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The genus Micropsalliota (Agaricaceae) in Kerala State, India
Mycol. Res. 95 ( 3 ) :341-346 (1991)
341
Printed in Greai Brifain
The genus Micropsalliota (Agaricaceae) in Kerala State, India
P. HEINEMANN Faculth des Sciences Agronomiques i7 B. 5030 Gembloux, Belgium and lardin Botanique national de Belgique 2 B. 1860 Meise, Belgium K. M. LEELAVATHY Mycology Research Laboratory, Department of Bofany, University of Calicuf, Kerala 673 635, India
Thirteen species of Micropsalliota, three of which are new, are considered. Full descriptions and illustrations are given for the new species, namely M . albosericea, M . pulverulenta and M . ptldica as well as for two collections attributed tentatively to M . cinnamomeopallida and to M . globocysfis, the others benefiting from complementary notes on the spores examined in SEM. A key to the species treated is given as an appendix.
The genus Micropsalliota v. Hoehn. emend Pegler is widely distributed in the tropical zones. It seems more represented in Asia, less in Africa and America but this is perhaps due to incomplete investigation. Wherever the small basidiomes are systematically investigated, several species are discovered, and nearly fifty species are now known. From Kerala, Heinemann & Little Flower (1983) described six new species, not mentioning some species known from other countries. Since then, new collections have been made by one of us (K.M.L.) and as a result thirteen species are known from Kerala. They all have been collected from the campus of the Calicut University which lies in the Malabar coast of southern India. The species are treated alphabetically. In our treatment, we give descriptions to the new species and two collections tentatively named, viz. M. cinnamomeopallida and M. globecystis. However, for the other species, existing descriptions are completed with results obtained from SEM examination of the spores. All data are taken from types unless otherwise stated. The spores measurements follow the procedure proposed by Heinemann & Rammeloo (1985).
Micropsalliota alba Heinem. & Little Flower, Bull. lard. Bot. Nat. Belg. 5 3 : 75, fig. 1 (1983). SEM: Spores nearly
(Fig. 6d)
smooth with very fine asperities
( < 0.1 pm) especially towards both apex and base ( x 20 000).
Specimen exnmined: Kerala: Calicut, 19 Aug. 1980, Little Flower 567 (BR holotypus, Calicut University Botany Herbarium (CUBH) isotypus).
Micropsalliota albosericea Heinem. & Leelavathy, sp. nov. (Figs I, 6s) Minusculus. Pileus albus, sericeus. Stipes graciIis, albus, tomentosus, annulo praeditus. Lamellae albidae deinde roseo-brunneae. Sporae
Fig. 1. Micropsalliofa albosericea (holotypus). (a) Cheilocystidia, x 1000; (b) spores, x 2000; (c) ca~loc~stidia, x 1000. 4.0-5.0 x 2.8-3.2 pm, plus minusve cymbifomes. Cheilocysfidia clavata, piriformia vel subcapitata apice usque 10 pm lato.
Fruitbody very slender and fragile. Pileus less than 0.5 cm diam, convexo-campanulate to plano-convex with a small umbo; surface pure white, silky smooth. Stipe 0.8-1.2 x ca 0.05 cm, white, hairy; annulus superior, membranous. Lamellae free, white to pinkish brown. Flesh white, not changing colour on bruising. Exsiccaturn ochre brownish. Spores 4.0452-5.0 x (2.6) 2.8-2.96-3.2 um (Q: (1.36) 1.39-1-53-1.66 (1.73); n = 30), brownish, shortly ovoid or f cymbiform, the apex obscurely conical; wall fairly thin; endosporium slightly thickened at the apex. Basidia 15-20 x 5-7.5 pm, clavate, hyaline, with 4 sterigmata. Cheilocystidia (10) 20-25 x 6-5-9-0 (12.5) pm, clavate-pyriform, rarely ventricose with a capitate head up to 10 Vm diam. Trama subregular; hyphae with swollen cells, up to 15 pm diam. Cuticle of radial hyphae, 5-10 pm diam, suberect in places with the endsf capitate. Stipe with tufts of caulocystidia (8) 15-28 x 5-8.5 um, clavate or f capitate. Clamp-connexions none. SEM: Spores nearly smooth to lowly verrucose; verrucae irregular, less than 0.05 um diam.
