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Anamorphs of the Bolbitiaceae (Basidiomycota, Agaricales)

     Friedrich-Schiller-Universität Jena, Dornburger Str. 159, D-07743 Jena, Germany

Michael Weiß

     Eberhard-Karls-Universität Tübingen, Botanisches Institut, Lehrstuhl für Spezielle Botanik und Mykologie, Auf der Morgenstelle 1, D-72076 Tübingen, Germany

	ABSTRACT

TOP ABSTRACT INTRODUCTION MATERIAL AND METHODS RESULTS DISCUSSION LITERATURE CITED

 

We describe and illustrate thallic conidiogenesis in 14 species of the Bolbitiaceae sensu Singer studied in culture. Conidiogenesis of 12 species is shown for the first time. Bolbitius vitellinus and the investigated species of Conocybe (C. albipes, C. appendiculata, C. magnicapitata, C. semiglobata, C. subovalis, C. subpubescens, C. sulcatipes and C. teneroides) possessed a similar mode of conidiogenesis. Species of both genera formed mostly coiled conidiogenous hyphae arising sympodially from differentiated conidiophores. The anamorphs of the Agrocybe species were not uniform and predominantly differed from those of Conocybe and Bolbitius. The conidia of Agrocybe dura, A. firma and A. praecox developed by the simple fragmentation of normally branched hyphae. Sympodially proliferating conidiophores occurred in Agrocybe arvalis and A. aegerita. Secretory cells of different size and shape were found in Agrocybe and in Conocybe. Our results corroborate a close phylogenetic relationship between Bolbitius and Conocybe as well as the polyphyly of the Bolbitiaceae as currently treated, which is consistent with recent molecular phylogenetic studies. Consequently we emend the family concept based on anamorphic characters.

Key words: Agrocybe, basidiomycetes, Bolbitius, conidia, conidiogenesis, Conocybe, culture characters, secretory cells

	INTRODUCTION

TOP ABSTRACT INTRODUCTION MATERIAL AND METHODS RESULTS DISCUSSION LITERATURE CITED

 
Anamorphs of the Bolbitiaceae sensu Singer (1986) have been documented only sparsely in the literature. A comprehensive description exists only for the anamorph of Agrocybe pediades (Vandendries 1937). Watling (1976) mentioned that conidia occur in cultures of many species of the Bolbitiaceae and illustrated the development of conidiophores in Conocybe farinacea Watling, C. percincta P.D. Orton and Agrocybe pediades (as A. semiorbicularis). In addition a few papers dealing with anamorphs of the Agaricales or Homobasidiomycetes include details about one or two species of Conocybe and Agrocybe anamorphs (Brefeld 1889, Oddoux 1955, Lamoure 1960, Galland 1968, Watling 1979).

In Walther et al (2005) we gave a survey of the modes of conidiogenesis in Agaricales and discussed the systematic relevance of anamorphs in this order. With respect to the Bolbitiaceae we found that anamorphic characters are only in part consistent with Singer’s classification of this family (Singer 1986), comprising the genera Agrocybe, Bolbitius, Conocybe, Descolea, Galerella and Pholiotina. A similar mode of conidiogenesis strongly suggests the monophyly of a clade including Bolbitius and Conocybe (including Pholiotina), while most Agrocybe species studied formed a deviating anamorph.

These results are consistent with recent molecular phylogenetic analyses (Moncalvo et al 2002, Walther et al 2005), which support the monophyly of a core group of the Bolbitiaceae consisting of Bolbitius and Conocybe, probably extended by Descolea (Moncalvo et al 2002, Gulden et al 2005) and indicate that Agrocybe might be distant to this Bolbitiaceae core group (Moncalvo et al 2002, Walther et al 2005, Gulden et al 2005).

In the present paper anamorphs of 14 species of the Bolbitiaceae belonging to Agrocybe, Bolbitius and Conocybe (including Pholiotina) are described in detail. The infrageneric sampling depended on the ability of haploid mycelia to grow in culture. The genera Galerella and Descolea were not studied because they do not occur in our sampling area. The majority of strains described and illustrated in this paper are included in the molecular phylogenetic analysis of Walther et al (2005).

	MATERIAL AND METHODS

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Morphological study.— – The strains were isolated from basidiospore deposits of freshly collected basidiocarps and cultivated on malt-extract agar (2.5%) in Petri dishes at room temperature. To obtain single-spore cultures, several dilutions of basidiospore suspensions were plated on malt-extract agar (2.5%) in Petri dishes. The outgrowing mycelia were isolated. The cultures were examined macroscopically and microscopically for the first time when they had reached about 3–4 cm diam. At least eight single-spore mycelia and one dikaryotic mycelium isolated from the same basidiocarp were studied. If none of these nine strains revealed conidia, three of them were studied a second time after 2–3 mo. The material for microscopic examinations was mounted with cotton blue in lactic acid. Drawings were prepared with a camera lucida.

The specimens used for isolating haploid and dikaryotic mycelia are deposited at the Staatliches Museum für Naturkunde Görlitz (GLM; Germany). One of the nine strains isolated from each basidiocarp is kept either at Bayer HealthCare (BAYER; Wuppertal, Germany), the Centraal-bureau voor Schimmelcultures (CBS; Utrecht, Netherlands), the Deutsche Sammlung von Mikroorganismen und Zellkulturen (German Collection of Microorganisms and Cell Cultures, DSMZ; Braunschweig, Germany) or the Hans-Knöll-Institut (HKI) at Jena (Germany). Herbarium and strain numbers are given in the results section. Several strains died before they could be deposited in a permanent collection. In these cases, the respective isolates are referred to in the text with herbarium numbers. Names of species that cannot be related to current nomenclature are given in quotation marks.

Germination tests of the conidia.— – Richly sporulating mycelia were rinsed with sterile distilled water. The resulting conidial suspensions were mixed 1:1 (v/v) with malt-extract solution (2.5%) and kept at room temperature. Every day a drop from the conidial suspension was examined microscopically for germinating conidia.

	RESULTS

TOP ABSTRACT INTRODUCTION MATERIAL AND METHODS RESULTS DISCUSSION LITERATURE CITED

 
Thallic conidiogenesis in the Bolbitiaceae.— – Apart from Conocybe mesospora (Kühner) Kühner & Watling (GLM 45894), Conocybe rickeniana P.D. Orton (GLM 45896) and Agrocybe vervacti (Fr. : Fr.) Singer (GLM 45870, BAYER G618), none of which produced an anamorph, all species studied showed thallic conidiogenesis. As in all other chromosporic agaric families, the thalloconidia of the Bolbitiaceae were released rhexolytically (see Walther et al 2005). The plasma contraction was relatively distinct and occurred after the conidia initials were delimited by septa. In early stages the septation proceeded in basipetal succession; septum formation occurred more randomly at later stages of development. Previously delimited larger segments often subsequently disarticulated into two or more conidia. In Bolbitius and Conocybe mainly pairs of conidia arose in this way, which were later recognizable by their position close to each other (see e.g. FIG. 1B). In general mature conidia of the family were one-celled, hyaline and thin-walled.

View larger version (16K): [in this window] [in a new window]   FIG. 1. Conidiogenesis in Bolbitius and Conocybe. A. Bolbitius vitellinus. B. Conocybe albipes. C. Conocybe magnicapitata. D. Conocybe appendiculata. Bar = 10 µm.

Bolbitius and Conocybe species not only showed the same mode of conidiogenesis but also were similar with respect to the macro- and micromorphology of vegetative mycelia. Especially substrate hyphae consisted of short, strongly inflated elements. The swelling of these elements probably caused the tuberculate appearance of the mycelia because mycelial patches were forced to separate from the substrate. Substrate mycelia mostly had an orange-ochraceous color. The whitish aerial mycelia generally were sparsely developed.

Below we describe the specific characters of each species. The amount of conidiogenous hyphae (ch) forming a distinct cluster by sympodial branching is used to measure the differentiation of the conidiophores (cph). In some cases the conidiogenous hyphae were coiled to such an extent that their number could exactly be determined only for young conidiophores. In older conidiophores only conidiogenous hyphae with a clearly recognizable connection to the conidiophoral stipe were taken into account so that their actual number probably was higher. The description ‘‘slightly mucous’’ is used in cases where the conidia formed only loose accumulations, while clearly ‘‘mucous’’ conidia adhered in compact masses.

Bolbitius vitellinus (Pers. : Fr.) Fr. (FIG. 1A)

Material studied: GLM 45874, HKI ST 27320; GLM 46048, BAYER G469.

Macroscopic: aerial mycelium nearly absent; substrate mycelium cream, pellicular.

Microscopic: ch coiled, up to 13 per cluster; cph stipes often long and branched; conidia curved, 1.5–5.0 x 1.0–1.6 µm, mucous.

Conocybe albipes (G.H. Otth.) Hausknecht, syn.: C. lactea (J.E. Lange) Métrod (FIG. 1B)

Material studied: GLM 45893.

Macroscopic: aerial mycelium whitish, floccose; substrate mycelium ochraceous, tuberculate.

Microscopic: ch straight or slightly curved, up to 10 per cluster; cph stipes often short; conidia straight, 2.1–6.0 x 1.6–2.6 µm, mucous.

Conocybe appendiculata J.E. Lange & Kühner ex Watling, syn.: C. velata (Velen.) Watling, Pholiotina velata (Velen.) Hauskn. (FIG. 1D)

Material studied: GLM 45892.

Macroscopic: aerial mycelium white, cottony to floccose; substrate mycelium only poorly differentiated.

Microscopic: ch straight to strongly coiled, up to nine per cluster, but mostly only three or four condensed in a geniculate cph part; cph stipes often long and branched; conidia straight or curved, cylindrical or sometimes slightly rounded, 2.5–8.3 x 1.1–2.3 µm, with one, two or several medium-sized droplets, dry; haploid and dikaryotic mycelia sporulating.

Conocybe magnicapitata P.D. Orton (FIG. 1C)

Material studied: GLM 45894.

Macroscopic: aerial mycelium whitish, cottony; substrate mycelium orange-ochraceous, tuberculate.

Microscopic: ch straight to slightly curved, up to 17 per cluster; cph swollen, especially at the apices of the cph stipes where the ch proliferated; conidia straight or slightly curved, 3.0–6.5 x 1.1–1.7 µm, slightly mucous.

Conocybe semiglobata (Kühner) Kühner & Watling (FIG. 2A)

View larger version (17K): [in this window] [in a new window]   FIG. 2. Conidiogenesis in Conocybe. A. Conocybe semiglobata. B. Conocybe subpubescens. C. Conocybe subovalis (parts from Walther et al 2005; permission granted). D. Conocybe sulcatipes. Bar = 10 µm.

Macroscopic: aerial mycelium sparse, whitish; substrate mycelium pale ochraceous, tuberculate in the central area.

Microscopic: ch coiled, up to six per cluster; apices of the cph stipe less distinctly geniculate compared with other species; conidia curved, 2.1–7.1 x 1.0–2.7 µm, mucous; conidia of mycelia isolated from GLM 45897 with small to medium-size droplets.

Conocybe subovalis (Kühner) Kühner & Watling (FIG. 2C)

Material studied: GLM 45898; GLM 46045, HKI ST 27332; GLM 46046, BAYER G261.

Macroscopic: aerial mycelium sparse, whitish; substrate mycelium pale ochraceous, tuberculate.

Microscopic: ch coiled, up to seven per cluster; apices of the cph stipes swelling with increasing sympodial branching, often strongly geniculate; conidia mostly curved, (1.3–)2.3–7.5(–9.0) x 1.1–2.0 µm, mucous; dikaryotic mycelia sparsely sporulating; ch of dikaryotic mycelia straight to coiled, with basal clamps.

Conocybe subpubescens P.D. Orton (FIG. 2B)

Material studied: GLM 45899, BAYER G262; GLM 46047, HKI ST 27342.

Macroscopic: aerial mycelium white, floccose to cottony; substrate mycelium pale orange.

Microscopic: ch curved to strongly coiled, up to 12 per cluster; conidia mostly curved, 2.0–5.5 x 1.0–2.0 µm, mucous.

Conocybe sulcatipes (Peck) Kühner (FIG. 2D)

Material studied: GLM 45900, BAYER G703.

Macroscopic: aerial mycelium sparse, whitish, floccose; substrate mycelium brownish gray, tuberculate.

Microscopic: ch nearly straight to coiled; conidia straight or curved, 3.2–7.5 x 1.0–2.1 µm, mucous.

Conocybe teneroides (J.E. Lange) Kits van Wav.

Material studied: GLM 45901.

Macroscopic: aerial mycelium nearly absent; substrate mycelium ochraceous to brown, smooth to velutinous, tuberculate.

Microscopic: mycelia sparsely sporulating; only few young cph observed; ch strongly coiled, up to 3 per cluster of the young cph; conidia mucous.

Thallic conidiogenesis and cultural characters in Agrocybe.— – The studied Agrocybe species differed in their conidiogenesis modes. Agrocybe dura, A. firma and A. praecox formed conidia simply by fragmentation of normally branched hyphae. In contrast Agrocybe arvalis developed sympodially proliferating conidiophores resembling those of Conocybe and Bolbitius, but the distance between the arising conidiogenous hyphae was somewhat greater. In Agrocybe aegerita differentiated conidiophores also occurred, but generally the fragmentation of undifferentiated hyphae dominated.

Contrary to Bolbitius and Conocybe, dikaryotic mycelia sporulated intensively in many cases (e.g. A. aegerita, A. dura, A. firma and A. praecox). The conidia of A. dura and A. praecox sometimes slightly swelled during maturation. Conidia that developed on differentiated conidiophores were mucous, those formed on undifferentiated conidiophores were dry.

All Agrocybe species produced luxurious aerial mycelia in culture. Substrate mycelia were hardly differentiated. In addition A. arvalis, A. firma and A. praecox formed coiled, rarely branched and acuminate skeletoid hyphae that were thinner and more cyanophilous than normal vegetative hyphae and possessed thicker cell walls.

Agrocybe arvalis (Fr. : Fr.) R. Heim & Romagn. (FIG. 3A)

View larger version (13K): [in this window] [in a new window]   FIG. 3. Conidiogenesis in Agrocybe. A. Agrocybe arvalis. B. Agrocybe aegerita. C. Agrocybe dura. D. Agrocybe firma. E. Agrocybe praecox. Bar = 10 µm.

Macroscopic: mat white, cottony, with sclerotia-like bodies.

Microscopic: ch curved to coiled, proliferating sympodially from differentiated cph, up to six per cluster, but often only two or three concentrated, disarticulating into 4–8 conidia; conidia straight to curved, 2.5–5.1 x 1.5–2.0 µm, mucous.

Agrocybe aegerita (V. Brig.) Singer, syn.: A. cylindracea (DC.) Gillet (FIG. 3B)

Material studied: GLM 46049, CBS 358.1.

Macroscopic: mat white to light brown, cottony; dikaryotic mycelium producing fruit bodies in culture.

Microscopic: haploid and dikaryotic mycelia sporulating; conidiogenesis highly variable; ch arising either as undifferentiated hyphae with straight or coiled tips or sympodially on differentiated cph; ch of differentiated cph straight, up to eight per cluster, fragmenting into four to many conidia; apices of the cph stipes occasionally geniculate; conidia cylindrical, straight to slightly curved, 3.5–9.5 x 1.5–2.1 µm, mucous, with one to several minute to medium-sized droplets (predominantly with two medium-sized droplets); dikaryotic mycelium sparsely sporulating; only hyphal tips sporulating, therefore clamps rarely involved.

Agrocybe dura (Bolton) Singer (FIG. 3C)

Material studied: GLM 45866, BAYER G820.

Macroscopic: mat white to yellowish, cottony.

Microscopic: haploid and dikaryotic mycelia sporulating; haploid mycelia completely disarticulating; conidia developing by fragmentation of undifferentiated, normally branched hyphae, cylindrical, slightly swollen to distinctly rounded in older mycelia, dry, homogenous, 3.5–22.5 x 1.5–4.1(–6 in older mycelia) µm, haploid and dikaryotic mycelia occasionally forming pseudoclamp-like projections; dikaryotic mycelia moderately sporulating; clamps becoming part of the conidia or forming conidia themselves.

Agrocybe firma (Peck) Singer (FIG. 3D)

Material studied: GLM 45867, DSMZ 15787.

Macroscopic: mat cream to yellowish, plain, cottony; dikaryotic mycelium produced primordia.

Microscopic: haploid and dikaryotic mycelia luxuriantly sporulating; the entire mycelia disarticulating, conidia developing by fragmentation of undifferentiated, normally branched hyphae, cylindrical, 2.1–15.0 x 1.6–3.5 µm, dry, mostly with one or two large droplets; ch of dikaryotic mycelia clamped regularly; clamps becoming part of the conidia or forming conidia themselves.

Agrocybe praecox (Pers. : Fr.) Fayod (FIG. 3E)

Material studied: GLM 45869, DMSZ G739.

Macroscopic: mat white to cream, cottony.

Microscopic: haploid and dikaryotic mycelia sporulating; conidia developing by fragmentation of undifferentiated normally branched hyphae; noticeably large distances between two mature conidia; frequently several retraction septa present between two conidia caused by successive plasma contractions; sometimes irregular conidiogenesis leaving out large hyphal parts; conidia occasionally swelling, cylindrical or rounded, 3.8–19.5 x 1.1–2.4(–4) µm, dry, with or without minute to small droplets; haploid and dikaryotic mycelia forming pseudoclamp-like projections, clamps becoming part of the conidia or forming conidia themselves.

Germination of the conidia.— – To test their function as diaspores, we performed germination tests in four species. The conidia of Conocybe subovalis and C. subpubescens were germinable, but the germination rate was below 1%. The conidia of Agrocybe dura and A. praecox germinated easily.

Secretory cells.— – Secretory cells were common in the studied genera. We found them in Agrocybe firma, A. vervacti (FIG. 4A), Conocybe appendiculata (FIG. 4B), C. magnicapitata, C. rickeniana, C. semiglobata, C. subovalis, C. subpubescens and C. sulcatipes. Clear differences in shape and size of these cells existed between both genera. Secretory cells of Conocybe spp. possessed larger capitate apices than Agrocybe spp. The droplet of fluid secreted by the heads of the secretory cells was frequently recognizable in Conocybe, but we observed it only once in Agrocybe.

View larger version (4K): [in this window] [in a new window]   FIG. 4. Secretory cells in Agrocybe vervacti and Conocybe appendiculata. A. Agrocybe vervacti. B. Conocybe appendiculata. Bar = 10 µm.

	DISCUSSION

TOP ABSTRACT INTRODUCTION MATERIAL AND METHODS RESULTS DISCUSSION LITERATURE CITED

In comparison to Conocybe, less is known about the anamorphs of Agrocybe. Anamorph descriptions of Agrocybe firma given by Oddoux (1955) and Galland (1968) are concordant with our observations. Sede and López (1999) found large, elliptic to globose and irregularly shaped, slightly thick-walled conidia (‘‘chlamydospores’’) produced terminally or intercalary from subtending thinner hyphae in dikaryotic mycelia of Agrocybe aegerita (as A. cylindrica). We did not observe this type of conidia in our cultures.

Anamorphic characters in Agrocybe are heterogeneous and do not support the monophyly of this genus. However further studies including more species are necessary to clarify this question. That anamorphs of three out of five Agrocybe spp. studied here clearly deviate from the characteristic anamorph found in Conocybe and Bolbitius indicates that these Agrocybe spp. do not belong to the core group of the Bolbitiaceae, which is consistent with results of molecular phylogenetic analyses (Moncalvo et al 2002, Walther et al 2005, Gulden et al 2005). Based on these analyses and the differences in anamorph morphology we propose this emendation:

Bolbitiaceae Singer (1948), emend. G. Walther & M. Weiß

Members of the Agaricales with agaricoid or secotioid basidiomata and spore prints predominantly with red tones (rusty, ochraceous, ferruginous-fuscous) to tobacco brown in some Bolbitius spp. (Singer 1986); basidiospores smooth or ornamented, pileipellis hymeniform throughout, pleurocystidia absent; sympodially proliferating conidiophores formed on haploid mycelia in culture, apices of older conidiophoral stipes geniculate; conidiogenous hyphae often coiled, mostly fragmenting into four to eight conidia, conidia straight cylindrical or curved, mostly mucous, adhering in compact masses.

This emended Bolbitiaceae includes Bolbitius, Conocybe (including Pholiotina), Descolea, Gastrocybe, and tentatively Galerella, Galeropsis and Setchelliogaster. Agrocybe is excluded from the family.

Secretory cells have been described for Conocybe albipes (Hutchison et al 1996, as C. lactea) and Agrocybe pediades (Vandendries 1937, Lamoure 1960). Vandendries (1937) misinterpreted broken capitate apices as blastoconidia. However similar secretory cells have been reported for various only distantly related taxa such as Psilocybe (Heim and Wasson 1958), Galerina (Kühner 1946), Pleurotus (e.g. Barron and Thorn 1987) or Schizophyllum (Brefeld 1889). Consequently such cells probably possess no systematic relevance at lower taxonomic levels.

	ACKNOWLEDGMENTS

 
We thank Heinrich Dörfelt and Walter Gams for helpful discussions of diverse problems concerning anamorphs and Bayer HealthCare for financially supporting this study. We are also grateful to Cambridge University Press for the kind permission to reproduce parts of FIG. 2C, which originally appeared in Walther et al (2005).

	FOOTNOTES

 
Accepted for publication August 28, 2006.

² Present address: CBS Fungal Biodiversity Centre, P.O. Box 85167, NL-3508 AD Utrecht, The Netherlands.

¹ Corresponding author. E-mail: grit_walther44{at}yahoo.de

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This Article Abstract Full Text (PDF) Services Similar articles in this journal Alert me to new issues of the journal Download to citation manager Permissions Citing Articles Citing Articles via Google Scholar Google Scholar Articles by Walther, G. Articles by Weiß, M. Search for Related Content PubMed Articles by Walther, G. Articles by Weiß, M. Agricola Articles by Walther, G. Articles by Weiß, M.