Gams) are described as new species. Doubtful and excluded species originally described in Paecilomyces and Spicaria are listed and two new combinations, Verticillium sub­fasciculatum (Petch) Samson & W. Gams (= Oospora subfasciculata Petch) and Sco­pulariopsis parva (Brown & Smith) Samson (= Paecilomyces parvus Brown & Smith) are proposed.

 

 

1. Introduction

 

The genus Paecilomyces was introduced by Bainier (1907), who described it as being closely related to Penicillium Link ex Fr., but differing in the absence of green coloured colonies and by the short cylindrical phialides which taper into long necks. Previously, similar species were often placed [p. 2] in Spicaria Harting (1846) (type species S. solani = Gliocladium solani (Harting) Petch), a genus mis-interpreted by Harz (1871), who character­ized the species by verticillate conidiophores and catenate conidia. Saccardo (1881) popularized Harz's interpretation, and the conception of Spicaria became confused. Hughes (1951) and Brown and Smith (1957) suggested that the name Spicaria is best forgotten, and transferred several species to Paecilomyces. After Paecilomyces had been little known for many years, Brown and Smith (1957) monographed the genus. They re­cognized the occurrence of similar conidiogenous structures as in the type species, Paecilomyces variotii, in some species described in Isaria and Spicaria, and transferred them to Paecilomyces. One of these species is Isaria farinosa (Holm ex S. F. Gray) Fr. (1832). Clements and Shear (1931), Mains (1955) and Morris (1963) considered this species as the lectotype of the genus Isaria Pers. ex Fr. This choice would mean that, because of priority, Isaria should be used for all Paecilomyces species. Petch (1934) and de Hoog (1972) discussed the choice of I. farinosa and other species as lectotype. Von Arx (1970) and subsequently de Hoog (1972) chose for Isaria felina DC. ex Fr. in order to provide a generic name for synnematous Beauveria-like fungi.

Paecilomyces as conceived by Brown and Smith (1957) contains 23 species; some of them seem to fall beyond the scope of the genus, because of deviating conidiogenous structures. Paecilomyces fusisporus Saksena (1953) has partly brown, warty conidiophores and often laterally prolifer­ating flask-shaped phialides; it has therefore been placed in the genus Acrophialophora (Samson and Tariq Mahmood, 1970). The phialides of another species, Paecilomyces elegans (Corda) Mason & Hughes, are almost awl-shaped, producing conidia in imbricate chains or in slimy heads; the species is therefore transferred to the genus Mariannaea Arnaud (1952). Paecilomyces herbarum Brown and Smith and P. berolinensis Olįh & Ackermann (1969) have basitonously verticillate conidiophores and awl-shaped phialides. They are regarded as belonging to the genus Septofusidium W. Gams.

The generic concept of Paecilomyces was broadened by Onions and Barron (1967), who included 10 monophialidic species, which have ortho­tropic awl-shaped phialides. Gams (1971b) transferred these monophialidic species, with the exception of Paecilomyces inflatus, to Acremonium. Subramanian (1972) placed some of these species together with Gliomastix murorum in a new genus, Sagrahamala.

Brown and Smith (1957) suggested that Spicaria rileyi (Farlow) Charles (= Nomuraea prasina Maublanc) should probably be excluded from Paecilomyces. This entomogenous species differs from the typical Paecilomyces species by the densely verticillate phialides which have a very short neck; for this species and the related Isaria atypicola Yasuda, the genus Nomuraea Maublanc (1903) should be retained.

In this paper 16 species of the genus Paecilomyces are described on account of the morphology and the structure of the phialides and conidiophores. In addition 6 species are described as new and for 9 species new [p. 3] combinations are proposed. The genera Mariannaea (2 species and one variety), Nomuraea (2 species), Septofusidium (3 species) and Gabarnaudia (4 species) are also revised.

 

 

2. Material and methods

 

This study is mainly based on living cultures maintained in the CBS col­lection. Numerous cultures were obtained from other culture collections, especially from NRRL (Peoria, USA) through the kindness of Miss Dorothy Fennell. The abbrevations indicating the origin of the cultures are given in the 28th edition of the List of Cultures (Centraalbureau voor Schimmelcultures, 1972). A number of strains were received from private collections or were sent in for identification. Several entomogenous species were collected and isolated by Dr H. C. Evans in cocoa plantations and rainforests in Ghana. Specimens obtained from herbaria are indicated by the abbrevations according to the Index Herbariorum.

For the descriptions of colony appearance and microscopic features, cultures growing on 2% maltextract- and oatmeal-agar were used. Some of the entomogenous fungi were grown on starch-, rice- or mealworm-agar or on boiled rice. Starch-agar was prepared according to Cohen (1967). Rice-agar contains an extract of 200 gr rice (boiled for 2 hours), 20 gr agar and 1000 ml tap water. Mealworm-agar contains an extract of 200 gr ground dried mealworms (Tenebrio molitor) (boiled for 3 hours), 20 gr sucrose, 20 gr agar and 1000 ml tap water (Samson and Evans, 1974). On these media an abundant formation of synnemata and conidia could often be observed.

The cultures of the mesophilic species were grown at 250 C in darkness in an incubator. Some species showed a better sporulation in diffuse daylight than in darkness. The thermophilic species were grown at their optimal temperature (between 30ŗ and 40ŗ C).

Capitalized colour names are according to Ridgway (1912) and Rayner (1970).

The terms indicating the branching of the conidiophores and the structure of the phialides are taken from Gams (1971b).

 

 

3. Delimitation and relationship to other genera

 

The genus Paecilomyces is restricted to species with verticillate conidio­phores bearing divergent whorls of branches and phialides. The phialides have a cylindrical or inflated basal portion and a long distinct neck, and produce one-celled hyaline conidia in basipetal chains. The species compiled in Paecilomyces can be grouped into two sections: [p. 4]

Section Paecilomyces, including the type P. variotii, containing mesophilic, thermotolerant and thermophilic species, often accompanied by a Talaromyces, Byssochlamys or Thermoascus ascigerous state, and colonies with yellow-brown to brownish colours.

Section Isarioidea, containing mesophilic species without ascigerous states, and colonies with purple, pink, green or yellow colours.

 

Paecilomyces elegans (Corda) Mason & Hughes is separated from Paecilomyces, because the phialides are more slender and the conidia are produced in slimy heads or in obliquely imbricate chains. For this and a new species the genus Mariannaea Arnaud is retained and validated.

The genus Nomuraea Maublanc is maintained for Spicaria rileyi (Farlow) Charles, which differs from Paecilomyces by a green or purple colour of the colony and by verticillate conidiophores forming dense clusters of branches and phialides below the septa; the phialides have a very short neck.

Acrophialophora Edward is distinguished from Paecilomyces on account of the brown warty conidiophores and the often proliferating phialides.

Paecilomyces berolinensis and P. herbarum differ from all other Paecilomyces species by the basitonously branched conidiophores with rather slender phialides. The two species are classified in Septofusidium W. Gams.

Oospora betae Delacr., Oospora tholispora Arnaud and the conidial state of Sphaeronaemella fimicola Marchal are classified in the new genus Gabarnaudia Samson & W. Gams. This genus differs from Paecilomyces by the large, usually basitonously branched conidiophores with long awl-shaped phialides, which are often integrated in septate branches. The large, elongate conidia are produced in chains, which partially may slime down.

Paecilomyces is distinguished from Acremonium Link ex Fr. by verticil­late conidiophores and flask-shaped phialides. Paecilomyces inflatus (Burn­side) Carmichael can be regarded as an intermediate form, because the phialides are often solitary and orthotropic; the inflated basal portion and the distinct neck classify it, however, in Paecilomyces. Gams (1971b) considered Acremonium vitellinum, A. bacillisporum, A. fusidioides and A. ochraceum as intermediate species because of the sometimes verticillate conidiophores. The species have awl-shaped phialides and are better placed in Acremonium.

The generic distinction between Paecilomyces and Penicillium is difficult. According to Brown and Smith (1957), Paecilomyces is characterized by the shape of the phialides, the irregular or verticillate branching of the conidiophores and the divergent conidial chains; true green conidial pigments are absent; another reason for the delimitation should be that Byssochlamys is the perfect state of Paecilomyces, while Eupenicillium (= Carpenteles) and Talaromyces are those of Penicillium. Stolk and Samson (1971, 1972) have shown, however, that species of Talaromyces can have either a Paecilomyces or a Penicillium conidial state.

With regard to the shape of the phialides the genus Penicillium is heterogeneous, and it might seem justified to split it into a number of [p. 5] genera. However, other morphological characters, such as the densely peni­cillate phialides and the usually green colour, are consistently combined, so that the present concept of the genus Penicillium is best maintained. If Paecilomyces was included in Penicillium, the genus would grow out into a very large and even more heterogeneous complex. Biourge (1923) considered this idea and transferred all species of Isaria and Spicaria, which he considered to have phialides, to Penicillium.

Paecilomyces can be separated from Penicillium by its phialides which taper into a long distinct neck and by its divergent or tangled conidial chains. The phialides are borne on verticillate or irregularly branched conidiophores; in Penicillium the phialides are usually flask-shaped with a distinct but short neck and they are arranged in penicillate clusters. Other delimiting characters are the presence of chlamydospores in several species of Paecilomyces, whereas these structures are absent in Penicillium.

Penicillium lilacinum, placed together with Spicaria violacea (= Paecilomyces marquandii) in a special series of Penicillium by Raper and Thom (1949) has phialides with a long slender neck borne on stiff verticillate conidiophores and a purple colony colour. The two species can therefore better be classified in Paecilomyces (section Isarioidea). Some similar but green species of the Penicillium janthinellum-series show a long neck, but the phialides are arranged in monoverticillate clusters on divaricate conidio­phores.

The brown and olive green species of the section Paecilomyces resemble those of the Penicillium cylindrosporum-series (Stolk and Samson, 1972) and Penicillium digitatum Sacc. These species are delimited from Paecilo­myces by the compact penicilli and the shorter neck of the phialides.

 

The genus Akanthomyces Lebert is known as being entomogenous, producing conspicuous synnemata. The phialides are swollen or cylindrical, usually with a distinct neck, resembling the phialides of the species of the sect. Isarioidea. Akanthomyces can be distinguished from Paecilomyces by the arrangement of the phialides in a hymenium-like layer along the outer hyphae of the synnemata. Akanthomyces gracilis forms a link between the two genera (Samson and Evans, 1974).

The slender, flask-shaped phialides and catenate conidia of Nalanthamala Subram. are similar to those produced in Akanthomyces and Paecilomyces. Nalanthamala can be separated from these genera by the formation of sporodochia.

No type material of the type species of Phialocladus Kreisel, P. zsoltii, was designated by Kreisel (1972) and the species is therefore not validly published. The fungus produces wide, irregularly branched conidiophores, solitary flask-shaped phialides and catenate conidia. It resembles the genus Trichoderma from which it differs by the catenate conidia. Phialo­cladus is distinguished from Paecilomyces by the broad, irregularly branch­ed conidiophores.

Barron (1968) stated that the genus Phialotubus Roy & Leelavathy shows a close relationship to Paecilomyces because of the shape of the [p. 6] phialides and the catenate conidia. The only species, P. microsporus, is characterized by cylindrical phialides with a short or no neck and by conidial chains aggregated in columns. Phialotubus shows more similarity to Metarrhizium than to Paecilomyces. It can be separated from that genus by the irregularly shaped conidia, which are covered by a mucilaginous layer.

Although the species of Scopulariopsis have a conidiogenous cell with an annellated, elongating neck, this character is sometimes very difficult to observe. Particularly the species of the Scopulariopsis sphaerospora and S. brumptii-series (Morton and Smith, 1963) produce conidiogenous cells without distinct annellations. An elongation of the conidiogenous cells and the basally truncate conidia indicate that the cell is not phialidic.

The genus Sesquicillium W. Gams is distinct from Paecilomyces by the verticillate conidiophores with flask-shaped phialides and a subterminal conidiogenous cell with a lateral neck (pleurophialide). The conidia are produced in imbricate chains, which are often united in columns.

The phialides of the genus Tolypocladium W. Gams (1971a) consist of a swollen or cylindrical basal portion, tapering into a distinct neck, usually bent away from the main axis of the cell. In these characters it resembles Trichoderma, but the slow-growing colonies are of entirely different structure. The genus differs from Paecilomyces by the production of the conidia in heads. Tolypocladium may be related to the nematode-parasitic genus Harposporium, which usually has curved conidia.

The genus Verticillium Nees ex Wallr. is characterized by verticillate conidiophores, awl-shaped phialides and conidia in slimy heads. Some species of the Section Prostrata (Gams 1971b) have, however, slightly swollen Paecilomyces-like phialides and catenate conidia.

 

 

4. Perfect states

 

Three genera of Ascomycetes with a Paecilomyces conidial state are known. The imperfect states are all classified in the Sect. Paecilomyces. All species are homothallic.

Byssochlamys Westling - Svensk bot. Tidskr. 3: 134. 1909.

Type species: Byssochlamys nivea Westling

 

The genus is characterized by almost naked ascomata in which from croziers globose asci are formed with ellipsoidal, smooth-walled ascospores. The ascoma initials consist of swollen antheridia and coiled ascogonia. All Byssochlamys species have a Paecilomyces conidial state. Stolk and Samson (1971) revised the genus and recognized 3 species: B. nivea, B. fulva and B. zollerniae. The latter species is only known from its type culture, which no longer produces ripe ascomata. [p. 7]

 

Talaromyces C. R. Benjamin - Mycologia 47: 681. 1955.

Type species: Talaromyces flavus (Klöcker) Stolk & Samson

 

Talaromyces is related to Byssochlamys, but differs by its soft ascomata with a distinct covering, by catenate asci and by globose or ellipsoidal, usually ornamented ascospores. The conidial states belong to Penicillium or Paecilomyces. The latter are described by Stolk and Samson (1972) for two species, T. leycettanus and T. byssochlamydoides.

 

Thermoascus Miehe - Die Selbsterhitzung des Heues, p. 70. 1907.

Type species: Thermoascus aurantiacus Miehe

 

The genus is characterized by its thermophilic nature, ascomata with distinct parenchymatous wall and asci produced from croziers. The conidial state may be absent or belongs to Polypaecilum or Paecilomyces. Though the 3 species of the genus have differing conidial states, the morphology of the ascomata, the asci and the ascospores are similar. For these reasons these species are best maintained under one generic name Thermoascus (cf. Apinis and Chesters, 1964; Stolk, 1965; v. Arx, 1970; Fennell, 1973).

 

Other genera

 

In the literature some Isaria and Spicaria species are reported as im­perfect states of the ascomycetous genera Cordyceps and Torrubiella. Mostly the imperfect state was observed in association with the ascomycete on the same host. For a long time Paecilomyces farinosus was considered as the imperfect state of Cordyceps militaris. Cultural studies of Petch (1934) and Kobayasi (1941), however, showed that the conidial state of C. militaris is Verticillium-like (cf. Gams, 1971b).

In his revision of the genus Cordyceps, Kobayasi (1941) discussed the connection between ascomycetous and conidial states and concluded, that it is difficult to find the taxonomic relationship of species merely from the conidial states. Their mutual relationships should be proven by culturing single ascospores and conidia; this however, is very difficult and in some cases impossible to accomplish.

From material of entomogenous fungi collected by Dr H. C. Evans in Ghana, numerous isolates were made from ascospores of Cordyceps and Torrubiella species and associated conidial states. No indication was found, that Paecilomyces conidial states are connected with these Ascomycetes. Some Torrubiella species which were isolated in pure culture developed into a Verticillium state.

Chesters (1934) described Cephalotheca sulphurea with a Paecilomyces-like state. Without cultural studies it is doubted whether these conidial states belong to Paecilomyces. [p. 8]

 

 

5. Key to the described genera and some relatives

 

1a.

Phialides awl-shaped, mostly solitary, arising from undifferentiated hyphae

Acremonium

1b.

Phialides on irregularly branched conidiophores or in whorls on distinctly differentiated conidiophores

2

 

 

 

2a.

Phialides awl-shaped or slenderly flask-shaped

3

2b.

Phialides with inflated basal portion, flask-shaped or cylindrical

9

 

 

 

3a.

Conidiophores with regular verticils, or penicillate clusters of branches and/or phialides

4

3b.

Conidiophores irregularly, often basitonously branched, or phialides solitary or in whorls on prostrate hyphae

6

 

 

 

4a.

Conidiophores brush-like with compacted whorls of phialides, conidial mass forming imbricate columns or globose heads

Clonostachys (Gliocladium)

4b.

Conidiophores verticillately branched with divergent whorls of phialides

5

 

 

 

5a.

Conidia in imbricate chains or sliming down; phialides slenderly flask-shaped

Mariannaea

5b.

Conidia in slimy heads; phialides awl-shaped

Verticillium s. str.

 

 

 

6a.

Phialides solitary or in whorls on prostrate, simple aerial hyphae

Verticillium sect. Prostrata

6b.

Phialides in erect basitonously branched conidiophores

7

 

 

 

7a.

Phialides thin-walled, integrated in septate conidiophore branches; colonies white, rather fast-growing

Gabarnaudia

7b.

Not combining the above characters

8

 

 

 

8a.

Phialides in whorls, rather thick-walled, often rough-walled or ornamented; colonies white to brownish, slow-growing; conidia catenate,cylindrical or fusiform

Septofusidium

8b.

Phialides mostly in irregular conidiophores; colonies in various shades, slow to fast-growing; conidia of various shape, in heads

Acremonium sect. Nectrioidea

 

 

 

9a.

Pleurophialides below the terminal phialides present

Sesquicillium

9b.

Pleurophialides absent

10

 

 

 

10a.

Phialides with a more or less swollen basal portion, tapering abruptly into a long thin neck

11

10b.

Phialides flask-shaped with a short neck, or of different shape

13

 

 

Paecilomyces

11a.

Conidia in chains

 

11b.

Conidia in slimy heads

12

 

 

 

12a.

Conidia mostly curved; nematophagous

Harposporium

12b.

Conidia globose to cylindrical; saprophytes or entomogenous

Tolypocladium

 

 

 

13a.

Conidiophores aggregated in sporodochia or synnemata

14

13b.

Conidiophores not sporodochial, and if synnematous, phialides not in a hymenium-like layer, but verticillate

17

 

 

 

14a.

Conidial chains combined in columns [p. 9]

15

14b.

Conidial chains more or less divergent

16

 

 

 

15a.

Conidia ellipsoidal-cylindrical; entomogenous or saprophytic

Metarrhizium

15b.

Conidia not ellipsoidal, covered by a mucilaginous sheat; saprophytic

Phialotubus

 

 

 

16a.

Sporodochia present; not entomogenous

Nalanthamala

16b.

Synnemata present, with phialides in a hymenium-like layer; entomogenous

Akanthomyces

 

 

 

17a.

Conidiophores penicillate

Penicillium

17b.

Conidiophores not penicillate, but verticillate or irregularly branched

18

 

 

 

18a.

Dark warty hyphae and conidiophores present

Acrophialophora

18b.

Dark warty hyphae and conidiophores absent

19

 

 

 

19a.

Conidiophores comparatively long, forming dense whorls of branches and phialides at the septa; phialides with very short neck

Nomuraea

19b.

Not combining the above characters

20

 

 

 

20a.

Conidiophores broad; phialides borne directly on irregularly arranged branches, which are more or less cylindrical and swollen

Phialocladus

20b.

Conidiophores thin, 2-4 µm wide (when broad, synnematous); phialides in divergent clusters on irregularly or verticillately branched conidiophores or sometimes solitary

Paecilomyces

 

 

6. The genus Paecilomyces

 

Paecilomyces Bainier - Bull. trimest. Soc. mycol. Fr. 23: 26. 1907.

Spicaria auct. [non Harting 1846].

Isaria Pers. ex Fr., 1832, pro parte.

? Alphitomyces Reissek - Sber. Akad. Wiss. Wien 21: 326. 5856.

Corollium Sopp - Videnskapsselskap. Skrifter Kristiania, I. Math.-Nat. Kl. 11: 98.1912.

? Spicariopsis Heim apud Heim & Bouriquet - Revue Path. vég. Ent. agric. Fr. 26: 25. 1939.

? Coremiopsis Sizova & Suprun - Vest. mosk. gos. Univ. 2: 55. 1957.

 

 

Generic description

 

Colonies usually growing moderately well on artificial media. Hyphae hyaline to yellowish, septate, mostly smooth-walled. Conidiogenous structures synnematous or mononematous, mostly consisting of verticillate or irregularly branched conidiophores, bearing terminally on each branch whorls of conidiogenous cells, which also may be solitary on the fertile hyphae. Conidiogenous cells phialidic, consisting of a cylindrical or swollen basal portion, tapering often abruptly into a long distinct neck. Conidia in dry, [p. 10] usually thick-walled, divergent or tangled, basipetal chains, one-celled (rarely two-celled), hyaline or slightly pigmented, smooth-walled or echinulate, of various shapes. Chlamydospores usually thick-walled, borne singly or in short chains, smooth-walled or ornamented, or absent.

Ascigerous states: Byssochlamys Westling, Talaromyces C. R. Benjamin, Thermoascus Miehe.

Type species: Paecilomyces variotii Bainier.

 

 

Discussion of the synonyms

 

Brown and Smith (1957) considered the genus Spicaria Harting (1846) as a nomen ambiguum and followed Hughes (1951), who stated ‘that the type species, Spicaria solani, does not seem to have been described and that therefore it is best for the time being not to use the name’. Petch (1944b) transferred S. solani to the genus Gliocladium. Recent studies on Gliocladium and related genera (Gams, pers. commun.) showed that this transfer may be justified. The species described in Spicaria sensu Harz (1871) and sensu Saccardo (from 1881 on) and others are now accomodated in Pae­cilomyces or are considered as doubtful.

The species described in the genus Isaria producing phialides with a distinct neck, catenulate conidia and verticillate conidiophores are classified in the sect. Isarioidea.

According to the description of Alphitomyces schroeteri Reissek (1856), this species might be identical with Paecilomyces farinosus (Holm ex S. F. Gray) Brown & Smith. No type material of A. schroeteri is available (at W) and the monotypic genus is therefore considered as a possible synonym.

The synonymy of the genus Corollium Sopp (1912) was suggested by Thom (1930) and accepted by Raper and Thom (1949) and Brown and Smith (1957).

The genus Spicariopsis was described by Heim and Bouriquet (1939) without a latin diagnosis. No type material is available, but judging from the description, Spicariopsis tropicalis might be an atypical form of Pae­cilomyces variotii and Spicariopsis is considered as a possible synonym of the genus Paecilomyces.

The description and the illustrations of the genus Coremiopsis Sizova & Suprun suggest a species of the section Isarioidea, but without examination of the type the identity cannot be established.

 

 

Key to the species of Paecilomyces

 

1a.

Colonies in yellow brown to brownish colours; ascigerous state often present; mesophilic to thermophilic; odour if present, sweet aromatic [p. 11]

sect. Paecilomyces 2

1b.

Colonies white or in other bright colours; ascigerous state absent; mesophilic; odour not sweet aromatic

sect. Isarioidea 11

 

 

 

2a.

Ascigerous state or ascoma initials present

3

2b.

Ascigerous state or ascoma initials absent

8

 

 

 

3a.

Ascigerous state belonging to Byssochlamys

4

3b.

Ascigerous state belonging to Talaromyces or Thermoascus

6

 

 

 

4a.

Ascospores 5.2-6.5 x 3.2-4 µm; chlamydospores absent; conidia cylindrical with both ends flattened

P. fulvus

4b.

Ascospores smaller; chlamydospores present; conidia not cylindrical

5

 

 

 

5a.

Ascospores 4-5.5 x 1.5-3.5 µm; chlamydospores smooth-walled or slightly roughened; conidia 3-5.7 x 2.2-4 µm

P. niveus

5b.

Ascospores 3-4.5 x 2.5-3 µm; chlamydospores ornamented; conidia 2.5-4 x 1.5-3 µm

P. zollerniae

 

 

 

6a.

Ascigerous state belonging to Thermoascus

P. crustaceus

6b.

Ascigerous state belonging to Talaromyces (sect. Emersonii)

7

 

 

 

7a.

Ascomata pink to red; ascospores smooth-walled, 3.7-4.5 x 3.5-4 µm

P. byssochlamydoides

7b.

Ascomata green-yellow; ascospores ornamented with 2 to 6 ridges, 3.2-4 x 2.2-3 µm

P. leycettanus

 

 

 

8a.

Conidiogenous cells elongating with age; conidia with a truncate base

cf. Scopulariopsis sphaerospora-series

8b.

Conidiogenous cells of constant length; conidia with a rounded or apiculate base

9

 

 

 

9a.

Colonies in grey-green colours; conidiophores simple, conidia surrounded by a mucilaginous layer; reverse green

P. aerugineus

9b.

Not combining the above characters

10

 

 

 

10a.

Conidiophores usually repeatedly branched; conidia of different sizes and shapes, mostly subglobose to broadly ellipsoidal

P. variotii

10b.

Conidiophores simple; conidia long cylindrical to clavate

P. clavisporus

 

 

 

11a.

Conidia finely echinulate; colonies white to pink; reverse usually dark green

P. carneus

11b

Not combining the above characters

12

 

 

 

12a.

Phialides mostly solitary on undifferentiated hyphae or arranged in short, irregular conidiophores; conidia lemon-shaped

P. inflatus

12b.

Phialides in differentiated conidiophores

13

 

 

 

13a.

Conidiophores with a penicillate, divergent cluster of branches and phialides; conidia cylindrical, 6-7.6 x 4-5 µm, chlamydospores hyaline, ellipsoidal to pyriform

P. penicillatus

13b.

Not combining the above characters

14

 

 

 

14a.

Conidiogenous structures sulphur-yellow; conidia minute, cylindrical, 2.2-2.7 x 0.8-1.0 µm

P. sulphurellus

14b.

Not combining the above characters

15

 

 

 

15a.

Vegetative hyphae cinnamon-brown to ashy brown; conidiophores slightly pigmented, especially near the base; conidia fusiform, 5.2-7.5 x 1.7-2.5 µm; [p. 12] on scale insects

P. cinnamomeus

15b.

Not combining the above characters

16

 

 

 

16a.

Conidial heads purple or vinaceous

17

16b.

Conidial heads white, pink, cream or green

18

 

 

 

17a.

Conidiophores pigmented, rough-walled; reverse uncoloured or purple; chlamydospores absent

P. lilacinus

17b.

Conidiophores hyaline, smooth-walled; reverse yellow; chlamydospores usually present

P. marquandii

 

 

 

18a.

Conidial heads or mycelium green

19

18b.

Conidial heads not green

20

 

 

 

19a.

Conidia globose, 2.5-3.2 µm; yeast-like state present

P. viridis

19b.

Conidia reniform, 4.5-5.5 x 2-3.5 µm; yeast-like state absent

P. reniformis

 

 

 

20a.

Colonies white, floccose; dark chlamydospores present

P. puntonii

20b.

Colonies white or in other colours; chlamydospores absent

21

 

 

 

21a.

Conidiogenous cells elongating with age; base of conidia truncate and equal to the width of the conidium-bearing neck

cf. Scopulariopsis brumptii-series

21b.

Conidiogenous cells of constant length; conidia mostly with rounded ends and the base smaller than the width of the conidium-bearing neck

22

 

 

 

22a.

Colonies spreading rapidly; conidia triangular in outline; colony margin arachnoid

P. niphetodes

22b.

Colonies usually growing slowly; conidiophores mono- or synnematous

23

 

 

 

23a.

Conidia cylindrical or narrowly fusiform, sometimes curved

24

23b.

Conidia of other shape

27

 

 

 

24a.

Conidia cylindrical, usually curved

25

24b.

Conidia cylindrical to narrowly fusiform, straight

26

 

 

 

25a.

Conidia 2-7 x 1.2-2.5 µm, synnemata 1-3.5 cm long; on lepidopterous pupae and larvae

P. tenuipes

25b.

Conidia 3.5-8 x 1.5-3.5 µm; synnemata large, 25-50 mm long; on Cicadae

P. cicadae

 

 

 

26a.

Colonies usually pink; conidia fusiform to cylindrical, 3-4 x 1-2 µm

P. fumosoroseus

26b.

Colonies white to cream; conidia fusiform, sometimes cylindrical, 4-6 x 1-1.5 ↨0m

P. javanicus

 

 

 

27a.