Micropsalliota in Kerala Specimen examined: Kerala: Calicut, University Campus, on the ground, I2 Nov. 1983, Leelavathy F 142 (BR holotypus, CUBH, isotypus). This species differs principally from M . plumaria (Berk. & Br.) v. Hoehn. by its cymbiform spores; other species have similar spores but their pileus is not white.
Micropsalliota arginea (Berk. & Br.) Pegler & Rayner,
IC!72
Fig. 2. Micropsalliota cinnamomeopallida, spores, x ZOO0 (a) Leelavathy F38; (b) holotypus.
Bull. 2 3 : 367 (1969). Agaricus argineus Berk. & Br., lour. Linn. Soc., Bot. 1 1 : 551 (1871). Stropharia arginea (Berk. & Br.) Sacc., Syll. Fung. 5 : 1026 (1887). Psalliofa arginea (Berk. & Br.) v. Hoehn., Akad. Wiss. Wien Math.-Nafumiss. Klasse 1 1 8 : 12, fig. 3 (1909). SEM: Spores with roundish verrucae, 0.084.5 pm large, distant, smaller at the apex ( x 20 000). Kerala: Calicut, University Campus, on the ground among litter, 30 Oct. 1982, Leelavathy F43 and F44 (CUBH, BR). The species was described from Sri Lanka and further defined by Petch (1917) from the same country and by v. Hoehnel(1909) from Java. It has also been recorded by Pegler (1977) from East Africa but the published sketch does not fit the habit of the species. There is also some doubt about the the two collections from Kerala State, for they show slight differences in spore form.
Micropsalliota avellanea Heinem. & Little Flower, Bull. lard. Bof. Nat. Belg. 5 3 : 77 (1983).
(Fig. 6j)
SEM: Spores verrucose, with roundish vermcae, 0.05-0.15 pm, rather spaced on a rugose background. Specimens examined: Kerala: Calicut, 8 Jun. 1980, Little Flower 483 (BR holotypus, CUBH isotypus); id., 23 Oct. 1980, Little Flower 617 (CUBH, BR).
SEM: Spores irregularly verrucose, with roundish 0.054.15 pm large verrucae (rarely and locally much larger), progressively smaller towards the apex which is sometimes nearly smooth ( x 20 000). Specimens examined: Kerala: Calicut University Campus, 29 Oct. 1982, Leelavathy F38 (CUBH, BR). The above description is exclusively based on the Kerala collection. We have seen the holotypus of M. cinnamomeopallida and found the following discrepancies with the Kerala collection: (a) spores slightly larger: 4.8-5.30-5.7x 2.9-3.10-3.3 pm (Q: 1.59-1.71-1.84; n = 31). In SEM the ornamentation pattern is similar but perhaps more regular; (b) cheilocystidia are more of a 'classic' form though they are intermixed with clavate, broad elements not signalized in the original description; (c) the pigmentation of the pileal surface is clearly membranal but we suspect that both types of pigmentation may occur on the same fruitbody, with variable intensity according to the place examined. A new species is not proposed on such slight differences but the simple assimilation might be hazardous. Micropsalliota species, though pantropical, seem to exhibit a certain endemism, at least at a continental level. M. cinnamome~~allida was described from central America (Costa Rica).
Micropsalliota cymbispora Heinem. & Little Flower, Bull. Micropsalliota cfr cinnamomeopallida Singer apud Singer & Gomez, Brenesia 19-20: 37 (1982).
(Figs 2, 6f)
Pileus 0.5-1.2 cm diam, campanulate with broad umbo or convex, white with the disk pinkish brown owing to minute light brown scales, at first with a white floccose covering. Sfipe 1.5-2.5 x 0.15-0.2 cm, cylindrical, white, nearly smooth; annulus white, cottony, sometimes fugacious. Lamellae free, white to dull pinkish brown, crowded, with lamellulae; edge serrate. Flesh white, not changing colour on bruising. Exisccafum: pileus ochre brownish, darker in the centre, stipe and lamellae light brown. Spores 4.1-462-5.1 x 2.5-2.87-3.1 ym ( Q : (1.46) 1.49-1.62-1.74 (1.78); n = 31), brownish, ellipsoid with the adaxial surface more applanate than the abaxial; wall rather thin, slightly thickened at the apex. Basidia 4-spored, 10-15 x 5-6 ym. Cheilocysfidia polymorphous, lanceolate, generally capitate, 20-30 x 5-7 pm, with the capitellum 4-6 ym diam, sometimes thickened or with digitate projections at the apex; some light brownish. Cuticle formed by cylindrical sometimes branched hyphae, 3-6 (10) ym diam; pigment probably vacuolar, brownish, $-soluble in NH,OH in which it discolours olivaceous. Surface of sfipe with parallel cylindrical hyphae, some with capitate apices, e.g. 4.7 pm diam.
lard. Bof. Naf. Belg. 5 3 : 78 (1983).
(Fig. 6 h)
SEM: Spores verrucose, with roundish, rather irregular verrucae, 0.05-0.3 pm in diam. Specimens examined: Kerala: Calicut, 5 Aug. 1980, Little Flower 555 (BR holotypus, CUBH isotypus); id., I0 Jun. 1980, Leelavathy in Little Flower 428 (CUBH, BR).
Micropsalliota cfr globocystis Heinem., Bull. lard. Bof. Nat. Belg. 5 0 : 5 7 (1980).
(Figs 3, 6a)
Pileus 1.5-2.4 cm diam, conical or convex then plane, umbonate; surface brown or light brown with the umbo dark purplish brown, fibrillose - tomentose, shaggy at the centre; margin becoming striate. Sfipe 3.5-4.6 x 0.1-0.15 cm, cylindric, probably solid then hollow, &cartilaginous, concolorous to the pileus. Lamellae free, 0.14.2 cm wide, moderately crowded, with lamellulae, date brown; edge serrate; annulus membranous, concolorous to the stipe, not persistent. Flesh thin, dull white then greyish brown. Spore-print brown. Exsiccatum deep brown sometimes with a violet hue; stipe twisted. Spores 5.9-6.74-7.8 (8.2) x 3.5-3.98-4.3 pm (Q: 1.48-1.69-2.00; n = 48), brownish, ellipsoid or feebly amygdaliform; wall fairly thick; with a light apical thickening.
P. Heinemann and K. M. Leelavathy
Fig. 3. Micropsalfiota cfr globocystis. (a) and (c) Cheilocystidia, x 1000; (b) and (a-c) Little Flower 419; (d-e) Little Flower 420).
(4 spores, x 2000; (e) hyphae of pileipellis, x 500.
Basidia (2) 4-spored, clavate, hyaline, 12.5-16.5 x 6-7.5 pm. Micropsalliota plumaria (Berk. & Br.) v. Hoehn., Akad. Wiss. Wien. Math.-Nafumiss, Klasse 1 2 3 : 32 (1914), auct. pl. Cheilocysfidia (20) 30-37.5 x 7-10 pm, capitate, the capitellurn (5) 9-1 1 prn diam. Pleurocysfidia probably none. ~ ~ m e n o ~ h o r a l non v. Hoehn. franta regular to subregular, the hyphae 4.5-12 pm diam; Agaricus plumarius Berk. & Br., Journ. Linn. Soc. Bot. 1 1 : 552 subhymenium narrow. Cuticle of ferect hyphae, 7-16.5 (1871). pm diam, somewhat constricted at the septa; wall somewhat SEM: Spores nearly smooth to lowly verrucose; verrucae thickened, with yellow brown incrustations. Sfipe surface with irregular, less than 0.05 pm diarn., smaller towards the apex cylindric hyphae, 5-9 pm diam. (Kerala collection). SEM: Spores finely verrucose with verrucae, 0.050.1 ym diam., smaller or absent at the apex ( x 20000; both Specimen examined: Kerala: Calicut, University Campus, on the soil, among litter, 9 Nov. 1982, Leelavathy F. 62 (CUBH, BR). collections). The Kerala collection corresponds fairly well with the Specimens examined: Kerala: Calicut 9 Sept. 1980, Little Flower 419 and 420 (CUBH, BR). species described by Berkeley & Broome but not with its interpretation by v. Hoehnel, whose fungus is said to have a The description is based upon the two Kerala collections finely squamulose pileus becoming wine red and brown and which are probably represented by fairly old fruitbodies. The spores 6-7 x 3 ym. The latter is evidently too large when comassimilation with M. globocysfis remains dubious some features pared with the spores of the type. The type (K. Thwaites 191) of it having not been seen on the Kerala material, especially has spores regularly ellipsoid with sometimes a slight apical a yellow and red discolouration of the context on bruising, but thickening, measuring: 4.5-497-5.5 (5.7) x 3.1-3.44-3.7 pm this may be the result of ageing. Furthermore, the dimensions (Q: (1.22) 1.26-1.45-1.63 (1.66); n = 31); SEM: spores with of the microscopical elements are somewhat smaller in the distant, roundish, 0.05-0.15 pm large verrucae, progressively Kerala collections. smaller towards the apex which is nearly smooth. The Kerala collection fits well this pattern. Micropsalliota malabarensis Heinem. & Little Flower, Bull. (Fig. 6i) lard. Bof. Naf. Belg. 53: 8 1 (1983). SEM: Spores verrucose with irregular, roundish verrucae, 005-0.15 ym diam. ( x 20000). Specimen examined: Kerala: Calicut, 4 Sept 1980, Little Flower 571 (BR holotypus, CUBH isotypus). Micropsalliota pleurocystidiata Heinern. & Little Flower, (Fig. 6e) Bull. lard. Bot. Naf. Belg. 5 3 : 81 (1983). SEM: Spores verrucose, with dense, roundish, 0.1-0.2 ym large verrucae which are often confluent especially near the apiculus. Specimens examined: Kerala: Calicut, 13 Jun. 1980, Leelavathy in Little Flower 449 (BR holotypus, CUBH isotypus).
Micropsalliota pudica Heinem & Leelavathy sp, nov. (Figs 4, 6 b) Carpophorw parvus, gracilis, albus, lutescens et mbescens. Pileus convexo-campanulatus, laevis, margo appendiculatus. Stipes elongatus, laevis; annulus absens. Larnellae liberae, albidae deinde brunneae. Sporae 5.2-6.4 x 3.4-4.1 pm, amygdalifomes, apice incrassatae et tmncatae, microscopio electronico leviter vermcosae. Cheilocystidia capitata, 40-50 x 5-10.5 pm, capitulo 4-65 pm diam. Fmifbody white, becoming yellow and orange red on bruising. Pileus up to 1.5 cm &am, convexo-campanulate; white, silky smooth; margin appendiculate. Stipe 3-3-5 x 0.15-0-2 cm, cylindrical, hollow; surface white,
Micropsalliota in Kerala
Fig. 4. Micropsalliota pudica (holotypus). (a) Cheil~c~stidia, x 1000; (b) spores, x 2000; (c) basidiurn and basidiole, x 1000; (d) ~ossibly a pleurocystidium, x 1000.
smooth; annulus absent. Lamellae free, up to 0.2 cm broad, dull white then brown; edge serrate. Flesh up to 0.15 cm thick near centre, white. Exsiccatum greyish brown. Spores 5-2-5.786.4 x 3.4-3.77-4.1 pm (Q: 1.41-1.54-1.68; n = 32), pinkish brown, amygdaliform with the apex obscurely truncate; wall thick with an apical thickening. Basidia &pored, ca 20-24 x 6 4 5 pm. Ckeilocystidia abundant, capitate with ventricose base, hyaline, (26) 40-50 (54) x 5-10.5 (12.5) ym, the head 4.5-6.5 (8.5) ym diam; wall smooth or somewhat incrusted. Pleurocystidia none, except perhaps near the lamella. of cylindrical hyphae, edge (one seen 40 x 12 ~ m ) Cuticle 3.5-7 ym diam, hyaline, without incrustations. Clamp-connexions not detected.
SEM : Spores nearly smooth to finely and irregular vermcose; verrucae 0.05 Vm or less diam, frequently smaller or even missing near the apex, the latter obviously truncate ( x 20000). Specimen examined: Kerala: Calicut, University Campus, in soil, isolated or two together, 4 Nov. 1983, Leelavathy F126 (BR holotypus, CUBH isotypus). This species is characterized, among the slender white species, by the apically truncated spores. An orange red colour oozes out into water when (fresh) material is crushed or cut.
(1.46); n = 231, yellowish brown, subcymbifonn; wall fairly thick, with an apical thickening. Basidia 4-spored. Cheilocysfidia cylindrical, some with ventricose base, capitate or subcapitate, 34.5-57.5 x 3.5-10 pm, the capitellurn ca 5 ym; content hyaline or brown near the apex. Cuticle with globose cells, 11-25 ym diam, slightly thickened wall, interspersed with pileocystidia; pigmentation membranal, Pileo- and caulocysfidia analogous to cheilocystidia. Clamp-connexions none.
SEM: Spores verrucose with roundish or irregular verrucae, 0.06-0.25 pm diam. ( x 20 000). Specimen examined: Kerala; Calicut, on the ground, 31 Oct. 1982, Leelavathy F39 A (CUBH, BR); id., 19 Sep. 1984, Leelavathy F271 (holotypus BR, isotypus CUBH). The description is based only on the holotypus, but a clear understanding of the relationship between the spherocyts and the pileal context was not discernible. The other collection shows abnormal variability of the spores. This is a puzzling species: the pileal globose cells would lead to Hymenagaricus Heinem. subgen. Xnnfhagaricw but the spores and cheilocystidiae retain it in Micropsalliota.
Micropsalliota pulverulenta Heinem. & Leelavathy, sp. nov. (Fig. 5) Minutus. Pileus albus; pulverulentus, mbescens. Stipes breviusculus, tornentosus, albus, ~bescens,annulo praeditus. Lameiiae liberae, ventricosae, albae deinde roseobmnneae. Sporae 4.5-5.2 x 3.5apici incrassatae. Cheil~c~stidia, pileocystidia 4.1 prn, s~bc~rnbiformes, et caulocystidia capitata vel subcapitata, 34.5-575 x 3.5-10 pm. Cutis pilei cellulis globosis praedita.
Pileu 0.3-0.6 cm diam, convex to plane, with a slight umbo; surface pruinose, pure white becoming dull vinaceous on drying, cracking up in some. Sfipe 0 - 4 4 . 7 x 0.034.05 cm, tomentose, white becoming vinaceous; annulus superior, white, evanescent. Lamellae free, up to 0.1 cm broad, ventricose, white becoming pinkish brown, moderately spaced, some forked near the margin, with lamellulae; edge fimbriate. Spores 4.5-4.82-5.2 x 3.5-3.83-4.1 Wm ( Q = 1.13-1.26-1.40
Fig. 5. Micropsalliota pulverulenta (holotypus). (a) Spherocyts of piieipellis, x 1000; (b) spores, x 2000; (c) cheilocystidia, x 1000.
P. Heinemann a n d K. M. Leelavathy
345
Fig. 6. