Studies in Mycology, No. 2

1 November 1972

 

 

The Genus Talaromyces

Studies on Talaromyces and related genera II.

 

 

AMELIA C. STOLK and R. A. SAMSON

 

Centraalbureau voor Schimmelcultures, Baarn

 

Summary

The genus Talaromyces is re-defined. It is restricted to species producing asci in chains. The genus is divided into 4 sections, Talaromyces, Emersonii, Thermophila and Purpurea, wich are primarily based on their conidial states. Descriptions, figures and scanning electron micrographs are given of the species belonging to the genus. The species, transferred by Benjamin and Udagawa from the imperfect genus Penicillium to the perfect genus Talaromyces are validated by providing latin diagnoses. Two species of Gymnoascus, G. flavus Klcker and G. luteus Sacc. and three species of Arachniotus, A. trachyspermus Shear, A. intermedius Apinis and A. purpureus Mller & Pacha-Aue are transferred to Talaromyces. Two new species, T. byssochlamydoides and T. udagawae, and two new varieties, T. flavus var. macrosporus and T. helicus var. major are described. A key to the genera with Paecilomyces or Penicillium conidial states and to the sections of Talaromyces is given. Species are keyed out in each section and in an additional key based on ascospore characters. A new series of Penicillium, the Penicillium cylindrosporum-series is proposed.

 

 

Introduction

 

When Benjamin (1955) introduced his genus Talaromyces with the type species, T. vermiculatus (= T. flavus (Klcker) Stolk & Samson, comb. nov.), he included in it 10 species with asci borne singly as well as species in which asci are produced in chains. In a first contribution Stolk and Samson (1971) transferred two species with single asci produced from croziers to a new genus Hamigera, thus leaving in Talaromyces only species with asci occurring in chains.

Benjamin (1955) and Udagawa (1966) transferred some of the perfect Penicillia belonging to Raper & Thom's P. luteum-series (1949) to the ascomycetous genus Talaromyces and published these perfect names as new combinations. However, according to Art. 59 of the International Code of Botanical Nomenclature (Stafleu, 1972), the combination of the specific epithet of a name typified by an imperfect state (in this case Penicillium) with a name of a genus characterized by a perfect state (in this case Talaromyces) shall be considered not validly published as a new combination but [p. 2] shall be considered the name of a new taxon and it shall be attributed to the author of that name and to him alone. This means that the names of the perfect states of the Penicillia mentioned above should be ascribed only to Benjamin or Udagawa. Unfortunately Benjamin and Udagawa did not provide the necessary latin diagnoses. To validate these names, the latin descriptions are supplied in this article and types are indicated.

 

Generic description

 

Ascomata globose to subglobose, soft, superficial, discrete or confluent, of indeterminate growth. Ascomatal coverings varying from scanty to dense, consisting of a network of hyphae, which may range from very loose-textured to closely knit, usually surrounded by a weft of thin, usually encrusted radiating hyphae, straight or twisted depending on the species. Ascomatal initials of various shape. Asci evanescent, 4-, 6- or 8-spored, globose to subglobose or slightly ellipsoidal, borne in chains. Ascospores globose or ellipsoidal, smooth or showing various ornamentations, yellow, rarely becoming reddish.

 

Conidial state: Penicillium Link ex Fr., Paecilomyces Bain.

Type species: Talaromyces vermiculatus (Dangeard) C. R. Benjamin (= T. flavus (Klcker) Stolk & Samson, comb. nov.).

 

Morphology and delimitation of the genus Talaromyces

 

Though, according to this conception, Talaromyces is now restricted to species with asci borne in chains, it is in many ways still heterogeneous.

The ascomatal coverings of its species consist of a network of hyphae, differing markedly in their density. They may be scanty and inconspicuous (e.g. T. byssochlamydoides), distinct and loose-textured (e.g. T. flavus) or thick-walled and closely knit, becoming pseudoparenchymatous in age (e.g. T. thermophilus).

Ascomatal initials develop inside loose tufts of very straight or occasionally twisted hyphae.. Usually several initials develop within one hyphal tuft. In 3 species the initials consist of a pair of gametangia. In the other species antheridia are completely lacking and initials may consist of subglobose cells (e.g. T. emersonii), coiled hyphae (e.g. T. leycettanus), swollen irregular septate hyphae producing coiled branches (e.g. T. wortmannii) or wide branched, gnarled hyphae (e.g. T. trachyspermus). Only in a few species are initials one-celled and chlamydospore-like and produce ascogenous hyphae directly; otherwise the initials become septate and then begin to produce branches. By their further profuse branching a compact mass is formed consisting mainly of ascogenous hyphae. After having produced septa, these ascogenous hyphae develop directly and consecutively into asci, which are thus being formed in chains. Chains of asci may be branched because of the continued branching of the ascogenous hyphae. In [p. 3] but most species the fertile parts of the ascogenous hyphae and subsequently the ascal chains are curved or coiled. The terminal cell of a chain of young asci in these species may be turned back upon itself, thus giving superficially the appearance of a crozier. However, end to end arrangement of the asci excludes the intervention of a crozier of the convential type as occurring in Byssochlamys and Hamigera, in which genera typically only the penultimate cell develops into an ascus (Fig. 1, p-q). Only in T. thermophilus chains of asci are straight or slightly curved and the terminal cell of a chain of asci is not turned back upon itself. Because of the continued branching of the ascogenous and covering hyphae, ascomata of indeterminate growth develop.

Simultaneously with the development of the ascogenous hyphae a weft of surrounding hyphae, for the greater part supplied by neighbouring hyphae, is produced, developing into the ascomatal covering. This is surrounded by a loose weft of mainly radiating, in most species heavily encrusted hyphae, arising from the original hyphal tuft in which the initials developed.

All species described in this paper are homothallic.

Ascospores show various patterns. They may be globose, subglobose or ellipsoidal, their walls being smooth, spinulose or provided with ridges. With the exception of a few species, there seems to be very little variation in the sizes of the ascospores produced in different strains of one Talaromyces species. In T. flavus a considerable variation in the sizes of the ascospores occurs.

The conidial states of the species of Talaromyces show much variation. In T. leycettanus and T. byssochlamydoides the imperfect states belong in Paecilomyces, those of the other species are to be classified in Penicillium, though they belong to quite different series of this genus.

The genus Paecilomyces is separated from Penicillium by the shape of the phialides, consisting of a cylindrical or swollen basal portion, tapering into a long distinct neck. The conidial structures are usually divergent, verticillate or irregularly arranged. Moreover the cultures do not show the true green colours like in most species of Penicillium.

For the description of the penicillus the terminology of Raper and Thom (1949) is used. Only the term `branch' is replaced by 'ramus' and instead of the incorrect term `sterigma', the term 'phialide' is used for the conidiogenous cell of Penicillium. The term `fractional' is used for atypical, reduced penicilli with a diminished number of metulae, sometimes occurring with a few phialides in one verticil or consisting only of very small verticils of phialides (occasionally only two).

Temperature relationships in Talaromyces range from mesophilic to strongly thermophilic. The terms thermotolerant and thermophilic are used in the sense of Cooney and Emerson (1964).

 

In view of its heterogeneous character, it might seem justified to split Talaromyces into a number of genera, which may be based on the pattern of the ascospores, on the structure of the ascomatal initials, on the texture [p. 4] of the ascomatal coverings, on the nature of the conidial state or on the temperature- relationships. However, since no consistent correlation between these different characteristics could be demonstrated, we prefer to keep all species in one genus: Talaromyces. Nevertheless, a division of the genus into 4 sections, primarily based on the structure of the conidial state seems desirable:

 

section Talaromyces

section Emersonii

section Thermophila

section Purpurea

 

Talaromyces differs from the other perfect penicillate genus Eupenicillium by its soft ascomata. The latter genus is characterized by sclerotioid ascomata, that are very hard to gritty when young. Talaromyces can easily be distinguished from the related genera in the Eurotiaceae: Byssochlamys Westling, Hamigera Stolk & Samson and Thermoascus Miehe by its catenate asci.

In the Gymnoascaceae four genera produce ascomata approximating those of Talaromyces, since their coverings consist, like in Talaromyces, of hyphal networks, which may be inconspicuous (occasionally even absent) or well-developed, viz. Arachniotus Schroeter, Narasimhella Thirum. & Mathur, Amauroascus Schroeter and Arachnotheca von Arx. Talaromyces can easily be separated from these genera by its conidial state. In the 4 gymnoascaceous genera mentioned above, the imperfect state is lacking or represented by arthro- or aleurioconidia (Kuehn, 1958; Apinis, 1964; von Arx, 1971). In Talaromyces, on the other hand, phialoconidia-bearing penicilli occur. Moreover Talaromyces differs from the latter 3 genera in developing asci in chains, while in Narasimhella, Amauroascus and Arachnotheca asci are produced singly from croziers (Fig. 1, i-k). As to Arachniotus, examination of species belonging to this genus revealed that asci often are borne in very short chains. Because of the compact structure of the elements comprising the primordia of Arachniotus, the exact development of these chains could not be demonstrated with certainty. It appears, that they are produced through crozier-like structures, which become septate in the usual way. It might be possible that the asci develop from all cells of these croziers (Fig. 1g), thus resembling the ascus development in Talaromyces. In the latter genus the ascogenous hyphae may also be crozierlike when young (Fig. 1a). The main difference between the development of the ascomatal fertile parts of Arachniotus and Talaromyces seems to be that the crozier-like ascogenous hyphae of Arachniotus do not continue to grow and branch profusely as they do in Talaromyces. [p. 5]

 

 

Fig. 1. Development of asci in Talaromyces and some related genera in the Eurotiaceae and Gymnoascaceae. a-b. Talaromyces flavus; c. Talaromyces leycettanus; d. Talaromyces thermophilus; e. Talaromyces purpureus; f-h. Arachniotus ruber, f. gametangia, g-h. development of asci; i. Narasimhella poonensis; j. Amauroascus aureus; k. Arachnotheca glomerata; l. Pseudogymnoascus bhattii; m. Gymnoascus reesii; o. CBS 114.72 = Orr o-3156; p. Byssochlamys fulva; q. Hamigera avellanea. [p. 4]

 

In general, one of the main characteristics for separating the family of the Eurotiaceae from the Gymnoascaceae is considered to be the imperfect state (Kuehn, 1958; von Arx, 1970). In the Eurotiaceae phialoconidia occur, whereas these should be lacking in the Gymnoascaceae. Recently we received an interesting culture from Dr G. F. Orr (Dugway, USA), labeled [p. 6] 0-3156 (= CBS 114.72), representing a new genus and linking in this respect the Gymnoascaceae with the Eurotiaceae. The structure of the ascomata, surrounded by loose dark networks of hyphae with spine-like appendages, would classify it in the Gymnoascaceae near Gymnoascus, whereas the conidial state, composed of Acremonium-like phialides, shows a close relationship with the Eurotiaceae. Moreover the initials which consist of simple, somewhat curved hyphae, producing coiling branches are different from the usual initials of the Gymnoascaceae (Fig. 1f). They agree much better with those of Talaromyces. In addition the development of the asci, which are borne in chains, developing directly from the ascogenous hyphae, also relates it with Talaromyces (Fig. 1o). For these reasons, Orr's species should in spite of its covering, be classified as a somewhat deviating species in the Eurotiaceae and the most reliable character for distinguishing the Gymnoascaceae from the Eurotiaceae still seems to be the conidial state.

Because of the similarity of their ascomata and the fact that their authors overlooked the penicilli, which in some Talaromyces strains occur only scantily, species of Talaromyces have often been confused with Arachniotus (Ghosh et al., 1961) or even with Gymnoascus (Orr et al., 1963).

Many species tend to produce mycelial sectors, usually consisting of a floccose to funiculose mycelium, occasionally bearing conidial structures. This is most pronounced in T. flavus and T. helicus, which species may become quite mycelial after a few transfers. A few species, when degenerating, produce only a submerged mycelium, developing occasionally a few ascomata and penicilli (e.g. T. purpureus). Other species may keep their capacity to produce ascomata for many years (e.g. T. wortmannii; its type strain still produces ascomata after having been maintained on artificial media for almost 70 years).

 

 

Methods

 

For the description of the microscopic features only cultures growing on 2% malt agar were used. In a few species, more numerous and more charateristic penicilli are produced on hay-infusion agar and on Czapek agar with 20% sucrose. In these cases penicilli occurring on these media are described, as mentioned in the text.

The influence of temperature was tested by cultivating the strains at temperatures between 10 and 55C with intervals of 5C. For the identification of the species belonging to the sect. Talaromyces and Purpurea usually a temperature between 25 and 30C is desirable. Higher temperatures (30-45C) for the species of the other sections are recommended.

Sizes of ascospores provided with longitudinal ridges are given inclusive of these ridges, whereas measurements of the other ascospores refer only to the ascospore body.

The scanning micrographs were made with a JEOL JSM-U 3 scanning [p. 7] electron microscope at an accelerating voltage of 5 - 15 kV. For examination with the electron microscope, ripe ascomata were crushed between two coverslips. The material, mainly containing ascospores, was attached to the standard specimen stubs using squares of double-side adhesive tape. The specimens were coated with carbon and gold in a vacuum evaporator. A lyophilization of the fungus material, as described by Clarke and Griffiths (1970) appeared to be unnecessary. Sufficient dryness of the material could be obtained by maintaining the specimens in the coating unit for 45 minutes before coating.

 

 

Key to the perfect genera with a Paecilomyces or Penicillium conidial state and to the sections of Talaromyces

 

1a.

Ascomata soon developing a thick ascomatal wall

2

1b.

Ascomata without distinct walls or covered with hyphal networks, in a few species more dense, becoming pseudoparenchymata-like in old colonies

3

 

 

 

2a.

Ascomata sclerotioid, usually hard when young, ripening from the centre outwards; asci not produced from croziers; conidial state Penicillium

Eupenicillium (Scott, 1968)

2b.

Ascomata not sclerotioid, orange brown, walls pseudoparenchymatous; asci produced from croziers; conidial state Paecilomyces; thermophilic

Thermoascus crustaceus (Apinis & Chesters) Stolk (Stolk, 1965, Apinis, 1967)

 

 

 

3a.

Asci borne from croziers

4

3b.

Asci borne in chains

Talaromyces 5

 

 

 

4a.

Ascomatal coverings scarce or lacking; conidial state Paecilomyces, Byssochlamys (Stolk & Samson, 1971)

 

4b.

Ascomatal coverings composed of well-developed hyphal networks; conidial state Penicillium

Hamigera (Stolk & Samson, 1971)

 

 

 

5a.

Conidial state Paecilomyces or belonging to the Penicillium cylindrosporum-series, brown to creamish; weakly thermotolerant to strongly thermophilic

section Emersonii

5b.

Conidial state Penicillium, but belonging to other series, green, greyish or olive-brown

6

 

 

 

6a.

Conidial state predominantly monoverticillate, belonging to the Penicillium restrictum-series, greyish to olive-brown; ascomata yellow

section Purpurea

6b.

Conidial state different

7

 

 

 

7a.

Penicilli typically biverticillate-symmetrical, often fractional, or monoverticillate, occasionally reduced to solitary phialides, green; mesophilic

section Talaromyces

7b.

Penicilli divaricate, grey-green to brown; ascomata not produced on agar media; strongly thermophilic

section Thermophilia

 

[p. 8]

 

 

 

Talaromyces section Talaromyces

 

Ascomatal coverings usually distinct, often yellow, occasionally white, creamish, pinkish or reddish. Ascospores yellow, in strains producing abundant red pigment occasionally red. Conidial state: Penicillium of the biverticillate-symmetrical type, often fractional, in some strains reduced to atypical monophialides. Phialides usually lanceolate, in a few species showing a wider base.

 

Species belonging to this section show their best development on malt and oatmeal agar. On Czapek agar they develop better than the species belonging to the other sections. In many strains the conidial state is lacking or occurs scantily. Cultivation of these strains on hay-infusion agar and Czapek agar + 20% sucrose may stimulate the production of penicillate structures.

The species belonging to this section are mesophilic, with optimum temperatures about 25C and maximum temperatures not exceeding 40C, with the exception of T. trachyspermus, which still grows slowly at 40C, attaining a diameter of about 4 cm within 3 weeks. Some strains of T. flavus show also a slight development at 40C. None of them grows at 45C.

The ascomatal coverings range from somewhat scanty and thin (T. wortmannii) to more closely knit, simulating slightly a pseudoparenchymatous wall (T. trachyspermus and T. intermedius).

Ascomatal initials are of various types. Paired gametangia occur in 3 species of this section: T. flaves, T. helicus and T. stipitatus. In the other species initials develop as branches or as intercalary portions of hyphae. Asci are produced in short coiled chains, they are usually 8-spored, occasionally 6-spored (T. rotundus). Ascospores are generally ellipsoidal (only in T. rotundus they are globose), their walls showing various ornamentations.

Typically the conidial state produced in the species of this section consists of penicilli of the biverticillate-symmetrical type, consisting of verticillate metulae, bearing whorls of usually lanceolate phialides. Besides these typical penicilli, most species produce also fractional structures with metulae and phialides occurring in one verticil or consisting only of small verticils of phialides. In addition also solitary, sometimes atypical phialides occur, composed of a wider basal portion tapering more or less abruptly to a long conidium-bearing tube, e.g. T. helicus. In some species, e.g. T. helicus, strains occur which produce well-developed, predominantly symmetrical penicilli, whereas in other strains of the same species only fractional, sometimes very much reduced penicilli and solitary phialides were observed, therefore the production of fractional penicilli and solitary phialides is apparently only due to variation or degeneration. The most reduced conidial state observed in this section is produced by the type strain of T. intermedius. In this strain mainly solitary phialides are present, resembling those occurring in the genus Acremonium (Gams, 1971). They are usually tubular, tapering [p. 9] only occasionally to long conidium-bearing necks. On hay-infusion agar penicillate structures were observed consisting of 2 to 3 phialides. Since these phialides agree with the solitary and sometimes atypical phialides present in most species of Talaromyces and since the species is closely related to T. trachyspermus, it is placed in the section Talaromyces.

In most species of this section all elements of the penicilli are smoothwalled, hyaline to slightly yellowish. Only in T. helicus conidiophores and penicilli show a conspicuous green colour. The conidia are produced in divergent chains.

 

 

Key to the species

 

1a.

Ascospores globose, spinulose, 4.5-5.2 um

T. rotundus

1b.

Ascospores ellipsoidal

2

 

 

 

2a.

Ascospores provided with ridges

3

2b.

Ascospores without ridges

6

 

 

 

3a.

Ascospores with transverse ridges

4

3b.

Ridges of ascospores not transverse

5

 

 

 

4a.

Ascospores with 4-6 irregularly transverse ridges, 4.5-5.5 x 3-3.7 m

T. luteus

4b.

Ascospores with 3-5 nearly parallel transverse ridges, often spirally arranged, 3.5-4.5 x 2.2-3.0 m

T. udagawae

 

 

 

5a.

Ascospores with a single equatorial ridge, 3.2-4 (-4.5) x 11.7-2 (-2.5) m, ascomata yellow, occasionally salmon

T. stipitatus

5b.

Ascospores with 2 to 6 irregular, for the greater part longitudinal ridges, 2.5-3.5 x 2-2.4 m, ascomata yellow

T. ucrainicus

 

 

 

6a.

Initials consisting of wide ascogonia around which thin antheridia are tightly coiled; ascospores spinulose or smooth

7

6b.

Initials consisting of single hyphae; ascospores spinulose

10

 

 

 

7a.

Ascogonia clavate to vermiform, not coiled terminally; ascospores spinulose; ascomata usually yellow, occasionally pinkish to purple red

8

7b.

Ascogonia terminating in conspicuous coils; ascomata yellow, occasionally becoming pinkish on Czapek and oatmeal agar

9

 

 

 

8a.

Ascospores ranging within the sizes 3-5 x 2.2-3.5 m

T. flavus var. flavus

8b.

Ascospores ranging within the sizes 5-6.5 x 3.5-5.2 m

T. flavus var. macrosporus

 

 

 

9a.

Ascospores ornamented with a few delicate spines, most of them apical, sometimes appearing nearly smooth, 2.5-3 (-3.7) x 1.5-2.2 m

T. helicus var. helicus

9b.

Ascospores smooth, 3-4.5 x 2-2.7 m

T. helicus var. major

 

 

 

10a.

Ascomata yellow to orange; initials consisting of thick, irregularly septate hyphae, producing coiling branches; ascospores 3.7-4.7 (-5) x 2.5-3

T. wortmannii

10b.

Ascomata white, creamish or pinkish; initials consisting of swollen [p. 10] hyphae producing profusely branching gnarled hyphae

11

 

 

 

11a

Ascomata white to creamish; ascospores 3-3.5 x 2-2.5 m

T. trachyspermus

11b.

Ascomata creamish to pinkish; ascospores 5-6 x 4-4.5 m

T. intermedius

 

1. Talaromyces flavus (Klcker) Stolk & Samson, comb. nov., var. flavus - Fig. 2 a-l.

 

Gymnoascus flavus Klcker - Hedwigia 41: 80. 1902 (basionym).

Talaromyces vermiculatus C. R. Benjamin - Mycologia 47: 684. 1955 [Talaromyces vermiculatus (Dangeard) C. R. Benjamin].

Arachniotus indicus Chattopadhyay & Das Gupta - Trans. Br. mycol. Soc. 42: 72. 1959.

Arachniotus indicus Chattopadhyay & Das Gupta var. major Chattopadhyay & Das Gupta - Trans. Br. mycol. Soc. 42: 73. 1959.

 

Status conidialis: Penicillium vermiculatum Dangeard - Botaniste 20: 123. 1907 (described inclusive of the perfect state).

 

Penicillium liani Kamyschko - Notul. syst. Inst. Cryptog. Horti bot. petropol. 15: 86. 1962 (described inclusive of the perfect state).

 

Colonies on malt agar spreading broadly, attaining a diameter of 7 to 8 cm within 2 weeks at 25C, consisting of a basal felt in which numerous ascomata are embedded near the agar surface, usually forming a continuous, thick layer, conspicuous yellow, ranging from Lemon Yellow to Empire Yellow (Ridgway, 1912, Pl. 4; Rayner, 1970, 23, 2b) towards the centre often overgrown by a loose aerial network, giving the central areas of some strains, e.g. CBS 261-55, a conspicuous pinkish shade approximating Orange-Vinaceous (Ridgway, Pl. 27; Rayner, 5"b). Very few strains, e.g. CBS 194.72 and CBS 352.72, grow less rapidly, about 4.5-5.5 cm in diameter within two weeks and produce paler yellow shades, becoming Cream-Buff in age (Ridgway, Pl. 30; Rayner, 19"d). Conidial state scanty, usually not affecting the colony appearance, but in some strains, e.g. CBS 195.72, more pronounced, giving the conidial areas a greyish green shade near Gnaphalium Green (Ridgway, Pl. 47; Rayner, 29 ""d). After having been maintained for some time on artificial media the species has a strong tendency to produce white to greyish green sectors consisting of a floccose to funiculose mycelium bearing penicilli more or less abundantly. Odour present, but unpronounced. Exudate lacking, in occasional strains present as small drops. Reverse ranging from orange red when young to orange brown in age, with the marginal areas pale ochraceous brown. In a few strains, e.g. CBS 261.55, the reverse shows conspicuous purple red shades with the pigment diffusing in the agar. In other strains the red pigment may be lacking completely. [p. 11]

Vegetative hyphae hyaline to yellow, often encrusted, 1-4 m in diameter.

Ascomata usually yellow, in some strains buff or pinkish to purple-red, globose, 200-700 m in diameter, commonly confluent but at the margin occasionally discrete, ripening within 2 weeks. In a few strains, e.g. CBS 261.55 and CBS 284.58, ascomata are predominantly discrete, forming a thinner layer than produced by the neotype culture. Covering consisting of a few layers of well developed networks of yellow pigmented hyphae, not becoming pseudoparenchymatous; surrounded by loose wefts of yellow, heavily encrusted, somewhat twisted, predominantly radiating hyphae, about 1 to 2 m in diameter. Initials consisting of club-shaped ascogonia, up to 250 m in length and 3.5-4.5 m in diameter, around which thin antheridia coil tightly several times. The antheridial apical cell is usually closely appressed against the ascogonium. It fuses with the ascogonium at about the central or somewhat higher portion of the latter cell. After fertilization, the ascogonia become septate. At first horizontal, later irregularly transverse and oblique septa develop. At this stage the primordium becomes enveloped in closely wound delicate hyphae, which develop into the covering. Most of the ascogonial cells produce branches, which continue to branch profusely. Asci usually 8-spored, broadly ellipsoidal to subglobose, 8-11 x 7.5-9 m, in some strains slightly smaller. Ascospores yellow, in strains producing abundant red pigment, e.g. CBS 261.55, becoming reddish in age, more or less broadly ellipsoidal, 3.5-5 x 2.5-3.2 m, in some strains somewhat smaller (table 1), thick-walled, spinulose, with spines ranging from very small to 0.5 m, rarely 1 m in length (Pl. 1. A).

Conidiophores arising primarily from the substratum, especially in marginal areas, occasionally borne also as short branches from aerial hyphae overgrowing the ascomata, usually erect, 24-250 x 1.5-2.5 m. Metulae 2 to 3 (-4) in the verticil, 10-15 x 1.7-2 m, occasionally lacking. In strains producing better developed penicilli, up to 6 metulae occur in the verticil, measuring 8-12 x 2-2.5 m. Phialides in whorls of 2 to 7, lanceolate, 8-12 x 1.7-2.5 m, in fractional penicilli often atypical, solitary phialides may be present. Conidia brownish green, ranging from subglobose to ellipsoidal, 2.2-3.5 x 2.0-2.5 m, in other strains slightly smaller or larger, with walls usually thick, smooth or nearly so.

CBS 310.38 is designated as neotype.

 

Material examined

 

Herbarium specimens

Arachniotus indicus in herb. CMI (type), IMI S7850, isolated from soil of paddy fields, W. Bengal, India.

Arachniotus indicus var. major in herb. CMI (type), IMI 57851, isolated from soil of paddy fields, W. Bengal, India.

 

Living cultures

CBS 310.38 = NRRL 2098, neotype culture, isolated by J. C. Neill in New Zealand, sent to the CBS as Penicillium luteum in 1938. [p. 12]

 

 

Fig. 2. Talaromyces flavus var. flavus, CBS 310.38. a. different types of conidiogenous structures; b. conidia; c-d. development of ascogonia and antheridia; e. fusion between the 2 gametangia and subsequent septation of the ascogonium; f. development of horizontal, transverse and oblique septa; g-h. production of branches of the ascogonial cells, which will develop into the ascogenous hyphae: i. chains of asci; j-1. ascospores, j. CBS 352.72; k. CBS 387.48; l. CBS 310.38. Talaromyces flavus var. macrosporus, CBS 317.63m. chain of asci; n. ascospores. [p. 13]

 

CBS 387.48 = NRRL 1019, isolated by K. D. Butler in Tucson (Arizona, USA), received from NRRL.

CBS 261.55, isolated from Clematis spec., Boskoop. CBS 283.58, isolated from jute, Delft.

CBS 284.58, isolated at the Plantenziektenkundige Dienst, Wageningen.

CBS 351.61, isolated from chicken meat, Amersfoort.

CBS 437.62, isolated from compost, Bonn.

CBS 225.66, type culture of Penicillium liani Kamyschko, isolated from soil in China.

CBS 194.72 and CBS 195.72, isolated by J. H. van Emden from soil, Wageningen.

CBS 351.72 = NRRL A 2566; CBS 352.72 = NRRL A 3339; CBS 354.72 A = NRRL A 3538; CBS 354.72 B = NRRL A 3949; CBS 354.72 C = NRRL A 2174; CBS 354.72 D = NRRL A 2361, received from D. I. Fennell, Peoria.

CBS 552.72, isolated from soil in France, 1971, received from J. Nicot as number 2002.

CBS 582.72 A = IFO 7232 - NHL 6063; CBS 582.72 B = IFO 7233 = NHL 6064; CBS 582.72 C = IFO 8896; CBS 582.72 D = IFO 8897; CBS 582.72 E = IFO 8898; CBS 582.72 F = IFO 8959; CBS 582.72 G = IFO 8960; CBS 582.72 H = IFO 9044, received from K. Tubaki, Tokyo.

 

Klcker (1902) presented only a brief morphological description of this species. Ascomata were described as yellow surrounded by a loose aerial network, and the ascospores as pale yellow, spinulose, oval to ellipsoidal, 5 to 6 m in length. He placed this species in the genus Gymnoascus since he did not recognize the observed conidial structures as phialides bearing chains of conidia, and thus representing extremely reduced penicilli. However, in 1903, when describing his P. wortmannii, he discussed in a footnote on p. 99 the possibility that the conidial state of G. flavus might represent a reduced Penicillium, but he persisted in classifying it in Gymnoascus. Though he illustrated besides solitary phialides only one verticil of two phialides, his figures 1-5 (Tafel II) of G. flavus do not leave any doubt that his species represents a Talaromyces. It is therefore transferred to that genus. Unfortunately neither a type culture nor herbarium material of G. flavus is preserved.

Fragmentary penicilli and solitary phialides, as reported by Klcker (1902) are occasionally observed in the strains of the fungus described above as T. flavus, especially in fresh isolates consisting mainly of ascomata. Usually after a few transfers these strains produce also more characteristic penicilli. The sizes of ascospores and conidia given in our description are slightly smaller than those provided by Klcker (1902) especially those of the conidia. This may be due to inaccuracy of measuring. As to the conidia, those described by Klcker may also have been larger because he described them only from liquid media. According to Klcker's description of P. wortmannii the ascospores of this species are nearly identical with those of T. flavus.

When describing his new species P. vermiculatum, Dangeard (1907) gave an accurate report of the penicilli of this species, including the size of the conidia, while he described in great detail the development of the ascomatal initials. The ascospores of this species are described as ellipsoidal and slightly spinulose, but no sizes are given. Benjamin (1955) transferred Dangeard's species to Talaromyces. Orr et al. (1963) considered G. flavus [p. 14] and T. vermiculatus as synonyms, a view with which we concur. The ascospores as described by Dangeard (1907) agree with those of Klcker's species, while some of the conidial structures figured by Dangeard (1907) (Pl. 18, fig. I) approximate to those illustrated in Klcker's fig. 4. The name P. vermiculatum remains valid for the conidial state.

Ghosh et al. (1961) proved that Arachniotus indicus Chattopadhyay & Das Gupta and its variety major Chattopadhyay & Das Gupta represented small-spored strains of T. vermiculatus.

Examination of the type culture of P. liani Kamyschko showed also this species to be synonymous with T. flavus.

T. flavus is the most common species of Talaromyces. It is frequently isolated from soil, but it may also occur upon organic materials undergoing slow decomposition. The species is world-wide in its distribution.

 

Talaromyces flavus var. flavus

 

 

strains

ascospore sizes in m

 

 

IMI 57850 (type) Arachniotus indicus

3.0-3.7 x 2.2-2.7

IMI S7851 (type) Arachniotus indicus var. major

3.0-4.0 x 2.2-2.7

CBS 352.72

3.0-3.7 x 2.2-2.7

CBS 351.72

3.2-4.2 x 2.2-3.0

CBS 387.48

3.5-4.2 x 2.5-3.0

CBS 225.66 (type) Penicillium liani

3.5-4.2 x 2.5-.0

CBS 354.72A

3.5-4.2 x 2.5-3.0

CBS 283.58

3.5-4.5 x 2.5-3.0

CBS 284-58

3.5-4.5 x 2.5-3.2

CBS 261.55

3.5-4.7 x 2.5-3.0

CBS 354.72B

3.7-4.7 x 2.5-3.0

CBS 354.72D

3.5-5.0 x 2.5-3.0

CBS 310.38 (neotype)

3.5-5.0 x 2.5-3.2

CBS 351.61

3.7-5.0 x 2.5-3.2

CBS 437.62

3.7-5.0 x 2.5-3.2

CBS 194.72

3.7-5.0 x 2.7-3.2(3.5)

CBS 354.72 C

3.7-5.0 x 3.0-3.5

Talaromyces flavus var. macrosporus

CBS 350.72

4.7-6.2 x 3.2-4.0

CBS 226.72

5.0-6.0 x 3.5-4.2

CBS 317.63 (type)

5.0-6.2 x 4.2-5.0

CBS 353.72

5.2-6.2 x 4.2-5.0

CBS 580.72

5.0-6.5 x 4.2-5.0

CBS 117.72

5.2-6.5 x 4.2-5.2

 

Table 1. Ascospore sizes of strains examined of T. flavus var. flavus and T. flavus var. macrosporus.

[p. 15]

 

T. flavus is an extremely variable species. Different strains may vary in colour, in the amount of red pigment produced, in the number of penicilli [p.15] and in the size of ascospores, which are ranging in the strains examined between 3 and 5 m in length and 2.2-3.5 m in diameter (Table 1). Slight differences in the shape of the ascospores were also noted.

As stressed by Klcker (1903), Gymnoascus flavus and P. wortmannii resemble one another strongly. Klcker separated them mainly on the basis of their conidial states which were quite different in the strains he studied; however, they may be more similar in other strains. He mentioned also a slight difference in the colour of the cultures. G. flavus is merely reported as yellow, whereas in P. wortmannii the shades were said to merge from yellow into orange. Thom (1930) considered P. wortmannii as a synonym of P. vermiculatum. It was Emmons (1935) who distinguished the two species on the basis of the quite different structure of their ascomatal initials and this is still, besides some slight differences in cultural appearance and rate of growth, considered the main character on which the two species can be distinguished.

Table 1, giving the measurements of the ascospores of the cultures examined, shows a distinct line of separation between strains with ascospores smaller than 5 m and strains with ascospores larger than 5 m in length. No real overlapping in sizes between the examined ascospores of these two groups has been observed. Within both groups a marked variation in spore measurements occurs, these size ranges overlap within the groups. All strains with ascospores ranging between 3 and 5 m in length and 2.2 to 3.5 m in diameter are considered to represent Klcker's G. flavus (= T. flavus), while the strains with ascospores ranging between 5 to 6.5 (7) m in length and 3.5 to 5.2 m in diameter are considered as a new variety of T. flavus.

 

 

2. Talaromyces flavus (Klcker) comb. nov., var. macrosporus var. nov. - Fig. 2, m-n.

 

Coloniae habitus, ascomatum initialia, ascomata et penicilli cum typo congruunt. Asci 13-18 x 11-14.5 m, ascosporae flavae, ovoideae vel late ellipsoideae, 5-6.2 x 4.2-5 m, pariete ad 0.5 m crasso, spinis ad 1 m longis obtectae.

Typus: CBS 317.63, isolatus e fructibus, Stellenbosch.

 

Status conidialis: Penicillium vermiculatum Dangeard pro parte

 

Colony appearance, rate of growth, structure of the initials, ascomata and penicilli agree completely with those of the species. Asci 13-18 x 1114.5 m. Ascospores yellow, ovoidal to broadly ellipsoidal 5-6.2 x 4.2-5 m, with walls up to 0.5 m thick and spines up to 1 m in length.

 

Material examined

 

CBS 317.63, type culture, isolated from fruit, Stellenbosch, S-Africa. [p. 16]

CBS 117.72 = NRRL 2101, isolated from cotton fabric, Panama, received from D. I. Fennell, Peoria.

CBS 226.72, isolated by M. C. Papendorf from soil, Potchefstroom, S-Africa. This strain has a tendency to degenerate, producing colonies consisting mainly of submerged mycelium bearing a thin layer of penicilli and occasionally a few scattered ascomata, sometimes in sectors. This culture produces a conspicuous red pigment.

CBS 350.72 = NRRL A 3232, isolated from top soil near Lanre, Gold Coast, and CBS 353.72 = NRRL A 466, isolated from tentage, New Guinea, both received from D. I. Fennell, Peoria. CBS 350.72 is characterized by more ellipsoidal ascospores showing more variation in size than those in the other strains.

CBS 580.72 = IFO 7132, received from K. Tubaki, Tokyo, as T. rotundus.

 

Like the var. flavus, the var. macrosporus shows a marked variation in colony appearance and in the size of the ascospores, which in the strains examined range between 5 to 6.5 (-7) m in length and 3.5 to 5.2 m in diameter. Moreover the length of the spines of the ascospores varies in different strains. Up till now the strains belonging to this variety come from warmer areas.

 

 

3. Talaromyces helicus C. R. Benjamin, spec. nov., var. helicus - Fig. 3

 

Talaromyces helicus (Raper & Fennell) C. R. Benjamin - Mycologia 47: 684. 1955 (as comb. nov.).

 

Diagnosis latina in Mycologia 40: 515. 1948, continetur.

Typus novae speciei status perfectus culturae CBS 335.48.

 

Status conidialis: Penicillium helicum Raper & Fennell - Mycologia 40: 515. 1948 (described inclusive of the perfect state).

 

Colonies on malt agar spreading fairly broadly, attaining a diameter of about 6 cm within 2 weeks at 25 C, consisting mainly of a comparatively thin layer of ascomata, produced near the agar surface, showing conspicuous yellow shades near Lemon Chrome, occasionally becoming Light Cadmium in age (Ridgway, Pl. 4; Rayner, 2b, 19b), embedded in and overgrown by a loose aerial network. Conidial state scanty, usually not affecting the colony appearance, but occasionally colouring central areas slightly greenish, especially in old mycelial sectors. Only in one of the examined strains penicilli occur abundantly (CBS 455.72). Reverse ranging from conspicuous yellow to pinkish buff, in predominantly conidial cultures green.

Colonies on oatmeal agar resemble those on malt agar, but the reverse occasionally shows dark greenish shades.

Vegetative hyphae hyaline, occasionally encrusted, 1-4 m in diameter. [p. 17]

Ascomata yellow, usually globose, 100-300 m in diameter, discrete to confluent, ripening within 1 to 2 weeks. Coverings consisting of thin interwoven hyphal networks, surrounded by loose wefts of yellow, heavily encrusted, slightly twisted, predominantly radiating hyphae, about 1-1.5 m in diameter. Initials resemble at first those of T. flavus, consisting of thick, club-shaped ascogonia, around which thin antheridia tightly coil. The antheridia remain restricted to the basal parts of the ascogonia, they terminate as slight enlargements closely appressed against the ascogonia. Soon the ascogonia grow out to coil terminally in a conspicuous helical pattern forming up to 8 coils. Usually the antheridium fuses with the ascogonium at the basal straight part of the latter structure. After fertilization, the ascogonium becomes septate and enveloped in closely wound delicate hyphae. Most of the cells of the coiled parts produce branches which continue to branch profusely, developing into the ascogenous hyphae. Asci broadly ellipsoidal to subglobose, 6-7 x 4.5-5.5 m. Ascospores ellipsoidal, 2.5-3 x 1.5-2 m, in some strains slightly larger, up to 3.7 x 2.2 m, very delicately spinulose, with the spines usually irregularly disposed on the surface, most of them occurring apically; in some strains spines very small, almost inconspicuous, in others distinct (Pl. 2. A-B).

Conidiophores arising from the aerial mycelium, 20-70 x 2-2.5 m, yellow-green, smooth or slightly roughened from dark encrustation, especially at the basal parts. Metulae in small verticils of 2 to 4, yellow-green, 8.5-18 x 2-2.5 m, often lacking. Phialides 2 to 6 in the verticil, occasionally solitary, usually yellow-green, 8-15 x 2-3 m, typically lanceolate, but when occurring in very small verticils or solitary, often consisting of a wider basal part with a conspicuous, sharply tapered, conidium-bearing tube. Conidia pale greenish, usually ellipsoidal to pear-shaped when young, becoming subglobose to globose at maturity, 3-4.5 x 2.5-4 m, smoothwalled.

 

Material examined

 

CBS 335.48 = NRRL 2106, type culture, isolated from soil from Sweden.

CBS 137.65, isolated by E. Mller from compost in Zrich.

CBS 134.67 and CBS 760.68, isolated by G. J. Bollen from greenhouse soil, Wageningen.

CBS 550.72 A, received from J. Nicot, Paris (strain 2032), isolated from salty soil, 1968.

The ascospores of the above-mentioned strains are nearly identical, those of the following ones are slightly larger, measuring 2.5-3.7 x 1.7-2.2 m, some of them are more distinctly spinulose.

CBS 455.72 = NRRL A 3059, received from D. I. Fennell, Peoria. This strain is predominantly conidial, producing well developed symmetrical penicilli and only a few ascomata. Ascospores distinctly spinulose. Reverse green.

CBS 550.72 B and CBS 550.72 C, received from J. Nicot, Paris (as strains 0 33 A and 2056), isolated from a wooden statue, 1969. Ascospores distinctly spinulose.

CBS 585.72 = IFO 7227; CBS 586.72 = IFO 7993, received from K. Tubaki, Tokyo. [p. 18]

 

 

Fig. 3. Talaromyces helicus var. helicus, CBS 335.48. a. different types of conidiogenous structures; b. conidia; c. young ascogonium with antheridium; d. ascogonium growing out, coiling terminally; e. septation of the ascogonium after fusion of the 2 gametangia; f. septation of the coiled part of the ascogonium, with a few cells growing out; g. optical section through an ascogonium producing branches, surrounded by thin hyphae; h. ascogonial cell producing branches; i. chains of asci; j. ascospores. CBS 455.72; k. ascospores. [p. 19]

 

T. helicus shows, because of the structure of its initials, a close relationship with T. flavus. It is distinguished from the latter species by its helicoidal ascogonia and the smaller sizes of its delicately irregularly spinulose ascospores. The 10 strains examined are similar in most characters. All of them produce fractional penicilli and occasionally solitary phialides. Only in CBS 335.48 and especially in CBS 455.72 many well developed symmetrical penicilli occur besides these reduced structures. In all of them conidiophores and penicilli are yellow-green. The ascospores show a slight variation in size.

P. sacchari Ray (1897) mentioned by Raper and Thom (1949) as a possible synonym of T. helicus is considered as a doubtful species (see p. 60).

 

 

4. Talaromyces helicus C. R. Benjamin, var. major Stolk & Samson, var. nov. - Fig. 4

 

Arachniotus aureus (Eidam) Schroeter sensu Apinis - Mycol. Pap. 96: 43, 1964 (misapplied).

 

Coloniae habitus, ascomatum initialia, ascomata et conidia cum typo congruunt. In coloniis vetustioribus ascomata nonnumquam rosea. Asci 6-11 x 5-7 m. Ascosporae ellipsoideae, demum fere irregulares, nonnumquam quasi reniformes, 3-4.5 x 2-2.7 m, leves. Penicilli conidiophoris brevibus portati, irregulares, summum e 3-4 phialidibus constant sicut in typo, saepe phialides solitariae irregulares adsunt. Conidiophora et penicilli hyalini, nonnumquam tarde flavo-virescentes.

Typus: CBS 652.66, isolatus ab A. E. Apinis e palude prope Attenborough in Anglia.

 

Status conidialis: Penicillium helicum Raper & Fennell pro parte

 

Colony appearance, rate of growth, ascomatal initials, ascomata and conidia agree completely with those of the var. helicus. In old cultures ascomata may occasionally become pinkish on Czapek and oatmeal agar. Asci 6-11 x 5-7 m. Ascospores ellipsoidal, in age occasionally somewhat more irregular in shape sometimes becoming somewhat reniform, 3-4.5 x 2-2.7 m, smooth-walled (Pl. 2.C). Penicilli, produced on short conidiophores, fractional, at the most monoverticillate, consisting of a verticil of about 3-4 phialides of the for T. helicus characteristic type, often reduced to solitary phialides which may be quite abnormal. Conidiophores and penicilli hyaline, occasionally becoming yellow-green in age. Conidia like the variety helicus.

 

Material examined

 

CBS 652.66 = BDUN 39, type culture, isolated from swamp near Attenborough and identified by A. E. Apinis (1964) as Arachniotus aureus (Eidam) Schroeter. In his figure 9i Apinis illustrated the first stage of an ascomatal initial. However, he overlooked the [p. 20] next phase, in which the ascogonia start to coil terminally. The conidia, described by Apinis (1964) as oidia, are phialoconidia and his fig. 9j represents an irregular penicillus. Penicilli occur only scantily in this strain, showing their best development on hay-infusion agar. They are often reduced to solitary phialides, which are sometimes tube-shaped, tapering only slightly at their apical parts, up to 21 m in length.

CBS 562.72 isolated from soil in Argentina by J. E. Wright. The ascospores of this strain agree exactly with those of CBS 652.66, but penicilli are produced in great numbers on all media tested. They are fractional in the same way, but consisting of more (up to 4) phialides in the verticil. They agree with the reduced penicilli occurring in some strains of the variety helicus. The conidia are slightly larger than those of CBS 652.66, measuring occasionally 6 x 5 m.

 

 

Fig. 4. Talaromyces helicus var. major, CBS 652.66. a. young and old conidiogenous structures; b. conidia; c. chain of asci; d. ascospores; CBS 562.72, e. young conidial structures; f. conidia; g. chain of asci; h. ascospores.

 

The variety major differs from the variety helicus in producing larger ascospores with smooth walls. Young conidiophores and penicilli do not show the characteristic yellow-green colour of the species, they are hyaline, but may become yellow-green in age. However, this difference may very well be due to the reduced state of the penicilli. In the strains examined of the variety helicus occasionally also solitary phialides occur which are hyaline when young.

 

 

5. Talaromyces intermedius (Apinis) Stolk & Samson, comb. nov. - Fig. 5

 

Arachniotus intermedius Apinis - Mycol. Pap. 96: 45. 1964 (basionym).

 

Status conidialis: Penicillium intermedium Stolk & Samson, stat. nov.

 

Penicilli pauci, optime in agaro extracto feni addito formati. Conidiophora 4-30 x 1.5-2 m, raro ramosa, levia. Penicilli summum e 2 - 3 phialidibus divergentibus constant. Phialides tubulares, 12-20 x 1.5-2 m, saepe solitariae, ad 30 Mm longae. Conidia hyalina vel fere hyalina, piriformia vel ellipsoidea, 2.7-4.5 x 2.2-3.5 m, levia.

Status ascigerus Talaromyces intermedius (Apinis) Stolk & Samson.

Typus: CBS 152.65, isolatus ab A. E. Apinis e terra, Attenborough, Nottinghamshire.

 

Colonies on malt agar spreading broadly, attaining a diameter of 8 cm within 2 weeks at 25C, consisting at maturity (3 to 4 weeks) of a dense layer of ascomata produced at or near the agar surface, showing conspicuous pink shades approximating Light Corinthian Red (Ridgway, Pl. 27; Rayner, 3"b) embedded in and covered by a floccose to funiculose, usually pinkish aerial mycelium. Conidial state lacking or very scanty, not affecting the colony appearance. Exudate in clear to pinkish drops. Odour spicy. The species has a tendency to produce creamish to yellowish sterile mycelial sectors. Reverse purple red, occasionally pinkish purple near Carmine and Dark Vinaceous (Ridgway, Pls. 1, 27; Rayner 1i, 1"b), yellow beneath the mycelial sectors.

Colonies on oatmeal agar attaining a diameter of 6 cm within 2 weeks, showing the same cultural characteristics and colours as on malt-agar.

Colonies on Czapek agar attaining a diameter of 4 cm within 2 weeks at 25C, consisting of a floccose to funiculose mycelium showing yellowish to pinkish shades, producing occasionally a few ascomata, slightly zonate. Reverse yellow to brown.

Vegetative hyphae hyaline to pinkish, occasionally encrusted, 1.5-4 (-7.5) m in diameter.

Ascomata at first white, then becoming pink, globose, variable in size, up to 1000 m in diameter, discrete or in clusters, ripening within 2 to 3 weeks. Covering distinct, consisting of a network of closely interwoven pinkish hyphae, simulating a thin wall, surrounded by a loose weft, mainly composed of radiating, pink, encrusted hyphae, about 1.5 m in diameter. Initials develop within 10 days as short branches or as intercalary portions of hyphae, which swell considerably, become strongly gnarled and branch [p. 22] profusely. Asci 8-spored, ovoidal to subglobose, 11-14.5 x 10-11 m. Ascospores thick-walled, conspicuously spinulose, broadly ellipsoidal, 5-6 x 4-4.5 m, spines about 0.2 m long.

Conidial state lacking or produced very scantily, best development on hay-infusion agar. Conidiophores, when present, arising as short branches from aerial mycelium, 4-30 x 1.5-2 m, rarely branched. Penicilli very irregular, atypical, at the most consisting of a verticil of 2 to 3 diverging phialides. Solitary phialides borne directly on aerial hyphae often present. Phialides tube-shaped, usually tapering only slightly at their apical parts, 12-20 x 1.5-2 m, especially the solitary phialides may be very long, measuring up to 30 m in length. Conidia hyaline or nearly so, pear-shaped to ellipsoidal, 2.7-4.5 x 2.2-3.5 m, smooth.

 

 

Fig. 5. Talaromyces intermedius, CBS 152..65. a. conidiogenous structures; b. conidia; c. initials; d. chains of asci; e. ascospores; f. ascomatal covering in surface view.

 

Material examined

 

CBS 152.65 = BDUN 267, type culture, isolated by A. E. Apinis from soil in Attenborough, Nottinghamshire, in 1949. [p. 23]

 

T. intermedius has the gnarled ascomatal initials in common with T. trachyspermus and T. ucrainicus. It is most closely related to T. trachyspermus because of its spinulose ascospores. It differs from the latter species in the sizes of the ascospores and in the colour of the ascomata. Because of these relationships and the presence of reduced penicillus-like structures the species is transferred to Talaromyces.

Apinis (1964) neither mentioned nor illustrated penicillate structures. However, he illustrated a hypha with a side branch (his fig. 11b) resembling a solitary phialide, producing a chain of conidia, which he interpreted as swollen cells (gemmae). The conidial state of T. intermedius is quite different from the biverticillate-symmetrical penicillus, occurring typically, though often only fractional, in the other species of the sect. Talaromyces. Its structure resembles more an Acremonium than a Penicillium. However, occasionally after having been maintained on agar media for a long time, Penicillia may degenerate and produce extremely reduced conidial structures with atypical phialides as occurring in T. intermedius. Moreover many species of the sect. Talaromyces (e.g. T. helicus) produce fractional penicilli, which occasionally are reduced to atypical solitary phialides, approximating those of T. intermedius. For these reasons the conidial state of T. intermedius is classified in Penicillium. However, only additional strains of this species can prove whether this placement is correct.

 

 

6. Talaromyces luteus (Sacc.) Stolk & Samson, comb. nov. - Fig. 6

 

Gymnoascus luteus Sacc. [Gymnoascus luteus (Zukal) Sacc.'] - Syll. Fung. 11: 437. 1895 (basionym) = Talaromyces luteus (Zukal) C. R. Benjamin - Mycologia 47: 681. 1955 (as comb. nov.).

 

Status conidialis: Penicillium luteum Zukal - Sber. Akad. Wiss. Wien 98: 42. 1889 (described inclusive of the perfect state).

 

Colonies on malt agar growing restrictedly, attaining a diameter of about 3 cm within 2 weeks at 25C, consisting of a tough, compact, for the greater part submerged mycelium, bearing numerous ascomata in a dense, continuous layer, bright yellow in colour, near Lemon Yellow (Ridgway, Pl. 4; Rayner, 23), commonly with only the central areas overgrown by a floccose to funiculose mycelium bearing conidial structures, giving these areas a green colour near Sage Green (Ridgway, Pl. 47; Rayner, 29"'b). Conidial state occasionally very scanty, but in young cultures growing at a slightly lower temperature than usual (about 22C) often produced abundantly, bluish green, near Celandine Green (Ridgway, Pl. 47; Rayner, 33"'b), in older cultures at 25C only present in central areas and in localized sections. Margin at first regular, later becoming lobed. Odour aromatic, sweetish. Reverse ranging from orange to red-brown [p. 24] in age, approximating Mars Yellow (Ridgway, Pl. 3; Rayner, 15i), with a pale orange to occasionally purple-red pigment, diffusing throughout the surrounding agar.

On Czapek agar the conidial state is more pronounced, the reverse is redbrown. On oatmeal agar colonies agree with those on malt, but they are occasionally zonate and the reverse is pinkish orange.

Cultures maintained for many years on agar media tend to loose the capacity to produce ascomata. These cultures often have a wet appearance, since they consist mainly of submerged hyphae producing only in central areas a floccose to strongly funiculose mycelium, occasionally bearing a few penicilli.

Vegetative hyphae 2-4.5 m in diameter, smooth or encrusted with yellow granules, submerged hyphae occasionally reddish, often showing inflations up to 7 m in diameter.

Ascomata yellow, globose, 100-400 m in diameter, confluent, occasionally discrete, ripening within 2 to 3 weeks. Coverings consisting of welldeveloped networks of interwoven hyphae surrounded by wefts of short, yellow-encrusted, twisted, branched hyphae, 1-1.5 m in diameter. Initials developing usually from atypical phialides, occasionally also directly from vegetative hyphae, consisting of relatively short swollen, compactly coiling hyphae, which immediately after their formation produce a side branch coiling around the first hypha. Later on, more branches develop, all of them coiling and thus producing a knot of fertile hyphae. No fusion between coiling hyphae has been observed. Asci globose to ellipsoidal, 12.5-15 x 8-10 m. Ascospores ellipsoidal, 4.5-5.5 x 3-3.7 m, thick-walled, with 4 to 6 conspicuous, irregularly transverse, locally interrupted, occasionally branched ridges, about 0.5 m wide (Pl. 1.H).

Conidial structures occur mainly in central areas, but occasionally a few may develop as tuft-like structures on an ascoma, thus resembling slightly the coremia described by Zukal (1889) and Wehmer (1893). No real coremia were observed. Conidiophores arising from aerial hyphae and ropes of hyphae, smooth, sometimes encrusted, usually 15-35 m, occasionally up to 100 m in length and 1.7-3 m in diameter. Metulae 2 to 3 in the verticil, 10-20 x 2-3 m, often lacking. Phialides in verticils of 2 to 4, usually lanceolate, but sometimes with a wider basal portion, 9-15 x 2-3 m, apices in old structures occasionally greenish; sometimes atypical with long conidia-bearing tubes or producing ascomatal initials instead of conidia. Solitary phialides, usually very long, are also present. Conidia ovoidal to ellipsoidal, 3-4 x 2.2-3.7 m, smooth, green in mass.

CBS 348.51 is designated as neotype.

 

Material examined

 

CBS 348.51 = IMI 89.305, neotype culture, isolated by J. H. Warcup from soil, Great Britain (culture no: BB228). [p. 25]

CBS 333.59, received from A. Dupuis, Laboratoire de Recherches Physiques, Charleroi, Belgium.

CBS 230.60 = ATCC 10466 = QM 7594 = NRRL 1010 = Thom 5357.208, sent by K. B. Raper; used for the description of P. luteum (Raper and Thom, 1949, p. 600-602). It shows a poor, mainly submerged growth, producing few aerial hyphae and ropes of hyphae, occasionally bearing penicilli; ascomata produced scantily, usually sterile. The initials and conidia are identical with those of the neotype.

CBS 865.71 = NRRL 2235, sent by D. I. Fennell, Peoria. CBS 588.72 = IFO 6896, received from K. Tubaki, Tokyo.

 

 

Fig. 6. Talaromyces luteus, CBS 348.51. a. conidiogenous structures; b. conidia; c. initials; d. chains of asci; e. ascospores.

 

Is has been generally assumed that only one species of Talaromyces produces ascospores with transverse to spiral ridges: T. luteus. However, recently a second species has been found characterized by ascospores with transverse (spiral) ridges which is described in this paper as T. udagawae. [p. 26]. The latter species has also characters in common with Zukal's description of P. luteus. Unfortunately no type material of P. luteus is known to be preserved. Nevertheless the species described above is still considered to represent Zukal's species. Its irregular, locally interrupted ridges suggest the ridges with wart-like projections as reported by Zukal (1889), while the sizes of the ascospores agree very well with those given by Zukal (4.8 x 3.3 m). Moreover the penicilli produced in tuft-like structures resemble the coremia described by Zukal and the spirally twisted hyphae covering the ascomata agree completely with those in Zukal's isolate. The red-coloured hyphae mentioned by Zukal as occurring in P. luteus have also been observed in some of the cultures examined, since most of them produce a red pigment. Only the observed initials do not agree with those described by Zukal. However, Zukal's description of the initials is rather confused and his drawings are very schematic.

Zukal (1889) classified T. luteus in the genus Penicillium, which he considered to belong to the Gymnoascaceae. Wehmer (1893) gave a few additional details especially of the penicilli. Besides the characteristic biverticillate - symmetrical penicilli, he illustrated fractional penicilli and even a solitary phialide. Saccardo (1895) transferred the species to the perfect genus Gymnoascus, publishing its name as a new combination, Gymnoascus luteus (Zukal) Sacc. The correct citation of the name is G. luteus Sacc. Since the species does not belong in Gymnoascus but represents a species of Talaromyces, it is transferred to that genus. The name Penicillium luteum Zukal remains valid for the imperfect state.

According to Derx (1925, 1926), T. luteus should be heterothallic. The development of its initials, however, excludes the possibility of heterothallism.

Ascomatal initials consisting of a pair of short coiled hyphae as described by Emmons (1935) have not been observed, nor has the formation of asci from croziers. Asci are borne in very short coiled chains.

Kominami et al. (1952) pointed out the resemblance between the ascospores of T. luteus and Trichocoma paradoxa Junghuhn, both being ornamented with transversely arranged ridges. Moreover they reported the isolation of a biverticillate-symmetrical Penicillium from an ascoma of the latter species. The authors did not obtain ascomata in their cultures. Branches of Machilus thunbergii, the host of Trichocoma paradoxa in Japan, were infected with conidia of this isolate, but no ascomata developed on them. The culture, CBS 247.57, isolated by Kominami et al. (1952) has been maintained at the CBS for about 15 years. It produces restricted creamish colonies, developing reduced penicilli, consisting mainly of a few atypical phialides. It is not identical with P. luteum. Because of the degenerated state of the culture an identification is not possible. Kominami et al. (1952) concluded that Talaromyces luteus and Trichocoma paradoxa are very closely related, and suggested that T. luteus might even represent a degenerated form of T. paradoxa. However, the latter hypothesis seems discussable, since the ascospores of T. luteus are yellow and those of Trichocoma paradoxa are brown. In addition, there are differences in ascospore sizes. [p. 27] Though the ascomata are very different in structure, the presence of a Penicillium-like conidial state may indicate some relationship.

T. luteus differs from T. udagawae in producing larger ascospores ornamented with irregular, locally interrupted ridges, whereas those of T. udagawae show regular, spiral ridges. Moreover the initials are different.

The ascomatal initials resemble slightly those of T. rotundus. In both species they develop predominantly from phialides.

 

 

7. Talaromyces rotundus C. R. Benjamin, spec. nov. - Fig. 7.

 

Talaromyces rotundus (Raper & Fennell) C. R. Benjamin - Mycologia 47: 683. 1955 (as comb. nov.).

 

Diagnosis latina in `Mycologia' 40: 518. 1948, continetur.

Typus novae speciei status perfectus culturae CBS 369.48.

 

Status conidialis: Penicillium rotundum Raper & Fennell - Mycologia 40: 518. 1948 (described inclusive of the perfect state).

 

Colonies on malt agar growing slowly, attaining a diameter of about 2,5 cm within 2 weeks at 25C, consisting at first of a compact tough orange felt occasionally furrowed in radial pattern, in which after 2 to 3 weeks numerous ascomata develop, showing bright orange shades near OrangeBuff and Capucine Yellow (Ridgway, Pl. 3; Rayner, 13d-15d, 13-15b), surrounded by a broad uncoloured growing margin, largely composed of submerged mycelium; conidial state scanty, usually not affecting the colony appearance; sometimes white to greyish green sectors occur, consisting of a floccose to funiculose mycelium bearing more or less numerous penicilli. Reverse pinkish orange, approximating Capucine Orange (Ridgway, Pl. 3; Rayner, 13 d).

Vegetative hyphae hyaline, occasionally orange and encrusted, 1-2.5 m in diameter.

Ascomata orange, usually globose, 50-300 m in diameter, discrete or confluent, ripening within 2-3 weeks. Covering consisting of a thin network of loosely interwoven hyphae, surrounded by a weft of radiating, yellow to orange, heavily encrusted hyphae, about 1.5 m in diameter. Initials developing either directly from vegetative hyphae or from atypical phialides, consisting of swollen, somewhat twisted hyphal elements, which become irregularly septate and produce coiling branches thus developing a compact mass of ascogenous hyphae. Asci 6-8-spored, globose to ovoidal, 12-14 x 9.5-13 m. Ascospores globose, 4.5-5.2 m in diameter, thick-walled, spinulose with spines up to 0,5 m in length.

Conidial state produced scantily on all media tested, best development on Czapek + 20% sucrose, on which medium conidial structures occur [p. 28] abundantly, giving the cultures a greyish green appearance. Conidiophores arising both from submerged and aerial hyphae (on malt agar only from aerial hyphae), short, 25-75 x 1.5-2.2 m. Metulae in small verticils of 2 to 3, but often lacking, 10-17 x 1.5-2.5 m. Phialides lanceolate, about 2 to 6 in the verticil, sometimes solitary, 10-16 x 1.5-2.2 m, occasionally atypical, producing ascomatal initials instead of conidia. Conidia hyaline, greenish in mass, fusiform, 3-5 x 1.5-2 m, smooth-walled.

 

 

Fig. 7. Talaromyces rotundus, CBS 369.48. a. conidiogenous structures; b. conidia; c. initial developing from an atypical phialide; d. initials developing directly from vegetative hyphae; e. asci produced in chains; f. ascospores.

 

Material examined

 

CBS 369.48 = NRRL 2107, type culture, isolated in 1946 by G. W. Martin from wood collected in Panama. [p. 29]

CBS 587.72 = NHL 6122, = IFO 9142, isolated in 1968 by S. Udagawa from pine forest soil, Sinisip, Philippines.

 

The species can be distinguished from the other species of the sect. Talaromyces by its spinulose, globose ascospores. In shape the ascospores are similar to those of T. bacillisporus (sect. Emersonii). They are, however, slightly larger. Moreover, T. rotundus differs from T. bacillisporus in the conspicuous orange colour of the ascomata and in the nature of the conidial state.

T. rotundus shows some relationships with T. wortmannii, since the ascomatal initials of both species are of the same type, though those of T. wortmannii develop always directly from the vegetative hyphae. The two species differ mainly in the shape of the ascospores.

 

 

8. Talaromyces stipitatus C. R. Benjamin, spec. nov. - Fig. 8

 

Talaromyces stipitatus (Thom) C. R. Benjamin - Mycologia 47: 684. 1955 (as comb. nov.).

 

Diagnosis latina status perfecti in `Mycologia' 27: 138. 1935, sub Penicillium stipitatum Thom.

Typus novae speciei status perfectus CBS 375.48.

 

Status conidialis: Penicillium stipitatum Thom in Emmons Mycologia 27:138.1935

 

Colonies on malt agar growing fairly rapidly, attaining a diameter of 5 to 5.5 cm within two weeks, consisting mainly of a dense layer of ascomata produced near the agar surface, showing usually yellow shades ranging from Barium Yellow to Citron Yellow (Ridgway, Pl. 16; Rayner, 23'd, 23'b) (in occasional strains salmon coloured, e.g. CBS 349.72), embedded in and overgrown by a loose aerial hyphal network; margin irregular, lobed; conidial state scanty, not affecting the colony appearance, odour unpronounced. Reverse yellow to orange yellow near Light Cadmium (Ridgway, Pl. 4; Rayner, 19b) with the pigment diffusing throughout the surrounding medium.

On Czapek agar the reverse is orange brown, while on oatmeal agar it is only slightly coloured, usually showing pale yellow to pinkish shades.

Vegetative hyphae hyaline to yellowish, occasionally encrusted, 2-6 m in diameter.

Ascomata usually yellow, in some strains pinkish to salmon-coloured, globose or nearly so, 150-400 m in diameter, discrete or confluent, ripening within 10 to 14 days. Coverings distinct, consisting of a few layers of yellow, closely interwoven hyphae, simulating a pseudoparenchymatous [p. 30] wall; surrounded by a loose weft of heavily encrusted mainly radiating hyphae, about 1-1.5 m in diameter. Initials consisting of ascogonia and antheridia, usually arising as similar or barely differentiated branches from separate hyphae, but occasionally produced from the same hypha. The two gametangia coil around each other and after about two coils they fuse. After fertilization the ascogonia grow out to form conspicuous sometimes somewhat irregularly swollen, usually aseptate, rarely septate hyphae, about 20-40 x 3-4 m. At their apices they produce a few gnarled branches which continue to branch profusely thus developing the ascogenous hyphae. On oatmeal and cornmeal agar the fertilized ascogonia often produce more straight, slightly longer, 1- to 4-septate prolongations before branching apically, this is most striking in the type culture. Asci 8-spored, broadly ellipsoidal to subglobose, 5.5-7.5 x 4-5.5 m. Ascospores yellow, flattened ellipsoidal, with a single equatorial ridge, 3.2-4 x 1.7-2.2 m (ridge 0.2 to 0.5 m), in some strains up to 4.5 x 2.5 m, smooth-walled or nearly so (Pl. 1.E).

Conidiophores arising primarily from aerial hyphae, 10-100 x 2-2.5 m. Metulae 11-16 x 2.2-3 m, in small verticils of about 2 to 3, sometimes lacking. Phialides about 2 to 6 in the verticil, typically lanceolate, 12-15 x 2-3 m, occasionally up to 25 m in length with a long neck, solitary phialides and reduced penicilli as occurring in T. intermedius rarely observed. Conidia ellipsoidal, ovoidal to pyriform, 2-7.5 x 2-4 m, hyaline to pale greenish, smooth-walled.

CBS 375.48 is regarded as the type culture of both states.

 

Material examined

 

CBS 375.48 = NRRL 1006, type culture, isolated from rotting wood in Louisiana in 1931. This is one of the strains studied by Emmons (1935) on which he based his description of P. stipitatum. The ascogonial hyphae growing out after fertilization may become much longer (up to 100-150 m in length) before producing branches and are more straight than those occurring in the other strains. On malt agar the pigment diffusing in the agar shows rather pinkish than yellow shades. This strain was used by Birkinshaw et al. (1942) and Segal (1957, 1959) in their study on the production of stipitatic and stipitatonic acid.

CBS 292.53, isolated from a beech-wood board in 1953, Delft, the Netherlands.

CBS 236.60, isolated from a chicken crop in 1960, Amersfoort, the Netherlands. The reverse of this strain shows occasionally pinkish red shades in age, especially on Czapek agar.

CBS 227.72 = NRRL 2105, isolated from soil in Minnesota in 1946. The four strains mentioned above agree very well in the colour of the ascomata and in the sizes of the ascospores.

CBS 349.72, isolated by M. S. Pavgi from blighted flowers of Tagetes patula in Varanasi, India. This strain is characterized by quite a different cultural appearance on all media tested. Its ascomata as well as its aerial mycelium show pinkish to salmon shades near Salmon Color (Ridgway, Pl. 14; Rayner, 7'd-9'd). Only occasionally central areas are overgrown by yellow mycelium while a few of the ascomata are creamish. The reverse shows pinkish orange shades when young, becoming reddish brown in age. Its microscopic characters agree with those of T. stipitatus, only its ascospores are slightly larger, 3.5-4.2 (-4.5) x 1.7-2.5 m (ridges about 0.2 m wide). The pinkish to orange colours displayed [p.32] by this strain are also mentioned by Emmons (1935) in his description of P. stipitatum (strain CBS 375.48) as occurring in old cultures. However, in this strain, these colours are now lost. A biochemical study of the red-brown colour of CBS 349.72, revealed that the pigment mainly consists of the anthraquinone derivative erythroglaucine (van Eijk, in preparation).

CBS 581.72 A = IFO 7230 = NHL 6057; CBS 581.72 B = IFO 8669 = NHL 6091, received from K. Tubaki, Tokyo. The sizes of the ascospores agree with CBS 349.72, but the cultures show rather yellow than salmon shades. [p.31]

 

 

Fig. 8. Talaromyces stipitatus, CBS 375.48. a. conidiogenous structures; b. conidia; c. coiling gametangia; d. outgrowing ascogonium, producing apically branched coiling hyphae on oatmeal agar; e. chains of asci; f. ascospores; g. fragment of ascomatal covering; CBS 227.72. h. initial on malt agar. [p. 32]

 

T. stipitatus can easily be distinguished from the other species of Talaromyces by the unique pattern of the ascospores and initials. Because of the presence of paired gametangia it shows relationships with T. flavus and T. helicus.

 

 

9. Talaromyces trachyspermus (Shear) Stolk & Samson, comb. nov. - Fig. 9

 

Arachniotus trachyspermus Shear - Science, N.Y. II, r6: 138. 1902 (basionym).

Talaromyces spiculisporus (Lehman) Benjamin - Mycologia 47: 683. 1955 (as comb. nov.).

 

Status conidialis: Penicillium spiculisporum Lehman - Mycologia 12: 268. 1920 (described inclusive of the perfect state).

 

Colonies on malt agar growing rapidly, attaining a diameter of about 3.5 cm within 2 weeks at 25C, consisting of a basal felt in which after one week numerous white to creamish ascomata develop, overgrown by a floccose to funiculose mycelium, surrounded by a colourless growing margin, composed of submerged mycelium. Conidial structures usually limited and developing in the central part of the colony, occasionally abundantly produced and giving a more greenish appearance to the colony, near Court Gray (Ridgway, Pl. 47; Rayner, 31f. Odour sweet aromatic. Exudate absent. Reverse in yellow shades.

Vegetative hyphae hyaline, smooth, septate, 1.5-4 m in diameter.

Ascomata at first white, becoming creamish to yellowish in age, globose, 50-350 m in diameter, usually confluent, ripening within 2 weeks. Covering distinct, consisting of a network of closely interwoven hyphae, simulating a pseudoparenchymatous wall, surrounded by a loose weft of radiating hyphae, usually encrusted with granules, 1.5-2.5 m in diameter. Initials consisting of swollen or intercalary portions of aerial hyphae, which become strongly gnarled and branch profusely. Asci globose to ovoidal, 6-7.5 x 6-6.5 m. Ascospores ellipsoidal, 3-3.5 x 2-2.5 m, spinulose with spines about 0.5 m long (Pl. 1.B).

Conidiophores usually short and borne as branches from aerial hyphae, [p.33] 17-45 x 1.5-2.5 m. Metulae, in small verticils of 2 to 3, 11-15 x 1.5-2.2 m, sometimes lacking. Phialides about 3 to 7 in a verticil, lanceolate, 12-20 x 2-2.5 m. Conidia ellipsoidal to ovoidal, 2.5-3.5 x 1.5-2.7 m, smoothwalled, green in mass.

 

 

Fig. 9. Talaromyces trachyspermus, CBS 373.48. a. conidiogenous structures; b. conidia; c. initials; d. chains of asci; e. ascospores; Type collection USDA 5798. f. ascospores; Herbarium specimen 7b. g. conidiogenous structures with conidia; h. ascospores.

 

Material examined

 

Herbarium specimens

 

Arachniotus trachyspermus in herb. BPI (type), USDA 5798, on Vaccinium macrocarpon, Jamesbury, N.J., Sept. 1901, collected and determined by C. L. Shear; 10 additional authentic specimens all collected on or isolated from Vaccinium macrocarpon by C. L. Shear from different localities in the USA.

The labeling of some additional specimens of Arachniotus trachyspermus (in herb. BPI) is somewhat confusing. One specimen labeled `A. trachyspermus specimen from pure culture on cornmeal agar no. 7a and 7b, June 2', is also indicated as type. However, this specimen was not isolated from the type (USDA 5798), but collected at Washington Market in 1902. Another specimen, labeled as `A. trachyspermus 7b', is provided with a handwritten note: 'Contam. with Penicillium? Penicillium evidently conid. form'. Another [p. 34] specimen labeled: `A. trachyspermus, on Vaccinium macrocarpon, New Jersey, Sept. 1905, coil. & det. C. L. Shaer', did not represent A. trachyspermus, but a Byssochlamys species. Unfortunately this material is too scanty for a more exact identification.

 

Living cultures

 

CBS 373.48 = NRRL 1028 = ATCC 10497 = IMI 40,043. This culture was sent by C. W. Emmons to the NRRL; Raper and Thom used it for the description of P. spiculisporum.

CBS 346.54, isolated by the Centre Technique Forestier Tropical, Nogent sur Marne. CBS 282.58, isolated by M. B. Schol-Schwarz from jute treated with copper-naphthanate.

CBS 112.64, from soil treated with cyanamide and sent by D. Knsel, Stuttgart-Hohenheim. These strains agree completely with one another.

 

T. trachyspermus is characterized by its conspicuously spinulose, ellipsoidal ascospores and its distinct ascomatal covering. It is closely related to T. intermedius, because of the structure of its initials and the spinulose ascospores. Shear (1902) classified his species in Arachniotus since he did not observe any penicilli, which indeed are lacking in the type specimen (USDA 5798). However, he had observed a Penicillium in his specimen 7b. This Penicillium proved to be identical with P. spiculisporum. Fresh isolates of this species may produce penicilli very scantily and consequently they may easily be overlooked. Most of Shear's material consists mainly of ascospores, but in a few specimens, chains of asci as occurring in Talaromyces can be recognized. The ascospores observed in the type specimen of A. trachyspermus are identical with those of P. spiculisporum Lehman. Moreover Shear's description of the ascomata agrees completely with those found in P. spiculisporum. Therefore A. trachyspermus is considered as the perfect state of P. spiculisporum and consequently Shear's species is transferred to Talaromyces.

Because of the white to creamish colour of the ascomata and the scantily produced, often fractional penicilli, T. trachyspermus has occasionally been misidentified as a species of Byssochlamys. However, T. trachyspermus can easily be distinguished from Byssochlamys by the disposition of its asci, the distinct wall-like ascomatal covering and the spinulose ascospores.

 

 

10. Talaromyces ucrainicus Udagawa spec. nov. - Fig. 10

 

Talaromyces ucrainicus (Panasenko) Udagawa - Trans. mycol. Soc. Japan 7: 94. 1966 (as comb. nov.).

 

Diagnosis latina in `Mycologia' 56: 59. 1964, continetur.

Typus novae speciei status perfectus culturae CBS 162.67.

 

Status conidialis: Penicillium ucrainicum Panasenko - Mycologia 56: 59. 1964 (described inclusive of the perfect state). [p. 35]

 

Colonies on malt agar growing restrictedly, attaining a diameter of about 3 cm within 2 weeks at 25C, consisting of a basal felt in which numerous yellow ascomata develop, overgrown by a floccose, occasionally funiculose mycelium, showing bright yellow colours approximating Maize Yellow and Buff-Yellow (Ridgway, Pl. 4; Rayner, 19 f, 19 d). Conidial state scantily produced and not affecting the colony appearance. Odour sweet or not pronounced. Exudate sometimes present as clear drops. Reverse in yellow to reddish shades.

Vegetative hyphae hyaline, septate, 0.7-2 m in diameter.

Ascomata yellow, globose, 50-500 m in diameter, discrete or confluent, ripening within 2 weeks. Covering consisting of a network of loosely interwoven yellow hyphae, surrounded by a loose weft of hyphae, encrusted with yellow granules, about 2 m in diameter. Initials consisting of swollen branches or intercalary portions of aerial hyphae, which become strongly gnarled and branch profusely. Asci 6-8-spored, globose to ovoidal, 6.5-7.5 x 5-6.5 m. Ascospores ellipsoidal, ornamented with thin, somewhat jagged, irregular, for the greater part longitudinal ridges, 2.5-3.5 x 2-2.4 m (Pl. 2.E-G).

Conidiophores arising from aerial hyphae, up to 80 m in length and 1.5-2.5 m in diameter. Metulae in small verticils of about 2 to 4, 11.5-15 x 2.5-3 m, sometimes lacking. Phialides lanceolate, about 2 to 4 in verticils, occasionally occurring singly, 9-18 x 2-3 m. Conidia subglobose to ellipsoidal, 2.5-3.5 x 1.5-3 m, grey green in mass, smooth-walled.

When Udagawa (1966) described T. ucrainicus (Panasenko) Udagawa, he based his description on two cultures NHL 6086 and NHL 6087. However, he did not designate a type for the perfect state. CBS 162.67 = NHL 6086 is now indicated as the type of the perfect state of this species.

 

Material examined

 

CBS 127.64, isolated by D. Knsel, Stuttgart-Hohenheim, from cyanamid-treated agricultural soil.

CBS 162.67 = NHL 6086, type culture of T. ucrainicus, isolated from soil in Nishinasuno, Tochigi Pref., Japan and sent to the CBS by S. Udagawa.

CBS 626.67 = BKM F-907 = type culture of Penicillium ucrainicum Panasenko, isolated from potato starch. No ascus production could be observed in this strain. The conidial state is not or very sparsely produced.

CBS 583.72 A = IFO 8395; CBS 583.72 B = IFO 8529; CBS 583.72 C = IFO 8891;

CBS 583.72 D = IFO 8895, received from K. Tubaki, Tokyo.

 

Because of its typically ornamented ascospores, this species shows relationships with T. thermophilus and T. leycettanus. However, these two species are thermotolerant or thermophilic, and they also produce different conidial states. The ascomatal initials of T. ucrainicus, described above, are identical with those of T. intermedius and T. trachyspermus, and although [p.36] ascospore ornamentation is quite different in these species, they are regarded as being closely related.

 

 

Fig. 10. Talaromyces ucrainicus, CBS 162.67. a. conidiogenous structures; b. conidia; c. initials; d. asci produced in chains; e. ascospores.

 

 

11. Talaromyces udagawae Stolk & Samson, spec. nov. - Fig. 11.

 

Status conidialis: Penicillium udagawae Stolk & Samson, stat. nov.

 

Coloniae in agaro maltoso 25C lente crescunt, flavae; lentius in agaro Czapekii. Hyphae vegetativae hyalinae, leves, 1.5-4 m crassae. Ascomata lutea ad aurantiaca, globosa, 200-400 m diametro, confluentia vel nonnumquam solitaria, 15 diebus maturantia. Reticulo tenui laxe intricatarum hypharum involuta hyphis radiantibus circumdata. Asci subglobosi vel ellipsoidei, 9-11 x 6.5-8 m. Ascosporae ellipsoideae, 3.54.5 x 2.2-3 m, 3 ad 5 cristis transversalibus saepe spiralibus ornatae.

Status conidialis abest in agaro maltoso, parcus in agaro extracto feni addito. Conidiophora erecta, hyalina, levia, 50-200 x 2.5-3.5 m; metulae binae vel quaternae verticillatae, 7.5-10 x 2.2-2.4 m. Phialides ternae ad septenae verticillatae, lanceolatae, 12-15 x 2.2-2.5 m. Conidia hyalina, subglobosa vel ellipsoidea, 3-4 x 2.2-3 m, levia.

Typus: CBS 579.72, isolatus a S. Udagawa, e terra, Misugimura, in Japonia. [p. 37]

Etym.: The specie is named after the Japanese mycologist Dr S. Udagawa

 

Colonies on malt agar growing very restrictedly, attaining a diameter of 1.5 to 1.7 cm within 2 weeks at 25C, consisting of a thick interwoven felt, in which numerous ascomata are embedded near the agar surface, usually forming a dense layer showing bright yellow colours ranging from Lemon Yellow at the margin to Empire Yellow in central areas (Ridgway, Pl. 4; Rayner, 23, 21 b) surrounded by a thin, regular, submerged margin. Exudate in clear to orange drops. Odour not pronounced. Reverse orange, near Orange-Buff, becoming Ochraceous Orange in the centre (Ridgway, Pl. 15; Rayner, 15'b, 15'). Pale-yellow rods are produced abundantly under the mycelium and in the surrounding agar.

Colonies on oatmeal agar resemble those on malt agar, but ascomata are less abundantly overgrown by aerial mycelium and the submerged colony margin is much broader, showing pale orange shades. Reverse as on malt agar.

Colonies on Czapek agar grow slightly slower than those on malt agar; colony texture, ascomata and colours are as described above for colonies on malt.

Vegetative hyphae hyaline to yellow, smooth, sometimes heavily encrusted, I.5-4 m in diameter.

Ascomata yellow to orange, globose or nearly so, about 200-400 m in diameter, usually confluent, occasionally discrete, ripening within 2 weeks. Covering consisting of thin networks of loosely interwoven hyphae with irregular swellings, surrounded by wefts of predominantly radiating, branched, heavily yellow-encrusted hyphae, about 1-1.5 m wide. Development of the initials starts with the terminal or intercalary swelling and branching of hyphae, occasionally forming a small plexus of swollen cells, from which long, single, coiling aseptate hyphae develop, 1.5-2 m in diameter, producing several loose coils in a helical pattern. These coiling initials may also develop from single swollen hyphal cells. No fusion has been observed. Soon the coiling hyphae become septate and produce branches which continue to branch profusely, thus developing into ascogenous hyphae. Asci 8-spored, broadly ellipsoidal to subglobose, 9-11 x 6.5-8 m. Ascospores ellipsoidal, 3.5-4.5 x 2.2-3 m, ornamented with 3 to 5 regularly transverse, nearly parallel ridges, 0.5 m wide (rarely I m), often spirally arranged (Pl.1.F-G).

Conidial state not produced on malt agar, abundantly developed on hay-infusion agar at the colony margin. Conidiophores predominantly arising from the substratum, occasionally from floccose to funiculose hyphae, hyaline, erect, smooth, 50-200 x 2.5-3.5 m. Metulae 2 to 4 in the verticil, 7.5-10 x 2-2.4 m. Phialides in whorls of 3 to 7, lanceolate, 12-15 x 2.2-2.5 m. Conidia hyaline, subglobose to ellipsoidal, 3-4 x 2.2-3 m, smooth-walled. [p. 38]

 

 

Fig. 11. Talaromyces udagawae, CBS 579.72. a. conidiogenous structures; b. conidia; c. swollen and branching, intercalary hyphae, from which the initials will develop; d. initials consisting of coiling hyphae; e. septation and subsequent branching of an initial; f. development of asci in chains; g. ascospores; h. fragment of ascomatal covering.

 

Material examined

 

Herbarium specimen

 

Sporotrichum malagense in herb. W (type), Mycotheca Universalis no. 1173; in herb. L. 910.251-467, Herb. myc. oec. 630. Both specimens on mouldy raisins, collected by F. de Thmen, Klosterneuburg, Aug. IS 78.

 

Living culture

 

CBS 579.72 = NHL 6089 = IFO 8808 = type culture, isolated in 1964 by S. Udagawa from soil, Misugimura, Japan and sent to the CBS by K. Tubaki, Tokyo. [p. 39]

 

T. udagawae is closely related to T. luteus, in both species the ascospores are ornamented with transverse bands. However, T. udagawae produces smaller ascospores than does T. luteus, while the ascospore ridges of the first species are thinner and often spirally arranged, and appear more regularly transverse than in the latter species. Moreover, the initials of the two species are different. In T. udagawae they consist of very long, single, strongly helicoidal hyphae, forming many loose coils, which at first are not branched. In T. luteus on the other hand, the initials are very compact, consisting of short, irregularly coiling and branching hyphae.

Examination of herbarium material of Sporotrichum malagense Thmen (Herb., myc. oec. 630) showed this specimen to consist mainly of asci and ascospores of T. udagawae. However, in the type specimen of S. malagense (Myc. Universalis no. 1173) neither ascospores of T. udagawae nor conidia of S. malagense were observed. Therefore S. malagense is considered as a doubtful species.

 

 

12. Talaromyces wortmannii C. R. Benjamin, spec. nov. - Fig. 12

 

Talaromyces wortmannii (Klcker) C. R. Benjamin - Mycologia 47: 683. 1955 (as comb. nov.).

 

Ascomata lutea vel aurantiaca, globosa, 75-300 m diam., reticulo tenui laxo cooperta, e quo hyphae contortae, 0.7-2 m crassae radiant. Asci 7.5-10 m diametro. Ascosporae ellipsoideae, 3.7-4.7 (-5) x 2.5-3.5 m, spinulosae, spinis ad sum longis, luteae. Stat. conid. Penicillium wortmannii Klcker.

Typus: CBS 391.48, isolatus ab A. Klcker e terra in Dania, 1903.

 

Status conidialis: Penicillium wortmannii Klcker - C. r. Trav. Lab. Carlsberg 6: 100. 1903 (described inclusive of the perfect state).

 

Colonies on malt agar growing restrictedly, attaining a diameter of 3 to 4 cm within 14 days at 25C, consisting of a basal felt with numerous ascomata, showing bright yellow orange shades near Orange-Buff to Deep Chrome (Ridgway, Pl. 3; Rayner, 13d-15d, 17b), in some strains more yellow colours are present, approximating Apricot Yellow to Empire Yellow (Ridgway, Pl. 4; Rayner, 19b, 2b). Conidial structures sparsely produced and not affecting the colony appearance or only occasionally produced abundantly in some sectors. Exudate sometimes present as orange drops. Reverse in yellow to orange shades near Orange to Mars Yellow (Ridgway, Pl. 3; Rayner, 13b-15b, 15i).

Vegetative hyphae hyaline or encrusted with orange granules, 1-3 m in diameter.

Ascomata yellow to orange, globose, 75-300 m in diameter, discrete or confluent, ripening within 14 days. Covering composed of a thin network of loosely interwoven hyphae, surrounded by radiating, twisted hyphae, which are encrusted, 0.7-2 m in diameter. Initials consisting of terminal or [p. 40] intercalary, swollen, somewhat twisted elements of the vegetative hyphae, which become irregularly septate and produce coiling branches, which continue to branch profusely, producing ascogenous hyphae. Asci globose to ovoidal, 7.5-10 m in diameter. Ascospores ellipsoidal 3.7-4.7 (-5) x 2.5-3.5 m, spinulose with spines occasionally up to 1 m long.

Conidiophores usually arising from submerged hyphae, up to 150 m long, 1.8-2.5 m in diameter, smooth-walled, occasionally encrusted with yellow granules. Metulae in verticils of 4 to 8, measuring 8-12 x 2-2.5 m. Phialides about 4 to 8 in the verticil, lanceolate, 10-12.5 x 1.5-2.2 m. Conidia ellipsoidal, but mostly fusiform, pointed, 2.5-4 x 1.7-2.2 m, smooth to finely spinulose, hyaline to greenish.

 

 

Fig. 12. Talaromyces wortmannii, CBS 391.48. a. conidiogenous structures; b. conidia; c. initials; d. chains of asci; e. ascospores.

 

Material examined

 

CBS 235.38, sent by B. Tubbingen.

CBS 391.48 = NRRL 1017 = IMI 40,047 = ATCC 10517 = IFO 7738 = type culture isolated by A. Klcker from soil in Denmark in 1903.

CBS 293.53, isolated by G.A. de Vries from a soil sample from Lorenzo Marques, S. Africa. [p. 41]

CBS 319.63, isolated at the CBS from air.

CBS 387.67, sent by W. Kerner, Berlin, as no V 74.

CBS 553.72, received from J. Nicot, Paris (strain 2003), isolated from soil in France, 1968.

 

T. wortmannii has often been confused with T. flavus, since the cultures and ascospores may strongly resemble one another. Fortunately the type culture of T. wortmannii is still preserved in the CBS collection, thus defining this species. The two species can easily be distinguished by their ascomatal initials. In addition T. wortmannii differs from T. flavus by its restricted growth, the more orange colour of the colonies, especially of the reverse and by the slightly larger and more definitely pointed conidia. In T. wortmannii and T. rotundus the same type of ascomatal initials can be observed. However, the initials in T. rotundus develop directly from the vegetative hyphae or from atypical phialides. In T. wortmannii the initials are always formed directly from the vegetative hyphae.

 

 

Talaromyces section Emersonii Stolk & Samson, sect. nov.

 

Involucrum ascomatum parcum vel distinctum. Ascosporae flavae. Status conidialis: Penicillium cylindrosporum-series vel Paecilomyces. Species thermophilae vel thermotolerantes.

 

Species typica: Talaromyces emersonii Stolk.

 

Ascomatal coverings scanty to distinct. Ascospores yellow. Imperfect state: Penicillium cylindrosporum-series or Paecilomyces. Thermophilic or thermotolorant.

 

The four species comprising this section develop very well on malt and oatmeal agar. On Czapek agar growth is more restricted, while penicilli and ascomata are produced in limited numbers. Cultures of T. bacillisporus develop more aerial mycelium on Czapek agar than those of the other three species of this section; in the latter species colonies are very thin, and the mycelium is for the greater part submerged on Czapek agar.

Optimum temperatures of the four species are relatively high, ranging from 35C to 45C depending on the species. All of them grow well at 40C. T. bacillisporus is slightly thermotolerant, T. leycettanus thermotolerant to thermophilic, the other two species are thermophilic.

In T. emersonii and in the new species T. byssochlamydoides, the ascomatal coverings are very scanty, whereas in T. leycettanus and especially in T. bacillisporus the coverings are much better developed. [p. 42]

No paired gametangia occur in the four species belonging to this section. Initials show different structures. Asci occur in short, curved or coiled chains, they are usually 8-spored. Ascospores are ellipsoidal or globose, their walls being smooth or showing various ornamentations.

Both T. leycettanus and T. byssochlamydoides have a Paecilomyces conidial state. The imperfect states of T. emersonii and T. bacillisporus constitute together with the two imperfect Penicillia, P. cylindrosporum G. Smith (1957) and P. argillaceum Stolk et al. (1969) a new series in the imperfect Penicillia: the Penicillium cylindrosporum-series, differing from all series, introduced by Raper & Thom in 1949. This new series is characterized by biverticillate-asymmetrical, somewhat appressed penicilli, with all elements typically rough-walled, by phialides consisting of a cylindrical basal portion and tapering abruptly to a short but distinct tip, and by predominantly cylindrical conidia, colouring the colonies brown or creamish depending on the species. Because of the colour of the colonies, the nature of the phialides and the cylindrical conidia, the P. cylindrosporum-series connects the genus Penicillium with Paecilomyces. It differs clearly from this latter genus by the much shorter conidium-bearing necks, the compact structure and the roughened walls of its penicilli. Moreover, the conidial chains are never divergent as in Paecilomyces; in T. emersonii they are even produced in columns.

 

Key to the species

 

1a.

Conidial state Paecilomyces

2

1b.

Conidial state Penicillium cylindrosporum-series

3

 

 

 

2a.

Ascomata pink to reddish; ascospores smooth, 3.7-4.5 x 3.5-4 m; strongly thermophilic

T. byssochlamydoides

2b.

Ascomata greenish yellow; ascospores with 2 to 6 irregular, for the greater part longitudinal ridges, 3.2-4 (-4.5) x 2.2-3 m; thermotolerant to thermophilic

T. leycettanus

 

 

 

3a.

Ascomata orange brown, ascospores smooth, 3.5-4 x 2.7-3.5 m; strongly thermophilic

T. emersonii

3b.

Ascomata pale yellow, ascospores globose, 3.5-4.5 m, spinulose; weakly thermotolerant

T. bacillisporus

 

 

13. Talaromyces bacillisporus C. R. Benjamin, spec. nov. - Fig. 13

 

Talaromyces bacillisporus (Swift) C. R. Benjamin - Mycologia 47: 684. 1955 (as comb. nov.).

 

Diagnosis latina in `Bull. Torrey bot. Club' 59: 221. 1932, continetur. Typus novae speciei status perfectus culturae CBS 296.48.

 

Status conidialis: Penicillium bacillisporum Swift - Bull. Torrey bot. Club 59: 221. 1932. [p. 43]

 

Colonies on malt agar, attaining a diameter of 5.5 to 7.5 cm within 14 days at 30C, usually zonate, consisting of a dense layer of ascomata developing near the agar surface, showing pale-yellowish to ochraceous shades, ranging from Ivory Yellow to Massicot Yellow, becoming Colonial Buff in age (Ridgway, Pls. 30, 16; Rayner, 21"f, 21'f, 21"d), embedded in and covered by a floccose to funiculose aerial mycelium bearing abundant penicilli not affecting the colony appearance; occasionally, greenish zones occur due to the green colour of the reverse shining through. Reverse typically intensely green, ranging from Deep Malachite Green to Cress Green (Ridgway, Pls. 32, 31; Rayner, 35"b, 29"k), usually with yellow or orange brown zones; central areas becoming dark brown.

Colonies on oatmeal agar usually showing paler shades, in age becoming Vinaceous-Buff (Ridgway, Pl. 40; Rayner, 17'''d) with the green reverse shining through. Reverse conspicuous green approximating to French Green, becoming Danube Green (Ridgway, Pl. 32; Rayner, 35"b, 35"m) with yellow to greenish yellow zones which become greenish brown in age.

Vegetative hyphae hyaline, encrusted with yellow or green granules, 1.5-4 m in diameter.

Ascomata yellow, globose, 80-150 m in diameter, discrete or confluent, ripening within two weeks at 30C. Covering consisting of a network of interwoven yellowish, often encrusted hyphae, surrounded by radiating, twisted, encrusted hyphae, about 1 m wide. Development of the ascomatal initials starts with the production of intercalary, occasionally terminal cells which swell considerably and grow out to produce wide, coiled, profusely branching hyphae which develop into the ascogenous hyphae. Asci globose to subglobose, 9-12 x 9-10.5 m. Ascospores globose, 3.5-4.5 m in diameter, finely spinulose (spines 0.2-0.5 m long).

Conidial state creamish, occasionally greyish. Conidiophores arising primarily as perpendicular branches from trailing hyphae, or from ropes of hyphae, hyaline to pale yellow, septate, smooth to roughened, 25-60 x 2-3 m. Penicilli typically biverticillate, consisting of verticils of 2 to 3 (4) metulae, bearing whorls of 3 to 6 parallel phialides, but very often monoverticillate. All elements of the penicillus hyaline to yellowish. Metulae smooth to rough-walled, 8-15 x 2-2.5 m. Phialides smooth or nearly so, 9-12.5 x 2-2.5 m, consisting of a cylindrical base, tapering abruptly to a conspicuous conidium-bearing tip, 1.5-2.5 x 1.2 m. Conidia cylindrical, occasionally ellipsoidal, 3-4.5 x 1.2-1.5 (-3) m, smooth, hyaline, produced in long parallel chains, at first forming loose. columns, which usually become slightly tangled in age.

The species is weakly thermotolerant, minimum temperature 17C, optimum 35C, maximum about 45C.

 

Material examined

 

CBS 296.48 = NRRL 1025 = type culture, isolated by M. C. Swift, from Begonia leaf, New York City, 1935.

CBS 136.45, received from N.C.T.C., London as NCTC 6601, October 1945. [p. 44]

CBS 158.67 = NHL 6085, isolated by S. Udagawa, from soil in Tochigi, Japan, August 1964.

CBS 580.68, isolated by T. Nilsson, from birch chips, Stockholm.

 

 

Fig. 13. Talaromyces bacillisporus, CBS 296.48. a. conidiogenous structures; b. conidia; c. swollen cells from which the initials will develop; d. initials, producing branches; e. developing asci; f. chain of asci; g. ascospores.

 

The ascomatal coverings of T. bacillisporus are much better developed than those of the other species of the sect. Emersonii. In this respect it resembles the species of the sect. Talaromyces, thus connecting the sections Emersonii and Talaromyces. Ascomatal initials consisting of a pair of short coiled hyphae, as described by Emmons (1935) and Udagawa (1966), have not been observed. However, it is very difficult to establish the exact development of the initials, due to the rapid branching of the coiling hyphae, which quickly produce very complicated primordia. Raper and Thom (1949) placed the conidial state of T. bacillisporus in the Biverticillata Symmetrica on account of the structure of the phialides. However, we found the phialides to be of the same type as those occurring in the P. cylindrosporum-series. In addition, the elements of the penicillus show the same tendency to be closely appressed, while the walls of both conidiophores and penicilli are somewhat roughened. Therefore, P. bacillisporum can better be classified in the P. cylindrosporum-series. It differs from the other species of this series by producing less complicated and less roughened penicilli. [p. 45] Moreover the colour of the conidial state is not brown but creamish.

T. bacillisporus can easily be distinguished from T. rotundus, the only other species of Talaromyces with globose, spinulose ascospores, by its different conidial state and the different size of the ascospores.

 

 

14. Talaromyces byssochlamydoides Stolk & Samson, spec. nov. - Fig. 14

 

Status conidialis: Paecilomyces byssochlamydoides Stolk & Samson, stat. nov.

 

Coloniae in agaro maltoso 40C repandae, primum ochraceae, deinde roseae. Coloniae in agaro Czapekii lentissime crescunt. Hyphae vegetativae hyalinae, leves, 1.5-5 (-6.5) m diametro. Ascomata rubella, globosa, 30-90 m diametro, plerumque discreta, non numquam confluentia, sex diebus maturantia. Involucrum sparsum, e hyphis laxis compositum, e quibus nonnullae hyphae tenues radiant. Asci subglobosi vel fere ellipsoidei, 9-12.5 x 6.5-7.5 m. Ascosporae globosae vel subglobosae, 3.7-4.5 x 3.5-4 m, leves.

Conidiophora hyalina vel dilute flava, ad 300 m longa, 3-5 m crassa, levia. Penicilli e 2 ad 8 metulis subirregulariter dispositis in apice conidiophororum, phialidibus verticillatis terminatis constant, omnino hyalini vel dilute flavi, leves. Metulae 11.5-15 x 2.5-3.5 m. Phialides 10-24 x 3-3.5 m, binae vel quaternae verticillatae vel singulae, e parte basilari cylindrica subito angustata et collo longo conspicuo, 4-7.5 x 1-1.5 m constant, apice plerumque inspissato. Conidia hyalina vel dilute flava, cylindrica vel ellipsoidea, 4-8 x 1-2.5 m, catenis divergentibus disposita. Chlamydosporae raro adsunt, globosae vel subglobosae, fere 4 m diametro.

Species thermophilica.

Typus: CBS 413.71, isolatus e terra sicca sub Pseudotsuga menziesii, Horse Rock Canyon, Oregon, USA, 1962.

 

Colonies on malt agar spreading rapidly at 40C, attaining a diameter of 9 cm within 7 days, composed of a thin layer of numerous ascomata, produced adjacent to the agar surface, intermixed and overgrown by a floccose to funiculose mycelium bearing conidial structures; when young showing ochraceous shades near Cinnamon-Buff, but with the ripening and subsequent colouring of the ascomata they develop more pinkish shades, ranging from Pinkish Cinnamon to Vinaceous-Cinnamon and Apricot Buff, becoming Vinaceous-Russet in age (Ridgway, Pls. 29, 14, 28; Rayner 17"b, 15"b, 13"b, 7"i). Odour not pronounced. After a few transfers the species has a tendency to become more conidial, appearing brownish near CreamBuff (Ridgway, Pl. 30; Rayner, 19"d). No exudate produced, Reverse of colonies yellow brown.

Colonies on Czapek agar growing very restrictedly, attaining a diameter of 2 cm within 7 days at 40C, thin, predominantly conidial, producing only a few ascomata; buff-coloured. Reverse colourless.

Colonies on oatmeal agar attaining a diameter of 9 cm within 7 days at 40C; structure as on malt agar, with colours ranging from buff to pinkish buff, becoming Pinkish Cinnamon in age, with the central areas more brownish near Wood Brown, due to the predominantly conidial state in these areas (Ridgway, Pls 29, 40; Rayner, 15"b-17"b, 17"'b). Exudate present, [p. 46] sometimes abundantly produced in large clear drops. Reverse brownish.

Vegetative hyphae hyaline, smooth-walled, 1.5-5 (-6.5) m in diameter.

Ascomata reddish, globose, 30-90 m in diameter, usually discrete, occasionally confluent, ripening within 6 days. Covering very scanty, consisting of loose-textured, branched hyphae, surrounded by an inconspicuous weft of thin, mainly radiating hyphae, about 1 m in diameter. Development of initials starts with the intercalary or terminal swelling and subsequent septation of hyphae. From these cells develop wide, somewhat irregularly coiling, gnarled branches, which continue to branch profusely, resulting in the formation of ascogenous hyphae. Simultaneously with the ascogenous hyphae, thinner, somewhat twisted hyphae develop forming later the ascomatal covering. Asci subglobose to slightly ellipsoidal, 9-12.5 x 6.5-7.5 m. Ascospores globose to subglobose, 3.7-4.5 x 3.5-4 m, thickwalled, smooth or nearly so, often partially covered by material which may represent the remnants of a gelatinous covering (Pl. 1.C).

Conidiophores arising primarily from submerged hyphae, but also as branches from aerial hyphae, hyaline to yellowish, septate, up to 300 m in length, 3-5 m in diameter, smooth-walled. Penicilli irregular, consisting of 2-8 metulae, somewhat irregularly disposed on the apical part of the conidiophore with whorls of phialides. All elements of the penicillus are hyaline to slightly yellowish and smooth-walled. Metulae 11.5-15 x 2.5-3.5 m. Phialides 10-24 x 3-3.5 m, in whorls of 2 to 4 or borne singly, consisting of a cylindrical basal portion, tapering abruptly to a long distinct neck, 4-7.5 m in length and 1-1.5 m wide, with the apical wall usually thickened. Conidia hyaline to pale yellow, usually cylindrical, 4-8 x 1-2.5 m, sometimes becoming ellipsoidal at maturity. Chlamydospores rarely produced, globose or subglobose, about 4 m in diameter.

The species is strongly thermophilic, minimum temperature about 25 C, optimum 40-45C, maximum slightly above 50C.

 

Material examined

 

CBS 413.71 = NRRL 3658 = type culture, isolated from a sample of dry soil under Douglas fir collected at Horse Rock Canyon, Oregon, USA, in 1962, sent to the CBS by D. I. Fennell, Peoria.

CBS 533.71, isolated by A. von Klopotek from piles of peat near Oldenburg, 1970.

The two strains are essentially alike, but the second one produces slightly larger ascomata than the type, while its conidial state is more conspicuous; most of the penicilli are larger, occasionally consisting of many metulae and phialides covering a larger portion of the conidiophore. Moreover the optimum temperature of CBS 533.71 (40-50C) is slightly higher than that of the type.

 

The ascomata of T. byssochlamydoides are similar to those of T. emersonii, but smaller, while their coverings are even more scanty. The ascospores of both species resemble one another closely, differing but slightly [p. 47] in shape and size. However, T. byssochlamydoides differs markedly from T. emersonii by its conspicuous Paecilomyces conidial state.

 

 

Fig. 14. Talaromyces byssochlamydoides, CBS 413.71. a. conidiogenous structures; b. conidia; c. initials; d. asci produced in chains; e. ascospores; f. loose-textured hyphae covering the ascoma; g. chlamydospores.

 

Because of the shape of the phialides and the divergent structure of its penicilli, the conidial state of T. byssochlamydoides belongs to Paecilomyces. The larger size of the penicilli and conidia are believed to relate it to P. crustaceus Apinis & Chesters (conidial state of Thermoascus crustaceus). It shows also relationship with the smaller-sized imperfect species P. [p. 48] variotii. P. byssochlamydoides is distinct from the latter species because of the Talaromyces perfect state and the cylindrical conidia. In P. variotii and P. crustaceus the conidia are usually ellipsoidal.

 

15. Talaromyces emersonii Stolk - Fig. 15

 

Talaromyces emersonii Stolk - Antonie van Leeuwenhoek 31: 262. 1965.

Byssochlamys sp. fide Cooney & Emerson - Thermophilic Fungi p. 155. 1964.

 

Misapplied name

Talaromyces dupontii (Griffon & Maublanc) Apinis sensu Apinis - Nova Hedwigia 5: 72. 1963 (as comb. nov.).

 

Status conidialis: Penicillium emersonii Stolk - Antonie van Leeuwenhoek 31: 262. 1965.

 

Colonies on malt agar growing rapidly, attaining a diameter of 9 cm within 7 days at 40C, mainly composed of a dense layer of numerous ascomata, produced near the agar surface; when young showing yellow shades near Colonial Buff and Deep Colonial Buff becoming reddish-brown in age (Ridgway, Pl. 30; Rayner, 21"d, 21"b), usually intermixed with and overgrown by a predominantly funiculose mycelium bearing conidial structures, giving especially the central areas a pale brown appearance, ranging from Dark-Olive Buff to Avellaneous (Ridgway, Pl. 40; Rayner, 21"'f, 15"'b-17"'b). Odour sweet aromatic. Reverse of colonies yellow brown, in some strains green.

Mature colonies on oatmeal agar show more pinkish colours, approximating to Onion-Skin Pink and Vinaceous-Tawny (Ridgway, Pl. 28; Rayner, 11'b, 11"b).

Vegetative hyphae hyaline, 1-3 m in diameter.

Ascomata reddish to orange brown, globose to subglobose, 50-300 m in diameter, often confluent, usually ripening within 7 days. Coverings very scanty, consisting of inconspicuous networks of hyphae, surrounded by loose wefts of radiating, branching, twisted, yellowish, encrusted hyphae, about 1 m in diameter. Initials consist of conspicuous swollen cells resembling chlamydospores. They are produced at the end of hyphae, sometimes occurring in small clusters. Each initial is commonly surrounded by a few thin branching hypha. No copulation between the swollen cells with the surrounding hyphae could be observed. At the base each initial produces an outgrowth which develops into a cluster of profusely branching, coiling ascogenous hyphae. The initial may remain visible as an empty cell for a short time. Asci subglobose to slightly ellipsoidal, 8-10.5 x 6.5-8 m. Ascospores thick-walled, smooth, subglobose to ovoidal, 3.5-4 x 2.73.5 m, often partially covered by material which may represent the remnants of a gelatinous coating (Pl. 1.D). [p. 49]

 

 

Fig. 15. Talaromyces emersonii, CBS 393.64. a. conidiogenous structure; b. foot-cell; c. conidia; d. initials; e. primordium consisting of profusely branching hyphae; f. fragment of ascomatal covering; g. asci in chains; h. ascospores.

 

Conidiophores arising primarily from submerged hyphae, but also as perpendicular branches from aerial hyphae or ropes of hyphae, pale yellow, septate, 35-150 m in length, 2.5-4 m in diameter, with coarsely roughened walls, rarely smooth-walled. Penicilli biverticillate-asymmetrical, typically consisting of a to 2 rami in addition to the main axis, metulae in verticils of 3 to 5, each bearing whorls of 5-10 phialides. All elements of the penicillus hyaline to slightly yellowish, appressed, usually rough-walled, rarely smooth, with both rami and metulae swollen at their apices. Rami 12-30 x 2.5-3 m, sometimes lacking. Metulae 10-17 x 2.5-3.5 m. Phialides parallel, 8.5-10 x 2-2.5 m, consisting of a cylindrical base, tapering abruptly to a short conidium-bearing tip, 1-2 m in length. Conidia smooth, hyaline to brownish, cylindrical when young, sometimes swelling at maturity and becoming ellipsoidal, usually 3.5-5 x 1.5-2.7 m, [p. 50] rarely up to 10 m in length, produced in long chains, up to 100 m in length, usually forming columns.

The species is strongly thermophilic, minimum temperature somewhat below 30C, optimum 40-45 C, maximum 55 C.

 

Material examined

 

CBS 393.64, type culture isolated by Mrs. A. J. van der Plaats-Niterink, from compost from Italy, 1961.

CBS 394.64, isolated by R. Emerson from compost, Berkeley, California, 1962 (see Cooney & Emerson, 1964: p. 155).

CBS 395.64 = IMI 96474, isolated by H. O. W. Eggins from heaps of oilpalm kernels from Port Harcourt, Nigeria, 1963 (cf. Eggins and Coursey, 1964).

CBS 396.64, isolated by G. A. de Vries from soil collected near Baarn, 1964.

CBS 397.64 = BDUN 272, isolated by A. E. Apinis from soil near Nottingham, G.B., and described by him as Talaromyces dupontii (Griffon & Maublanc) Apinis.

CBS 180.68, isolated by H. C. Evans from Betula litter on mine tips at Silverdane, G.B., June, 1967 (Evans, 1971). This strain consists of a floccose to funiculose yellow aerial mycelium obscuring the ascomata, which are not only produced on the agar surface, but also occur in limited numbers in the aerial mycelium. Penicilli occur only scantily and do not affect the colony appearance. Originally the reverse of the colonies showed green colours as in T. bacillisporus. However, after a few transfers this green colour has been lost and the reverse is now yellow-brown as in the other strains of this species.

CBS 814.70, isolated by R. A. Hill from sugar cane begasse, Reading, G.B., 1964; in colony appearance identical with CBS 80.68, but the colour of the aerial mycelium is more intensely bright yellow.

CBS 266.71 = NRRL 5118, isolated by T. Nilsson from wood chips in Sweden, 1970, and sent to the CBS by D. I. Fennell, Peoria, 1971.

CBS 698.71, isolated by A. von Klopotek from piles of peat near Oldenburg, 1970.

CBS 759.71, isolated by A. Ofosu-Asiedu, from spruce-pine wood pile, Vancouver and sent to the CBS by R. S. Smith as Byssochlamys emersonii (Stolk) Apinis (det. A. E. Apinis, Nottingham).

 

The different strains of T. emersonii vary in some minor features, such as number of penicilli, roughness of the walls of conidiophores and penicilli and amount of aerial mycelium. Occasionally strains occur, which develop a conspicuous floccose to funiculose bright yellow mycelium. Though the colour of the reverse is usually yellow-brown, rarely a green reverse as in T. bacillisporus has been observed (e.g. CBS 180.68). Ascomata, ascospores and structures of the penicilli are in all strains identical.

T. emersonii is closely related to the new species T. byssochlamydoides. These species can be separated from one another by the quite different conidial states and the shape and size of the ascospores.

As in T. bacillisporus, the conidial state belongs to the P. cylindrosporum-series. It differs from that of T. bacillisporus, however, in producing larger and more complicated penicilli, while the walls of conidiophores and penicilli are usually more conspicuously roughened.

T. emersonii appears to be an ubiquitous and relatively common fungus. [p. 50]

 

 

16. Talaromyces leycettanus Evans & Stolk - Fig. 16

 

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

 

Status conidialis: Paecilomyces leycettanus (Evans & Stolk) Stolk & al. - Persoonia 6: 342. 1971.

 

Penicillium leycettanum Evans & Stolk - Trans. Br. mycol. Soc. 56: 45. 1971.

 

Colonies on malt agar spreading rapidly at 40C, attaining a diameter of about 7 cm within 7 days; largely composed of a thin layer of ascomata, showing greenish yellow shades near Sea-foam Green (Ridgway, Pl. 31; Rayner, 25"f-27"f), usually intermixed with and overgrown by a somewhat floccose to funiculose mycelium, bearing abundant penicilli, giving especially the central parts a more brownish colour near Avellaneous (Ridgway, Pl. 40; Rayner, 15"'b-17"'b); central areas slightly raised and somewhat wrinkled. Reverse orange brown approximating Kaiser Brown (Ridgway, Pl. 14; Rayner, 9'k) with the surrounding agar coloured in lighter tints of the same shade.

Vegetative hyphae ranging from hyaline to yellow or brown, smooth, 1-3 m in diameter.

Ascomata greenish-yellow, globose, 50-175 m in diameter, discrete, sometimes confluent, ripening within 7 days. Coverings consisting of thin hyphal networks (at the base of the ascomata brown) surrounded by radiating, twisted, branching, pale yellow, often encrusted hyphae, 1-2.5 m in diameter. Initials consisting of coiled hyphae, borne as branches from vegetative hyphae inside hyphal tufts. After having formed septa, branches arise, which develop into the strongly branching, coiled ascogenous hyphae. Asci subglobose to slightly ellipsoidal, 7.5-9 x 5.5-6.5 m. Ascospores ellipsoidal, 3.2-4 (-4.5) x 2.2-3 m, ornamented with 2-6 thin, somewhat jagged, irregular, for the greater part longitudinal ridges of very different length, about 0.2-1 m wide (Pl.2.D).

Conidiophores smooth, rarely encrusted with brown granules, septate, usually arising from submerged hyphae, but also borne as branches of aerial hyphae, 80-350 m in length and 3-4 m in diameter. Penicilli often complicated, irregular, usually consisting of 1 to 3 verticils of 2 to 4 metulae with whorls of 2 to 6 phialides; occasionally monoverticillate. All elements of the penicillus hyaline to yellowish, smooth. Rami occasionally present, 10-15 x 2.5-3.5 m. Metulae 8-12 x 1.5-2.5 m. Phialides 10-15 x 1.5-2.5 m, consisting of a cylindrical basal portion, tapering abruptly to a thin, long neck, 4-5 m in length and about 1 m wide, with the wall of the apex sometimes thickened. Conidia yellowish, yellow-brown in mass, cylindrical, 4-6 x 1-1.5 m, smooth, in dry divergent, sometimes tangled chains. Chlamydospores thick-walled, smooth, globose or nearly so, yellowish, 4-4.5 m in diameter.

The species is thermotolerant to thermophilic, minimum temperature 18 C, optimum about 40C, maximum 55 C. [p. 52]

 

 

Fig. 16. Talaromyces leycettanus, CBS 398.68. a. complicated conidiogenous structure; b. monoverticillate penicillus; c. conidia; d. initials; e. development of asci; f. chains of asci; g. ascospores; h. fragment of ascomatal covering; i. chlamydospores.

 

Material examined

 

CBS 398.68 = type culture, CBS 275.70, CBS 276.70, all isolated by H.C. Evans from Leycett coal spoil tips in Staffordshire, G.B.

 

The ascospores of T. leycettanus and T. thermophilus are almost identical. The two species can easily be separated by their different ascomata, those of T. leycettanus being small, greenish yellow with thin coverings, whereas those of T. thermophilus are very large, creamish and thick-walled. The ascomata of T. leycettanus develop very well on various agar media, those of T. thermophilus are formed only on sterilized oats and grass etc. In addition, their imperfect states are quite different.

The conidial state of T. leycettanus was described by Evans and Stolk (1971) as a Penicillium. They suggested a relationship with the P. janthinellum-series [p. 53] on account of the phialide shape. However, a reexamination of this species proved the phialides with very long necks to be more of the Paecilomyces than of the Penicillium-type. Moreover, chlamydospores, as occurring in some species of the Paecilomyces variotii-group are present and the conidial state shows a brown colour. For these reasons we have transferred this species to Paecilomyces (Stolk and Samson, 1971).

On the other hand, the penicilli are less divergent than in a typical Paecilomyces, resembling more those of a Penicillium, though they may be somewhat more irregularly branched. Therefore Paecilomyces leycettanus seems to represent an intermediate form between Penicillium and Paecilomyces, but it is better classified in Paecilomyces.

P. leycettanus shows affinities to P. variotii Bain., e.g. in colour and growth. It is also very close to P. fulvus Stolk & Samson (conidial state of Byssochlamys fulva), but it is distinct from this species by its Talaromyces perfect state, the presence of chlamydospores and its cylindrical conidia with rounded ends. In P. fulvus the conidia are larger and cylindrical with both ends flattened.

 

 

Talaromyces section Thermophila Stolk & Samson, sect. nov.

 

Ascomata 45C in granis avenae neque in substratis agarosis formantur, pariete crasso e dense intricatis hyphis composito circumdata peridii veri simili. Status conidialis ad Penicillium pertinet. Species thermophilae.

Species typica: Talaromyces thermophilus Stolk.

 

Ascomata at 45C usually not produced on agar media, but abundantly on sterilized oat grains, walls thick, consisting of closely knit hyphae simulating a true peridium. Imperfect state Penicillium. Thermophilic.

 

The sect. Thermophila differs from the other sections of Talaromyces primarily in not or very rarely producing ascomata on agar media. The ascomata have well-developed, thick, parenchymatous walls. Asci are not produced in helicoidal chains as occurring in most species of Talaromyces, but in rather straight and branched chains. In addition the conidial state of the only species is classified in the Asymmetrica-Divaricata, near the P. janthinellum-series, thus being different from the imperfect states of all other sections.

 

17. Talaromyces thermophilus Stolk - Fig. 17

 

Talaromyces thermophilus Stolk - Antonie van Leeuwenhoek 31: 268. 1965. [p. 54]

Penicillium dupontii Griffon & Maublanc emend. Emerson in Raper & Thom, The Penicillia, p. 573. 1949 = Talaromyces dupontii (Griffon & Maublanc) Emerson, incidententally mentioned by Fergus - Mycologia 56: 277. 1964.

 

Status conidialis: Penicillium dupontii Griffon & Maublanc - Bull. trimest. Soc. mycol. Fr. 27: 73. 1911.

 

? Citromyces sphagnicola Mal'chevskaya - Trudy Pushkin sel'skokhoz. Inst. 13: 23. 1939, fide Cooney & Emerson - Thermophilic Fungi, p. 28. 1964.

 

Colonies on malt agar attaining a diameter of 5 to 7 cm within 7 days at 45C; floccose, occasionally slightly funiculose, fairly deep, especially in central areas up to 2 to 3 mm; consisting of loosely interwoven and trailing hyphae bearing numerous penicilli; grey green near Pea Green when young, becoming brown in age, approximating Wood Brown (Ridgway, Pls 47, 40; Rayner, 29"'b, 17"'b); sterile sectors white or pinkish. Ascomata not produced. Reverse brownish.

Colonies on sterilized oat grains at 45C at first grey-green because of conidial development, later producing deep overgrowths, and at maturity consisting of a thick, tough, close-textured felt of hyphae, showing white, pinkish, greyish and brownish colours, bearing large creamish ascomata, occurring scattered or in clusters.

Growth on Czapek agar at 45 C poor.

Vegetative hyphae hyaline, 1.5-3 m in diameter with numerous conspicuous inflations, up to 8 m wide.

Ascomata usually discrete, globose or nearly so, 400-1300 m in diameter, covered by a thick wall composed of many layers of rather closely knit hyphae, becoming pseudoparenchymatous at maturity. Initials consisting of irregularly swollen cells, often chlamydospore-like, producing one or two outgrowths; these develop into wide, coiling, branching, at first aseptate later septate hyphae, thus forming a compact mass of ascogenous hyphae. No copulation has been observed. Asci in straight to slightly curved and branched chains, 8-spored, globose to subglobose, 7.5-9 m in diameter. Ascospores ellipsoidal, 3.5-4.5 x 2.2-3.5 m, ornamented by 2 to 6 somewhat jagged, irregular, usually longitudinal ridges, about 0,2 m wide (Pl. 2.H-I).

Conidiophores arising as short, more or less perpendicular branches from aerial hyphae, smooth, 3-30 x 1.5-2.5 m. Penicilli irregular, ranging from monoverticillate to biverticillate, typically consisting of a verticil of 2 to 4 metulae, with whorls of 2 to 4 phialides; occasionally one or two rami are present; most elements somewhat divergently arranged and smoothwalled. Rami and metulae 5-10 x 1.5-2 m. Apices of conidiophores, rami and metulae slightly inflated in young penicilli, often becoming conspicuously swollen when old. Phialides 7-10 x 1.5-2 m, usually consisting of a cylindrical basal portion which tapers abruptly to a distinct neck about 1.5 m in length. Conidia smooth, ellipsoidal, 2.5-4.5 x 1.5-2.5 m, forming short divergent chains. Chlamydospores occurring abundantly infertile [p. 55] areas of cultures on oat grains, only in limited numbers on agar cultures, thick-walled, globose or nearly so, 4.5-6.5 m in diameter.

The species is strongly thermophilic, minimum temperature somewhat below 30C, optimum about 45C, maximum about 57C.

 

 

Fig. 17. Talaromyces thermophilus, CBS 236.58. a. conidiogenous structures; b. conidia; c. initials; d. primordium consisting of profusely branching hyphae; e. chains of asci; f. ascospores; g. fragment of ascomatal covering; h. chlamydospores; i. vegetative hyphae, showing inflations. [p. 56]

 

Material examined

 

CBS 236.58 = NRRL 2155 = type culture, isolated by R. Emerson from retted Parthenium argentatum in California, 1945.

CBS 292.61, isolated at the 'Proefstation voor de Champignoncultuur', Horst (L.), the Netherlands, from composted straw, 1961.

CBS 889.70 and 890.70, isolated by R. A. Hill, from hay, Reading, G.B., 1970.

CBS 161.71, isolated by M. Luykx from soil of Zuid-Flevoland, the Netherlands, 1970.

CBS 116.72, isolated by M. C. Papendorf from compost consisting mainly of leaves of Celtis africana, Potchefstroom, S-Africa, 1971.

 

Griffon and Maublanc (1911) isolated the conidial state of T. thermophilus from manure and damp hay and described it as Penicillium dupontii. Emerson found it again on retted guayule shrub in 1945. He obtained also the perfect state and published an emended description of P. dupontii (Raper and Thom, 1949, P. 573). Fergus (1964) mentioned the name Talaromyces dupontii (Griffon & Maublanc) Emerson, but Emerson never published this new combination. The conidial state of the fungus designated by Apinis (1963) as 'Talaromyces dupontii (Griffon & Maublanc) Apinis' is not identical with the fungus described by Griffon & Maublanc as P. dupontii (Stolk, 1965).

Raper and Thom (1949 P. 567) classified P. dupontii in the Biverticillata Symmetrica section, because they considered the phialides to be lanceolate or acuminate. The penicilli are described as `often irregular and asymmetric, but typically biverticillate and sometimes symmetrical'. However, the phialides are characterized by distinct necks as occurring in the P. janthinellum-series. We prefer to place P. dupontii in the Asymmetrica-Divaricata on the basis of the phialide shape and the divergent irregular penicilli. It is not assigned to any of its series, but it is considered to be closely related to the P. janthinellum-series. Because of the distinct conidium-bearing necks of the phialides, the presence of chlamydospores and the brown colour of the old conidial areas, the fungus shows also relationships with Paecilomyces. However, the necks of the phialides are short as compared with those of Paecilomyces and the colour of the young colonies is definitely green. Moreover, the structure of the penicilli agrees bettet with Penicillium than with Paecilomyces.

The ascospores of T. thermophilus resemble those of T. leycettanus. As mentioned under T. leycettanus, the two species can easily be separated by their ascomata and penicilli.

 

 

Talaromyces section Purpurea Stolk & Samson, sect. nov.

 

Involucrum ascomatum conspicuum, e reticulo tenui laxo hypharum flavarum compositum [p. 57]. Ascosporae flavae, deinde rubrae tinctae pigmento diffundente. Status conidialis ad Penicillium restrictum-seriem pertinet.

Species typica: Talaromyces purpureus (Mller & Pacha-Aue) Stolk & Samson.

 

Covering of ascomata distinct, consisting of a thin, loose-textured network of hyphae, yellow. Ascospores yellow, becoming reddish from diffusing red pigment. Conidial state Penicillium restrictum-series.

 

 

18. Talaromyces purpureus (Mller & Pacha-Aue) Stolk & Samson, comb. nov. - Fig. 18

 

Arachniotus purpureus Mller & Pacha-Aue - Nova Hedwigia 15: 552, 1968 (basionym).

 

Status conidialis: Penicillium purpureum Stolk & Samson, stat. nov.

 

Scopulariopsis state fide Mller & Pacha-Aue (1968).

 

Coloniae dilute griseae vel olivaceo-brunneae ascomatibus flavis satae. Conidiophora e hyphis aeriis oriuntur, brevia, 5-25 x 1-1.7 m, pariete levi vel ruguloso, apice inflato ad 3.5-4 m. Penicilli plerumque monoverticillati, nonnumquam unum, rarius duos ramos quasi metulas proferunt; phialidibus divergenter verticillatis terminati. Rami (metulae) 5-8 x 1-2 m, sursum inflati, leves. Phialides binae ad duodecenae verticillatae, nonnumquam singulae, utrinque angustatae, sursum in collum angustum constrictae, 5.5-8 x 2-2.7 m. Conidia subglobosa, ovoidea vel fere ellipsoidea, 3.2-4 x 2.5-3 m, olivaceobrunnea, maturitate crassitunicata, cristis transversalibus, saepe spiralibus, conspicuis obtecta.

Status ascigerus Talaromyces purpureus (Mller & Pacha-Aue) Stolk & Samson. Typus: CBS 475.71, isolatus e terra prope Esterel in Gallia.

 

Colonies on malt agar growing somewhat restrictedly, attaining a diameter of 4 cm within 2 weeks at 30C, composed of a submerged, compact basal mycelium, forming a tough pellicle covered with a very scanty aerial mycelium, which may show a pinkish colour near Light Brownish Vinaceous (Ridgway Pl. 39; Rayner 5"'d), slightly zonate, at first appearing largely mycelial, but after about 10 days ascomata develop on the agar surface, usually forming a continuous layer, giving mature colonies a conspicuous yellow colour, ranging from Citron Yellow to Chalcedony Yellow (Ridgway Pls 16, 17; Rayner 23'b, 25'b), sometimes occurring only in sectors; conidial state greyish, olive brown in age, often not affecting the colony appearance. Reverse of colonies purplish red, approximating Oxblood Red and Maroon (Ridgway Pl. 1; Rayner 1k, 3m) with the surrounding agar usually coloured throughout in lighter tints of the same shades.

Ascoma production is most abundant on malt agar at 30C, but even on that medium ascomata may be scarce. They are also produced in limited numbers on oatmeal and cornmeal agar. On hay-infusion agar only the [p. 58] conidial state occurs, especially at lower temperatures, colouring the surface of the colonies greyish, while the red pigment is lacking.

Vegetative hyphae hyaline, occasionally red because of diffusing red pigment, 1-3 m in diameter.

Ascomata yellow, globose to subglobose, small, 20-60 m in diameter, often confluent, ripening within 3 weeks. Coverings consisting of a thin, loosely interwoven network of yellowish hyphae with irregular swellings up to 2.5 m in diameter, surrounded by a loose weft of somewhat twisted, very thin, mainly radiating hyphae, about 0.5-1.5 m in diameter. Initials consisting of coiled, aseptate hyphae, borne as branches from vegetative hyphae, soon becoming septate and producing short, septate branches, with most cells developing into asci. Asci borne in short chains, 4-6 spored, globose to subglobose, 9-12 m in diameter, usually occurring in small numbers. Ascospores yellow, often becoming reddish in age, thickwalled, spinulose over their entire surface, ellipsoidal, often with one side flattened, 6.5-8 x 4.5-5.5 m.

Conidiophores borne as short branches from aerial hyphae, usually 5-25 x 1-1.7 m, with walls smooth or slightly roughened and apices swollen to about 3.5-4 m. Penicilli small, mostly monoverticillate but occasionally developing one (rarely two at the same level) divergent metula-like rami in addition to the main axis, each of them bearing divergent clusters of about 2-12 phialides; solitary phialides may be borne directly on the vegetative hyphae. Rami (metulae) 5-8 x 1-2 m, inflated at the apex to about 3.5 m, smooth-walled. Phialides 5.5-8 x 2-2.7 m, covering the sometimes irregularly swollen apices and occasionally also the subterminal portions of conidiophores and branches, showing the P. restrictum type, narrowed at both ends, with thin conidium-bearing tips of about 1.5 m in length. Conidia subglobose, ovoidal or slightly ellipsoidal, 3.2-4 x 2.5-3 m, hyaline and smooth when young, becoming olive-brown, thick-walled when mature, showing conspicuous transverse bands (often spirally-arranged) conidial chains divergent to somewhat tangled, up to 200 m in length, with small hyaline connectives linking the conidia.

Minimum temperature 10C, optimum temperature 30C, maximum temperature about 37C.

 

Material examined

 

CBS 475.71, type culture, isolated by Ch. Stoll from soil near Esterel, France 1965. [p. 59]

 

 

Fig. 18. Talaromyces purpureus, CBS 475.71. a. conidiogenous structures; b. conidia; c. initial surrounded by thin hyphae; d. initials; e. asci produced in chains; f. ascospores; g. loosely branched hyphae covering the ascoma.

 

Mller and Pacha-Aue (1968) described this fungus as a species of Arachniotus because they considered the conidial state to belong to the genus Scopulariopsis. However, an examination of the type culture proved that no annellated zone of the conidiogenous cells is present, but that the conidia are produced on phialides arranged in the manner characteristic of Penicillium. Moreover asci are not borne singly as indicated by Mller and Pacha-Aue (1968) but in short chains as occurring in Talaromyces. For these reasons the species is transferred to the genus Talaromyces. It differs markedly from the other species of Talaromyces by its conidial state, which is classified in the P. restrictum-series, because of the predominantly monoverticillate penicilli, the shape of the phialides and the olive-brown colour of the conidia. Conidia showing the same ornamentation occur only in P. daleae Zaleski. According to Raper and Thom (1949) the latter species belongs to the P. janthinellum-series, but in our opinion (Stolk and Malla, 1971) P. daleae can much better be placed in the P. nigricans-series, which is very closely related to the P. restrictum-series; the two series differ mainly the structure of the penicilli, those of the P. nigricans-series being much more complicated than the small, predominantly monoverticillate penicillate structures of the P. restrictum-series. Both have olive-brown conidia and the same type of phialides. In none of the described species of Talaromyces [p. 60] does an imperfect state belonging to the P. restrictum-series occur. T. purpureus differs also from most species of Talaromyces in producing ascospores which become red in age. Since the ascospores are at first yellow just as in the other species of Talaromyces, it is assumed that the red discolouration is brought about by diffusing red pigment abundantly produced in the agar. The same phenomenon is seen in strains of T. flavus with a dark red reverse (e.g. CBS 261.55).

For these reasons, but especially because of the structure of the conidial state, T. purpureus is considered to represent a separate section within the genus Talaromyces.

 

 

Doubtful and excluded species

 

Penicillium aureum van Tieghem - Bull. bot. Fr. 24: 158. 1877 [non Penicillium aureum Corda - Prachtflora p. 37-38. 1839].

According to the description this species has golden yellow ascomata. The initials described consist of two identical hyphae coiling twice around one another. This may point towards T. stipitatus, but the oval, smooth ascospores, measuring 5 x 3 m, are quite different from this species. The conidial state is a Penicillium.

 

Talaromyces avellaneus (Thom & Turesson) C. R. Benjamin - Mycologia 47: 682. 1955 - Hamigera avellanea Stolk & Samson - Persoonia 6: 345. 1971.

 

Talaromyces cejpii Mil'ko - Nov. Sist. niz. Rast. 1964: 208. 1964.

This species does not belong to Talaromyces because of the structure of its peridium, consisting of a single layer of thin-walled, hyaline, polygonal cells. The conidial state is represented by a Polypaecilum. Scott (1970) transferred this species to Dichotomomyces.

 

Penicillium petchii Sartory & Bain. - Annls mycol. 11: 272. 1913.

Penicillium petchii is characterized by yellow ascomata, 150-200 m in diameter and by symmetrical penicilli. The asci, 12-13 m in diameter, contain 6 spinulose ascospores, measuring 6 m. The initials figured do not agree with any of the recognized species of Talaromyces. The ascospores approximate to those of T. flavus and T. wortmannii, but the structures of the initials and of the ascomatal wall do not agree with any of these species.

Penicillium sacchari Ray - Revue gen. Bot. 9: 298. 1897.

Because of the small size of the ascospores (3-2.5 m), Raper and Thom (1949) mentioned this species as a possible synonym of T. helicus. However, [p. 61] according to Ray (1897), the conidia of P. sacchari measured 2 x 1 m, being much smaller than those of T. helicus or any other species of Talaromyces. Therefore the microscopical measurements given by Ray (1897) might be inaccurate. Ray's description and his figures 23 to 27 demonstrate that P. sacchari must have represented a species of Talaromyces, but his description is too scanty for an accurate identification.

 

Talaromyces striatus (Raper & Fennell) C. R. Benjamin - Mycologia 47: 682. 1955. = Hamigera striata Stolk & Samson - Persoonia 6:347. 1971.

 

 

Key to the species based on ascospore characters

 

1a.

Ascospores ellipsoidal, provided with ridges

2

1b.

Ascospores without ridges

7

 

 

 

2a.

Ascospores with transverse or spiral ridges

3

2b.

Ridges of ascospores not transverse

4

 

 

 

3a.

Ascospores with 4-6 irregularly transverse ridges, 4.5-5.5 x 3-3.7 m

T. luteus

3b.

Ascospores with 3-5 regular, nearly parallel, transverse ridges, often spirally arranged, 3.5-4.5 x 2.2-3 m

T. udagawae

 

 

 

4a.

Ascospores with a single equatorial ridge, 3.2-4 (-4.5) x 1.7-2.2 (-2.5) m; ascomata yellow, occasionally salmon

T. stipitatus

4b.

Ascospores with 2 to 6 irregular, for the greater part longitudinal ridges

5

 

 

 

5a.

Ascospores 2.5-3.5 x 2-2.4 m; ascomata yellow; mesophilic species

T. ucrainicus

5b.

Ascospores larger; ascomata creamish or greenish yellow, thermo-tolerant of thermophilic species

6

 

 

 

6a

Ascomata large, creamish, with thick walls, mainly produced on oat grains, not or sparsely on agar media; ascospores 3.5-4.5 x 2.2-3.5 m; penicilli divaricate

T. thermophilus

6b.

Ascomata small, greenish yellow, abundantly produced on agar media; coverings thin; ascospores 3.2-4 (-4.5) x 2.2-3 m; conidial state Paecilomyces

T. leycettanus

 

 

 

7a.

Ascospores smooth or nearly so, never showing spines

8

7b.

Ascospores ornamented

10

 

 

 

8a.

Ascomata yellow, becoming pinkish on some media; ascospores ellipsoidal, 3-4.5 x 2-2.7 m, conidial state usually not affecting the colony appearance; penicilli of the symmetrical type but usually fractional; ascogonia coiled terminally; mesophilic species

T. helicus var. major

8b.

Ascomata orange, brown or pink; conidial states brown; thermophilic species [p. 62]

9

 

 

 

9a.

Ascomata orange brown; ascospores smooth, 3.5-4 x 2.7-3.5 m; conidial state belonging to the P. cylindrosporum-series; conidia 3.55 x 1.5-2.7 m

T. emersonii

9b.

Ascomata pink to reddish; ascospores smooth, 3.7-4.5 x 3.5-4 m; conidial state Paecilomyces; conidia 4-8 x 1-2.5 m

T. byssochlamydoides

 

 

 

10a.

Ascospores ornamented with a few very delicate spines, most of them occurring apically, very small ascospores appearing nearly smooth; 2.5-3 (-3.7) x 1.5-2.2 m; ascogonia coiled terminally; ascomata yellow

T. helicus var. helicus

10b.

Ascospores usually conspicuously spinulose

11

 

 

 

11a.

Ascospores globose

12

11b.

Ascospores subglobose or ellipsoidal

13

 

 

 

12a.

Ascomata orange; ascospores 4.5-5.2 m; conidial state green, biverticillate-symmetrical; penicilli often fractional

T. rotundus

12b.

Ascomata pale yellow; ascospores 3.5-4.5 m; conidial state belonging to the P. cylindrosporum-series, creamish

T. bacillisporus

 

 

 

13a.

Ascomata white, creamish or pinkish

14

13b.

Ascomata usually yellow, occasionally yellow orange, buff or pinkish to purple red

15

 

 

 

14a.

Ascomata white to creamish; ascospores 3-3.5 x 2-2.5 m

T. trachyspermus

14b.

Ascomata creamish to pinkish, ascospores 5-6 x 4-4.5 m

T. intermedius

 

 

 

15a.

Ascospores ellipsoidal, usually with one side flattened, 6.5-8 x 4.55-5 m, ascomata yellow; conidial state mostly monoverticillate (P. restrictum-series); conidia olive-brown, showing transverse ridges, 3.24 x 2.5-3 m

T. purpureus

15b.

Ascospores regularly ellipsoidal; conidial state typically biverticillate-symmetrical; penicilli often fractional, conidia without transverse ridges

16

 

 

 

16a.

Ascomata yellow to orange; initials consisting of thick, irregularly septate hyphae, producing usually coiled branches; ascospores 3.74.7 (-5) x 2.5-3 m; colonies restricted

T. wortmannii

16b.

Ascomata usually yellow, occasionally yellow orange, buff or pinkish to purple red; initials consisting of vermiform ascogonia around which thin antheridia are tightly coiled; colonies usually spreading broadly

17

 

 

 

17a.

Ascospores ranging within the sizes of 3-5 x 2.2-3.5 m

T. flavus var. flavus

17b.

Ascospores ranging within the sizes of 5-6.5 x 3.5-5.2 m

T. flavus var. macrosporus

[p. 63]

 

 

Acknowledgements

 

The authors are very grateful to all those, who contributed to this paper by providing living cultures and herbarium material. We wish to thank Dr J. A. von Arx for his encouragement and Dr M. A. Donk for his valuable advices on problems of nomenclature. Moreover we are much indebted to Dr W. Gams for his helpful suggestions and cooperation and to Dr K. M. Old for correcting the English text. Finally Mr S. Henstra and Mr F. Thiel of the Technical and Physical Engineering Research Service (Wageningen) are acknowledged for the scanning electron micrographs.

 

 

References

 

Apinis, A. E., 1963 - Occurrence of thermophilous microfungi in certain alluvial soils near Nottingham. - Nova Hedwigia 5: 57-78.

Apinis, A. E., 1964 - Revision of British Gymnoascaceae. - Mycol. Pap. 96: 1-56.

Apinis, A. E., 1967 - Dactylomyces and Thermoascus. - Trans. Br. mycol. Soc. 50: 573-582.

Arx, J. A. von, 1970 - The genera of fungi sporulating in pure culture. - J. Cramer, Lehre.

Arx, J. A. von, 1971 - On Arachniotus and related genera of the Gymnoascaceae. - Persoonia 6:371-380.

Benjamin, C. R., 1955 - Ascocarps of Aspergillus and Penicilliium. - Mycologia 47: 669-687.

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Explanation of plates 1 and 2

 

Scanning electron micrographs of ascospores.

 

Plate I. A. T. flavus var. flavus, CBS 310.38 (T), x 5000; B. T. trachyspermus, CBS 373.48, x 4000; C. T. byssochlamydoides, CBS 413.71 (T), x4000; D. T. emersonii, CBS 393.64 (T), x 4000; E. T. stipitatus, CBS 375.48 (T), x 4000; F. - G. T. udagawae, CBS 579.72 (T), F. x 4000, G. x 5000; H. T. luteus, CBS 348.51 (T), x 5000.

 

Plate 2. A. T. helicus var. helicus, CBS 134.67, x 4000; B. T. helicus var. helicus, CBS 550.72 B, x 4000; C. T. helicus var. major, CBS 562.72, x 4000; D. T. leycettanus, CBS 398.68 (T), x 5000; E. - G. T. ucrainicus, CBS 127.64, x 4000; H. - I. T. thermophilus, CBS 236.48 (T), x 4000.