Conidia less than 3.5 m long

28

27b.

Conidia longer than 3.5 µm

29

 

 

 

28a.

Conidia ellipsoidal to fusiform; colonies white to yellow; synnemata on the natural substrate white to yellow

P. farinosus

28b.

Conidia subglobose to ellipsoidal, often irregular in shape; colonies white to red; synnemata on the natural substrate darker red

P. amoeneroseus

 

[p. 13]

 

29a.

Synnemata typically strongly branched; conidia pyriform, apiculate, 3.2-4.5 x 1.5-2.7 µm

P. ramosus

29b.

Not combining the above characters and conidia larger

30

 

 

 

30a.

Conidia subglobose to broadly fusiform, 3.5-5.0 x 2.5-3 µm

P. ghanensis

30b.

Conidia not broadly fusiform

31

 

 

 

31a.

Conidia broadly ellipsoidal, 5.7-8.5 x 2.5-3 µm (on the natural substrate 4.5-5.5 µm long); synnemata conspicuous and stout

P. coleopterorum

31b.

Conidia subglobose to ellipsoidal or broadly cylindrical; synnemata loose or absent

32

 

 

 

32a.

Conidiophores simple; conidia subglobose to ellipsoidal, 4.5-7 x 3.0-4.5 µm

P. xylariiformis

32b.

Conidiophores repeatedly branched; conidia ellipsoidal to cylindrical, 4.5-12 x 3.0-4.0 µm

P. suffultus

 

 

Paecilomyces section Paecilomyces

 

Colonies mostly spreading broadly, in yellow-brown, olive-brown or other dark colours. Odour sweet aromatic. Mycelium hyaline to yellow, thin- to thick-walled; hyphae broad, especially when submerged. Conidiophores mononematous, usually rather complex, consisting of verticillately and/or irregularly arranged branches, bearing whorls of several phialides. Phialides consisting of a cylindrical basal portion, tapering abruptly into a long cylindrical neck, which is sometimes bent away from the main axis. This apex of the phialides is sometimes thickened. Conidia in long, diver­gent, dry chains, hyaline to yellow, smooth-walled, one-celled. Chlamydospores present in several species, thick-walled, dark, smooth-walled to ornamented, occasionally in short chains. Thermotolerant to thermophilic. Saprophytic, not entomogenous.

Perfect states: Byssochlamys, Talaromyces, Thermoascus.

Type species: Paecilomyces variotii Bainier.

 

Discussion

 

The colonies have usually a rapid growth, only some P. variotii strains grow restrictedly. The colour of the colonies can be sometimes affected by the colour of the cleistothecia, e.g. pink in P. byssochlamydoides or green-yellow in P. leycettanus. The colonies of P. niveus are white to cream, but when the Paecilomyces state is abundantly produced, they are near avellaneous. The sweet aromatic odour is usually most pronounced in old cultures. [p. 14]

The phialides can be of different shape, but mostly they have a cylin­drical basal portion which tapers abruptly into a long cylindrical neck. In some strains of P. variotii the phialides can be strongly inflated and may occasionally proliferate laterally. These proliferations are, however, not a constant character. This feature is regarded as an atypical development of some strains. This may have given rise to the description of Spicariopsis tropicalis Heim as a new species.

The conidia are usually produced in long divergent chains and may form columns in strains with abundant sporulation and densely compacted conidiophores. The shape of the conidia is variable, especially in P. variotii, depending on colony age and the extent of sporulation. Smaller conidia are produced in abundantly sporulating strains, whereas larger conidia can be observed under less suitable conditions. Chlamydospores are produced in P. variotii, P. niveus, P. leycettanus, P. byssochlamydoides, P. clavisporus and P. zollerniae. Those of the last species are warty.

The species of the sect. Paecilomyces show optimal growth between 30° and 45°C. P. leycettanus, P. byssochlamydoides, and occasionally also P. variotii are thermophilic; other species are mesophilic to thermotolerant.

 

 

1. Paecilomyces variotii Bainier - Fig. 1, 2.

 

Paecilomyces variotii Bainier - Bull. trimest. Soc. mycol. Fr. 23: 26. 1907 = Penicillium variotii (Bain.) Sacc. - Syll. Fung. 22: 1273. 1913.

Penicillium divaricatum Thom - Bull. Bur. Anim. Ind. U.S. Dep. Agric. 118: 92. 1910 = Spicaria divaricata (Thom) Gilman & Abbott - Iowa St. Coll. J. Sci. 1: 301. 1929 = Spicaria divaricata (Thom) Ma - Lingnan Sci. J. (Suppl.) 12: 115. 1933.

Corollium dermatophagum Sopp - Videnskapsselskap. Skrifter Kristania, Math. Nat. Kl. 11: 99. 1912.

? Penicillium repandum Bainier & Sartory - Bull. trimest. Soc. mycol. Fr. 29: 367. 1913.

Spicaria elegans (Corda) Harz var. flava P. See - Les Maladies du Papier pique, p. 33. 1919.

Penicillium mandshuricum Saito - Rep. S. Manchur. Rly. Co. cent. Lab. 6: 11. 1921 = Paecilomyces mandshuricus (Saito) Thom - The Penicillia, p. 550. 1930.

Penicillium arenarium Shaposhnikov & Manteifel - Trudy nauch. khim.-farm. Inst. Mosk. 5: 3. 1923.

Penicillium aureo-cinnamomeum Biourge - La Cellule 33: 213. 1923 = Paecilomyces aureo-cinnamomeus (Biourge) Thom - The Penicillia, p. 547. 1930.

Eidamia catenulata Horne & Williams - Ann. Bot. 37: 398. 1933.

Paecilomyces mandshuricus (Saito) Thom var. saturatus Nakazawa, Takeda & Suematsu - J. agric. Chem. Soc. Japan 10: 102. 1934.

Spicaria taurica Naumoff & Kiryalova - Trudy bot. Inst. Erevan 2: 363. 1934.

? Spicariopsis tropicalis Heim apud Heim & Bouriquet - Revue Path. vég. Ent. agric. Fr. 26: 25. 1939 (without latin diagnosis).

Monilia formosa Sakaguchi, Inoue & Tada - Zentbl. Bakt. ParasitKde (Abt. 2) 100: 302. 1939.

Paecilomyces dactylethromorphus Batista & Maia (as 'dactylethromorphis') - Anais Soc. biol. Pernamb.15:152. 1957.

Paecilomyces variotii Bainier var. antibioticus Abe, Takeuchi & Yonehara - J. Antibiot., Tokyo 12: 202. 1959 (without latin diagnosis). [p. 15]

Paecilomyces variotii Bainier var. brunneolus Inagaki - Trans. mycol. Soc. Japan. 4: 4. 1962.

Spicaria divaricata (Thom) Gilman & Abbott var. heterospora Rownak & Hedayetellah - Pakist. J. scient. ind. Res. 7: 36. 1964 (without latin diagnosis).

Spicaria asymmetrica K.B. Deshpande & K. S. Deshpande - Sydowia 19: 156. 1965 (without indication of the type)

Paecilomyces subglobosus Misaki, Yasui & Okabe apud Sawada et al. - Agric. Biol. Chem. 32: 64. 1968.

Paecilomyces lecythidis Ram (as 'lecythisii') - Nova Hedwigia 16: 307. 1968.

Paecilomyces maximus Ram - Nova Hedwigia 16: 306. 1968.

Paecilomyces indicus Rai, Tewari & Mukerij - Mycopath. Mycol. appl. 38: 25. 1969.

 

 

Fig. 1. Paecilomyces variotii, different types of conidiogenous structures. [p. 16]

 

 

Fig. 2. Paecilomyces variotii, a. atypical phialides with proliferations; b-g. conidia, b. CBS 370.70, c. CBS 990.73, d. CBS 371.70, e-g. 3 typical strains; h. chlamydospores.

 

Colonies on malt-agar usually growing rapidly, attaining a diameter of 8 cm within 14 days at 25° C; consisting of a dense felt of numerous conidiophores, giving a powdery appearance, sometimes floccose or funiculose or tufted, in older strains mostly with overgrowth of white aerial mycelium; colour variable depending on the strain, ranging from Deep Olive Buff to Dark Olive Buff (Ridgway, Pl. 40; Rayner, 21"'b, 21"') or Light Yellowish Olive, Yellowish Olive to Ecru-Olive (Ridgway, Pl. 30; Rayner, 23"i,23"k, 21"i), with age changing to darker shades, in some strains with a dark cast due to abundant chlamydospores; odour sweet aromatic, pronounced with age. Exudate absent or consisting of colourless drops; reverse yellow to yellow-brown, in some strains dark-brown to almost black, from abundant dark chlamydospores. Vegetative hyphae hyaline, mostly thick-walled, smooth-walled or sometimes encrusted with yellow granules, 3.0-5.5 µm wide, submerged hyphae strongly inflated up to 20 µm. Conidiophores consisting of dense whorls of verticillately or irregularly arranged branches, each bearing 2 to 7 phialides, smooth-walled, slightly roughened or en­crusted with granules, 35-90 x 3.5-7.0 µm, but occasionally up to 150 µm in length. Phialides in whorls or solitary, variable in size, mostly 12-20 x 2.5-5.0 µm, but also up to 35 µm long, consisting of a cylindrical or ellipsoidal basal portion, tapering abruptly into a long cylindrical, 1.0-2.0 µm wide neck, sometimes slightly bent away from the main axis; the phialides have apically an internal wall-thickening. In some strains the phialides may [p. 17] proliferate and form 2 to 3 lateral necks (see fig. 2). Conidia hyaline to yellow, yellow-brown in mass, smooth-walled, variable in shape, mostly subglobose to ellipsoidal, in some strains ellipsoidal to cylindrical or clavate, dimensions varying from 3.2-5.0 x 2.0-4.0 µm in some strains to 3.5-15.0 x 2.0-5.0 µm in others. Chlamydospores usually present, especially in old cultures, singly or in short chains, mostly brown or dark brown, smooth-walled to slightly roughened, globose, subglobose to pyriform, thick-walled, 4.0-8.0 µm, sometimes up to 10 µm in diameter. Perfect state not produced.

Optimal growth occurs at 35ŗ C, some strains show a higher optimum at about 40ŗ C (see also Knösel and Rész, 1973); minimum temperature about 5ŗ C; maximum near 50° C.

 

Material examined

 

Herbarium specimens

Paecilomyces subglobosus, dried subculture of the type (TPR-3811), isolated from air in Japan, sent by Jiro Sawada, Tokyo, Japan.

Spicariopsis tropicalis, isolated from soil, Yangambi, Congo, by J. Meyer (IMI 83980).

 

Living cultures

CBS 323.34 = IMI 58411 = type culture of Paecilomyces mandshuricus var. saturatus, received from R. Nakazawa.

CBS 265.35, isolated by G. Goidįnich from oak-wood, Italy.

CBS 284.48 = IMI 40025 = ATCC 10121 = NRRL 1115, type culture of Penicillium divaricatum, isolated from mucilage bottle with library paste (cf. Thom. 1930). CBS 338.51, isolated by H. Lüthi from fruit-juice, Wädenswil, Switzerland. CBS 339.51, isolated from leather at Progil S.A., Lyon, France.

CBS 251.55 = IMI 65752 = ATCC 11971, type culture of Paecilomyces dactylethromor­phus, isolated as a contaminant of acetic acid solution, Brazil, received from A. C. Batista, Recife, Brazil.

CBS 628.66 = IMI 114930 = CEB 3292, isolated from tanned sheep leather in France and used for testing mould proofing.

CBS 368.70 = IMI 82348, isolated by C.L.A. Leahy from quinine-containing medicine, Great Britain.

CBS 370.70 = IFO 7563, type culture of Paecilomyces variotii var. brunneolus, isolated by N. Inagaki from food in Japan.

CBS 371.70 = IMUFPe 2192, type culture of Paecilomyces maximus, isolated by C. Ram from Annona squamosa in Brazil.

CBS 372.70 = IMUFPe 2191, type culture of Paecilomyces lecythidis, isolated by C. Ram from wood of Lecythis unsitata Brazil.

CBS 990.73a = NRRL 1123, type culture of Penicillium viniferum, and CBS 990.73b = NRRL 1282, type culture of Monilia formosa, both isolated by Sakaguchi et al. (1939) in Japan.

CBS 102.74 = NRRL 1116, probably type culture of Paecilomyces variotii, sent to C. Thom in 1922 as strain no. 436 from France by O. da Fonseca (D. I. Fennell, pers. commun.).

CBS 991.73, isolated from air by E. Stix in Munich, Germany.

CBS 992.73 = ATCC 24330, isolated from necropsy specimens of a Weimaraner dog by S. S. Jang et al. (1971). [p. 18]

 

In the Herbarium IMI P. variotii is represented on the following substrates: soil and air (several specimens collected in different countries of various climates); man (157734); aborted calf foetus (57409, 177443); stomach contents of 1 week old calf with mycotic ulcers (61878); rat caecum (125830); cooked rice (62571); sugarcane (123162); fermented cocoa beans (77368); inoculation product of salami sausage (131711); silage (62077); hay (96227, 58451, 113741); photographic paper (53037); synthetic rubber (108007); jute fibre (16196); kerosene storage tank (112572); water exposed to alcoholic atmosphere (134313); shale (132632); on the following plants; Arachis hypogaea, Carica papaya, Coffea species, Elaeis guineensis, Hordeum species, Malus pumila, Nicotiana tabacum, Oryza sativa (roots), Pinus elliotii, Pinus radiata, Prunus domestica, Quercus species, Raphanus sativus, Ricinus communis, Saccharum officinarum, Sorghum species, Theobroma cacao, Vicia faba,Zea mays (stored) (cited numbers are all Herb. IMI no's).

 

About 50 NRRL-strains of P. variotii were received from Miss D.I. Fennell (Peoria). Most of them were isolated from soil or air; the strains originated from chest abscess (A-2029); brain and lung of a turkey (Emmons 1171-1172); dried eggs (Watson 8); flour (A-1S410); starch (A-15542); peach fruit (A-5405); refrigerated wheat flour (A-12401); ground corn (A-13831); mouldy feed sample (A-17886); mouldy cigars (A-S968); rotten wood (A-6867); decomposing agaric (A-6919); creosoted wood (A-3639); individual protective cover (CWS 280); mouldy leather instrument case (A-279); water resistant matches (A-626); fountain pen (A-2177); 15% ethanol solution (A-3926); Shaeffer's ink (A-2952); pharmaceutical emulsion (A-6885).

 

Discussion

 

The description of P. variotii has been broadened to accomodate also some deviating strains. All strains are connected with each other by intermediate forms. The conidia especially can vary in shape and size, depending on the colony age, the extent of sporulation and the temperature of in­cubation. The rate of growth can vary depending on the strain.

Raper and Thom (1949) and Brown and Smith (1957) listed several synonyms of P. variotii. The type cultures of Paecilomyces dactylethromorphus Batista & Maria and P. lecythidis Ram also fit in the present description. P. variotii var. brunneolus Inagaki differs from the typical P. variotii strains by the restricted growth of the colonies. The colony characters and the conidiogenous structures are, however, normal. P. maximus Ram differs only by the production of large, more or less clavate conidia; such large conidia are not uncommon in P. variotii. Only a dried culture of P. subglobosus Misaki & al. could be studied, which is sufficient to prove its identity. No type material of P. variotii var. antibioticus Abe & al., Spicaria divaricata var. heterospora Rownak & Hedayetellah, Spicaria asymmetrica Deshpande & Deshpande and P. indicus Rai & al. could be obtained, but from the descriptions their synonymy with P. variotii is obvious. No type material of Spicariopsis tropicalis Heim was available for study. A specimen in Herb. IMI identified as Spicariopsis tropicalis proved to be identical with P. variotii. The morphological characters fit in the description of Spicariopsis tropicalis.

 

Occurrence: P. variotii is a rather common fungus in the air, in soil (subtropical and tropical climates), in compost (Knösel and Rész, 1973), in [p. 19] self-heated substrates, on wood (Cartwright, 1937), and it is frequently recorded as a moderately thermophilous organism. The species is reported as common in food-stuffs, such as cocoa beans (Maravalhas, 1966), riceflour (Kurata and Ichinoe, 1967), in fruit juices (Senser and Rehm, 1965; Senser et al., 1967a and b; Eschmann, 1971), in manioc loaves (Goncalves de Lima and Magali de Araujo, 1969). Bonner and Fergus (1959) and Gladoch (1973) isolated P. variotii from animal feeding stuffs.

 

Metabolites: Several organic acids are reported to be produced by P. variotii or species now considered as synonymous (Shaposhnikov and Manteifel, 1923; Loesecke, 1945; Sakaguchi et al. 1939) The antibiotic variotin showing low toxicity to mice and antifungal properties was discovered by Takeuchi et al. (1959). For a detailed review of this antibiotic see Tanake (1967). Further metabolites reported from P. variotii are fusigen, a sideramine (Dieckmann, 1967) and indole-3-acetic acid (Voinova-Raikova et al., 1969).

 

Biodeterioration: The species is reported as the causal agent of biodeterioration of palm oil (Coursey, 1959; Cornelis et al. 1965), of leather (Sopp, 1912; Orthmann and Highly, 1929; Lazįr and Brįtulescu, 1971), of cosmetics (Navarre and Baley, 19S7), of jute (Macmillan and Basu, 1947; Basu, 1948), of paper (Lavers and Illman, 1946), and of pharmaceutical emulsions (Eggins and Walker, 1963). A yellow staining of oakwood by P. variotii is known as `golden oak', (Williamson, 1923; Davidson, 1935 Cartwright, 1937; Brown and Smith, 1957).

 

Pathogenicity: Although P. variotii is generally considered as a sapro­phyte, it can apparently become pathogenic in man and animals, causing the so-called paecilomycosis (Baker, 1971). A case of infective endocarditis caused by P. variotii growing on an aortic allograft was observed by Silver et al. (1971). Uys et al. (1963) reported endocarditis following cardiac surgery due to a Paecilomyces species. A similar fungus was isolated by Henig et al. (1971) from an infected lacrimal sac of a 67 years old woman. The fungi described in the two latter papers are probably Paecilomyces variotii. Jang et al. (1971) and Patnaik et al. (1972) isolated P. variotii strains causing a systemic mycosis in dogs.

According to the literature, P. variotii should play a rather important role in the mycotoxicosis in several animals. Turresson (1916) already reported that rabbits died after ingesting conidia and mycelium of Penicil­lium divaricatum. Jungherr and Gifford (1944) isolated some strains of P. variotii from internal organs of a turkey, which had been attacked by an unknown disease. Forgacs and Carl (1955, 1962), Forgacs et al. (1958) observed a toxicosis in swine and death in chickens after ingestion of food-stuffs contaminated with P. variotii. Guallini (1968) and Re­daelli et al. (1968) isolated a P. variotii strain from a horse, which had succumbed to a kidney disease. This strain proved toxic in animal tests. [p. 20]

 

 

2. Paecilomyces aerugineus Samson, spec. nov. - Fig. 3.

Spicaria silvatica Oudemans sensu Apinis = Scopulariopsis silvatica (Oudem.) Apinis - Nova Hedwigia p: 73. 1963.

 

Coloniae in agaro maltoso celeriter crescunt, ad 8 cm diam. 14 diebus 35° C, fere dense intricatae, primum griseolae, deinde griseo-virides; reversum obscure viride. Conidiophora simplicia, plerumque e hyphis submersis oriuntur, 30-75 x 2.5-4.0 µm, pluries verticillata, quoties binas vel ternas phialides ferunt. Phialides 12-19 x 2.5-4 µm, e basi fere cylindrica in collum distinctum apice inspissato angustatae. Conidia in catenis divergentibus connexa, luteo-brunnea, primum mucilagine circumdata, deinde sicca et verrucosa, ellipsoidea vel fusiformia, utrinque modice truncata, 6-8.5 x 2.5-3.5 µm.

Temperatura optima prope 35° C, maxima 40° C, minima 10° C.

Typus CBS 350.66, isolatus e palude Glyceriae maximae prope Nottingham, G.B.

 

 

Fig. 3. Paecilomyces aerugineus, a-b. conidiophores; c. conidia.

 

Colonies on malt-agar growing rapidly, attaining a diameter of 8 cm  within 14 days at 35° C, composed of a rather densely matted felt, with a thin overgrowth of mycelium, at first grey, changing into more grey-green colours near Dark American Green (Ridgway, pl. 41; Rayner, 33"' k) due to the dark green reverse. Exudate and odour not produced. Reverse typically dark green near Meadow Green (Ridgway, Pl. 6; Rayner 35k), the pigment mostly diffusing into the surrounding agar. Vegetative hyphae thin-walled, septate, usually encrusted with green granules or smooth-walled, 2-4 µm in diameter. Conidiophores simple, mostly arising from the submerged hyphae, 30-75  x 2.5-4.0 µm, consisting of a few verticillate branches, bearing groups of 2 to 3 phialides. Occasionally the phialides  [p. 21] are solitary. Phialides 12-19 x 2.5-4 µm, consisting of a more or less cylindrical basal portion, tapering into a distinct neck which is usually internally thickened at the apex. Conidia in dry divergent chains, yellow-brown, when young surrounded by a mucilaginous layer, later dry and verrucose, ellipsoidal to fusiform, with both ends somewhat flattened, 6-8.5 x 2.5-3.5 µm. Chlamydospores and perfect state not observed.

The species has its temperature optimum near 35° C; maximum at 40° and minimum at about 10° C.

 

Material examined

 

CBS 350.66 = IMI 105412 = BDUN 276 = type culture, isolated by A. E. Apinis from Glyceria maxima swamp near Nottingham, Great Britain.

 

Discussion

 

Apinis (1963) proposed the new combination Scopulariopsis silvatica (Oudem.) Apinis for a microthermophilic fungus isolated from a Glyceria maxima swamp near Nottingham. No type material is left of Spicaria silvatica Oudem., but Oudemans and Koning (1902; original drawings in L) described and depicted S. silvatica with grey-green colonies growing on gelatine and with rather large, smooth-walled, ellipsoidal conidia produced on Penicillium-like conidiophores. This fungus may be identical with Penicillium digitatum Sacc., which occasionally produces conidia measuring 6-12 x 4-6 µm. The conidia of Apinis' isolate are usually 6-8.5 x 2.5-3.5 µm and surrounded by a mucilagenous layer.

 

3. Paecilomyces clavisporus Hammill - Fig. 4.

Paecilomyces clavisporus Hammill - Mycologia 62: 109. 1970.

 

Colonies on malt-agar growing rapidly, attaining a diameter of 6 cm within 14 days, composed of a densely matted felt with a loose, floccose to funiculose overgrowth of aerial mycelium, giving the surface of the colony buff shades near Pale Olive Buff and Olive Buff (Ridgway, Pl. 40; Rayner, 21"'f, 21"'d). Reverse yellow-brown to olive. Odour and exudate not pronounced or absent. Vegetative hyphae hyaline to yellow, smooth-walled, 2-3.5 µm wide. Conidiophores usually short, about 80 µm in length and 3-4 µm in diameter, smooth-walled, hyaline to yellow, con­sisting of short branches bearing one or two phialides; phialides may also be borne directly and singly on trailing hyphae. Phialides 13.5-25 x 2.5-4.6 µm, consisting of a basal cylindrical portion, tapering abruptly into a long neck, 4.2-10.5 µm in length and 1.5-2.0 µm in diameter. Occasionally the coni­diogenous cells produce apically chains of chlamydospore-like cells, which do not separate. Conidia in divergent or tangled chains, variable in size [p. 22] and shape, broadly ellipsoidal to pyriform or clavate, smooth-walled, or slightly roughened, hyaline to yellow, yellow-brown in mass, 7.0-11.5 x 1.7-4.3 µm. Chlamydospores borne singly, terminally or intercalarily, thick-walled, smooth-walled, globose, up to 8 µm in diam.

The species has its temperature optimum near 35° C; minimum at about 10° C; maximum at 40° C.

 

 

Fig. 4. Paecilomyces clavisporus, a-b. conidiophores; c. catenate chlamydospores; d. chla­mydospores; e. conidia.

 

Material examined

 

CBS 329.70, type culture, isolated by T. M. Hammill from forest soil in Clarke County, Georgia, USA.

 

Discussion

 

The species is only known from the type culture. It is distinct from other species by the clavate conidia and the presence of thick-walled chlamydo­spores. The clavate conidia of P. clavisporus resemble those of P. maximus Ram, [p. 23] which is regarded as a synonym of P. variotii. P. clavisporus differs from P. variotii by the restricted growth, the less branched conidiophores and the uniformly clavate conidia.

 

 

4. Paecilomyces fulvus Stolk & Samson - Fig. 5.

 

Paecilomyces fulvus Stolk & Samson - Persoonia 6: 354. 1971.

 

Ascigerous state: Byssochlamys fulva Olliver & G. Smith - J. Bot. 72: 197. 1933

 

Colonies on malt-agar spreading broadly, attaining a diameter of 9 cm within 7 days at 30° C; fulvous, near Olive Buff or Deep Olive Buff (Ridgway, Pl. 40; Rayner, 21"'d, 21"'b). Conidiophores short, smooth-walled, hyaline, up to 150 µm in length, bearing branches with two or three phialides, occasionally with densely verticillate conidiogenous cells. Phialides 12.5-17.0 x 2.5-3.5 µm. Conidia in divergent or tangled chains, usually cylindrical, with truncate ends, smooth-walled, yellow, 4.0-8.7 x 1.5-5.0 µm. Chlamydospores absent.

The species is thermotolerant with an optimal temperature between 30° and 35° C; temperature minimum about 10° C; maximum at 45° C.

A detailed description and list of material examined are given by Stolk and Samson (1971).

 

 

Fig. 5. Paecilomyces fulvus, a-b. conidiophores; c. conidia. [p. 24]

 

Occurrence: Byssochlamys fulva and B. nivea have been reported as the cause of spoilage in canned food products, because of their heat re­sistance, their ability to grow at low oxygen partial pressure (King et al., 1969) and optimum growth at low pH (Hull, 1939). Incidence of spoilage outbreaks in heat processed acid foods caused by these two fungi seems to be increasing and the physiology and biochemistry of these organisms is being studied by many investigators (King et al., 1969; Downing, 1969; Splitts­toesser et al., 1969, 1971; Meyer, 1971).

 

Heat resistance: The ascospores of B. fulva and B. nivea possess a high heat resistance, they survive the exposure to temperatures of 80° to 90° C for 10 minutes (King et al. 1969; Peter, 1965; Schumann and Werner, 1973). Partsch et al. (1969) assumed that the ascospores are protected against these external influences by the rather thick cell wall with a distinct electron transparent layer between the outer cell wall and the cyto­plasmic membrane. Since thermal treatment is often not sufficient for killing Byssochlamys ascospores, the effects of gamma irradiation (Partsch and Altmann, 1970), of germicides (Ito et al., 1972) or of antifungal agents (Takano and Yabuki, 1964) were investigated. These agents had little effect.

 

Byssochlamic acid: Raistrick and Smith (1933) discovered large amounts of mannitol and byssochlamic acid in B. fulva, the structure of which was elucidated by Baldwin et al. (1963, 1965). The acid is formed only under aerobic conditions, and partial or complete exclusion of air prevents its formation. It is produced from a wide variety of sugars at various con­centrations and at pH range of 2.5-8.0 (Meyer and Rehm, 1967; Meyer, 1971). Schmidt and Rehm (1968, 1971) reported methods of detection and quantitative analysis from culture filtrate and fruit juices. Toxicity of byssochlamic acid to mice was shown by Raistrick and Smith (1933). The toxin is weakly hepatotoxic to guinea pigs (Gedek, 1971). The solubilized acid is moderately toxic when injected intraperitoneally to mice (King et al., 1972). When large amounts of dried mycelium were fed to rats for 90 days, only an increase in liver weight, and a decrease in body weight and haemoglobin concentration could be observed. Meyer and Rehm (1967, 1969) showed that byssochlamic acid inhibited the growth of Sinapis alba seedlings and the fermentation of Saccharomyces cerevisiae.

 

Enzymes: The production of pectolytic enzymes by B. fulva was first demonstrated by Olliver and Rendle (1934). Beaven and Brown (1949) reported that the disintegration of fruit by B. fulva was due to the action of a protopectinase as well as a disaggregation enzyme. According to Reid (1950, 1951), B. fulva should produce polygalacturonase and pectinesterase. Chu and Chang (1973) have shown that B. fulva produces a variety of pectolytic enzymes, including pectinesterase, polygalacturonase, poly­methylgalacturonase and pectate lyase. A rennin-like enzyme in B. fulva was reported by Knight (1966), Reps et al. (1969) and Chu et al. (1973). [p. 25]

 

 

5. Paecilomyces niveus Stolk & Samson - Fig. 6.

 

Paecilomyces niveus Stolk & Samson - Persoonia 6: 351. 1971.

? Spicaria multicola Naumoff & Kiryalova - Trudy bot. Inst. Akad. Nauk SSSR 3: 262. 1935.

 

Ascigerous state: Byssochlamys nivea Westling - Svensk bot. Tidskr. 3: 134. 1909.

 

 

Fig. 6. Paecilomyces niveus, a-d. conidiophores, a. CBS 100.11, b-d. strains isolated from soil, collected in Surinam; e-g. conidia; h. chlamydospores. [p. 26]

 

Colonies on malt-agar spreading broadly, attaining a diameter of 9 cm within 7 days at 30° C; cream, near Cartridge Buff (Ridgway, Pl. 30; Rayner, 19"f), in predominantly conidial strains in buff shades near Olive Buff, Deep Olive Buff or Avellaneous (Ridgway, Pl. 40; Rayner, 21"'d, 21"'b). Conidiophores usually not distinct, smooth-walled, up to 300 µm in length and 2-3 µm in diam., bearing branches with 2 to 3 phialides, oc­casionally densely and verticillately branched. Phialides 12.5-20 x 2-3.5 µm. Conidia in divergent chains, globose to ellipsoidal with a flattened base, smooth-walled, hyaline to yellow, 3.0-5.7 x 2.2-4.0 µm. Chlamydospores abundantly produced, thick-walled, smooth-walled to slightly roughened, globose, ovoidal to pyriform, yellow-brown to brown, up to 10 µm in diam.

The species is thermotolerant with an optimal temperature between 30 and 35° C; temperature minimum about 10° C; maximum at 40° C.

A detailed description, list of material examined and discussion of the synonymy are given by Stolk and Samson (1971).

 

Discussion

 

Stolk and Samson (1971) regarded Byssochlamys nivea var. lagunculariae Ram as a synonym of B. nivea, in spite of its avellaneous colour of the colony. In the meantime several other isolates from soil of Surinam were obtained which fit the description of B. nivea var. lagunculariae. The avel­laneous colour of the colony is due to the abundant production of conidio­phores. The dimensions of the conidiogenous structures and the ascospores are somewhat smaller than the typical B. nivea strains, but they fall with­in the given range. Furthermore, in fresh isolates of B. nivea or cultures grown on media which influence conidium production, the colour of the colonies may also be avellaneous. For these reasons B. nivea var. lagun­culariae is regarded as conspecific with B. nivea.

Karrow and Foster (1944) and Kis et al. (1969) reported that B. nivea produces the mycotoxin patulin. Zaroogian and Curtis (1963, 1964) isolated a plant growth inhibitor from the culture filtrate of B. nivea and named it `asymmetrin'. Yates and Mooney (1968) demonstrated the presence of two pectic enzymes, endopolygalacturonase and endopolymethylgalacturonase. Other biochemical or physiological properties are given under Paecilomyces fulvus.

 

 

6. Paecilomyces zollerniae Stolk & Samson - Fig. 7.

 

Paecilomyces zollerniae Stolk & Samson - Persoonia 6: 356. 1971.

 

Ascigerous state: Byssochlamys zollerniae Ram - Nova Hedwigia 26: 312. 1968 [p. 27]

 

 

 Fig. 7. Paecilomyces zollerniae, a-b. conidiophores; c. conidia; d. chlamydospores.

 

Colonies on malt-agar spreading broadly, attaining a diameter of 8 cm within 14 days at 30° C; white at first, changing to Cream Buff (Ridgway, Pl. 30; Rayner, 19"d). Conidiophores smooth-walled, up to 150 µm in length, arising from the aerial hyphae, bearing short branches with two or three phialides. Phialides 8-20 x 2.0-2.5 µm. Conidia in divergent chains, globose to ellipsoidal, apiculate, smooth-walled, hyaline, 2.5-4.0 x 1.5-3.0 µm. Chlamydospores abundantly produced, singly or in short chains, smooth when young, later becoming warty, brown to dark brown, 5.o-10.5 µm in diam., usually globose.

The species is thermotolerant with a temperature optimum between 30 and 35° C; minimum at about 10° C; maximum at 45° C.

A detailed description and discussion of the perfect state is given by Stolk and Samson (1971).

 

 

Discussion

 

Paecilomyces zollerniae can be easily distinguished from the other species by its cottony growth and the abundant warty chlamydospores. According to Ram (1968) Byssochlamys zollerniae differs from the other two Bys­sochlamys species by the smaller ascospores and the different imperfect state. [p. 28]

 

 

7. Paecilomyces byssochlamydoides Stolk & Samson - Fig. 8.

 

Paecilomyces byssochlamydoides Stolk & Samson - Stud. Mycol. 2: 45. 1972.

 

Ascigerous state: Talaromyces byssochlamydoides Stolk & Samson - Stud. Mycol. 2: 45. 1972.

 

Colonies on malt-agar spreading broadly, attaining a diameter of 8 cm within 7 days, at first ochraceous near Cinnamon Buff, but with the ripening of the ascomata changing into pink shades near Pinkish Cinnamon to Vinaceous Cinnamon, becoming Vinaceous Russet in age (Ridgway, Pl. 29, 28; Rayner, 15"b, 7"i). Conidiophores smooth-walled, hyaline to yel­low, up to 300 µm long, arising primarily from submerged hyphae, but also as branches from aerial mycelium. Phialides 10-24 x 3.0-3.5 µm. Conidia usually cylindrical with rounded ends, sometimes becoming ellipsoidal at maturity, smooth-walled, hyaline to yellow, 4.0-8.0 x 1.0-2.5 µm. Chlamy­dospores rarely produced, globose, hyaline, smooth-walled, about 4 µm in diameter.

The species is strongly thermophilic with a temperature optimum between 40 and 45° C; minimum about 25° C; maximum slightly above 50° C.

A detailed description is given by Stolk and Samson (1972).

 

Discussion

 

Paecilomyces byssochlamydoides is characterized by its Talaromyces ascigerous state and its cylindrical conidia with rounded ends. It is closely related to P. crustaceus and P. variotii which usually have ellipsoidal conidia.

 

 

Fig. 8. Paecilomyces byssochlamydoides, a-b. conidiophores; c. conidia; d. chlamydospores. [p. 29]

 

 

8. Paecilomyces leycettanus (Evans & Stolk) Stolk, Samson & Evans - Fig. 9.

 

Penicillium leycettanum Evans & Stolk - Trans. Br. mycol. Soc. 56: 45. 1971. = Paecilomyces leycettanus (Evans & Stolk) Stolk & al. - Persoonia 6: 342. 1971.

 

Ascigerous state: Talaromyces leycettanus Evans & Stolk - Trans. Br. mycol. Soc. 56: 45. 1971

 

Colonies on malt-agar spreading broadly, attaining a diameter of about 7 cm within 7 days; green-yellow near Sea Foam Green (Ridgway, Pl. 31; Rayner, 27'f). Conidiophores smooth-walled, rarely encrusted with brown granula, 80-350 µm in length and 3-4 µm in diameter, usually arising from submerged hyphae, but also as branches of aerial hyphae. Phialides 10-15 x 1.5-2.5 µm. Conidia cylindrical with rounded ends, smooth-walled, yellow, yellow-brown in mass, 4.0-6.0 x 1.0-1.5 µm. Chlamydospores occasionally produced, globose, yellow, smooth-walled, 4-4.5 µm in diameter.

The species is thermotolerant to thermophilic with a temperature optimum at about 40° C; minimum at 18° C; maximum 55° C.

A detailed description is given by Stolk and Samson (1972).

 

 

Fig. 9. Paecilomyces leycettanus, a-b. conidiophores; c. conidia; d. chlamydospores. [p. 30]

 

Discussion

 

Paecilomyces leycettanus is distinct from the other species by its Talaromyces perfect state, the presence of smooth-walled chlamydospores and the cylindrical conidia with rounded ends. The species resembles P. byssochla­mydoides and P. fulvus. In P. byssochlamydoides the conidia are larger, while in P. fulvus the cylindrical conidia have truncate ends. P. leycettanus is regarded as an intermediate form between Paecilomyces and Penicillium, because of its less divergent conidiophores.

 

 

9. Paecilomyces crustaceus Apinis & Chesters - Fig. 10.

 

Paecilomyces crustaceus Apinis & Chesters - Trans. Br. mycol. Soc. 47: 428. 1964.

Ascigerous state: Thermoascus crustaceus (Apinis & Chesters) Stolk.

 

Dactylomyces crustaceus Apinis & Chesters - Trans. Br. mycol. Soc. 47: 428. 1964 = Thermoascus crustaceus (Apinis & Chesters) Stolk - Antonie van Leeuwenhoek 31: 272. 1965.

 

 

Fig. 10. Paecilomyces crustaceus, a-b. conidiophores; c. conidia.

 

Colonies on malt-agar agar spreading broadly, attaining a diameter of 8 cm within 7 days at 40° C; at first avellaneous, later changing to orange colours. Conidiophores arising from submerged hyphae or sometimes also as [p. 31] branches of the aerial hyphae, up to 100 µm in length and 4-10 µm in diameter. Phialides 11-23 x 2.5-4.0 µm. Conidia ellipsoidal to cylindrical, rounded at both ends, smooth-walled, hyaline to pale brown, 6-10 x 3-6 µm. Chlamydospores absent.

The speces is thermophilic with a temperature optimum at 40° C; minimum about 15° C; maximum about 55° C.

A detailed description is given by Apinis and Chesters (1964) and Stolk (1965)

 

Discussion

 

Paecilomyces crustaceus is characterized by the Thermoascus state and the ellipsoidal to cylindrical conidia. The species is related to P. bysso­chlamydoides and P. variotii (cf. Stolk and Samson, 1972). It appears to be a rather common thermophilic organism.

 

 

Paecilomyces section Isarioidea Samson, sect. nov.

 

Coloniae laete coloratae, albae, flavae, pallide virides, roseae, rubrae vel violaceae. Conidiophora mononematica vel synnematica, plerumque compluries verticillata. Phialides e basi cylindrica vel inflata in collum longum angustatae, conidia in longis catenis divergentibus connexa proferunt. Conidia continua, raro bicellularia, levia vel ornamen­tata, hyalina vel purpurea vel viridia. Chlamydosporae nonnumquam adsunt. Status perfectus in vitro ignotus. Complures species entomogenae. Omnino mesophilae.

 

Species typica Paecilomyces farinosus (Holm ex S. F. Gray) Brown & Smith.

 

Colonies in bright colours, white, yellow, pale green, pink, red or purple. Hyphae hyaline to slightly pigmented, rough- or smooth-walled. Conidio­phores mono- or synnematous, usually consisting of several verticillate branches, each bearing a dense whorl of phialides. Phialides consisting of a cylindrical or swollen basal portion, terminating in a thin often long neck, producing divergent conidial chains. Conidia one- or rarely two-celled, smooth-walled or ornamented, hyaline, purple or green. Chlamydo­spores present in some species.

Perfect state absent in culture; on the natural substrate some species can be associated with Torrubiella or Cordyceps. Several species are entomo­genous. Mesophilic.

Type species: Paecilomyces farinosus (Holm ex S. F. Gray) Brown & Smith.

 

Discussion

In the section Isarioidea several species formerly placed in Isaria with verticillate conidiophores and catenate conidia are accomodated. [p. 32] Paecilomyces marquandii and P. lilacinus form a rather deviating group and might be regarded as representing a distinct section. They fit in the sect. Isarioidea by the verticillate, swollen phialides with a distinct neck, but differ by pigmentation.

The entomogenous species of this section usually produce conspicuous synnemata on the host, which consist of loosely interwoven hyphae ter­minating with powdery conidiogenous structures. In pure culture the synnematous habit is often produced in fresh isolates, but subsequently lost. Cultures grown on media containing starch as carbon source (oatmeal-, cornmeal-, starch- or rice-agar) show a more pronounced synnematous growth and maintain it after several transfers. Also on mealworm-agar or on boiled rice conspicuous synnemata can be observed. Synnematous growth can also be stimulated by keeping cultures in diffuse daylight at 25° C.The length of the synnemata is influenced by the depth of the substrate. Strains which form synnemata of 2 to 3 cm on an insect can produce syn­nemata up to 5 or 6 cm on agar media or on boiled rice.

The hyphae are usually hyaline and smooth-walled. In P. tenuipes and P. cicadae rough-walled hyphae were occasionally observed. The vegetative hyphae of P. marquandii and P. farinosus may contain yellow granula. The conidiophores consist of several verticillate branches with whorls of 2 to 6 phialides. In P. inflatus the phialides are swollen, but mostly occur singly. On the natural substrate (insect) the elements of the verticillate conidiophores are strongly inflated. In pure culture they are still inflated in fresh isolates, but become more slender after repeated subculturing. The conidia produced are usually somewhat larger in pure culture than on the natural substrate. In P. tenuipes conidia may be two-celled in pure culture, whereas they are one-celled on the insect. A yeast-like state occurs in P. viridis. Chlamydospores are observed in P. marquandii, P. penicillatus and P. puntonii. They can be hyaline or brownish (P. puntonii) and are produced in the agar.

According to the literature some species of this section are regarded as imperfect states of the ascomycetous genera Cordyceps and Torrubiella. In the type specimens of Spicaria longipes and S. pulvinata (now consider­ed as synonymous with P. farinosus) perithecia of Torrubella gonylepticida and T. pulvinata were found. Paecilomyces cicadae may be accompanied by Cordyceps sobolifera. However the connection between the imperfect and the perfect states has not yet been proved with cultural experiments.

 

 

10. Paecilomyces farinosus (Holm ex S. F. Gray) Brown & Smith - Fig. 11.

 

Ramaria farinosa Holm - Kgl. Danske Vidensk. Selsk. Skrift. Ny. Saml. I: 279. 1781; Holmskiold - Beata ruris Otia Fungis Danicis a Theodoro Holmskiold impensa 1-2: 94. 1790 = Clavaria farinosa (Holm) Dickson - Plantar. Crypt. Brit. Fasc. 2: 25. 1790 = Corynoides farinosa (Holm) ex S. F. Gray - Nat. Arrang. Brit. Pl., p. 654. 1821 (basionym cited as 'Clavaria farinosa Dickson') = Isaria farinosa (Holm ex S. F. Gray) Fr. [p. 33] - Syst. mycol. 3: 271. 1832 (basionym cited as 'Ramaria farinosa Dickson') = Isaria farinosa (Holm ex S. F. Gray) Fr. - Syst. mycol. 3: 271. 1832 (basionym cited as `Ramaria farinosa Dickson') = Spicaria farinosa (Holm ex S. F. Gray) Vuill. - Bull. trimest. Soc. mycol. Fr. 27: 76. 1911 = Penicillium farinosum (Holm ex S. F. Gray) Biourge - La Cellule 33: 102. 1923 = Paecilomyces farinosus (Holm ex S. F. Gray) Brown & Smith - Trans. Br. mycol. Soc. 40: 50. 1957.

lsaria crassa Pers. - Comment. Fung. clavaeform. p. 99. 1797; ex S. F. Gray - Nat. Arrang. Brit. Pl., p. 562. 1821.

lsaria velutipes Link - Mag. Ges. naturf. Freunde, Berlin 3: 20. 1809; ex S. F. Gray - Nat. Arrang. Brit. Pl., p. 562. 1821.

? lsaria clavata Ditm. - Sturm Deutschl. Fl., Pilze 1: 113. 1817; ex Pers. - Mycol. eur. 1: 46. 1822.

Isaria bicolor Schw. - Trans. Am. Phil. Soc. 4: 305. 1834.

Isaria globosa Schw. - Trans. Am. Phil. Soc. 4: 305. 1834.

Isaria furcata Schw. - Trans. Am. Phil. Soc. 4: 304. 1834­

? Alphitomyces schroeteri Reissek - Sber. Akad. Wiss. Wien. 21: 326. 1856.

Verticillium corymbosum Lebert - Z. wiss. Zool. 9: 445. 1858.

Isaria muscigena Cooke & Muller - Grevillea 16: 81. 1887.

Coremium necans Oudem. - Archs néerl. Sci. 3: 918. 1903.

Isaria mitruliformis P. Henn. - Bot. Jb. 28: 329. 1906.

Spicaria verticilloides Fron - Bull. trimest. Soc. mycol. Fr. 26: 483. 1911 = Spicaria farinosa verticilloides Fron - Bull. trimest. Soc. mycol. Fr. 27: 153. 1912.

Isaria pattersonii Massee - Bull. misc. Inf. bot. Gdns Kew 1912: 358.

Isaria psychidae Pole Evans - Annls mycol. 10: 282. 1912 and Agric. J. Union S. Africa 1912: 35  = Penicillium psychidae (Pole Evans) Biourge - La Cellule 33: 505. 1923.

Spicaria aleyrodis Johnston - Mem. Soc. cubana Hist. nat. 1917: 79.

Coremium swantonii A.L. Smith - Trans. Br. mycol. Soc. 6: 156. 1919 = Spicaria (Isaria) swantonii (A. L. Smith) Petch - Naturalist, Hull 1931: 503.

Isaria cornea Lloyd - Mycol. Writings 6: 919. 1920.

Coremium breve Petch - Trans. Br. mycol. Soc. 11: 259. 1926.

Coremium gracile Petch - Trans. Br. mycol. Soc. 11: 260. 1926 = Spicaria gracilis (Petch) Perch - Trans. Br. mycol. Soc. 16: 241. 1932.

Spicaria longipes Petch - Trans. Br. mycol. Soc. 2I: 51. 1937 = Paecilomyces longipes (Petch) Brown & Smith - Trans. Br. mycol. Soc. 40: 75. 5957.

Paecilomyces subflavus Szilvinyi - Zentbl. Bakt. ParasitKde (Abt. 2) 103: 176. 1941.

Isaria takamizusanensis Y. Kobayasi - Sci. Rep. Tokyo Bunrika Daig. 5: 229. 1941.

Isaria farinosa (Holm ex S. F. Gray) Fr. var. citriformis Y. Kobayasi - Sci. Rep. Tokyo Bunrika Diag. 5: 235. 1941.

Spicaria pulvinata Mains - Mycologia 41: 305. 1949.

Penicillium alboaurantium G. Smith - Trans. Br. mycol. Soc. 40: 485. 1957.

?Isaria moriokaensis Aoki apud Aoki, Nakasato, Fujimoto & Suzuki - Bull. sericult. Exp. Stn Japan 14: 576. 1957.

 

Colonies on malt-agar growing moderately fast, attaining a diameter of 4-6 cm within 24 days at 25° C; consisting of a basal felt from which numerous conidiophores arise, in fresh isolates powdery and sometimes with conspicuous yellow synnemata, in older strains with floccose aerial mycelium; at first white, remaining so or turning bright yellow or cream. Reverse cream to yellow, in fresh isolates often brightly yellow. Odour in distinct. Vegetative hyphae smooth-walled, hyaline or containing yellow granula, 0.7-2.5 µm wide. Conidiophores mainly arising from the submerged mycelium, usually measuring 100-300 x 1.0-2.5 µm, in some strains with a floccose overgrowth arising as sidebranches from aerial hyphae; [p. 34] conidiogenous structures smooth-walled, hyaline, consisting of verticillate branches with whorls of 2 to 4 phialides. Phialides 5-15 x 1.2-2.5 µm, with a swollen basal portion tapering into a distinct neck, about 0.5 µm wide. Conidia ellipsoidal to fusiform, sometimes lemon-shaped, smooth-walled, hyaline, 2.0-3.0 x 1.0-1.8 µm. Chlamydospores absent.

 

 

Fig. 11. Paecilomyces farinosus, a-c. conidiophores, a. on the natural substrate, b-c in pure culture; d-f. conidia, d. Spicaria longipes, e. Spicaria pulvinata, f. various strains.

 

Madelin et al. (1967) observed clavate cells in slide-cultures appressed to the glass surface. These structures may have an appressorium-like function on insects.

On the insect the fungus produces a white mycelial felt from which numerous conidiophores arise in a dense powdery layer; usually several white to yellow synnemata arise from all parts of the insect body. The [p. 35] synnemata have a white clavate and powdery head and sometimes a sterile white to yellow stipe; they are cylindrical to clavate, simple or irregularly branched, 0.5-1.2 cm long, although specimens up to 3 cm are not uncom­mon. The conidiogenous structures are mostly more swollen than in pure culture.

 

According to the literature and to own observations, P. farinosus is an ubiquitous insect pathogen widely distributed throughout the world. The species is polyphagous and is collected from lepidopterous larvae and pupae, Diptera, Homoptera, Coleoptera, Hymenoptera and Arachnida (Petch, 1936; Harris, 1960; Soper and Olsen, 1963; Madelin, 1966 a; Dunbar et al., 1972). P. farinosus is also frequently isolated from forest soils.

Reviews of experiments on the biological control of insects with P. farinosus are given by Steinhaus (1949) and Müller-Kögler (1965).

 

 Material examined

 

Herbarium specimens

 

Numerous specimens on various insects listed under Isaria farinosa in the herbaria B, BPI, BR, CBS, CMI, K, L, PC, PH, U, UPS and others.

Isaria crassa, type, in herb. Persoon, no. 910.258-658 (L).

Isaria velutipes, type, in herb. Persoon, no. 910. 258-357; on lepidopterous pupa collected by Miquel, no. 910227-5 (L).

Isaria clavata, labeled as the type, on chrysalide, ex herb. Saccardo (PAD).

Isaria bicolor, type, in herb. Schweinitz `July 1824, 3072' (PHIL).

Isaria globosa, type, in herb. Schweinitz 'Betlehem, 3067' (PHIL).

Isaria furcata, type, in herb. Schweinitz 'Betlehem, 3068' (PHIL).

Isaria muscigena, type, on Hypnum serpens, collected by C. J. Muller, Eastborne (K).

Coremium necans, original drawings and notes by C. J. Koning (L).

Isaria mitruliformis, type, on removed grass tussocks, Ukinabey; no fungus material, but description and figures on the cover (B).

Isaria pattersonii, type, on scale insect, Newara viridula, collected by W. H. Patterson, West-Indies (K).

Isaria psychidae, type, on Acanthopsyche junodi, collected by I. B. Pole Evans, Natal, 1916 (BPI).

Coremium swantonii, type, on dead wasp (Vespa sylvestris), leg. E. W. Swanton, 1919 (K).

Isaria cornea, type, collected by J. Gossweiler, Angola, ex herb. Lloyd no. 42590 (BPI).

Coremium breve, type, on Lecanium sp., ex herb. Petch (K).

Coremium gracile, type, on a spider, Peradeniya, Ceylon, ex herb. Petch, R 6310 (K) and additional specimens as Spicaria gracilis, ex herb. Petch (K).

Spicaria longipes, type, on a spider, Mauritius, Les Mares; collected by P. Mauret, ex herb. Petch (K).

Isaria takamizusanensis, part of type, on Oncotympana maculaticollis, in herb. Y. Ko­bayasi, Tokyo.

Spicaria pulvinata, type, on an Opilionoidea species, together with Torrubiella pulvinata (MICH).

Isaria farinosa, formae variae, in J. E. Klotzsch, Herb. vivum mycol. no. 1740 (BR).

Isaria columnaris Tul., type of probably undescribed species, `in pupa infossa, Chaville, 1860' ex herb. L. R. Tulasne (PC).

Isaria damaecornu, type of probably undescribed species ex herb. D'Udekem, Belgium (BR).

Numerous specimens labelled as Isaria ssp. in numerous herbaria.

 

Living cultures

 

CBS 142.24, isolated by E. J. Butler from bagworm (S. Africa) and sent as Isaria psychidae.

CBS 250.29 = IMI 14420, isolated by E. W. Mason from Rose Feather Moth, Great Britain.

CBS 240.32, isolated by H. Diddens from insect pupa, Netherlands.

CBS 264.35 = IMI 58411, isolated by F. W. Beekhuis van Till from honey bee, Laren, Netherlands.

CBS 296.38, isolated by F. W. Beekhuis van Till from Philanthus triangulum, Nether­lands.

CBS 282.48, isolated from eggs and sent by M. Geiger-Huber, Basel, Switzerland.

CBS 262.58 = IMI 71622 = QM 7525 = IFO 8108 = ATCC 18236, type culture of Penicillium alboaurantium, isolated from garden soil near Harrow, Middlesex, Great­ Britain.

CBS 265.59, isolated as strain 238/59 from Kotochalia junodi by E. Müller-Kögler, Darm­stadt, Germany.

CBS 334.61, isolated by W. Liese from wood, Germany.

CBS 156.65, isolated by S. Urban from pupa of Thaumatopoea pityocampa, Zagrazden, Bulgaria.

Numerous strains isolated from insects and forest soils in the Netherlands.

 

Discussion

 

The authorship of Isaria farinosa is usually ascribed to Dickson (1790) in accordance with Fries (1832). The fungus was, however, first described by Holm (1781) (cf. Skou, 1967) and validated by S. F. Gray (1821). The species should therefore be cited as Paecilomyces farinosus (Holm ex S. F. Gray) Brown & Smith.

Because of its variable macroscopic habit and its common occurrence, P. farinosus has been described under many different names. Brown and Smith (1957) have already listed numerous synonyms. Some more are added after examination of the type collections and original descriptions. No type material was available of Isaria clavata Ditm., Alphitomyces schroeteri Reissek, Isaria mitruliformis P. Henn. and Isaria moriokaensis Aoki; the descriptions of these species are vague and therefore these names are regarded as possible synonyms.

As already pointed out by Petch (1934), Kobayasi (1941) and Brown and Smith (1957), there is no connection between P. farinosus and Cordy­ceps militaris. Spicaria longipes Petch and S. pulvinata Mains, now re­garded as synonyms of P. farinosus, have been described as the imperfect states of Torrubiella gonylepticida (Petch, 1937) and Torrubiella pulvinata Mains (1949) respectively. Both conidial states differ from the typical P. farinosus strains by the somewhat larger conidia, but the dimensions fall within the species variation. As long as no cultural confirmation exists of the connection between the Spicaria and the Torrubiella state, it should be regarded with caution.

P. farinosus can be distinguished from the other species by its small el­lipsoidal to fusiform conidia and white to yellow colonies.

The synnemata of P. farinosus are sometimes parasitized by Melanospora parasitica Tul. or by Tritirachium isariae (Petch) de Hoog (1973). [p. 37]

 

11. Paecilomyces amoeneroseus (P. Henn.) Samson, comb. nov. - Fig. 12.

 

Isaria amoene-rosea P. Henn. - Hedwigia 41 (Beiblatt): 66. 1902 (basionym).

? Coremium pulcherrimum Petch - Trans. Br. mycol. Soc. 11: 259. 1926 = Spicaria (Isaria) pulchella Petch - Annls Cryptog. exot. 6: 234. 1933 (as nom. nov.) [non Isaria pulcherrima Berk. 1875].

 

Colonies on malt-agar growing slowly, attaining a diameter of 2.5-3 cm within 14 days at 25°C, consisting of a floccose mycelium with powdery localized areas with numerous conidiophores; at first white, sometimes remaining so, but usually quickly changing to pink and red colours near Light Congo Pink (Ridgway, Pl. 28; Rayner, 7"d), Orange Vinaceous and Light Corinthian Red (Ridgway, Pl. 27; Rayner, 5"b, 3"b); in fresh isolates some conspicuous red synnemata, 8-10 mm long and 300-800 µm in diameter, are produced which disappear after some transfers. Reverse cream to red. Odour indistinct. Vegetative hyphae hyaline to reddish, smooth-walled, 1.5-2.5 µm wide. Conidiophores usually arising from the sub­merged mycelium, erect, 90-150 x 2.0-2.5 µm, consisting of complex verti­cillate branches with whorls of 2 to 4 phialides. Phialides 4.0-7.5 x 1.5-3.0 µm, consisting of a globose basal portion, tapering into a thin neck, 0.3-0.5 µm wide. Conidia small, subglobose to ellipsoidal or sometimes irregularly cylindrical, 2.5-3.5 x 1.7-2.2 µm, hyaline, smooth-walled.

 

 

Fig. 12. Paecilomyces amoeneroseus, a-c. conidiophores, a-b. on the natural substrate, c. in pure culture; d-e. conidia, d. on the natural substrate, e. in pure culture.

 

On the insect the fungus produces red synnemata up to 5 mm long and 300-500 µm in diameter, usually branched at the apex, appearing powdery [p. 38] due to the conidia. In dried condition the stipe is mostly red coloured near Ferruginous or Carrot Red (Ridgway, Pl. 14; Rayner, 9'i, 7’b). Conidiogenous structures are densely branched and swollen.

 

Material examined

 

Herbarium specimens

 

Isaria amoene-rosea, type, 'Aprillo 1897, Rio de Janeiro, Tijuca, Brasilia, in chrysalidibus, leg. E. Ule' ex herb. P. Hennings (B).

Coremium pulcherrimum = Spicaria (Isaria) pulchella Petch, nom. nov., mission R. De­cary, det. T. Petch (PC).

Collections by H. C. Evans, Ghana 1971-1972: R.S. 220 on Chelonethidae species (Pseudoscorpion); R.S. 226, on Stictococcus sjostedi (Homoptera); R.S. 234 on adult Coleoptera; R.S. 221 on Phrynoponera gabonensis (Ponerinae); R.S. 235 on Anomma species (Dorylinae); R.S. 236 on parisitic wasp; R.S. 233 on Crematogaster gabonensis; R.S. 229, 240, 174, 091 on lepidopterous larvae and pupae; R.S. 179 on unidentified Coleoptera (CBS).

 

Living cultures

 

CBS 107.73, from coleopterous pupa; CBS 729.73, from Nitidulidae species; CBS 730.73, from lepidopterous larva; CBS 737.73, from Pheidole species; CBS 738.73, from coleopterous larva, all isolated by H. C. Evans in Ghana.

 

Discussion

 

According to the description, Coremium pulcherrinum Petch may be identical with Paecilomyces amoeneroseus. The synonymy is, however, considered as doubtful, because the type material (in PC) proved to be sterile.

P. amoeneroseus is a common species in Ghana, occurring on various insects. The species is closely related to P. farinosus and P. fumosoroseus, but differs by its ellipsoidal conidia and the reddish colonies. On insects P. amoeneroseus is easily recognizable by the bright red synnemata.

 

 

12. Paecilomyces fumosoroseus (Wize) Brown & Smith - Fig. 13.

 

Isaria fumosorosea Wize - Bull. int. Acad. pol. Sci. Lett. (Classe Sci. math. nat.) 1904: 72 = Spicaria fumosorosea (Wize) Vassiljevski - Morbi Plant. p. 146. 1921 = Paecilo­myces fumosoroseus (Wize) Brown & Smith - Trans. Br. mycol. Soc. 40: 67. 1957.

Spicaria aphodii Vuill. - Bull. Séanc. Soc. Sci. Nancy, Sér. 3, 11: 24. 1910 = Penicil­lium (Spicaria) aphodii (Vuill.) Biourge - La Cellule 33: l00. 1923.

Spicaria cossus Portier & Sartory - C. r. Séanc. Soc. Biol. 79: 700. 1916. = Paecilo­myces cossus (Portier & Sartory) Brown & Smith - Trans. Br. mycol. Soc. 40: 74. 1957.

Paecilomyces hibernicus Kennelly & Grimes - Scient. Proc. R. Dubl. Soc. 19: 513. 1930.

Paecilomyces isarioides Inagaki - Trans. mycol. Soc. Japan 4: 2. 1962. [p. 39]

 

 

Fig. 13. Paecilomyces fumosoroseus, a-d. conidiophores, a-b. on the natural substrate, c-d. in pure culture; e-h. conidia, e. on the natural substrate, f-h in pure culture, h. Ghanaian strains.

 

Colonies on malt-agar growing moderately fast, attaining a diameter of about 4 cm within 14 days at 25°C, consisting of a basal felt with raised floccose overgrowth, powdery when freshly isolated; in some isolates pink synnemata occur which may be branched; white at first, remaining so or changing to pink shades, especially when sporulating abundantly. Reverse uncoloured or yellow. Odour absent. Vegetative hyphae smooth-walled, hyaline, 1.5-3.5 µm wide. Conidial structures mostly complex, consisting of erect conidiophores arising from submerged or laterally from aerial hyphae. Conidiophores mono- or synnematous, up to 100 µm in length, 1.5-2 µm wide, [p. 40] smooth-walled, hyaline, consisting of verticillate branches bearing whorls of 4 to 6 phialides. Phialides 5.7-8 x 1-2 µm, with a globose or el­lipsoidal basal portion which tapers into a long distinct neck, about 0.5 µm wide. Conidia cylindrical to fusiform, smooth-walled, hyaline to slightly pink, 3-4 x 1-2 µm. Chlamydospores absent.

On insects the fungus produces mononematous conidiophores or distinct but loose synnemata. Synnemata erect, branched, appearing pink and powdery due to the conidia, up to 3 cm in length and 400 µm in diameter. Conidiogenous elements inflated. Conidiophores 2.5-4 µm wide; verticillate branches more or less globose or broadly cylindrical, 4-9 x 3-4.5 µm. Phialides with a globose basal portion, 2.3-3.5 µm wide.

 

Paecilomyces fumosoroseus appears to be rather common on various in­sects (Vassiljevski, 1929; Siemaszko, 1937; Rozsypal, 1930; Madelin, 1966a). Sheng et al. (1965) reported a rather wide host range in China: Luciana separata, Pyrausta nubilalis, Carposina niponensis and various flies. P. fumosoroseus is also rather common on silkworms (Bombyx mori) in Japan (Aoki et al., 1951). Aoki (1967) and Aoki and Yanase (1970a and b) investigated the physiology of P. fumosoroseus in connection with its oc­currence on silkworms.

 

Material examined

 

Herbarium specimens

 

Paecilomyces fumosoroseus, several specimens on various insects in K and IMI; on un­identified insect, collected by J. Daams, Kortenhoef, Netherlands (CBS). Collections by H. C. Evans, Ghana 1972: R.S. 0034, on Stictococcus species; R.S. 0224, on Diptera species; R.S. 0225, 0231, 0239 on lepidopterous larvae; R.S. 0223 on mealy bugs; R.S. 0228 on hymenopterous pupae; R.S. 0237 on Anachetus africanus; R.S. 0230 on Cataulacus guineensis (CBS).

Isaria leprosa, on insect pupa, Mulgrave Woods, 1930, ex herb. T. Petch, R 726 (K).

 

Living cultures

 

CBS 107.10 = IMI 58433, received as Spicaria aphodii from P. Vuillemin, probably type culture.

CBS 243.31 = IMI 58439, isolated from Loxostege sticticollis, received from A. Bondar­zew, Leningrad.

CBS 244.31 = IMI 58413, type culture of Paecilomyces hibernicus, isolated from Irish butter.

CBS 337.52, isolated by C. Vago from cocoon of Thaumatopoea pityocampa and sent by J. Nicot (Paris).

CBS 339.54, isolated from sputum, Netherlands.

CBS 264.58, isolated from air, Netherlands.

CBS 309.59, isolated by E. Müller-Kögler from cocoon of Thaumatopoea pityocampa, Darmstadt, Germany.

CBS 106.66, isolated by K. H. Veen from Melolontha melolontha, La Miničre, Versailles, France.

CBS 101.73, isolated by J. H. van Emden from agricultural soil, Wageningen, Netherlands. [p. 41]

CBS 721.73A from lepidopterous larvae; CBS 721.73B from mealy bugs; CBS 721.73C from hymenopterous pupa; CBS 721.73D from Anachetus species, collected and isolated by H. C. Evans, Ghana.

CBS 37S.70 = IFO 7562, type culture of Paecilomyces isarioides, isolated from wood in Japan.

 

Discussion

 

The synonymy of P. hibernicus Kennelly & Grimes and Spicaria aphodii Vuill. is discussed by Brown and Smith (1957). These authors considered Spicaria cossus Portier & Sartory as a distinct species, because of the white to pale cream colonies and the more complex conidiogenous structures. Since the typical pink colour of P. fumosoroseus can be absent, especially when the cultures are grown in darkness, and since complex conidiophores are typical in fresh isolates, P. cossus is considered as a further synonym. Petch (1933a) remarked on a specimen which he assigned to Isaria leprosa Fr., that it may be an Isaria farinosa parasitized by a Sporotrichum; this specimen (in K) was found to be P. fumosoroseus. Unfortunately no type material of Isaria leprosa is left (in UPS) and the species is doubtful.

Paecilomyces fumosoroseus is distinguished from the other species by its pink colour and the cylindrical conidia, measuring 3-4.6 x 1-2 µm. In the specimens collected in Ghana the conidia are of somewhat different shape; they are cylindrical to narrowly fusiform, intermediate between P. fumo­soroseus and P. javanicus. The specimens and the pure culture from Ghana have the typical pink colour of P. fumosoroseus.

Beside its occurrence on insects, P. fumosoroseus has been frequently isolated from soil (Domsch and Gams, 1972) and from food-stuffs.

 

 

13. Paecilomyces javanicus (Friederichs & Bally) Brown & Smith - Fig. 14.

 

Spicaria javanica Friederichs & Bally - Meded. Koffiebessenboeboek-Fonds 6: 146. 1923 = Paecilomyces javanicus (Friederichs & Bally) Brown & Smith - Trans. Br. mycol. Soc. 40: 65. 1957.

Spicaria formosensis Sawada - Spec. Publ. Coll. Agric. natn. Taiwan Univ. 8: 188. 1959.

 

Colonies on malt-agar growing slowly, attaining a diameter of 3 cm in 14 days at 25° C, consisting of a basal felt with a floccose to cottony overgrowth, in fresh isolates more powdery, occasionally producing a few synnemata of about 2 cm in height. At first white, in age becoming cream. Odour and exudate lacking. Reverse uncoloured to yellow. Vege­tative hyphae smooth, septate, hyaline, 0.5-2.2 µm wide. Conidial structures consisting of erect conidiophores arising from the aerial hyphae, up to 50 µm in length and 1.5-2.5 µm in diameter, or solitary phialides borne [p. 42] directly on trailing hyphae; conidiophores forming verticillate branches with phialides in whorls of 2 to 3. Phialides 8-14 x 2-2.8 µm, consisting of a cylindrical basal portion, tapering into a thin distinct neck, 2-4.5 µm in length and 0.7-1.4 µm in diameter. Conidia usually cylindrical to fusiform, smooth-walled, hyaline, 5-7.4 x 1.4-1.7 µm. No chlamydospores observed. On the insects the fungus usually produces a white powdery felt with numerous conidiophores. The conidiophores are usually inflated with almost globose phialides. The conidia are somewhat smaller than on agar media and fusiform, measuring 3-3.5 x 1.2-1.7 µm.

 

 

Fig. 14. Paecilomyces javanicus, a-b. conidiophores; c. conidia.

 

Material examined

 

Herbarium specimens

 

Spicaria javanica, several specimens collected on various insects in Ceylon by T. Petch (K).

Paecilomyces javanicus, R.S. 0170, on lepidopterous pupa and cocoons, collected by H. C. Evans in Begoro, Ghana (CBS).

 

Living cultures

 

CBS 134.22 = type culture, sent by W. Bally and isolated from Stephanoderis hampei, Java, Indonesia.

CBS 174.25, isolated from pupa of Brachartona species in Bogor, Indonesia, by M. B. Schol-Schwarz.

CBS 994.73 and CBS 995.73, both isolated from lepidopterous pupae by H. C. Evans, Ghana (R.S. 0170). [p. 43]

 

Discussion

 

Sawada (1959) described Spicaria formosensis as occurring on larvae of Ocinara varians and Archips ingentara and pupae of Euproctis staudingeri in Taiwan. No material was available for study, but from the description the fungus is regarded as P. javanicus.

The conidiogenous structures of Paecilomyces javanicus in pure culture differ from those produced on the natural substrate. These structures are more complex and inflated, producing globose phialides. Although Paecilo­myces javanicus is usually mononematous on insects, synnematous coni­diogenous structures were observed in CBS 994.73 in pure culture. The synnemata are white to yellow and about 2 cm high. The phialides produce cylindrical to fusiform conidia. These characters show that P. javanicus is closely related to P. tenuipes. The latter species differs, however, by the curved, cylindrical conidia and the synnematous habit, whereas the conidia of P. javanicus are straight and usually fusiform.

 

 

14. Paecilomyces reniformis Samson & Evans, spec. nov. - Fig. 15.

 

Hospes a mycelio tenui, hyalino, levi obtectus; hyphae 2.5-4 µm crassae, multa conidio­phora viridia proferunt. Conidiophora erecta, mononematica, 75-120 x 2-4.5 µm, hyalina, levia, ramulos et quoties binas ad ternas phialides ferunt. Phialides 4.5-6 x 3.7-5 µm, e basi inflata globosa in collum 0.5-1 µm crassum angustatae. Conidia continua, reniformia, levia, hyalina, 4.5-5.5 x 2.0-3.5 µm.

Typus R.S. 0029, in Tettigoniida (locusta), lectus a H. C. Evans, Tafo, in Ghana (CBS).

 

 

Fig. 15. Paecilomyces reni f ormis, a-b. conidiophores; c. conidia.

 

The description is based on herbarium material:

Mycelium covering the host with thin-walled, hyaline, smooth-walled hyphae, 2.5-4 µm wide, from which numerous conidiophores arise, giving a pale green appearance. Conidiophores erect, mononematous, 75-120 x 2-4.5 µm, hyaline, smooth-walled, bearing short branches with whorls of 2 to 3 phialides or phialides directly on the stalk of the conidiophores in whorls at the septa. Phialides 4.5-6 x 3.7-5 µm, consisting of a swollen globose basal portion, tapering into a thin neck, 0.5-1 µm wide. Conidia one-celled, reniform, smooth-walled, hyaline, 4.5-5.5 x 2-3.5 µm.

 

Material examined

 

Paecilomyces reniformis, type, R.S. 0029, on a grasshopper (Tettigoniidae, Orthoptera), collected by H. C. Evans, Tafo, Ghana (CBS).

 

Discussion

 

The conidiogenous structures of Paecilomyces reniformis are reminiscent of those of Paecilomyces tenuipes, differing by the pale green colour and the less compact conidiophores. Moreover, the phialides occur also directly in whorls on the conidiophore stalk. P. reniformis resembles also Isaria amorpha Höhnel (1909) (= Isaria orthopterorum Petch (1933 b)), which has sympodial conidiogenous cells (de Hoog, 1972).

P. reniformis appeared to be a rare species in Ghana, since it was only collected once. Unfortunately isolation on agar media failed, so that the species could be examined only from the type specimen.

 

 

15. Paecilomyces ramosus Samson & Evans, spec. nov. - Fig. 16.

 

Synnemata erecta, cylindrica, 2.5-5.0 cm, nonnumquam ad 6-7 cm longa, 0.3-1 mm cras­sa, recentia aurantia, sicca lutea, pulverulenta alba, frequenter ramosa. Conidiophora plerumque secundum synnema sparsa, 45-90 µm longa, 2.5-3 µm crassa, compluries ver­ticillata, quoties binas ad quaternas phialides ferunt. Phialides 11-20 x 2.5-3.5 µm, e basicylindrica subito in collum longum, 0.7-1.2 µm crassum angustatae. Conidia pyriformia, apiculata, hyalina, levia, 3.2-4.5 x 1.5-2.7 µm.

Typus R.S. 0177, in larvis chrysalidibusque lepidopterorum, Mt. Atewa, Begoro, in Ghana, lectus a H. C. Evans, 1971 (CBS).

 

The description is based on herbarium material:

Synnemata erect, measuring 2.5-5.0 cm in length, but sometimes up to 6 or 7 cm, 300-1000 µm in diameter, orange when freshly collected, yellow when dried, with white powdery heads, cylindrical with many side branches. Conidiophores mostly scattered along the synnema, 45-90 µm in length and 2.5-3 µm in diameter, consisting of some verticillate branches [p. 45] with whorls of 2 to 4 phialides. Phialides 11-20 x 2.5-3.5 µm, consisting of a cylindrical portion, tapering abruptly into a long neck of 0.7-1.2 µm. Conidia pyriform, apiculate, hyaline, smooth-walled, 3.2-4.5 x 1.5-2.7 µm.

 

 

Fig. 16. Paecilomyces ramosus, a-b. conidiophores; c. conidia; d. synnema.

 

Material examined

 

Paecilomyces ramosus, type, R.S. 0177, on lepidopterous pupae and larvae, Mt. Atewa, Begoro, Tafo; R.S. 0178, 0182 and 094 on lepidopterous pupae and larvae, Atewa, Begoro.

All specimens collected by H. C. Evans, Ghana, 1972 (CBS).

 

Discussion

 

The specific epithet of the new species refers to the typically branched synnemata. The conidiophores of P. ramosus resemble strongly those pro­duced in the genus Penicillium. The species is, however, placed in Paecilo­myces because of the white colour, the synnematous habit, and the phialides which terminate into a long thin neck. [p. 46]

 

 

16. Paecilomyces ghanensis Samson & Evans, spec. nov. - Fig. 17.

 

Coloniae in agaro maltoso lente crescunt, ad 2 cm diam. 14 diebus 25° C, albae, deinde cremeae vel flavo-brunneae; in coloniis nuper isolatis synnemata conspicua flavo-brunnea, 4-15 mm longa, 0.5-1 mm crassa formantur. Conidiophora erecta, hyalina, levia, 90-180 x 2.5-3.5 µm, compluries verticillata, ramis phialidibus quoties binis ad quaternis terminatis. Phialides 5.5-8 x 2-3.5 µm, e basi inflata, fere ellipsoidea in collum tenerum, 0.5-0.75 µm crassum, angustatae. Conidia continua, late fusiformia, hyalina, levia, 3.5-5.0 x 2.5­-3.0 µm.

Typus R.S. 0096, in chrysalide lepidopterorum, Mt. Atewa in Ghana, lectus a H. C. Evans, 1972 (CBS); vivus CBS 107.73.

 

Colonies growing restrictedly on malt-agar, attaining a diameter of about 2 cm within 14 days at 25° C, consisting of a white floccose mycelium and some powdery localized areas, where the conidiophores arise from the submerged mycelium; in age becoming cream or yellow-brown; in fresh isolates conspicuous yellow-brown synnemata, 4-15 mm long, 0.5-1 mm in diameter are produced. Reverse uncoloured. Odour absent. Vegetative hyphae hyaline, smooth-walled, 1.0-2.5 µm wide. Conidiophores erect, hyaline, smooth-walled, 90-180 x 2.5-3.5 µm, consisting of complex ver­ticillate branches with whorls of 2 to 4 phialides. Phialides 5.5-8 x 2-3.5 µm, consisting of a swollen, more or less ellipsoidal basal portion, tapering into a thin neck, 0.5-0.75 µm wide. Conidia one-celled, broadly fusiform, hyaline, smooth-walled, 3.5-5.0 x 2.5-3.0 µm. On oatmeal-agar showing similar structures, but here sporulation richer and reverse yellow.

 

On the insect the fungus produces one long, yellow-brown synnema, 15-30 mm long and 300-500 µm in diameter, with a sterile stipe, terminating in a branched powdery yellow head of numerous conidiophores. Conidio­phore structure similar as in pure culture, but all parts of the conidio­phores more inflated; the phialides have a more globose basal portion.

 

 

Fig. 17. Paecilomyces ghanensis, a-b. conidiophores; c. conidia. [p. 47]

 

Material examined

 

Herbarium specimens

Paecilomyces ghanensis, type, R.S. 0096, on lepidopterous pupa, collected by H. C. Evans at Mt. Atewa, 1972; R.S. 0184, on lepidopterous pupa, collected in Nobi, Ghana, 1972 (CBS).

 

Living cultures

 

CBS 105.73, isolated from type specimen R.S. 0096; CBS 996.73 from R.S. 0184.

 

Discussion

 

Paecilomyces ghanensis differs from similar species in shape and size of the conidia. The synnematous habit resembles P. tenuipes. The species seems to be uncommon.

 

 

17. Paecilomyces coleopterorum, Samson & Evans, spec. nov. - Fig. 18.

 

Coloniae in agaro maltoso lente crescunt, ad 2.5 cm diam. 14 diebus 25° C; in coloniis nuper isolatis synnemata conspicua formantur; primum albae, deinde cremeae vel flavo-brunneae. Conidiophora erecta, plerumque mononematica, e hyphis submersis oriuntur, 75-100 x 2.0-2.5 µm, compluries verticillata quoties binas ad quaternas phialides ferunt. Phialides 9-18 x 2.0-3.2 µm, e parte basilari cylindrica in collum 0.5-0.8 µm crassum angustatae. Conidia late ellipsoidea vel cylindrica, hyalina, levia, 5.7-8.5 x 2.5-3.0 µm.

Typus R.S. 0172A, in larva coleopterorum, Tafo in Ghana, lectus a H. C. Evans, 1971 (CBS); vivus CBS 110.73 praeservatur.

 

Colonies on malt-agar growing slowly, attaining a diameter of about 2.5 cm within 14 days, consisting of a raised floccose mycelial felt, which becomes powdery when sporulating, in fresh isolates showing conspicuous synnemata, which are lost after several transfers, at first white, turning creamish to yellow-brown with age. Reverse uncoloured. Odour absent. Vegetative hyphae hyaline, smooth-walled, 0.7-3.5 µm wide. Conidiophores erect, synnematous in fresh isolates, but usually mononematous, arising from the submerged mycelium, 75-200 x 2-2.5 µm, consisting of verticillate branches, with whorls of 2 to 4 phialides. Phialides 9-18 x 2.0-3.2 µm, consisting of a cylindrical basal portion, tapering into a thin neck of  0.5-0.8 µm width. Conidia broadly ellipsoidal to cylindrical, hyaline, smooth-walled, 5.7-8.5 x 2.5-3.0 µm.

 

On larvae the fungus produces several long cylindrical, simple synnemata, which are yellow-brown with a white to yellow head of numerous conidiogenous structures. Synnemata of variable size, usually measuring 1-8 mm x 150-300 µm, sometimes up to 3 cm in length. Conidiogenous [p. 48] structures similar to those produced in pure culture, but conidiophores mostly more inflated and phialides with a swollen globose basal portion. Conidia broadly ellipsoidal, measuring 4.5-5.5 x 2.5-3-3 µm.

 

 

Fig. 18. Paecilomyces coleopterorum, a-d. conidiophores, a-b. on the natural substrate, c-d. in pure culture; e-f. conidia, e. on the natural substrate, f. in pure culture.

 

Material examined

 

Herbarium specimens

 

Paecilomyces coleopterorum, type, R.S. 0172 A, on coleopterous larva, collected in Tafo; additional specimens R.S. 0076 and 0172 B-E, collected in Tafo and Begoro by H. C. Evans, Ghana, 1972 (CBS).

 

Living cultures

 

CBS 110.73, isolated by H. C. Evans from the type specimen R.S. 0172.

CBS 102.73, received as Isaria species from Mme J. Nicot, isolated by L. Faurel (1966) from coleopterous larva (Lampyride species), La Maboké, Republic of Central Africa.

 

Discussion

 

Paecilomyces coleopterorum shows affinities to P. tenuipes and P. cicadae. It can be distinguished by its broadly ellipsoidal conidia, while in the other species the conidia are cylindrical, usually curved and of differing sizes.

 

 

18. Paecilomyces tenuipes (Peck) Samson, comb. nov. - Fig. 19.

 

? Isaria truncata Pers. - Comment. Fung. clavaef., p. 99. 1797; ex Pers. - Mycol. eur. 2: 44. 1822.

Isaria tenuipes Peck - Rep. N. Y. St. Bot. 1879: 49 (basionym).

Isaria arachnicida Speg. - An. Mus. nac. Hist. nat. B. Aires 6: 345. 1889.

Isaria argentina Speg. - An. Mus. nac. Hist. nat. B. Aires 6: 345. 1889.

Isaria suffruticosa Cooke & Massee - Grevillea 19: 45. 1890.

Isaria dussii Pat. - Bull. Soc. mycol. Fr. 16: 187. 1900.

Isaria palmatifida P. Henn. - Hedwigia (Beiblatt) 41: 65. 1902.

Isaria ochracea Boudier - Bull. Soc. mycol. Fr. 29: 197. 1903 = Spicaria ochracea (Bou­dier) Vuill. - Bull, trimest. Soc. mycol. Fr. 27: 76. 1911 = Penicillium (Spicaria) ochra­ceum (Boudier) Biourge - La Cellule 33: 104. 1923.

Gibellula phialobasia Penz. & Sacc. - Icon. Fung. javan., p. 108. 1904.

Gibellula eximia Höhnel - Sber. k. Akad. Wiss. Wien 83: 37. 1907.

Isaria japonica Yasuda - Bot. Mag. Tokyo 29: 117. 1915.

Isaria peckoltii Lloyd - Mycol. Writings 6: 884. 1919.

Spicaria heliothis Charles - Phytopathology 28: 97. 1938 = Paecilomyces heliothis (Charles) Brown & Smith - Trans. Br. mycol. Soc. 40: 75. 1957.

Isaria yokohamensis Y. Kobayasi - Sci. Rep. Tokyo Bunrika Daig., Sect. B, 5: 237. 1941.

 

Colonies on malt-agar growing rather slowly, attaining a diameter of 2 to 3 cm within  14  days at 25° C, consisting of a thin mycelial felt with numerous conidiophores giving the colony a powdery appearance; at first white, remaining so or turning cream, or quickly changing to buff near Vinaceous Buff (Ridgway, Pl. 40; Rayner, 17"'d) or Pinkish Buff (Ridgway, Pl. 23; Rayner, 17"d); in fresh isolates conspicuous synnemata are produced, which can reach 3 to 5 cm in length. Reverse cream to yellow. Odour absent or mushroom-like. Vegetative hyphae smooth-walled, hyaline to yellow, 0.5-3.0 µm wide. Conidiophores arising primilarly from submerged hyphae, smooth-walled, hyaline, erect, mononematous or synnematous, 90-120 x 2.5-4.0 µm, consisting of verticillate branches with whorls of 2 to 6 phialides. Phialides 4.5-6.2 x 2.5-3.2 µm, consisting of a globose basal portion, tapering into a thin neck about 0.5 µm wide. Conidia cylindrical, mostly curved, smooth-walled, hyaline, in some strains strictly one-celled, measuring  3-7.5 x 2.0-2.5 µm, in other strains one- or two-celled, 6-12 µm long.

 

On the insect the fungus produces conspicuous synnemata which are simple, furcate or irregularly branched, usually 1-3.5 cm in length [p. 50] and 200-500 µm in diameter, but they may be up to 5 cm long; when freshly collected yellow, sometimes with a yellow-green stipe; in older specimens the stipe is mostly yellow-brown to brown and terminates in a white to yellow fertile head. Hyphae are occasionally rough-walled. Conidiogenous structures densely branched and swollen. The phialides have a globose basal portion and a thin neck. Conidia cylindrical, slightly to strongly curved, measuring 2.0-7.0 x 1.0-2.5 µm.

 

 

 Fig. 19. Paecilomyces tenuipes, a-c. conidiophores, a-b. on the natural substrate, c. in pure culture; d-h. conidia, d. type, e. Isaria japonica, f. Isaria peckoltii, g. CBS 104.72, h. CBS 226.60. [p. 51]

 

Petch (1933b) and Kobayasi (1941) assumed that Isaria dussii Pat., I. palmatifida P. Henn. and Gibellula eximia Höhnel, now considered as synonyms of P. tenuipes, are the imperfect state of Cordyceps polyarthra Möller.

 

Material examined

 

Herbarium specimens

 

Isaria truncata in herb. Persoon, labelled 'Beauv. in herb. Pers. Isaria truncata Syn. Fung.? Clavaria farinosa Dickson ?var.' no. 910.258-668 (L); in C. Roumegučre, Fungi selecti exs. no. 4395. ‘I. truncata Pers., I. farinosa var. truncata Fr., sur chrysalide dans les bois, Montech., France’ (BR).

Isaria tenuipes, type, collected by C. H. Peck, Center, USA (NYS); on unidentified in­sect, collected in the Boggy Woods, vicinity of Picairn, St. Lawrence County, 23960 (NYS); on lepidopterous pupa, Percy Warner Parks, Nashville, leg. F. T. Wolf and W. T. Allman, 754 (MICH); on lepidopterous pupa, collected by M. Lange, 26250 (MICH); some specimens identified by T. Petch, collected on lepidopterous pupae from different localities in Great Britain (K); on a larva collected by H. A. van der Aa, Esbeek, Nether­lands, R.S. 397 (CBS); on lepidopterous pupae and larvae, collected by H. C. Evans in Begoro, Atewa, Nobi, Ghana, R.S. 0095, 0180, 0184 (CBS).

Isaria arachnicida, type, labelled `s. Araneas, Buenos Aires, Santa Catalina, 1889' leg. C. Spegazzini, no. 11212 (LPS).

Isaria argentina, type, labelled `s. crisalida de Noctuidea? La Plata', leg. C. Spegazzini, no. 11205 (LPS) and additional specimen labelled `s. lepidoptero, Paraguay, Asuinaino, 1919', no. 1,206 (LPS).

Isaria palmatifida, type, `auf toten Puppen, Foresta da Tijuca' (B).

Gibellula eximia, 'auf Schmetterlingspuppen, Java, Buitenzorg, 1907-1908', ex herb. von Höhnel (K).

Isaria japonica, type, on pupa, Sendai, leg. A. Yasuda, ex herb. Lloyd no. 42616; on moth, leg. E. Yakushiji & E. Kumagawa, Botanic Gardens Koishikawa, 1929, Japan; on pupa of lepidoptera, Tan-Hsien, Hainan, leg. S. C. Teng, 1934 (BPI).

Isaria peckoltii, type, leg. C. Peckolt, Rio de Janeiro, Brazil, ex herb. Lloyd, no. 42589 (BPI).

Isaria suffruticosa, type, leg. A. R. Crawford, Australia (K).

Spicaria heliothis, type, on Heliothis obsoleta from hibernation cages, collected by F. F. Dicke, Arlington Exp. Farm, 1937, det. V. K. Charles (BPI); in addition dried cultures of the type on different media, and two specimens collected in 1937 and 1938 in Hoops­town, Ill., USA (BPI).

Isaria yokohamensis, part of type, on lepidopterous pupa, Yokohama City, Japan, ex herb. Y. Kobayasi, Tokyo.

Isaria elegans, type of probably unpublished species in herb. Arnaud, labelled `sur chry­salide de lépidoptčre, Bois de Metz, Versailles, 1946' no. 1723 (PC).

Isaria mattiroliana, type of a probably unpublished species, in herb. Sacc., labelled `in hort. bot. Aug. 1917, aff. I. farinosa' (PAD).

Isaria farinosa var. prolifera-ramosa de Thümen, Mycoth. univ. 2080, leg. J. Therry (L, BR); in Roumeguére, Fungi Gall. exs. 1361, leg. J. Therry (L). as Isaria crassa, several specimens collected in the Netherlannds and Belgium (L, BR). as Isaria farinosa and Isaria spp., numerous specimens in herbaria BPI, BR, L, LPS, U, UPS and others.

 

Living cultures

 

CBS 226.60, isolated by E. Müller-Kögler from Tipula larva as strain 325, Darmstadt, Germany. [p. 52]

CBS 109.73, isolated from Bombyx mori in Japan as strain 35 and sent as Spicaria species by K. Yanase (see also Aoki, 1971).

CBS 104.73 and CBS 739.73, isolated by H. C. Evans from lepidopterous larvae in Ghana.

CBS 997.73, isolated from lepidopterous larva, collected by H. A. van der Aa in Es­beek, Netherlands.

 

Discussion

 

Isaria tenuipes is transferred to Paecilomyces on account of the verticillate conidiophores and the phialidic conidiogenous cells with catenate conidia. Isaria truncata Pers. (1797) may be an older epithet but from the description its systematic position is uncertain. Persoon (1797) probably regarded I. truncata as a variety or deviating form of I. farinosa, because when describing it he named Ramaria farinosa Holmsk. as a synonym. A specimen labelled 'Isaria truncata Syn. Fung?' in herb. Persoon (L) is identical with P. tenuipes, but Persoon's notes on the cover indicate that Persoon doubted the identity of this specimen with his own collection (now lost). Isaria truncata is therefore considered as a doubtful synonym.

Most synonyms could be ascertained by studies of the type collections. The type specimens of Isaria peckoltii Peck and I. palmatifida P. Henn. showed somewhat smaller conidia, but fit within the given range of P. tenuipes. No material of I. ochracea Boudier, I. dussii Pat. and Gibellula phialobasia Penz. & Sacc. was available, but from the original description their identity is obvious.

In some strains (e.g. CBS 226.60, 997.73) two-celled conidia are pro­duced. Separation into two species was considered, but in subcultures of a recently collected specimen (R.S. 0397) with one-celled conidia, two-celled as well as one-celled conidia were produced.

On insects P. tenuipes may produce synnemata similar to P. farinosus and the species is therefore often confused with this species. Microscopically P. tenuipes can easily be recognized by the cylindrical, usually curved and larger conidia.

 

 

19. Paecilomyces cicadae (Miquel) Samson, comb. nov. - Fig. 20.

 

Isaria cicadae Miquel - Bull. Sci. phys. nat. néerl. 1838: 86 (basionym) = Cordyceps cicadae (Miquel) Massee - Ann. Bot. 9: 38. 1895.

? Sphaeria basili Taylor - New Zealand and its Inhabitants, p. 424. 1844 = Isaria basili (Taylor) Y. Kobayasi - Sci. Rep. Tokyo Bunrika Daig., Sect. B, 5: 244. 1941.

Sphaeria sinclairii Berk. - Fl. New Zealand  2: 338. 1855 = Cordyceps sinclairii (Berk.) Sacc. - Syll. Fung. 2: 577. 1883 = Isaria sinclairii (Berk.) Lloyd - Mycol. Writings 7: 1179. 1923.

Torrubia caespitosa Tull. - Set. Fung. Carpol. 3: 11. 1865 = Cordyceps caespitosa (Tul.) Sacc. - Michelia 1:320. 1878.

Isaria arbuscula Hariot - Bull. Soc. mycol. Fr. 8: 67. 1893.

Isaria hariotii Arnaud - Bull. trimest. Soc. mycol. Fr. 31: 20. 1915.

Isaria mokanshawii Lloyd - Mycol. Writings 5: 854. 1919. [p. 53]

Isaria cosmopsaltriae Yasuda - Bot. Mag. Tokyo 32: 263. 1918 - and Lloyd - My­col. Writings 5: 806. 1918.

? Isaria cryptotympanae Sawada - Spec. Publ. Agric. natn. Taiwan Univ. 8: 232. 1959.

 

The description is based on herbarium material:

 

Synnemata erect, simple or irregularly branched with branching near the apex, often clavate, growing from a white to pale yellow mycelium which covers the host, powdery and floccose near the apex due to the conidial mass, tan to dark brown, measuring 25-50 mm in length and 0.5-3.0 mm in diameter. Conidiophores densely branched, 3-6 µm wide, consisting of verticillate branches with whorls of 2 to 5 phialides. Phialides 4.2-6.5 x 2.5-3.5 µm, consisting of a swollen globose basal portion tapering into a thin neck about 0.5 µm wide. Conidia cylindrical, usually curved, smooth-walled, hyaline, 3.5-8.0 x 1.5-3.5 µm.

According to Petch (1933 b, 1942) and Mains (1951) often associated with Cordyceps sabolifera (Hill.) Sacc.

 

 

Fig. 20. Paecilomyces cicadae, a-b. conidiophores; c. conidia; d. synnema.

 

Material examined

 

Herbarium specimens

 

Isaria cicadae, Fungi malaisiani, on cicadae, Rnggano, Boeha-boeha; leg. W. G. Lutjeharms, det. K. B. Boedijn, no. 961245-154 (L); on cicadae, coll. by A. E. Mass, S. Sebastian, [p. 54] Jaliso, Mexico (BPI); on cicade, collected in China 1917, det. E. B. Mains (BPI); several specimens of Isaria cicadae identified by T. Petch (K).

Isaria sinclairii, labelled as `Cordyceps (Isaria) sinclairii, Caterpillars and Fungus, Poventry Bay, New Zealand, 1870' ex herb. Berkeley (K); on cicadae, Mission Waterlot, Madagascar, probably ex herb. Berkeley (PC); collection by W. P. Fang, Szechnan, 1928 (BPI); ex herb. Lloyd no. 42600 Nanking, China, H. H. Um (BPI); on pupa of cicadae, from drugstore Yannan, det. and coll. S. C. Teng (BPI).

Isaria cosmopsaltriae, authentic material sent by A. Yasuda, 1917, Sendai, Japan, ex herb. Lloyd no. 42596 (BPI); on Meimuna oparifera, coll. by E. Yakushiji and M. Kumazawa in Koishikawa Botanic Gardens, Japan (BPI).

Isaria mokanshawii, type, sent by N. Gist Gee, Soochow, China, ex herb. Lloyd no. 42611 (BPI).

Isaria arbuscula, type, Guatemala, leg. Mr. Raunaud, 1899 (PC).

Isaria spec., on cicade pupa, sent by N. Gist Gee, Soochow, China, ex herb. Lloyd no. 37697 (BPI).

 

Discussion

 

Although no type or authentic material of Isaria cicadae is left, it is evident from the original description and illustrations that the species is identical with Sphaeria sinclairii Berk., of which several authentic specimens have been preserved (in K) as Cordyceps (Isaria) sinclairii. Because of its large, stout synnemata, Isaria cicadae has sometimes been regarded as an immature Cordyceps. Two examples of this confusion are Torrubia caespitosa Tul. and probably Sphaeria basili Taylor (cf. Kobayasi, 1941).

The synonymy of I. hariotii Arnaud is concluded from the description. I. cryptotympanae Sawada was described as occurring on a Cicada species. It produces large, ellipsoidal to cylindrical conidia, 8-13 x 4-5 µm. The description is too vague to allow a definite conclusion.

Paecilomyces cicadae is not known in pure culture. The species is, how­ever, easily recognizable by the large synnemata and conidia. It seems to be restricted in its occurrence to Cicadae species. Rant (1925) reported the use of Isaria cicadae as a medicine for children in China, called `san dwhe'.

 

 

20. Paecilomyces xylariiformis (Lloyd) Samson, comb. nov. - Fig. 21.

Isaria xylariiformis Lloyd [as xylariaformis] - Mycol. Writings 7: 1200. 1923 (basionym).

 

The description is based on herbarium material:

 

Synnemata few, simple, erect, up to 1.8 mm length and 100 µm in diameter, more or less white, powdery from the conidia, at the base surrounded by a white mycelium, with a tan to brown stalk. Conidiophores 1.7-2.5 µm in diameter, hyaline, smooth-walled, septate. Phialides solitary or in clusters of 2, sometimes 3, measuring 3.6-15 x 2-3.7 µm, consisting of a swollen basal portion, tapering abruptly into a long distinct neck about 1 µm in diameter, slightly bent away from the main axis. Conidia in [p. 55] short chains, globose to broadly ellipsoidal, smooth-walled, hyaline, 4.5-7 x 3.0-4.5 µm.

 

 

Fig. 21. Paecilomyces xylariiformis, a-b. conidiophores; c. conidia.

 

Material examined

 

Herbarium specimen

 

Isaria xylariiformis, type, herb. Lloyd no. 42613, collected and sent to Lloyd by J. Rick, Brazil (BPI).

 

Discussion

 

The conidiogenous structures of this species fit into the Sect. Isarioidea, though they are less compact than those of species such as P. farinosus and P. tenuipes. The phialides are solitary or in groups of 2 to 3. The shape is more slender and the necks are slightly bent away from the main axis. Because of its large conidia, P. xylariiformis is somewhat reminiscent of P. suffultus, which has more complex conidiophores, phialides with a globose basal portion, and cylindrical conidia. P. xylariiformis seems to be a rare species.

 

 

21. Paecilomyces suffultus (Petch) Samson, comb. nov. - Fig. 22.

 

Cylindrodendrum suffultum Petch - Trans. Br. mycol. Soc. 27: 91. 1944 (basionym).

 

The description is based on herbarium material:

Mycelium forming white tomentose tufts on insect pupae, sometimes forming loose, white to yellow-brown synnemata, 1.8-2.0 mm high and [p. 56] about 100 µm in diameter, powdery from conidia. Conidiophores 15-120 x 1.7-2.5 µm in diameter, hyaline, smooth-walled, verticillate, bearing whorls of 2 to 5 phialides. Phialides 2.5-4 µm, consisting of a globose to ellipsoidal basal portion, tapering abruptly into a long neck, about 1 µm wide. Conidia in short chains, ellipsoidal to cylindrical, smooth-walled, hyaline, 4.5-12 x 3.0-4.0 µm.

 

 

 Fig. 22. Paecilomyces suffultus, a-b. conidiophores; c. conidia.

 

Material examined

 

Herbarium specimen

 

Cylindrodendrum suffultum, type, on dipterous pupae (Psychodidae), collected by E. A. Ellis in Norwich Trowse Marshes, and additional specimens from Home Marsh, Poor's Marsh and Wheatfen Broad, Great-Britain (K).

 

Discussion

 

Petch (1944a) described Cylindrodendrum suffultum without discussion on the taxonomy. He found that the conidia were not catenulate. In [p. 57] the type specimen, however, short chains were seen on phialides, which have a swollen basal portion and a distinct neck; therefore C. suffultum is placed in the genus Paecilomyces.

P. suffultus resembles P. cicadae, but differs by the straight and wider conidia and the small, loose synnemata.

 

 

22. Paecilomyces marquandii (Massee) Hughes - Fig. 23.

 

Verticillium marquandii Massee - Br. Fungus Fl. 3: 24. 1898 = Paecilomyces mar­quandii (Massee) Hughes - Mycol. Pap. 45: 30. 1951.

Spicaria violacea Abbott - J. Iowa St. Coll. Sci. 1: 26. 1926.

? Penicillium amagasakiense Kusai - A. Rep. scient. Wks. Osaka Univ. 8: 43. 1960 (without latin diagnosis and indication of the type).

 

 

Fig. 23. Paecilomyces marquandii, a-b. conidiophores, c. conidia; d. chlamydospores.

 

Colonies on malt-agar growing moderately fast, attaining a diameter of 5 to 7 cm within 2 weeks at 25°C, consisting of a dense felt with a floccose overgrowth of aerial mycelium, in fresh isolates sometimes producing short erect loose synnemata; at first white, becoming pale vinaceous [p. 58] to vinaceous near Dark Vinaceous Brown, changing with age to brown shades near Liver Brown (Ridgway, pl. 39; Rayner 5"'k, 7'm). Reverse usually bright yellow to orange-yellow, exudate usually diffusing into the surrounding agar, with age becoming yellow-brown. Odour absent. Vegetative hyphae hyaline, smooth-walled, 2.5-3.2 µm wide. Conidial structures variable, mostly verticillate, sometimes synnematous, especially in fresh isolates; conidiophores hyaline, smooth-walled, 50-300 x 2.5-3 µm, arising from submerged hyphae or formed as sidebranches on the aerial hyphae, consisting of verticillate branches with whorls of 2 to 4 phialides. Phialides 8-15 x 1.5-2 µm, consisting of a short cylindrical to ellipsoidal basal portion, tapering into a distinct neck, about i µm wide. Conidia in dry divergent chains, ellipsoidal to fusiform, smooth-walled to finely roughened, hyaline, pale vinaceous in mass, 3-3.5 x 2-2.2 µm. Chlamydospore-like structures usually present in the submerged mycelium, consisting of solitary, thin-walled, globose to ellipsoidal cells, about 3.5 µm in dia­meter.

 

Material examined

 

CBS 182.27 = IMI 39815 = NRRL 901 = ATCC 10525 = IFO 6120 = type culture of  Spicaria violacea Abbott, isolated from soil by E. V. Abbott, Iowa, USA.

CBS 282.53 = IMI 58450, isolated by C. T. Ingold from Camarophyllus sp., Sevenoaks, Kent, Great-Britain.

CBS 302.63, isolated from soil by F. Smit, Baarn, Netherlands.

CBS 448.64, isolated from rhizophere of Beta vulgaris by S. B. Mathur, Wageningen, Netherlands.

CBS 130.67, isolated from glasshouse soil under Lycopersicum esculentum by G. J. Bollen, Wageningen, Netherlands.

and numerous strains, isolated from soil and other substrates.

 

Discussion

 

According to the description, Penicillium amagasakiense might be identical with P. marquandii, but without examination of the type material the species is regarded as doubtful.

P. marquandii is distinguished from P. lilacinus by the yellow reverse and smooth-walled, hyaline conidiophores. Moreover, chlamydospore-like cells are usually present. Both fungi are rather common soil hyphomycetes (Domsch and Gams, 1972).

 

 

23. Paecilomyces lilacinus (Thom) Samson, comb. nov. - Fig. 24.

 

Penicillium lilacinum Thom – Bull. Bur. Anim. Ind. U.S. Dep. Agric. 118: 73. 1910 (basionym).

Graphium cicadicola Speg. - An. Mus. nac. Hist. nat. B. Aires 2o: 446. 1911.

? Penicillium amethystinum Wehmer sensu Biourge - La Cellule 33: 221. 1923. [p. 59]

? Penicillium (Scopulariopsis) rubellum Bainier sensu Biourge - La Cellule 33: 221. 1923.

Spicaria violacea Petch - Trans. Br. mycol. Soc. 16: 239. 1932 [non Abbott, 1926].

Spicaria rubidopurpurea Aoki - Bull. serie. Exp. Stn Japan 10: 440. 1941.

 

 

Fig. 24. Paecilomyces lilacinus, a-d. conidiophores, a-b. in pure culture, c. Ghanaian specimen (R.S. 0242), d. Graphium cicadicola; e-f. conidia, e. Graphium cicadicola, f. in pure culture.

 

Colonies on malt-agar growing rather fast, attaining a diameter of 5-7 cm within 14 days at 25° C, consisting of a basal felt with a floccose over­growth of aerial mycelium; at first white, but when sporulating changing [p. 60] to vinaceous near Light Vinaceous Drab (Ridgway, Pl. 45; Rayner, 5""b ) and Light Brownish Drab (Ridgway, Pl. 45; Rayner. 9""b). Reverse some­times uncoloured, but usually in vinaceous shades near Daphne Red to Vernonia Purple (Ridgway, Pl. 38; Rayner, 69"i). Exudate limited or absent. Odour indistinct. Vegetative hyphae smooth-walled, hyaline, 2.5-4.0 µm wide. Conidiophores arising from submerged hyphae, 400-600 µm in length, or arising from the aerial hyphae and half as long, or occasionally forming loose synnemata up to 2 mm high; stalks rather thick-walled, pigmented in yellow or purple shades with a roughened wall, 3-4 µm wide, consisting of verticillate branches with whorls of 2 to 4 phialides. Phialides 7.5-9.0 x 2.5-3.0 µm, consisting of a swollen basal part, tapering into a thin distinct neck, about 1 µm wide. Conidia in divergent chains, which sometimes become tangled, ellipsoidal to fusiform, smooth-walled to slightly roughened, hyaline, purple in mass 2.5-3.0 x 2.0-2.2 µm. Chlamy­dospores absent.

 

On insects the fungus produces a thin mycelial felt from which mono­nematous conidiophores arise, forming a purple layer, or synnemata which may be loose or stout and conspicuous, 6-20 mm long and 50-150 µm in diameter. Conidiogenous structures as in culture, but elements more swollen.

 

Material examined

 

Herbarium specimens

 

Graphium cicadicola, type, labelled `s. cicade sp., Chaco, 25-11-1901. leg.: Kermčs, no. 12273' (LPS).

Spicaria violacea, type, on tineid case, Nuwara Eliya , Ceylon, ex herb. Petch no. R 363  (K).

Paecilomyces lilacinus R.S. 0084, 0242, on Aethus species (Cynidae, Heteroptera), collected by H. C. Evans in Tafo, Oyoko, Ghana (CBS).

 

Living cultures

 

CBS 284.36 = IMI 27830 - QM 7592 = ATCC 10114 = NRRL 895 = Thom 2258, type culture of P. lilacinum, isolated by C. F. Atkinson and C. W. Edgarton from soil, Ithaca, N.Y., USA.

CBS 346.48, probably identical with type culture.

CBS 346.51 = IMI 58442, received as Spicaria rubidopurpurea Aoki, culture no. 4129, sent by K. Kominami.

CBS 204.57, isolated by H. L. Schmidt from soil in Germany.

CBS 279.58, isolated from wood pulp in the Netherlands.

CBS 630.67, isolated by R. Kenneth from Hordeum sativum in Israel.

CBS 225.73, strain 11232, isolated by R. Schneider from a larva of Aleurotrachilis species, Berlin, Germany.

CBS 226.73 A, isolated by J. Schreurs from roots of Persea americana, Hartel, Liberia. CBS 226.73 B, isolated by F. G. Poelma from intestine of Chamaeleo dilepsi, Utrecht.

CBS 940.73, isolated by H. C. Evans from Aethus species (Heteroptera) in Tafo, Ghana.

Numerous isolates from agricultural soils, Wageningen, isolated by J. H. van Emden and J. W. Veenbaas-Rijks. [p. 61]

 

Discussion

 

Paecilomyces lilacinus differs from P. marquandii by its pigmented coni­diophores, which are usually rough-walled, and the uncoloured or purple reverse.

Together with Spicaria violacea Abbott and Penicillium humuli van Beyma, Penicillium lilacinum was placed by Raper and Thom (1949) in the lilacinum-series of Penicillium. S. violacea was included in this series because of its close relationship with P. lilacinum. Raper and Thom (l.c.) stated that intergrading forms are not uncommon, and considered the possibility of transferring S. violacea to Penicillium. A new combination was, however, not proposed, since these authors found the generic description of Spicaria very confused. Brown and Smith (1957) accepted Paecilomyces marquandii (Massee) Hughes as the correct name for Spicaria violacea Abbott, but Penicillium lilacinum was left in Penicillium, because of its possible relationship to other species of the Penicillium janthinellum-series.

Re-examination of strains of Penicillium lilacinum studied by Raper and Thom (1949) and Brown and Smith (1957) and observations on numerous new isolates, mainly collected by J. H. van Emden, showed indeed the existence of intergrading forms between Penicillium lilacinum and Paecilo­myces marquandii. In such strains characters of both species can be ob­served. Penicillium lilacinum is here placed in Paecilomyces, in spite of its similarities to the Penicillium janthinellum-series, because of the stiff, verticillate (not divaricate) conidiophores, the phialides with a distinct neck and the purple or vinaceous colour of the conidial heads.

The formation of synnemata of P. lilacinus was described by Spegazzini (1911) in Graphium cicadicola on an Cicada species. On the type specimen these synnemata were still recognizable and they consist of pigmented, slightly rough-walled conidiophores producing small, fusiform conidia. Conspicuous stout synnemata were observed on insects collected in Ghana. In pure culture the isolate CBS 940.73 showed conidiogenous structures similar to P. lilacinus, but differed by having some localized areas with a yellow reverse. When first isolated the strain produced a few synnemata which disappeared after subculturing. The tendency to synnematous growth of the conidiophores was also occasionally observed in fresh isolates, col­lected from agricultural soil in the Netherlands.

Because of the similarity of the two purple species, a decision about synonymy of other described species is sometimes difficult. Brown and Smith (1957) placed Spicaria rubidopurpurea Aoki into synonymy with Penicillium lilacinum, but considered Spicaria violacea Petch as identical with P. marquandii. In the type specimen of S. violaceae Perch, however, pigmented and rough-walled condiophores were found. For this reason S. violacea Petch, a later homonym of S. violacea Abbott, is now placed in the synonymy of P. lilacinus. The systematic position of Penicillium amethystinum Wehmer and Penicillium rubellum Bainier is uncertain. From the description given by Biourge (1923) they may be identical with P. lilacinus (cf. Raper and Thom, 1949). [p. 62]

Tsuchiya et al. (1952) reported that strains of Penicillium lilacinum and Spicaria violacea (= P. marquandii) produce active extracellular dextran­ degrading enzymes. Shima (1955) and Shima and Suzuki (1955) showed that Spicaria rubidopurpurea, a common fungus on silkworms in Japan, produced the following metabolites: oxalic, dipicolinic and succinic acids, several unidentified amino acids and large amounts of D-mannitol. Arai et al. (1973) isolated a peptide antibiotic, leucostatin, from a strain identified as Penicillium lilacinum. Leucostatin is active against some gram-positive bacteria and a wide range of fungi. The authors mentioned also another water-soluble peptide antibiotic, lilacin, isolated from the culture fluid (cf. Yamano, 1970).

 

 

24. Paecilomyces cinnamomeus (Petch) Samson & W. Gams, comb. nov. - Fig. 25.

 

Verticillium cinnamomeum Petch - Trans. Br. mycol. Soc. 16: 233. 1932 (basionym) = Verticillium heterocladum Penzig sensu Fawcett - Fungi parasitic on Aleyrodes citri, p. 22. 1908. [non Penzig, 1882].

 

The description is based on herbarium material:

 

Mycelium covering the insect with thick-walled, hyaline to yellow, smooth-walled, to finely roughened hyphae, 3-7 µm wide, giving a cin­namon-brown to ashy brown colour. Conidiophores erect, mononematous, 150-240 x 2-3 µm, hyaline to yellow, especially near the base, smooth-walled, consisting of verticillate branches with whorls of 2 to 4 phialides or whorls of phialides directly on the stalk. Phialides 9-14 x 2.5-3.0 µm, flask-shaped. Conidia in chains, fusiform, smooth-walled, hyaline, 5.2-7.5x 1.7-2.5 µm.

 

Material examined

 

Herbarium specimens

 

Verticillium heterocladum on Citrus sp., received from A. J. Goddard, 1913, Abbeville, La., det. Fawcett; neotype, on Citrus sp., coll. E. S. Tucker, Baton Rouge La., 1913; on Ci­trus aurantium infested with `white fly', Iberia, La., 1913, coll. E. S. Tucker; on Aleyrodes citri on Citrus sp., Thibodaux, La., 1915; on Citrus sp., coll. R. Drango, Havanna, Cuba; on Aspidotus destructor coll. S. C. Bruner, Santiago de las Vegas, Cuba, 1922; on Cape Jessamine on white fly, 192.3, Moblie, Ala., det. V. K. Charles (all BPI).

Verticillium cinnamomeum on Aleyrodes citri on orange, Ocean Springs, Miss., 1920, coll. J. E. Miles; on insect on Gardenia, Winter Park, Fla., coll. & det. V. K. Charles; on Marcgravia insects, Alta Verapaz, Guatemala, coll. P. C. Standley & P. L. Lentz, 1941; on Camellia species ex Mexico intercepted at Fort Worth, Texas, by C. F. Daley; on Aleyrodes citri on Citrus sp., Highlands Mammock, Fla., 1937, coll. C. L. Shear (all BPI). Paecilomyces cinnamomeus, on Coccidae, collected by H. C. Evans in Kukurantumi, Ghana, 1972 (CBS). [p.63]

 

 

Fig. 25. Paecilomyces cinnamomeus, a-c. conidiophores; e. conidia.

 

Discussion

 

Verticillium cinnamomeum Petch is placed in Paecilomyces because of its rather stout conidiophores with flask-shaped phialides with a distinct neck and its catenate conidia. In the genus Verticillium catenate conidia have been described only in some species of the section Prostrata (Gams, 1971b). The species is reminiscent of Mariannaea, but differs by the straight conidial chains. P. cinnamomeus can be distinguished from the other species by its pigmented conidiophores and fusiform conidia. It is closely related to P. lilacinus, which has smaller conidia and purple conidial heads. P. cinnamomeus seems to be a common fungus on Citrus-inhabiting scale insects.

The only similarity between the present fungus and the description of [p. 64] Verticillium heterocladum Penzig is its occurrence on scale insects. No herbarium material of the latter species has been preserved. The original description (Penzig, 1882) and the illustrations in Saccardo (I88ķ) emphasize the polyblastic(?) conidiogenous cells. It is therefore very uncertain, whether V. heterocladum can be placed in synonymy with Verticillium lecanii (Zimm.) Viégas, as has often been suggested (cf. Gams, 1971b).

 

 

25. Paecilomyces viridis Segretain ex Samson, spec. nov. - Fig. 26.

 

Paecilomyces viridis Segretain, Fromentin, Destombes, Brygoo & Dodin – C.r. hebd. Séanc. Acad. Paris 259: 260. 1964 (without latin diagnosis).

 

Coloniae in agaro maltoso ad 4 cm diam. 14 diebus 25° C, primum cremeae, cito flavo­virides. Conidiophora erecta, mononematica, e hyphis submersis vel raro lateraliter e hyphis aeriis oriuntur, ad 120 µm longa, basi 1.5-2.6 µm crassa, ramos verticillatos et quoties binas ad quaternas phialides ferunt vel phialides in stipite ipso verticillatas. Phialides 4.0-9.0 x 2.5-3.1 µm, e basi globosa in collum 0.5-1.0 µm crassum angustatae. Conidia globosa vel subglobosa, nonnumquam apiculata, levia, hyalina, acervata, dilute viridia, 2.5-3.2 µm diam. In coloniis vetustis vel in mycelio submerso hyphae modo sac­charomycetis secedunt in cellulas globosas vel subglobosas.

Typus CBS 348.65 = IP 85o, isolatus e Chamaeleone laterali, capto in Madagascar.

 

 

Fig. 26. Paecilomyces viridis, a-b. conidiophores; c. conidia; d. yeast-like cells.

 

 

Colonies on malt-agar attaining a diameter of 4 cm within 14 days at 25°C, consisting of a thin basal felt from which numerous conidiophores arise, giving the surface a velvety appearance; at first cream, soon becoming [p. 65] yellow-green between Seafoam Yellow and Seafoam Green (Ridg­way, Pl. 31; Rayner, 25"f, 27"f). Reverse yellow, later yellow-brown. Vegetative hyphae smooth-walled hyaline, 1.5-3.0 µm wide. Conidiophores erect, mononematous, arising from the submerged hyphae or occasionally as sidebranches from the aerial hyphae, up to 120 µm long and 1.5-2.6 µm wide, consisting of verticillate branches with whorls of 2 to 4 phialides or verticillate phialides borne directly on the stalk. Phialides 4.0-9.0 x 2.5-3.1 µm, consisting of a globose basal portion tapering into a thin neck, 0.5-1.0 µm wide. Conidia globose to subglobose, sometimes apiculate, smooth-walled, hyaline, green in mass, 2.5-3.2 µm in diameter. Chlamydo­spores absent. Especially in older colonies yeast-like hyphal fragments occur submerged in the agar, consisting of globose to subglobose cells about the same size as the conidia.

 

Material examined

 

CBS 348.65 = IP 850, type culture; CBS 659.71, 660.71, 661.71, all strains isolated from different organs of Chamaeleo lateralis, sent to G. Segretain.

 

Discussion

 

The green colour of the colonies of Paecilomyces viridis indicates a rela­tionship to the genus Penicillium. The characteristic globose phialides with a distinct neck and the verticillate conidiophores point to the inclusion of this species in Paecilomyces. P. viridis can be distinguished from other Paecilomyces species by its green colour, the yeast-like fragmentation and the globose to subglobose conidia.

P. viridis was isolated from Chamaeleo lateralis, sent from Madagascar. The animal was suffering from spontaneous deep and fatal mycosis. The fungus was found in all organs of the animal and in the blood, but especial­ly in the liver and the spleen. In vivo it showed only the yeast-like form, but in pure culture the Paecilomyces mycelial state appeared (Segretain and Segretain, 1971). Betina et al. (1966), Barath and Betina (1972), Koman et al. (1972) and Barathovį et al. (1972) studied the effects of antibiotics on the dimorphism of P. viridis in vitro and showed that yeast-like growth is inducible by the antibiotics cyanein and azalomycin F.

 

 

26. Paecilomyces niphetodes Samson, spec. nov. - Fig. 27.

 

Coloniae in agaro maltoso fere celeriter crescunt, ad 6-7 cm diam. 14 diebus 25°  C, candidae, pulverulentae, margine arachnoidea. Conidiophora praecipue e hyphis submersis oriuntur, erecta, 90-120 x 3.5-4.0 µm, hyalina, levia, irregulariter ramosa, sed plerumque verticillata, ramis phialides quoties binas ad quaternas ferentibus. Phialides 7.5-12.5 x 3-4 µm, e parte fere cylindrica in collum distinctum 0.5-0.8 µm crassum angustatae. Conidia [p. 66] triangularia, nonnunquam late ellipsoidea vel subglobosa, hyalina, levia, 3.5-4.0 x 2.5-3.0 µm.

Typus CBS 229.73, lectus in ligno Fagi sylvaticae, Vogelenzang prope Haarlem in Hollandia, a W. Gams.

 

Colonies on malt-agar growing rather rapidly, attaining a diameter of 6-7 cm within 14 days at 25° C, consisting of a dense mycelial felt of numerous conidiophores, giving the colony a white powdery appearance, with an arachnoid margin. Reverse uncoloured. Odour absent. Vegetative hyphae hyaline, smooth-walled, 3.5-5.0 µm wide. Conidiophores primarily arising from the submerged hyphae, erect, mononematous, 90-120 x 3.5-4 µm, hyaline, smooth-walled, sometimes irregularly branched, but usually verticillate, bearing on each branch whorls of 2 to 4 phialides. Phialides in whorls or occasionally singly along the aerial hyphae, 7.5-12.5 x 3-4 µm, consisting of a globose to ellipsoidal basal portion, tapering into a distinct neck about 0.5-0.75 µm wide. Conidia triangular, sometimes broadly el­lipsoidal or subglobose, hyaline, smooth-walled, 3.5-4 x 2.5-3 µm. Chlamy­dospores absent.

 

 

Fig. 27. Paecilomyces niphetodes, a-b. conidiophores; c. conidia.

 

Material examined

 

Herbarium specimens

 

IMI 136368 and 139638, dried cultures isolated by P. A. G. Jenkins from Mercurialis perennis, Keele University, Keele, Great Britain. [p. 67]

 

Living culture

 

CBS 229.73, type culture, isolated from decaying wood of Fagus sp., collected by W. Gams near Vogelenzang, Netherlands,

 

Discussion

 

The triangular conidia and the arachnoid growth distinguish this species from others. The morphology of the conidiophores resembles those produced in Mariannaea elegans, in which the conidia are produced in imbricate chains by slenderly flask-shaped phialides.

 

 

27. Paecilomyces sulphurellus (Sacc.) Samson & W. Gams, comb. nov. - Fig. 28.

 

Verticillium sulphurellum Sacc. - Fungi ital. autogr. delin., tab. 641. 1881; Michelia 2: 554. 1882 (basionym).

 

The description is based on fresh collections on the natural substrate:

 

Mycelium in tufts or covering the substrate with a thin mycelial felt from which the conidiophores arise, sulphur yellow. Vegetative hyphae smooth-walled, hyaline to slightly yellowish, 2-2.5 µm wide. Conidiophores erect, mononematous, 35-60 x 2.0-2.5 µm, consisting of verticillate branches with whorls of 2 to 4 phialides. Phialides 4.5-7.0 x 1.9-2.5 µm, flask-shaped, with a slightly inflated base, tapering into a thin neck. Conidia at least partly in chains, cylindrical, smooth-walled, hyaline to slightly yellowish, 2.2-2.7 x 0.8-1.0 µm. Chlamydospores not observed.

 

 

Fig. 28. Paecilomyces sulphurellus, a-b. conidiophores; c-d. conidia. [p. 68]

 

Material examined

 

Paecilomyces sulphurellus, W.G. 1619, on decayed wood, 1969, and on decayed bark of Quercus petraea, 1970, coll. W. Gams, Sanddunes near Vogelenzang; on decayed bark of Quercus spec., coll. W. Gams, Meerveldse Bos near Uddel, Netherlands, 1974 (CBS).

 

Discussion

No fungus material was found in the type collection (PAD), but the Dutch specimens on which the new description is based, agree with the description and illustrations by Saccardo (1881, 1882). Verticillium sul­phurellum is transferred to Paecilomyces on account of its verticillate conidiophores, the phialides with a distinct neck and the catenate conidia. The species is easily recognizable by the sulphur yellow colour and the unusually small cylindrical conidia. In spite of repeated efforts, the fungus could not be grown in pure culture.

 

 

28. Paecilomyces puntonii (Vuill.) Nannizzi - Fig. 29.

 

Corethropsis puntonii Vuill. - C.r. hebd. Séanc. Acad. Paris 190: 1334. 1930 = Pae­cilomyces puntonii (Vuill.) Nannizzi - Tratt. Micopatol. umana 4: 245. 1934.

 

 

Fig. 29. Paecilomyces puntonii, a-b. conidiophores; c. conidia; d. chlamydospores.

 

Colonies on malt-agar growing rapidly, attaining a diameter of 4.5 cm  within 14 days at 25°C, consisting of a densely matted felt with floccose overgrowth, sometimes becoming funiculose. Reverse yellow. Odour in­distinct. Vegetative hyphae smooth-walled, hyaline, about 4 µm wide. [p. 69] Conidiophores sparingly produced, arising from aerial hyphae, consisting of verticillate branches with whorls of 2 to 3 phialides. Phialides also borne directly and singly on aerial hyphae, 8.5-20 x 1.7-2.3 µm, consisting of a cylindrical basal portion, tapering into a thin neck, 0.5-0.9 µm wide. Conidia in dry tangled chains, with some tendency to form loose heads in age, ellipsoidal to cylindrical, smooth-walled, hyaline, 3.5-4.0 x 1.7-2.0 µm. Chlamydospores abundantly produced, singly or in short chains, thick­walled, brown, subglobose to pyriform, up to 10 µm in diameter.

 

Material examined

 

CBS 250.37 = IMI 58409, type culture of Corethropsis puntonii, isolated by Puntoni from human skin and received from R. Ciferri.

 

Discussion

 

The description is based on the type culture, which shows poor sporula­tion. P. puntonii can be distinguished from other species by its chlamydo­spores, its hyaline conidiogenous structures and the ellipsoidal to cy­lindrical conidia. With its tendency to form heads in older colonies it resembles some Verticillium species.

 

 

29. Paecilomyces inflatus (Burnside) Carmichael - Fig. 30.

 

Myceliopthora inflata Burnside - Pap. Mich. Acad. Sci. 8: 82-84. 1928 = Paecilomyces inflatus (Burnside) Carmichael - Can. J. Bot. 40: 1148. 1962.

Paecilomyces flavescens Brown & Smith - Trans. Br. mycol. Soc. 40: 56. 1957.

 

 

Fig. 30. Paecilomyces inflatus, a-b. conidiophores; c. conidia. [p. 70]

 

Colonies on malt-agar growing slowly, attaining a diameter of 3.5 cm within 14 days at 25° C; when freshly isolated appearing powdery and velvety; after several transfers becoming more floccose to funiculose and tougher from an increase in vegetative hyphae. Vegetative hyphae hyaline, smooth-walled, 1.5-3 µm wide. Conidiogenous structures irregular, but usually consisting of whorls of 2 to 3 phialides on short branches of trailing hyphae, distinct conidiophores usually lacking. Phialides 7-15 x 2-3 µm, mostly occurring singly along aerial hyphae, consisting of a swollen basal portion, tapering abruptly into a long neck of about 1 µm in width. Conidia hyaline, lemon-shaped, smooth-walled or very finely roughened, 3-4 x 2-3 µm. Chlamydospores absent.

 

Material examined

 

CBS 259.39 = IMI 113168 = ATCC 6491 = ATCC 10843 = type culture of Mycelioph­thora inflata, isolated by J. A. Stevenson, from honeybee (mycelium from abdomen of adult workerbee), Washington DC., USA.

CBS 263.58 = IMI 45948  = type culture of Paecilomyces flavescens, isolated from soil, Ampthill Forest, Beds., Great-Britain.

CBS 183.65 = IMI 109890, isolated by G. L. Barron from forest soil under Thuja oc­cidentalis, Aberfoyle, Ontario, Canada.

CBS 487.67, isolated by W. Gams from decaying wood in glass-house, Botanical Garden Kiel, Germany.

CBS 228.73 = NRRL 7373a, isolated by G. L. Barron from soil, Iowa, USA. Several isolates from soil in the Netherlands.

 

Discussion

 

This is the only monophialidic species of Paecilomyces, described by Onions and Barron (1967), which Gams (1971b) left in Paecilomyces, because of the phialides which are flask-shaped with a somewhat inflated base. In addition the phialides are sometimes clustered in small groups on well-defined conidiophores. In some strains the phialides proliferate, resembling the conidiogenous cells of Acrophialophora. Dark pigmented, warty conidiophores are, however, lacking in P. inflatus.

P. inflatus is a rather common species, found especially in forest soil.

 

 

30. Paecilomyces carneus (Duché & Heim) Brown & Smith - Fig. 31.

 

Spicaria carnea Duché & Heim - Recl Trav. crypt. dédiés į Louis Mangin, p. 454.­1931 = Paecilomyces carneus (Duché & Heim) Brown & Smith - Trans. Br. mycol. Soc. 40: 70. 1957.

? Spicaria decumbens Oudemans - Archs néerl. Sci. (Set. 2) 7: 290. 1902.

Paecilomyces austriacus Szilvinyi - Zentbl. Bakt. ParasitKde, (Abt. 2) 103: 176. 1941.

Spicaria viridis Szilvinyi - Zentbl. Bakt. ParasitKde (Abt. 2) 103: 176. 1941.

Penicillium nopporoense Sasaki & Nakane - J. agric. Chem. Soc. Japan 19: 775. 1943.

Spicaria carnosa Miller, Giddens & Foster - Mycologia 49: 800. 1957. [p. 71]

 

 

Fig. 31. Paecilomyces carneus, a-b. conidiophores; c. conidia.

 

Colonies on malt-agar growing rather slowly, attaining a diameter of 14 to 22 mm within 2 weeks, consisting of a basal felt and a floccose over­growth, later with some tendency to funiculose or fasciculate growth; usually with a rich sporulation giving the surface of the colony a powdery appearance; at first white, soon becoming pink near Shell Pink (Ridgway, Pl. 28; Rayner, 11"f) with a green cast from the reverse. Reverse yellow brown, but mostly green to dark green, with age sometimes turning black; exudate sometimes present as green drops. Odour absent. Vegetative hyphae rather thin-walled, smooth, 1.5-2.5 µm wide. Conidiophores mono­nematous, erect, arising primarily from the submerged hyphae, l00-150 x 1.5-2.5 µm, consisting of verticillate branches, with whorls of 2 to 4 phialides. Phialides 9-18 x 1.5-2.5 µm, consisting of a cylindrical, some­times somewhat inflated base, tapering into a thin long neck, about 0.5 µm in width. Conidia in long divergent chains, subglobose to ellipsoidal, roughened to spinulose, 3-4 x 2-2.5 µm. Chlamydospores absent.

 

Material examined

 

CBS 239.32 = IMI 58434, type culture of Spicaria carnea, isolated by J. Duché from wet sandy soil, Hills of Vauville, France.

CBS 159.42 = IMI 58435, isolated by P. Bels from a fly in Limburg, Netherlands.

CBS 338.54, strain 21741, isolated by J. Meyer from soil in Congo. [p. 72]

CBS 399.59 = ATCC 13218, type culture of Spicaria carnosa, received from J. Miller and isolated from forest soil, Georgia, USA.

CBS 158.69 = ATCC 18336 = AHU 8023, type culture of Penicillium nopporoense, isolated by Sasaki and Nakane.

Numerous strains isolated from soil in the Netherlands.

 

Discussion

 

The synonymy of Spicaria decumbens, Paecilomyces austriacus and Spicaria viridis is based on the descriptions and already discussed by Brown and Smith (1957).

Paecilomyces carneus is a very common hyphomycete in soil (Domsch and Gams, 1972). It is easily recognizable by the echinulate conidia and the green reverse.

 

 

31. Paecilomyces penicillatus (Höhnel) Samson, comb. nov. - Fig. 32.

 

Spicaria elegans (Corda) Harz sensu Grove - J. Bot. Lond. 23: 165. 1885 = Spicaria elegans (Corda) Harz var. muscorum Grove ex Sacc. - Syll. Fung. 4: 167. 1886.

Spicaria penicillata Höhnel - Annls mycol. 2: 56. 1904 (basionym).

? Spicaria fuligonis F. Moreau - Bull. trimest. Soc. mycol. Fr. 32: 36. 1916.

 

Colonies on malt-agar growing rather fast, attaining a diameter of 4 cm within 14 days at 25° C, consisting of a basal felt with floccose over­growth; sporulation mostly in localized areas, white. Reverse uncoloured. Odour absent. Vegetative hyphae smooth-walled, hyaline, 3-6 µm wide. Conidiophores erect, mononematous, 150-600 µm long, 3.7-5.0 µm wide, consisting of a penicillate, divergent cluster of branches with whorls of 2 to 6 phialides. Phialides 9-14 x 3.0-4.5 µm, consisting of a cylindrical basal portion with distinct neck. Conidia in divergent chains, cylindrical, smooth-walled, hyaline, one-celled, 8-9 x 1.5-2.0 µm, on the natural sub­strate the conidia have truncate ends and are sometimes two-celled, 6.5-10 x 2.5-3.2 µm. Chlamydospores produced submerged in the agar, single, ellipsoidal to pyriform, one-celled, smooth-walled, hyaline, 6-7.5 x 4.0-5.0 µm.

For the description on the natural substrate cf. Hughes (1951) under Spicaria elegans var. muscorum.

 

Material examined

 

Herbarium specimens

 

Spicaria elegans, ex herb. Grove, on dead moss and rotting wood, Sutton, Great Britain (K). [p. 73]

Spicaria penicillata, type, on Arcyria punicea, coll. by F. von Höhnel, Wienerwald, 1903; on Arcyria cinerea, Wienerwald, 1916 (FH).

 

Living culture

 

CBS 448.69, isolated by J. van Holder from a rotting mushroom, Gent, Belgium.

 

 

Fig. 32. Paecilomyces penicillatus, a-b. conidiophores, a. CBS 448.69. b. type Spicaria penicillata. c-d. conidia, c. CBS 448.69, d. type; e. chlamydospores.

 

Discussion

 

The species is placed in the genus Paecilomyces, though the phialides when produced in pure culture have a rather short neck and are densely compacted resembling Penicillium. On the natural substrate however, the [p. 74] phialide necks are longer (cf. Hughes, 1951). Because of the hyaline conidiogenous structures and the divergent conidial chains the species is placed in Paecilomyces. It differs from Mariannaea elegans by the penicil­late conidiophores, the cylindrical conidia, which are produced in straight chains. The mis-interpretation of Grove (1885) of Spicaria elegans has been discussed by Hughes (1951).

Judging from the description, Spicaria fuligonis F. Moreau might be identical with P. penicillatus. However, no type material is available.

P. penicillatus differs from the other species by the presence of hyaline chlamydospores, the long mononematous conidiophores and the cylindrical conidia, which have truncate ends, when produced on the natural substrate.

 

 

7. The genus Mariannaea Arnaud ex Samson, gen. nov.

 

Mariannaea Arnaud - Bull. trimest. Soc. mycol. Fr. 58: 196. 1952 (without latin diagnosis).

 

Coloniae fere celeriter crescunt. Conidiophora ramos verticillatos et in apice phialides verticillatas proferunt. Phialides graciles, lageniformes. Conidia continua, oblique apicula­ta, in catenis imbricatis divergentibus connexa, hyalina, levia. Chlamydosporae nonnum­quam adsunt. Status perfectus ignotus.

Species typica Penicillium elegans Corda.

 

Generic description

 

Colonies grow moderately fast on artificial media. Conidiophores with verticillate branches and terminal whorls of conidiogenous cells. Conidio­genous cells phialidic, slender, flask-shaped. Conidia one-celled, hyaline, smooth-walled, produced in divergent, imbricate chains or in slimy heads. Chlamydospores sometimes present. Perfect state not known. Type species: Penicillium elegans Corda.

 

Discussion

The genus Mariannaea was proposed by Arnaud (1952) with the single species M. elegans. It was characterized as being distinct from Clonostachys by its divergent chains of conidia. According to the latter author, M. elegans resembled Spicaria (Penicillium) elegans Corda (1838), but should be distinct because in Corda's drawing the conidia are not formed in oblique chains. Hughes (1951) studied the type material of Penicillium elegans Corda and transferred the species to Paecilomyces. Brown and Smith (1957) followed him and considered Mariannaea elegans as a synonym. [p. 75] The genus Mariannaea is regarded as distinct from Paecilomyces on account of the conidia in imbricate chains or slimy heads on almost awl-shaped phialides.

Mariannaea is also closely related to Clonostachys (Gliocladium) which has brush-like conidiophores with conidial chains, which may be united in imbricate columns. The imbricate chains of conidia produced in Mariannaea species are also reminiscent of those in Sesquicillium W. Gams (1968). In this genus pleurophialides occur besides the normal phialides.

 

Key to the species

 

1a.

Colonies showing a yellow-brown reverse, conidia ellipsoidal to ovoidal, flattened on one side or concave; 5.0-7.0 x 2.0-2.5 µm

M. camptospora

1b.

Colonies with dark-brown or purple reverse; conidia almost symmetrical

2

2a.

Reverse dark brown

M. elegans var. elegans

2b.

Reverse red purple

M. elegans var. punicea

 

 

1. Mariannaea elegans (Corda) Samson, comb. nov. var. elegans. - Fig. 33; Pl. 1.

 

Penicillium elegans Corda - Icones Fung. 2: 17. 1838 (basionym) = Hormodendron elegans (Corda) Bonorden - Handb. allg. Mykol., p. 76. 1851 = Spicaria elegans (Corda) Harz - Bull. Soc. imp. Nat. Moscou 44: 238. 1871 = Paecilomyces elegans (Corda) Mason & Hughes apud Hughes - Mycol. Pap. 45: 27. 1951.

Spicaria elegans (Corda) Harz var. sorghina Sacc. - Syll. Fung. 4: 166. 1886.

Spicaria elegans (Corda) Harz var. microspora Jaczewki - Mater. Mikol. Fitopat. Ross. 4: 15. 1922.

Mariannaea elegans Arnaud - Bull. trimest. Soc. mycol. Fr. 68: 196. 1952 (not validly published).

 

Colonies on malt-agar growing moderately fast, attaining a diameter of 3-5 cm within 14 days at 25° C, consisting of a matted felt with numerous erect conidiophores, showing a strong zonation; in age becoming more funiculose to fasciculate, especially in the central part; at first white, later cream to yellow-brown, usually due to the dark brown cast of the reverse. Reverse yellow-brown to dark brown, caused by diffusion of the exudate into the surrounding agar; sometimes forming brown drops. Odour absent. Vegetative aerial hyphae hyaline to yellow, up to 10 µm wide, usually finely rough-walled or verrucose; submerged hyphae inflated, brown and up to 15 µm wide. Conidiogenous structures irregular and complex, but usually consisting of erect verticillate stalks, roughened or slightly verrucose, especially near the base, up to 400 µm in length and 3-7 µm in [p. 76] diameter, bearing short branches with whorls of 3 to 6 phialides. Phialides slenderly flask-shaped, hyaline, smooth-walled, 9-25 x 2-3 µm. Conidia ellipsoidal to fusiform, hyaline, smooth-walled, 4-6 x 1.5-2.5 µm, in dry, divergent, imbricate chains or partly in slimy heads, mostly in older cultures. Chlamydospores present, intercalary or terminal, produced singly or in short chains, thick-walled, roughened, yellow-brown, globose to subglobose or ellipsoidal, 7-9.5 µm in diameter.

 

 

Fig. 33. Mariannaea elegans, a. conidiophores; b. conidia; c-d. chlamydospores. [p. 77]

 

Material examined

 

Herbarium specimens

 

Penicillium elegans, type, in herb. Corda, on the inner surface of loose bark of rotten trunks of fir-trees, 1837, Brezina, Czechoslovakia (PR).

Mariannaea elegans ex herb. Arnaud, 'sur ecclat de bois mourrissant sur le sol, Parc de Versailles, S. et O., 19 mai 1946' (PC).

 

Living cultures

 

CBS 132.41 = IMI 58431, isolated as contaminant in a slant culture of Polyporus sp. by Miss Weintroub, S. Africa.

CBS 337.51 = IMI 58432, isolated by W. Laatsch from soil, Hamburg. According to Laatsch et al. (1952) this strain produces humic acids.

CBS 308.59 = DAOM 55208, isolated by S. J. Hughes from Pseudotsuga taxifolia, Kam­loops, British Columbia.

CBS 217.73 A, isolated from decaying bark of Pinus sylvestris, Schiermonnikoog, Nether­lands.

CBS 217.73 B, isolated from decaying bark of Pinus, Baarn, Netherlands.

CBS 217.73C, isolated by W. Gams from rotten Pinus trunk, Kreuzkrug bei Schlangen, Teutoburger Wald, Germany.

CBS 219.73 A = NRRL 5886, isolated from soil sample, collected in Brownfield Woods at the University of Illinois. This strain remains white in colour with a pale brown reverse. Further it shows the same characteristics as the other strains.

CBS 219.73 B = NRRL 5887, isolated by D. Wells from bark, Ottawa, Ontario, Canada. CBS 219.73 C = NRRL 5888, isolated from soil, collected in Hocking State Park, Ohio, USA.

CBS 219.73 D = NRRL 5889, isolated from soil, collected in Poland. The dilution plates were incubated at 34° C.

Many other cultures isolated from decaying wood in the Netherlands.

 

Discussion

 

The examination of the type material of Penicillium elegans Corda and Mariannaea elegans Arnaud showed that both species are the same. The identity of Spicaria elegans var. sorghina Sacc. and var. microspora Jac­zewski is concluded from the description.

M. elegans is mostly found on rotting coniferous bark or in forest soils (Domsch and Gams, 1972). According to Laatsch et al. (1952), M. elegans is a strong humic acid producer. Bose (1962) observed that it is a moderately strong soil-borne cellulose decomposer and that it is somewhat more thermo­philic than other species usually encountered on cellulose textiles.

The species is characterized by its zonate growth, its tendency to fasci­culate growth and its ellipsoidal to fusiform conidia. Brown and Smith (1957) gave a full detailed description of the fungus and included also a strain with crimson reverse, which showed no morphological differences with the other strains studied. During this study several other strains were obtained with a typical red-purple reverse. All these strains originated from soil. No significant difference in the microscopical characters was found with those showing a brown to dark brown reverse. However, the colonies with the red-purple reverse grew faster and formed a densely [p. 78] matted felt with a thin overgrowth of aerial mycelium. The colonies showed no tendency to funiculose or fasciculate growth, while the strong zonation was also absent. Because of these different macroscopical characters a new variety is proposed.

Identification of the type of Penicillium elegans with the brown variety is mainly based on the ecological difference.

 

 

2. Mariannaea elegans (Corda) Samson var. punicea Samson, var. nov.

 

Coloniae in agaro maltoso ad 6 cm diam. 14 diebus 25° C, reverso puniceo, mycelio aerio albo.

Typus CBS 239.56, isolatus e terra prope Yangambi, Congo.

 

Colonies on malt-agar growing rapidly, attaining a diameter of 6 cm within 14 days at 25°C, consisting of a densely matted felt with thin overgrowth of the aerial mycelium; at the margin sometimes slightly zonate; at first white, the reverse becoming purple red near Eugenia Red (Ridgway, Pl 13; Rayner, i') and Pomegranate Purple (Ridgway, Pl. 12; Rayner, 71 i). Submerged hyphae encrusted with red granules, inflated up to 12 µm. Conidiogenous structures similar to those of var. elegans.

 

Material examined

 

CBS 239.56, type culture, isolated by J. Meyer from soil, Yangambi, Congo.

CBS 105.66, isolated by J. H. van Emden from soil, Wageningen, Netherlands.

CBS 208.73, isolated by G. C. Bhatt from white cedar soil as strain UW 232, Puslinch, Ontario.

CBS 258.73 A = NRRL A-7737, isolated from soil, Forest Nursery, Albany, Georgia, USA.

CBS 218.73 B = NRRL A-5839, isolated from soil, collected by D. I. Fennell in Flo­rida, USA.

CBS 258.73 C = NRRL A-1572, isolated by E. S. Beneke from soil, Michigan, USA.

 

 

3. Mariannaea camptospora Samson, spec. nov. - Fig. 34.

 

Coloniae in agaro maltoso fere celeriter crescunt, ad 5-7 cm diam. 14 diebus 25° C, primum albae, deinde cremeae; reversum flavo-brunneum. Conidiophora erecta, septata, hyalina, verrucosa praecipue in parte basilari, ad 300 µm longa, 3-5 µm crassa, fere irregulariter ramosa, plerumque verticillatos ramos et quoties ternas ad senas phialides ferunt. Phialides 12.5-25 x 5.5-3.5 µm, anguste lageniformes. Conidia hyalina, levia, ellipsoidea vel ovoidea, una parte concava vel applanata, 5-6 x 2-2.5 µm. Typus CBS 209.73, lectus ab E. Jansen e terra sylvatica prope Arnhem in Hollandia.

 

Colonies on malt-agar growing moderately fast, attaining a diameter of 5 to 7 cm within 14 days at 25°C, consisting of a basal felt with floccose [p. 79] to funiculose overgrowth; due to the rich sporulation sometimes ap­pearing powdery; with age slightly zonate; at first white, later changing to cream. Reverse yellow-brown, with age sometimes turning brown. Odour absent. Vegetative hyphae hyaline, smooth-walled to verrucose, septate, 3-8 µm wide. Submerged hyphae yellow, inflated up to 50 µm. Coni­diogenous structures irregular and complex, but usually consisting of verticillate stalks, bearing short branches with whorls of 3 to 6 phialides or whorls of phialides in verticils along the main stalk; conidiophore stalks erect, septate, hyaline, verrucose especially near the base, usually up to 300 µm long and 3-5 µm wide. Phialides hyaline, smooth-walled, 12.5­-25 x 1.5-3.5 µm, slenderly flask-shaped. Conidia hyaline, smooth-walled, ellipsoidal to ovoidal, concave or flattened on one side, 5-7 x 2-2.5 µm. Chlamydospores not observed.

 

 

Fig. 34. Mariannaea camptospora, a-c. conidiophores; d-e. conidia. [p. 80]

 

Material examined

 

CBS 209.73 = type culture, isolated by E. Jansen from forest soil, Arnhem, Netherlands.

CBS 210.73, isolated by V. Holubovį-Jechovį from dead fallen trunk of Quercus sp., near Rhederoord, Netherlands.

CBS 215.73A-E, isolated by B. Söderström from soil (acid humus layer with a pH of 3 to 4), Meerdink, Netherlands.

CBS 216.73A and B, isolated by J. Grinbergs from decayed bark of Podocarpus spec., Puerto Varas, Chile.

CBS 231.73, isolated by J. Grinbergs from Aextoxicum punctatum, island Mocha, Chile.

CBS 942.73, isolated by J. C. Went (cellulolytic strain 5839) from soil, Arnhem, Netherlands.

 

Discussion

 

Mariannaea camptospora differs from M. elegans by its concave or flattened conidia. In addition the colonies of M. camptospora are more floccose to funiculose and have a yellow-brown reverse.

 

8. The genus Nomurea

 

Nomuraea Maublanc - Bull. Soc. mycol. Fr. 19: 296. 1903.

 

Generic description

 

Colonies grow slowly on artificial media. Vegetative hyphae hyaline to slightly pigmented, smooth-walled, septate, often inflated. Conidiophores mono- or synnematous, forming verticils of branches and phialides, occurring in compact clusters, encircling the stalk below the septa. Phialides in whorls, (broadly) cylindrical or sometimes with a swollen base, with a very short neck. Conidia in dry basipetal chains, one-celled, ellipsoidal or cylindrical, smooth-walled, hyaline or slightly pigmented. Perfect state unknown. Entomogenous.

Type species: Botrytis rileyi Farlow.

 

Discussion

 

The genus Nomuraea was erected by Maublanc (1903) on the single species N. prasina, which was found on the larvae of Pionea forficalis. Sawada (1919) transferred this species to Spicaria, probably because of the formation of the conidia in chains. Charles (1936) recognized the identity of Spicaria prasina with a fungus described by Farlow (1883) as Botrytis rileyi. [p. 81] Brown and Smith (1957) transferred most of the species of Spicaria to the genus Paecilomyces, except S. rileyi, since it differed from all other Paecilomyces species by the green colour and the shape of the phialides; the authors suggested that the species might belong to the Divaricata section of Penicillium. For the accomodation of S. rileyi and the related Isaria atypicola the retention of the genus Nomuraea is proposed by Kish et al. (1974).

The green colour and the phialide shape of N. rileyi indicate a close relationship with the genus Penicillium. In some species, e.g. P. nalgiovense Laxa and P. patulum Bain. (= P. urticae Bain.), the phialides also lack a distinct neck. Nomuraea differs from Penicillium by the arrangement of its conidial structures in numerous compact clusters along the conidiophore. Moreover, the two species of Nomuraea are insect parasites. The genus Nomuraea is also reminiscent of the entomogenous genus Metarrhizium Sorok. In this genus phialides are long cylindrical and closely compacted, borne on short, sporodochial conidiophores, while the conidia are produced in columns. The green species of Trichoderma are distinct from Nomuraea by the rapid growth, the divergent flask-shaped, mostly swollen phialides with a distinct neck and conidia produced in heads.

The phialide shape of the species of Nomuraea shows some resemblance to those occurring in the genera Akanthomyces (e.g. A. aranearum) and Gibellula. Akanthomyces can be easily distinguished from Nomuraea by the hymenium-like arrangement of the phialides, which cover the synnema densely or sometimes scattered along the outer synnema hyphae (Samson and Evans, 1974). In Gibellula the conidial structures form Aspergillus-like heads (cf. Samson and Evans, 1973).

 

Key to the species

 

1a.

Colonies green; conidiophores mononematous; conidia broadly ellipsoidal to cylindrical, 3.5-4.5 x 2-3.1 µm

N. rileyi

1b.

Colonies in purple shades; conidiophores mono- or synnematous, conidia cylindrical, slightly curved, 4.0-6.0 x 1.2-1.5 µm

N. atypicola

 

 

1 Nomuraea rileyi (Farlow) Samson - Fig. 35, a-b.

 

Botrytis rileyi Farlow - Rep. U.S. Comm. Agric. 1883: 121. = Spicaria rileyi (Farlow) Charles - Mycologia 28: 398. 1936 = Beauveria rileyi (Farlow) Gosswald - Arb. biol. BundAnst. Land- u. Forstw. 22: 434. 1939 = Nomuraea rileyi (Farlow) Samson apud Kish, Samson & Allen - J. Invert. Path. (in press).

Nomuraea prasina Maublanc - Bull. Soc. mycol. Fr. 19: 296. 1903 = Spicaria prasina (Maublanc) Sawada - Spec. Bull. Agric. exp. Stn Formosa 19: 606. 1919.

Spicaria stricta Petch - Trans. Br. mycol. Soc. 27: 89. 1944.

 

Colonies on malt-agar growing slowly, attaining a diameter of 0.7-1.2 cm within one month at 25° C; consisting of a basal felt from which erect [p. 82] conidiophores arise, usually closely together, occasionally covered with a floccose overgrowth of aerial mycelium and the conidial structures occurring only in localized areas. At first the colour of the colony is pale green near Pale Turtle Green (Ridgway, Pl. 32; Rayner, 3I"f), later changing to Olivine-Green (Ridgway, Pl. 32; Rayner, 23m) to Malachite Green (Ridgway, Pl. 32; Rayner, 35"b). Odour and exudate lacking. Reverse uncoloured or yellow. Vegetative hyphae smooth-walled septate, hyaline to slightly pigmented, 2-3 µm in diameter. Conidiophores erect, septate, arising from the submerged hyphae, up to 160 µm in length and 2-2.5 µm in diameter, forming dense clusters of branches, each bearing two or three compacted phialides, around the stalk. All conidiogenous structures are smooth-walled and hyaline to slightly green. The branches measure 4-6.5 x 2.2-3.5 µm. Phialides 4.7-6.5 x 2.3-3 µm, usually short cylindrical, occasionally with a swollen base, with neck absent or very short. Conidia in dry divergent chains, broadly ellipsoidal, sometimes cylindrical or nearly so, smooth-walled, pale green, green in mass, 3.5-4.5 x 2-3.1 µm.

In nature the fungus covers insect larvae with a rather thin white felt of hyphae from which compacted conidiophores arise, forming a somewhat pale green layer.

Because of its slow growth, N. rileyi is rather difficult to cultivate. On most media growth is very poor to scanty. On cornmeal-agar growth is also slow, but after one month the colonies sporulate very richly. The colour of the colonies on this medium is pale green, approximating Pale Turtle Green (Ridgway, Pl. 32; Rayner, 31"f). In poorly developed cultures atypical phialides may occur, producing conidia of variable shape and size; such phialides consist of a cylindrical base, tapering into a long distinct neck, resembling those of the genus Paecilomyces. However, fresh isolates and specimens on the natural substrate always show only phialides with a very short neck.

 

Material examined

 

Herbarium specimens

 

Botrytis rileyi, type, on Plusia brassicae, Washington, D.C., USA; 10 additional specimens, collected and determined by R. Thaxter from different localities in USA (accession nos 123, 130, 132, 133, 135, 138, 139, 145, 2724; FH).

Spicaria prasina, collected by H. K. Plank, Mayaguez, Puerto Rico, 1943, determined by V. K. Charles; collected by Jahle on a caterpillar, 1951 (BPI).

Spicaria (Botrytis) rileyi, collected by V. K. Charles on an unnamed caterpillar at Vadosta, 1935; collected by J. R. Johnston on grassworm, at Romano, Santo Domingo, 1913 (in herb. J. A. Stevenson); collected by E. A. Vaugon on Prodenia ornithogali, Orlando, 1924; collected by J. E. Jenkins in larvae on Asparagus ferns, Auburndale (BPI). as Spicaria javanica, collected by A. S. Rhoads as parasite of the velvet bean caterpillar (Anticarsia gemmatalis) in Florida (BPI).

Spicaria stricta, type, on a spider in Great Britain, ex herb. Petch (K). [p. 83]

 

Living cultures

 

CBS 424.51, received as Nomuraea prasina from K. Kominami (Japan).

CBS 194.65, isolated by K. H. Veen from Heliothis armigera, la Miničre, Versailles, France.

CBS 807.71 = IFO 8560, received as Spicaria prasina from the Institute of Fermentation, Tokyo (Japan).

CBS 806.71 = NRRL 5885, sent as Spicaria sp. by D. I. Fennell and isolated by C. R. Jackson from Trichoplusia ni.

CBS 732.73A and B, isolated by L. P. Kish from soybean loopers, Gainesville, Florida, USA.

CBS 733.73, isolated as strain 2843 by R. Kenneth from Spodoptera littoralis, Kfar Pines, Israel.

 

 

Fig. 35. Nomuraea rileyi, a. conidiophores; b. conidia; Nomuraea atypicola, c. conidiophores; d. conidia.

 

Discussion

 

Getzin (1961) reported N. rileyi as an entomogenous fungus, which caused an epizootic among the populations of the cabbage looper, Trichoplusia ni in Texas. The fungus killed early-instar Trichoplusia larvae, but not the later instars.

Experiments by Bell and Hammalle (1970) with N. rileyi for control [p. 84] of the cowpea curculio, Chalcodermus aeneus, showed, however, that the fungus caused no mortality on this insect, while other fungi, Beauveria bassiana and Metarrhizium anisopliae, killed the larvae after 6 days.

 

 

2. Nomuraea atypicola (Yasuda) Samson, comb. nov. - Fig. 35, c-d.

 

? Spicaria araneae Sawada - Bot. Mag. Tokyo 18: 309. 1914.

Isaria atypicola Yasuda - Bot. Mag. Tokyo 19: 117. 1915 (basionym) = Spicaria (Isaria) atypicola (Yasuda) Petch - Trans. Br. mycol. Soc. 23: 140. 1939.

Isaria pachylomera Kawamura - Jap. J. Bot. 4: 229. 1929.

Spicaria velutiformis Petch - Trans. Br. mycol. Soc. 16: 238. 1932.

 

Colonies on oatmeal-agar growing moderately fast, attaining a diameter of 6 cm within 4 weeks at 25° C, consisting of a compact crustose felt of numerous conidiophores, occasionally covered with a floccose overgrowth of aerial mycelium, zonate. At first white, soon becoming purple with conidium production, in age changing to dark purple colours. Odour and exudate absent. Reverse yellow. Vegetative hyphae smooth-walled, 2.5-4 µm wide, often inflated and then up to 15 µm in diameter. Conidiophores erect, densely grouped, arising from the submerged hyphae, up to 150 µm in length and 3.5-5 µm in diameter, forming dense clusters of branches, each bearing two or three compacted phialides around the stalk. Branches swollen, 6-7 x 3-5 µm. Phialides 3.5-5 x 2-2.5 µm, usually cylindrical, neck absent or very short. Conidia in dry divergent chains, cylindrical, slightly curved, smooth-walled, hyaline to slightly purple, 4-6 x 1.2-1.5 µm.

On the insect the fungus produces synnemata or covers it by a purple layer of conidiophores. Synnemata are grey-purple, consisting of a sterile cylindrical stipe, 20-30 x 2.7-3.5 mm, and a swollen fertile part, 15-30 x 3-4 mm. The hyphae are smooth-walled, hyaline to slightly pigmented, 2.5-4.5 µm in diameter. Conidial structures as in pure culture. Perfect state not observed.

 

Material examined

 

Herbarium specimens

 

Isaria atypicola, collected by E. Yakushiji and M. Kuwazawa on pupa of Kishinoyeus typicus, in the Botanical Garden Koishikawa, Tokyo, Japan (BPI).

Isaria species, in herbarium Lloyd, no. 37680, collected by S. Kawamura, Sendai (Japan), 1909 (BPI).

Spicaria velutiformis, type, on a spider collected in Vavuniya, Ceylon, Dec. 1923, and additional specimens collected by T. Petch from various localities in Ceylon (K).

 

Living cultures

 

CBS 744.73 = IFO 9205, sent as Isaria atypicola by K. Tubaki.

CBS 743.73, isolated by L. P. Kish from a spider, Gainesville, Florida, USA. Strains isolated by H. C. Evans, Tafo, Ghana: CBS 220.73, from a spider; CBS 731.73 A and C, from an Aethus species; CBS 731.73 B, from a spider. [p. 85]

 

Discussion

 

From the description of Isaria pachylomera Kawamura it seems evident that this species is synonymous with Nomuraea atypicola. Spicaria araneae Sawada (1934) was described as being closely related to S. prasina. Brown and Smith (1957) suggested that the species might be close to Penicillium lilacinum. No type material could be studied and it is therefore regarded as a doubtful synonym of N. atypicola.

Nomuraea atypicola is said to be common in gardens and on roadsides in Japan (Kobayasi, 1941).

 

 

9. The genus Septofusidium

 

Septofusidium W. Gams - Cephalosporium-artige Schimmelpilze, p. 147. 1971.

 

Generic description (from Gams, 1971b)

 

Acremonium-like, parasitic on leaf-inhabiting fungi, growing sparingly in pure culture. Conidiophores basitonously branched. Phialides awl-shaped with apical wall-thickening. Conidia in chains, spindle-shaped, one-celled or in culture 2-(to 4-) celled.

Type species: Septofusidium elegantulum (Pidopl.) W. Gams (for description cf. Gams, 1971b).

 

Discussion

 

Septofusidium elegantulum became known as a parasite of Meliolaceae and other leaf-inhabiting ascomycetes. Probably because of this pecularity, axenic growth was very poor and the pure culture of the neotype has been lost in the meantime. After long hesitation it was decided to accommodate the species Paecilomyces berolinensis and P. herbarum in this genus. This transfer necessitates the broadening of the generic concept. Both species are saprophytes with ornamented conidia, which are one-celled in P. herbarum and one- to more-celled in P. berolinensis. The genus Septofusidium resembles Gabarnaudia, which has thin-walled phialides integrated in septate conidiophore branches and white fast-growing colonies.

 

Key to the species

 

1a.

Conidia smooth-walled; colonies white to cream with poor but spreading growth in pure culture; parasitic on leaf-inhabiting ascomycetes [p. 86]

S. elegantulum

1b.

Conidia rough-walled to echinulate; colonies brownish; with good growth in pure culture; not parasitic on other fungi

2

 

 

 

2a.

Conidia one-celled with truncate ends; phialides smooth- or rough-walled

S. herbarum

2b.

Conidia one- to seven-celled with a truncate base; phialides warty

S. berolinense

 

 

1. Septofusidium berolinense (Olįh & Ackermann) Samson, comb. nov. - Fig. 36.

 

Paecilomyces berolinensis Olįh & Ackermann [as `berlinensis'] - Bull. trimest. Soc. mycol. Fr. 85: 494. 1969 (basionym).

 

 

Fig. 36. Septofusidium berolinense, a-b. conidiophores; c-d. conidia.

 

Colonies on malt-agar growing slowly, attaining a diameter of about 2.5 cm within 14 days at 25° C, consisting of a dense matted felt from which numerous erect conidiophores arise; slightly zonate, powdery; at [p. 87] first white, changing to Pale Cinnamon Pink (Ridgway, Pl. 29; Rayner, 13"f). Reverse red-brown from exudate, which also diffuses into the surrounding agar. Odour conspicuous, but not distinctive. Vegetative hyphae hyaline to yellow, 2.5-5.0 µm wide, smooth-walled or rarely warty. Conidiophores basitonously verticillate, mostly arising from the submerged hyphae, 50-220 µm in length. Phialides 20-35  x 2.0-4.0 µm, sometimes integrated in septate branches, awl-shaped, when young smooth-walled, becoming warty, hyaline to yellow. Conidia in long divergent chains, cylindrical, echinulate, one-celled or septate with up to 7 septa, all states of septation occurring in the same chain, hyaline to yellow, one-celled measuring 5.0-8.0 x 3.0-4.0 µm, when septate up to 16-19 x 4-5 µm. Chlamydospores not observed.

 

Material examined

 

CBS 731.70, type culture, isolated from air by H. W. Ackermann, Berlin, Germany.

 

Conidium ontogeny and ultrastructure of the hyphal wall of S. berolinense were investigated by Olįh (1972).

 

 

2. Septofusidium herbarum (Brown & Smith) Samson, comb. nov. - Fig. 37.

 

Paecilomyces herbarum Brown & Smith - Trans. Br. mycol. Soc. 40: 43. 1957 (basionym).

 

 

Fig. 37. Septofusidium herbarum, a-b. conidiophores; c. conidia. [p. 88]

 

Colonies on malt-agar growing slowly, attaining a diameter of 2 cm within 14 days at 25° C, consisting of a basal felt with thin overgrowth of aerial mycelium; conidial structures very sparse. Colonies on oatmeal-agar producing abundant conidial structures after 14 days; brown near Avellaneous, becoming Wood Brown (Ridgway, Pl. 40; Rayner, 17"'b), reverse yellow to brown. Exudate sometimes formed as colourless drops. Odour absent. Vegetative hyphae smooth-walled to roughened, hyaline to yellow, 0.7-2.5 µm wide. Conidiophores basitonously verticillate, mostly arising from the submerged hyphae, 50-120 µm in length. Phialides 15-30 x 3-4 µm, sometimes integrated in septate branches, awl-shaped, when young smooth-walled, sometimes becoming roughened, hyaline to slightly yellow. Conidia in long divergent chains, fusiform with truncate ends, roughened to verrucose, one-celled, 8-9.5 x 3-3.8 µm.

 

Material examined

 

Herbarium specimens

 

Paecilomyces herbarum on Heracleum, Gunnera scabra, Petasites ovatis and Urtica dioica (IMI) (cited by Brown and Smith, ķ957).

 

Living cultures

 

CBS 265.58 = IMI 53581 = type culture isolated by C. Booth from Urtica dioica, Mudgrave Woods, Yorks., Great Britain.

CBS 486.67, isolated by W. Gams from cut stems of Larix leptolepis, Gut Schierensee, Kr. Rendsburg, Germany.

CBS 948.73, isolated by W. Gams from leaf-sheath of Musa in a glass-house in Maartensdijk , Netherlands.

 

 

10. The genus Gabarnaudia Samson & W. Gams, gen. nov.

 

Coloniae albae, tenues, fere celeriter crescunt. Hyphae ad 4 µm crassae, leves, hyalinae. Conidiophora basitone vel mesotone ramosa, tenuitunicata, a hyphis vegetativis paulo distincta. Phialides aculeatae, discretae vel in ramis septatis integratae. Conidia plerumque longis catenis basipetalibus connexa, continua, cylindrica, fusiformia vel ellipsoidea, hyalina, levia. Chlamydosporae absunt.

Species typica: Gabarnaudia betae (Delacr.) Samson & W. Gams.

 

Generic description

 

Colonies white, thin, growing usually moderately fast on agar media. Hyphae up to 4 µm wide, smooth-walled, hyaline and septate. Conidiophores basitonously or mesotonously branched, thin-walled and little [p. 89] differentiated from vegetative hyphae. Phialides awl-shaped, discrete or integrated in plagiotropic septate branches, with apical internal wall-thickening. Conidia formed in basipetal succession, usually in long chains, sometimes sliming down, one-celled, cylindrical, fusiform or ellipsoidal, hyaline, smooth-walled. Typical chlamydospores absent.

Ascigerous state: Sphaeronaemella Karst.

Type species: Gabarnaudia betae (Delacr.) Samson & W. Gams.

The genus name is dedicated to the late French mycologist Gabriel Arnaud, who described two species very accurately.

 

Discussion

 

The genus Gabarnaudia resembles the genera Acremonium (sect. Nectrioidea) and Septofusidium by the awl-shaped phialides and the complex basitonously branched conidiophores. The latter genus is characterized by the rather slow, often poor growth in pure culture, by the catenate conidia which are one- to more-celled, and the sometimes slightly pigmented and ornamented phialides. The species of the sect. Nectrioidea of Acremonium usually form conidia in heads on shorter and narrow, mostly discrete phialides, which often also occur solitarily.

The perfect states of G. fimicola and G. humicola belong to the genus Sphaeronaemella Karst., as described by Seeler (1943) and Cain and Weresub (1957). In a more recent collection of the type species of Sphaeronaemella, S. helvellae Karst., conidia similar to those of S. fimicola were described by Seeler (l.c.). On the type specimen (H), however, no conidial state could be found. In S. humicola, the ascomata are non-ostiolate, but the structures of ascomata and asci agree perfectly with the ostiolate species. According to von Arx (1973) the presence or absence of an ostiolum is often not sufficient to distinguish genera.

 

Key to the species

 

1a.

Conidia fusiform, with a wall-thickening at the pointed apex

G. tholispora

1b.

Conidia elongate with a rounded apex

2

 

 

 

2a.

Sphaeronaemella state present in pure culture, ascomata non-ostiolate; conidiophores branching within the agar; soil-borne

G. humicola

2b.

Sphaeronaemella state absent or only formed near colonies of contaminant fungi; conidiophores branching above the agar surface

3

 

 

 

3a.

Conidiophores repeatedly branched; conidia regularly cylindrical, usually not giving rise to secondary conidia; on plant debris or parasitic on other fungi

G. betae

3b.

Conidiophores few times branched; conidia irregularly cylindrical, [p. 90] usually tapering in the lower half; germinating with short phialides, which form secondary almost rod-shaped conidia; coprophilous

G. fimicola

 

 

1. Gabarnaudia betae (Delacr.) Samson & W. Gams, comb. nov. - Fig. 38 a-b, 39 a-c.

 

Oospora betae Delacr. - Bull. Soc. mycol. Fr. 13: 166. 1897 (basionym) = Penicillium betae (Delacr.) Biourge - La Cellule 33: 100. 1923 = Paecilomyces betae (Delacr.) Cornford - Trans. Br. mycol. Soc. 43: 155. 1960.

Oospora lasiosphaeriae Arnaud - Bull. trimest. Soc. mycol. Fr. 68: 195. 1952 (without latin diagnosis).

 

 

Fig. 38. Original drawings by G. Arnaud. Oospora lasiosphaeriae, a. conidiophores; b. conidia. Oospora tholispora, c. conidiophores; d-e. conidia. Magnification 600x, except d: 1200x.

 

Colonies on malt-agar growing rather fast, attaining a diameter of 40-50 mm within 10 days at 20-24° C, consisting of a white, usually thin [p. 91] mycelium, which becomes floccose to funiculose with age. Odour sweetish (like Sistotrema cultures) or spermatic. Conidiophores arising from the submerged hyphae, repeatedly branched, septate. Phialides integrated, awlshaped, 30-42 x 2.5-3.0 µm, tapering to 1.5-2.0 µm at the tip, with internal wall-thickening. Conidia in chains or sliming down with age, regularly cylindrical or slightly ellipsoidal, with a truncate base and rounded apex, hyaline, smooth-walled, 8-13 x 3.5-4.5 µm. Conidia do normally not germinate with formation of short phialides and secondary conidia.

 

 

Fig. 39. Gabarnaudia betae, a-b. conidiophores; c. conidia. Gabarnaudia tholispora, e-f. conidiophores; g. conidia; h. chlamydospore-like cells.

 

Material examined

 

Herbarium specimens

 

Paecilomyces betae, neotype, IMI 72674, isolated by C. E. Cornford from Beta vulgans, Dunholm, Lines., Great Britain (IMI). [p. 92]

Oospora lasiosphaeriae, original drawings (in part reproduced here as Fig. 38 a, b) and description by G. Arnaud (PC).

 

Living cultures

 

CBS 733.69, isolated by W. Gams from decaying trunks of µmus sylvestris, together with Chalara spec., Deelerwoud, Netherlands.

CBS 350.70, isolated by W. Gams from Urtica dioica stems, Kiel-Kitzeberg, Germany.

 

Discussion

 

G. betae resembles G. fimicola, which has less frequently branched conidiophores and conidia of more irregular shape. The conidia in the latter species tend after germination to form phialides with narrower secondary conidia. These differences are difficult to recognize in older strains and species delimitation is not very sharp.

 

 

2. Gabarnaudia fimicola Samson & W. Gams, spec. nov. - Fig. 40.

 

Ascigerous state: Sphaeronaemella fimicola Marchal - Bull. Soc. R. Bot. Belg. 33: 12. 1894.

 

Coloniae in agaro maltoso fere celeriter crescunt, 10-28 mm diam. post 1o dies 20-24° C, albae, tenues, nonnumquam floccosae vel funiculosae. Conidiophora plerumque e hyphis submersis oriuntur, semel atque iterum ramosa, ad 200 µm alta. Phialides discretae vel in ramis septatis integratae, 45-80 x 3.5-4.0 µm, ad 1.5-2.5 µm attenuatae. Conidia catenata, hyalina, levia, fusiformia vel late ellipsoidea, basi truncata, apice rotundato vel modice acutato, 12-18 x 3.5-6.0 µm; germinando saepe conidia secundaria angustiora generant.

Typus: CBS 348.70, isolatus e stercore bovino in Germania.

 

Colonies on malt-agar growing moderately fast, attaining a diameter of 10-28 mm within  days at 20-24° C, consisting of a white, thin, occasionally floccose to funiculose mycelium, in age with a wet appearance. Odour not pronounced. Hyphae hyaline, smooth-walled, 1-4 µm wide. Conidiophores arising primarily from submerged hyphae, few times branched, up to 200 µm high. Phialides discrete or integrated in septate branches, 45-80 x 3.5-4.0  µm, tapering towards 1.5-2.5 µm, the tip hardly constricted and with internal wall-thickening; lateral proliferation sometimes occurs. Conidia in long chains, which may slime down in age, hyaline, smooth-walled, fusiform to broadly ellipsoidal, with a truncate base and rounded or slightly pointed apex, 12-18 x 3.5-6.0 µm. The conidia often germinate with short phialides and form secondary almost rod-shaped conidia, 1.5 -3.0 µm wide (Fig. 40 c).

For a detailed description of Sphaeronaemella fimicola see Pease (1948) and Cain and Weresub (1957). [p. 93]

 

 

Fig. 40. Gabarnaudia fimicola, a. conidiophore; b. conidia; c. germinating conidia with short phialides; d. secondary conidia; e. chlamydospore-like cells; f. ascomata of Sphaeronaemella fimicola; g. asci; h. ascospores.

 

Material examined

 

CBS 347.70, isolated from rabbit dung, Baarn, Netherlands.

CBS 348.70, type culture, isolated from Ascophanus spec., on cow dung, Schüttbrehm, Kr. Plön, Germany.

CBS 349.70, isolated from chamois dung, Karwendel, Tyrol, Austria.

CBS 493.72 and 113.72, isolated from pony dung, Dartmoor, Great-Britain. The last 3 strains were isolated from ascospores. Several other strains from dung, collected in the Netherlands. [p. 94]

 

Discussion

 

Sphaeronaemella fimicola seems to be a common fungus on dung of several herbivorous animals and is often accompanied by its Gabarnaudia state, which grows particularly on the undecomposed plant remains. In vitro S. fimicola usually produces only the imperfect state. Pease (1948) and Cain and Weresub (1957) have shown that fresh isolates produced the ascigerous state on agar media, when the fungus was cultivated under wet conditions or in association with other fungi, e.g. Eurotium repens or Microascus sordidus. Our attempts to obtain the ascigerous state with several CBS strains under the same conditions failed. Pure fresh isolates from ascospores of S. fimicola produced only the conidial state on agar media and sterilized dung; after contamination with bacteria and other dung fungi (e.g. Sporormia spec.), however, also ascomata appeared. The species is apparently homothallic.

S. fimicola differs from S. helvellae by the dark wall of the ascomata and the smaller, curved ascospores. In S. helvellae the ascomata are yellow-brown and the ascospores are cylindrical. The ascomata of this species are reminiscent of another fungus frequently found on decomposing agarics, Hyalopycnis blepharistoma (Berk.) Höhnel, which forms pycnidia with a long beaked apex (Seeler, 1943, Petch, 1943). In two strains (CBS 953.73, 954.73) of H. blepharistoma, isolated from rotten Russula spp., the conidium formation on branching conidiophores within pycnidia could be observed easily. No living cultures of S. helvellae were available. In the type collection (H) no asci could be observed, but groups of 8 ascospores. In all species of Sphaeronaemella the asci are evanescent.

 

 

3. Gabarnaudia humicola Samson & W. Gams, spec. nov. - Fig. 41; Pl. 2.

 

Ascigerous state: Sphaeronaemella humicola Samson & W. Gams, spec. nov.

 

Coloniae celeriter crescunt, 5-8 cm diam. post 14 dies 20-24° C, albae, tenues; odor

spermaticus. Ascomata superficialia vel in agaro submersa, tribus ad quattuor hebdamadibus maturantia, globosa, 80-250 µm diam., nonnumquam comam hypharum fasciculatarum velut ostiolum formant, 200-750 µm longam; peridium flavobrunneum vel rubrobrunneum, e compluribus stratis cellularum compositum. Asci fere globosi, 10-15 µm, evanescentes, plerumque octospori. Ascosporae continuae, pallide brunneae, late ellipsoideae, crassitunicatae, leves, 5-6 x 4-5 um. Species homothallica.

Conidiophora compluries ramosa, plerumque in agaro submersa, singulis vel binis ramis quotiescumque oriundis. Phialides aculeatae, discretae vel in ramis septatis integratae. Conidia catenulata vel saepe mucilagine compacta, cylindrica vel ellipsoidea, basi truncata et apice rotundata, 13-22(-27) x 4-5(-6.2) µm.

Typus CBS 115-72, isolatus a J. H. van Emden e terra agresti, Wageningen.

 

Colonies growing fast, attaining a diameter of 5-8 cm within 14 days at 20-24° C, consisting of a white, thin felt of mycelium which sometimes becomes funiculose or tufted, with a feathery margin. Odour spermatic. [p. 95]

 

 

Fig. 41. Gabarnaudia humicola, a-b. conidiophores; c. conidia; d. ascoma of Sphaeronaemella humicola; e. ascoma initial; f. asci; g. ascospores.

 

Conidiophores repeatedly branched, mostly within the agar, with 1-2 branches arising at each level. Phialides awl-shaped, discrete or integrated in septate branches. Conidia in chains, which slime down very quickly, cylindrical or ellipsoidal, with a truncate base and a rounded apex, 13-22 (-27) x 4-5 (-6.2) µm. Ascoma initials consisting of two swollen cells surrounded by a trichogyne which forms a dense cluster. Ascomata produced on the surface of the agar or submerged, ripening within 3-4 weeks, globose, 80-250 µm in diam., sometimes ending in an ostiole-like, 200-750 µm long tuft of loose hyphae; peridium yellow-brown or red-brown, [p. 96] composed of several layers of cells. Asci more or less globose, 10-15 µm, evanescent, usually 8-spored. Ascospores one-celled, brownish, broadly ellipsoidal, thick- and smooth-walled, 5-6 x 4-5 µm. The species is homothallic.

 

Material examined

 

CBS 115.72, type culture, isolated by J. H. van Emden as strain 700818/535 from agricultural soil, Wageningen. Two identical strains had been isolated in 1964 by W. Gams from agricultural soil at Kiel-Kitzeberg, but were lost.

 

Discussion

 

Even when the fungus was cultivated under various conditions of nutrients, light, temperature or humidity, the ascomata had no ostiolum, but sometimes an apical tuft of long hairs. Under pressure of a coverslip a cirrus of ascospores can be released at any part of the ascoma (Pl. 2 D). Although the fungus is non-ostiolate, it is described as Sphaeronaemella, because it is corresponding with the ostiolate species by the structure of the ascoma wall, the initials, the asci and by the Gabarnaudia conidial state.

G. humicola differs from the other species by the mostly verticillate branching of the conidiophores, which begins within the agar, and the presence of a Sphaeronaemella ascigerous state with broadly ellipsoidal ascospores.

 

 

4. Gabarnaudia tholispora Arnaud ex Samson & W. Gams, spec. nov. - Fig. 38 c-e; 39 e-f.

 

Oospora tholispora Arnaud - Bull. trimest. Soc. mycol. Fr. 68: 195. 1952 (without latin diagnosis).

Fusidium parasiticum Westend. - Bull. Acad. Belg. Brux. 18: 412. 1851; sensu Backus & Stowell - Mycologia 45: 836. 1953 [non Westend.].

 

Coloniae in agaro maltoso fere lente crescunt, 18-22 mm diam. post to dies 20-24° C, densum stratum conidiophororum formant, albae. Conidiophora praecipue e hyphis submersis oriuntur, ramos singulos vel verticillatos formant. Phialides aculeatae, discretae vel in ramis septatis integratae, 25-80 x 3.0-4.0(-5.0) µm, ad 1.5-2.0 µm angustatae. Conidia plerumque longis catenis connexa, fere fusiformia, basi truncata et apice acutato et inspissato, hyalina, levia, (7-)9-16 x 3-5(-6) µm. Conidia non dimissa velut chlamydosporae in phialidibus manent. Germinando conidia saepe conidia secundaria angustiora generant.

Typus CBS 351.70, isolatus e Russula nigricante putrida, Baarn.

 

Colonies on malt-agar growing rather restrictedly, attaining a diameter of 18-22 mm within 10 days at 20-24° C, consisting of a dense layer of conidiophores, white. Conidiophores primarily arising from submerged [p. 97] hyphae, with verticillate or simple branches. Phialides awl-shaped, discrete or integrated in longer branches, 25-80 x 3.0-4.0 (5.0) µm, tapering towards 1.5-2.0 µm. Conidia in long chains, partially sliming down to form balls or irregular masses with age, more or less fusiform with a truncate base and a pointed apex with a thickened wall, hyaline, smooth-walled, (7-)9-16 x 3-5(-6) µm, the first formed conidium with a thin-walled, round apex.

Conidia which are not shed may form chlamydospore-like cell which are smooth-walled, hyaline, ellipsoidal to pyriform, 6.5-13 x 4 5-7 5 µm, produced singly on conidiophores, which may proliferate subterminally. The conidia germinate with short phialides and produce secondary, narrower and less pointed conidia.

 

Material examined

 

Herbarium specimens

 

Oospora tholispora, original drawings and description by G. Arnaud, preserved in PC, in part reproduced here as Fig. 38 c-e.

 

Living cultures

 

CBS 192.60, isolated from Xylaria oxyacanthae and sent to the CBS as Fusidium parasiticum by M. B. Backus, Madison, Wisconsin, USA.

CBS 351.70, type culture, isolated by A. C. M. Hoozemans, from Russula nigricans, together with Calcarisporium arbuscula, Groeneveld, near Baarn, Netherlands.

CBS 675.71, on Nectria cinnabarina, Staverden near Ermelo, Netherlands.

CBS 676.71, on dead leaves of Typha latifolia, Staverden near Ermelo, Netherlands.

CBS 677.71, isolated together with Gliocladium solani from rotten potato, Baarn.

CBS 221.73, isolated from Elaphomyces sp., collected by G. A. de Vries, Hulshorst, Netherlands.

Further isolates were obtained from a wheat culm-base and rust-infected leaves of Tussilago farfara in Germany, and from Scleroderma citrinum in the Netherlands.

 

Discussion

 

The type material of Oospora tholispora (PC) is not recognizable. The original drawings and illustrations (labelled Fusidium tholiforme, pl. 2427) by G. Arnaud prove, however, that the species described here is identical with G. tholispora.

Fusidium parasiticum Westend. is probably the conidial state of Xylaria cornuta (Gams, 1971b). The strain CBS 192.60 isolated by Backus and Stowell  (1953) grows only on cornmeal-agar, but not on malt-agar.

G. tholispora can be easily distinguished from the other species by the fusiform conidia. [p. 98]

 

 

11. Doubtful and excluded species

 

Spicaria acridiorum (Brongniart & Delacroix) Vuill. - Bull. trimest. Soc. mycol. Fr. 27: 76. 1911 = Botrytis acridiorum Brongniart & Delacroix (a later homonym of B. acridiorum Trabut). → Probably a Beauveria with globose conidia (de Hoog, 1972). Botrytis acridiorum Trabut is Trichothecium acridiorum (Trabut) Madelin (1966 b).

 

Paecilomyces ampullaris Matsushima - Microf. Solomon Isl., Papua p. 42.1971.

In the type collection (MFC 2716) the conidiogenous cells show a sympodial elongation. More material should be examined to ascertain its systematic position.

 

Spicaria anomala (Corda) Harz - Bull. Soc. imp. Nat. Moscou 44: 138. 1871 = Penicillium anomalum Corda - Icones Fung. 2: 76. 1838.

No type left and description too vague; doubtful species.

 

Spicaria arachnoidea Sacc. & Therry - Syll. Fung. 4: 167. 1886. Fischer (1892: p. 281) synonymized this species with Mortierella arachnoides Therry & Thierry - Revue mycol. 4: 161. 1882.

In the type specimen (PAD) of S. arachnoidea no fungus material was left. Some drawings on the envelope suggest, that the species might be Mariannaea elegans.

 

Paecilomyces aspergilloides Pidoplichko - Mikrobiologiya 12: 35. 1950. No type available, but probably a degenerated or atypical strain of Aspergillus or the phialidic state of Acrospeira, Chlamydomyces or similar genera.

 

Paecilomyces baarnensis Fassatiovį - Ceskį Mykol. 21: 156. 1967. In the type culture (CBS 409.67) synnemata are produced with awl-shaped divergent phialides and catenulate conidia. The species is therefore excluded from Paecilomyces and is regarded as identical with Oospora subfasciculata Petch. Gams (1971b) regarded this species as synonymous with Verticillium griseum (Petch) W. Gams. The now available isolates indicate, that the species is better kept distinct as Verticillium subfasciculatum (Petch) Samson & W. Gams, comb. nov. (basionym: Oospora subfasciculata Petch - Trans. Br. mycol. Soc. 16: 63. 1931).

 

Paecilomyces burci (Pollacci) Thom - The Penicillia, p. 548. 1930 = Penicillium burci Pollacci - Atti Ist. bot. Univ. Lab. crittogam. Pavia 18: 128. 1921.

Brown and Smith (1957) accepted this species provisionally. No cultures were found again and the identity of the species is doubtful. [p. 99]

 

Paecilomyces buxi (Schmidt in Link) Bezerra - Acta bot. néerl. 12: 63. 1963 = Fusidium buxi Schmidt in Link - Linné Spec. Pl. Ed. 4(2): 97. 1825 → Sesquicillium buxi (Schmidt in Link) W. Gams (1968).

 

Spicaria cephalospora Kamyschko - Nov. Sist. nizsh. Rast. I6: 95. 1963. Without indication of type not validly published; the description suggests that this species might be Mariannaea camptospora.

 

Spicaria clavulifera Petch - Trans. Br. mycol. Soc. 16: 238. 1932. In the type (K) and in specimens collected by H. C. Evans in Ghana penicillate conidiophores were observed with cylindrical phialides and catenate conidia. On some insects the fungus produces conspicuous synnemata. S. clavulifera does not fit in Paecilomyces because of the cylindrical phialides, but should be compared with Gibellula (Samson and Evans, in preparation).

 

Paecilomyces coccosporus (Drechsler) Brown & Smith - Trans. Br. mycol. Soc. 40: 74. 1957 = Spicaria coccospora Drechsler - Phytopathology 31: 787. 1941.

The species is described as a nematophagous fungus on Plectus parvus; probably a good species of Paecilomyces, but no material available.

 

Spicaria colorans de Jonge - Recl Trav. bot. néerl. 6: 48. 1909 → Fusarium decemcellulare Brick, fide Booth (1971).

 

Spicaria crocea (Kunze ex Pers.) Oudem. - Enum. syst. Fung. 1: 281. 1919 [Ozonium croceum Kunze ex Pers.] = Alytosporium croceum (Kunze ex Pers.) Link - Linné Spec. Pl. Ed. 4(I): 24. 1824 = Sporotrichum croceum Kunze - Mykol. Hefte I: 81. 1817; fide Rogers and Jackson (1943) this species is identical with Corticium sulphureum sensu Burt = Corticium bicolor Peck. According to Jülich (1972) the latter species is Piloderma bicolor (Peck) Jülich, which is not synonymous with Ozonium croceum, a nomen dubium.

 

Spicaria delacroixii (Sacc.) Vuill. - Bull. Séanc. Soc. Sci. Nancy, Sér. 3, II: 153. 1910 → Beauveria bassiana (Bals.) Vuill., fide de Hoog (1972).

 

Paecilomyces cremeo-roseus Batista - Anais Soc. biol. Pernamb. 15:149. 1957.

The type culture (CBS 250.55 = IMI 113844 = ATCC 120.76) of P. cremeo-roseus represents a poorly growing species of Penicillium with monoverticillate structures. No adequate description can be obtained from this culture and therefore no new combination in Penicillium is proposed.

 

Spicaria densa (Link) Vuill. - Bull. Séanc. Soc. Sci. Nancy, Sér. 3, II: 25. 1910 → Beauveria bassiana (Bals.) Vuill. [p. 100]

 

Spicaria (Isaria) dubia (Delacr.) Petch - Trans. Br. mycol. Soc. 21: 297. 1938 → Isaria dubia Delacr. - Bull. Soc. mycol. Fr. 9: 264. 1893 and Revue mycol. 16: 18. 1894.

No type material available, but from examination of material collected by T. Petch (K) and by E. Müller-Kögler (Darmstadt) it is obvious that this species can be excluded from Paecilomyces. The accommodation in Syngliocladium or a new genus is considered.

 

Spicaria elegans (Corda) Harz var. macrostachya Speg. - An. Soc. cient. argent. 22: 208. 1886.

The type (LPS 3777) contains a Clonostachys species.

 

Paecilomyces ehrlichii (Klebahn) Delitsch - Systematik der Pilze, p. 90. 1943 → Eupenicillium ehrlichii (Klebahn) Stolk & Scott (1967).

 

Paecilomyces eriophytis (Massee) Leatherdale - J. Invert. Path. 7: 325. 1965 = Botrytis eriophytis Massee apud Taylor - J. econ. Biol. 4: 5. 1909 → Verticillium lecanii (Zimm.) Viégas, fide Gams (1971b).

 

Paecilomyces erectus Demelius - Verh. zool.-bot. Ges. Wien 72: 78. 1923

No type material is available, but the original drawings by Paula Demelius (W) show a green fungus with penicilli and lanceolate phialides. This fungus represents a Penicillium species of the Biverticillata-Symmetrica section.

 

Spicaria (Isaria) erotylis Petch - Trans. Br. mycol. Soc. 21: 41. 1937. Petch (1937) described this species on Erotylus species (Coleoptera) collected in Trinidad. No type material is left in K, but from its description it might be Akanthomyces pistillariaeformis (Pat.) Samson & Evans (1974).

 

Spicaria felina (DC.) Vuill. cited in Biourge - La Cellule 33: 102. 1923 → Isaria felina (DC.) Fr.

 

Paecilomyces fimetarius (Moesz) Brown & Smith - Trans. Br. mycol. Soc. 40: 75. 1957 = Spicaria fimetaria Moesz - Bot. Közl. 19: 58. 1921. → ? Isaria cicadae Miquel, but no type material available.

 

Spicaria fumosa Ellis & Everh. - Bull. Torrey bot. Club 10: 97. 1883. The type specimen (NY) contains only a myxomycete and no fungus agreeing with the vague description of S. fumosa.

 

Paecilomyces fuscatus Inagaki - Trans. mycol. Soc. Japan 4: 4. 1962 → Scopulariopsis gracilis Samson apud Samson & Klopotek (1972).

 

Paecilomyces fusidioides Nicot - Cah. Maboké 6: 17. 1968 → Acremonium fusidioides (Nicot) W. Gams (1971 b). [p. 101]

 

Paecilomyces fusisporus Saksena - J. Indian bot. Soc. 32: 188. 1953 = Spicaria fusispora (Saksena) Gilman - Manual of Soil Fungi, 2nd ed. p. 308. 1956 → Acrophialophora fusispora (Saksena) Samson apud Samson & Tariq Mahmood (1970); syn. Masoniella indica Salam & RamaRao - Curr. Sci. 29: 149. 1960.

 

Spicaria gongylophorae Sacc. - Michelia 2: 552. 1882 → ? Phialocladus zsoltii Kreisel (1972).

 

Spicaria griseola Sacc. - Michelia 2: 552. 1882.

In the type specimen (PAD) no fungus was found; the description is inadequate; doubtful species.

 

Paecilomyces griseo-viridis Onions & Barron - Mycol. Pap. 107: 22. 1967 → Acremonium griseo-viride (Onions & Barron) W. Gams (1971 b).

 

Paecilomyces humicola Onions & Barron - Mycol. Pap. 107: 20. 1967 = Sagrahamala humicola (Onions & Barron) Subramanian - Curr. Sci. 41: 49. 1972 → Acremonium humicola (Onions & Barron) W. Gams (1971 b). [p. 101]

 

Paecilomyces laeensis Matsushima - Microf. Solomon Isl. Papua, p. 42. 1971. The type (MFC 2721) of P. laeensis was identified by W. Gams as Acremonium hyalinulum (Sacc.) W. Gams.

 

Spicaria lateritia Ciferri - Zentbl. Bakt. ParasitKde. (Abt. 2) 71: 79. 1927; fide Ciferri (letter to Brown and Smith, 1957) a poor strain of Neurospora crassa.

 

Spicaria mori Hara - A List of Japanese Fungi, 4th Ed., p. 400. 1954. No type available and description inadequate; doubtful species.

 

Spicaria mucoricola Speg. - An. Mus. nac. Hist. nat. B. Aires 6: 333. 1899.

In the type specimen (LSP no. 11334) some conidiophores were seen, probably Verticillium lecanii.

 

Spicaria nivea Harz - Bull. Soc. imp. Nat. Moscou 44: 137. 1871. Doubtful species, see Brown and Smith (1957).

 

Paecilomyces ochraceus Onions & Barron - Mycol. Pap. 107: 15. 1967 → Acremonium ochraceum (Onions & Barron) W. Gams (1971 b).

 

Paecilomyces paranensis (Marchionatto) G. Müller - Wiss. Z. Humboldt-Univ. Berlin, math. nat. R. 14: 780. 1965 = Sporotrichum paranense Marchionatto - Physis, B. Aires II: 348. 1933. → Metarrhizium anisopliae (Metschn.) Sorok.

In the type culture of S. paranense (IMI 74660) the conidia are slightly curved and pale green in colour, but other characters fit into M. anisopliae. [p. 102]

 

 

Fig. 42. Scopulariopsis parva, a-b. conidiophores; c. conidia.

 

Paecilomyces parvus Brown & Smith - Trans. Br. mycol. Soc. 40: 5 8. 1957.

The species is identical with Scopulariopsis parvula Morton & Smith (1963). In the type cultures of both species the conidiogenous cells elongate with age. The conidia have a truncate base. P. parvus therefore has to be classified as Scopulariopsis parva (Brown & Smith) Samson, comb. nov. (Fig. 42).

 

Material examined

 

CBS 245.32 = IMI 58410, type culture of Paecilomyces parvus, isolated from man. CBS 209.61 = IMI 86943, type culture of Scopulariopsis parvula, isolated from soil.

 

For description see Brown and Smith (1957) and Morton and Smith (1936).

 

Spicaria perpusilla Speg. - An. Soc. cient. Argent. 10: 163. 1880. In the type collection (LPS 11332) no fungus was found agreeing with the description. The drawings on the cover of the specimens suggest a Cladosporium species.

 

Paecilomyces persicinus Nicot - Bull. trimest. Soc. mycol. Fr. 74: 227. 195 → Acremonium persicinum (Nicot) W. Gams (1971 b.)

 

Paecilomyces persimplex (Klebahn) Delitsch - Systematik der Pilze, p. 91. 1943 = Penicillium persimplex Klebahn - Ber. dt. bot. Ges. 51: 318. 1933.

Doubtful species, also fide Raper and Thom (1949). [p. 103]

 

Spicaria purpurogenes Nadzon & Zolkiewicz - Izv. glav. bot. Sada SSSR 21: 1. 1922.

Doubtful species, also fide Brown and Smith (1957).

 

Spicaria purpuroides Hara - A List of Japanese Fungi, 4th Ed., p. 400. 1954.

No type available and description inadequate; doubtful species.

 

Spicaria (Isaria) rectangularis Petch - Annls Cryptog. exot. 6: 234. 1933. Petch (1933 a) described this fungus on lepidopterous larvae. No type available, but from the description the species might be Paecilomyces tenuipes.

 

Paecilomyces roseolus G. Smith - Trans. Br. mycol. Soc. 45: 388. 1962. Sagrahamala roseola (G. Smith) Subramanian - Curr. Sci. 41 49. 1972 Acremonium roseolum (G. Smith) W. Gams (1971 b).

 

Spicaria rubra (Baquis) Dodge - Med. Mycol. p. 843.1935 = Verticillium rubrum Baquis - Annali Ottal. Clin. ocul. 34: 947. 1905 = Tritirachium rubrum (Baquis) Langer. - Annls Parasit. hum. comp. 22: 98. 1947.

Doubtful species, see de Hoog (1972).

 

Spicaria satoi Hara - A List of Japanse Fungi, 4th Ed., p. 400, 1954. No type available and description inadequate; doubtful species.

 

Spicaria shirako Hara - A List of Japanese Fungi, 4th Ed., p. 400. 1954. No type available and description inadequate; doubtful species.

 

Spicaria silvatica Oudem. - Archs néerl. Sci., Sér. 2, 7: 291. 1902. → ? Penicillium digitatum Sacc., see discussion under Paecilomyces aerugineus.

 

Paecilomyces simplex (Petch) Brown & Smith - Trans. Br. mycol. Soc. 40: 76. 1957 = Spicaria simplex Petch - Naturalist, Hull 1936: 59. - Probably Verticillium catenulatum (Kamyschko ex Barron & Onions) W. Gams (fide Gams 1971 b).

 

Spicaria simplicissima Oudem. - Ned. kruidk. Archf,Ser. 3, 2:763.1903 = Penicillium simplicissimum (Oudem.) Thom (1930).

 

Paecilomyces smilanensis (Wize) Brown & Smith - Trans. Br. mycol. Soc. 4o: 76. 1957 = Spicaria smilanensis Wize - Bull. int. Acad. pol. Sci. Lett., Cl. Sci. math.-nat. 1904: 72.

No type available and not identifiable from description.

 

Spicaria smithii Oudem. - Ned. kruidk. Archf, Ser. 3, 2: 753. 1903 No type left, but Oudemans' original drawings (L) show a fungus producing [p. 104] roughened conidia with a distinctly truncate base. According to the description, S. smithii is considered as Scopulariopsis brevicaulis (Sacc.) Bain.

 

Spicaria solani Harting - Nieuwe Verh. kon. Inst. Wetensch. Amsterdam 12: 203. 1846 → Gliocladium solani (Harting) Petch (1944).

 

Paecilomyces striatisporus Onions & Barron - Mycol. Pap. 107: 19. 1967 = Sagrahamala striatispora (Onions & Barron) Subramanian - Curr. Sci. 41: 49. 1972 →Acremonium striatisporum (Onions & Barron) W. Gams (1971 b).

 

Paecilomyces terricola (Miller et al.) Onions & Barron - Mycol. Pap. 107: 10. 1967 = Fusidium terricola Miller & al. - Mycologia 49: 796. 1957 = Acremonium terricola (Miller & al.) W. Gams (1971 b) → Acremonium implicatum (Gilman & Abbott) W. Gams (1974).

 

Spicaria unguis Weil & Gaudin - Arch. Med. exp. Anat. path. 28: 452. 1918.

No type material available and description inadequate; fide Lodder and Kreger-van Rij (1952) doubtful species.

 

Spicaria valdiviensis Speg. - Revta Fac. Agron. Univ. nac. La Plata 6: 181. 1910.

In the type material (LPS 26699) only some hyphal fragments were found resembling arthroconidia. The description is inadequate; doubtful species.

 

Spicaria verticillata (Corda) Harz - Bull. Soc. imp. Nat. Moscou 44: 138. 1871 = Penicillium verticillatum Corda - Icones Fung. AT: 21. 1837. No type left (in PR) and description inadequate; doubtful species.

 

Paecilomyces variabilis Barron - Can. J. Bot. 39: 1576. 1961 = Scopulariopsis diversispora van Beyma - Zentbl. Bakt. ParasitKde (Abt. 2) 96: 430. 1937 → Acremonium diversisporum (van Beyma) W. Gams (1971b).

 

Paecilomyces victoriae (Szilvinyi) Brown & Smith - Trans. Br. mycol. Soc. 40: 601. 1957 = Penicillium (Paecilomyces) victoriae Szilvinyi - Arch. Hydrobiol. 1936: 535

The type culture (CBS 274.36) forms conidiophores with whorls of lanceolate phialides. Usually the penicilli are monoverticillate, but occasionally biverticillate symmetrical penicilli with green conidial heads occur. The species is therefore better classified in Penicillium.

 

Spicaria viridans Sasaki & Nakane - J. appl. Mycol. Japan 1: 105. 1946. Fide Brown and Smith (1957) probably identical with Metarrhizium anisopliae (Metschn.) Sorok. [p. 105]

 

 

Acknowledgements

 

The author is very grateful to all those, who contributed to this work by providing living cultures and herbarium material, especially Miss D. I. Fennell, Dr Agnes H. S. Onions, Dr H. C. Evans and Ir J. H. van Emden. He thanks Mme J. Nicot and Mme G. Arnaud for making available the original drawings of G. Arnaud.

He wishes to express his sincere gratitude to Dr J. A. von Arx and Prof. Dr K. Verhoeff for valuable critism during the preparation of the manuscript. He is much indebted to Dr W. Gams for amicable cooperation and many valuable suggestions. Thanks are also expressed to Mrs M. Tulloch for her comments on the taxonomy and Miss A. C. Stolk for discussions on genus delimitation. Mrs J. H. Brouwer-Zeilmaker and Miss E. Mul are acknowledged for typing the manuscript and technical assistance. Mrs M. Tulloch and Mr D. Yarrow kindly corrected the English text. The scanning electron micrographs were made with the co-operation of Mr J. A. J. M. Stalpers and the SEM Research Group of the University of Amsterdam. [p. 106]

 

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Plate 1. Mariannaea elegans, Scanning electron micrographs. A. conidiophores (c. 1600 x): B. imbricate chains of conidia (c. 1300 x) C. phialides (c. 4100 x).

 

 

 

Plate 2.  A. Gabarnaudia humicola, conidiophores (480 x); B-D. Sphaeronaemella humicola, B. tuft of hairs at the top of the ascoma (920 x) C. part of ascoma, ascospores stained with aniline blue (940 x), D. cirrus of ascospores (530 x)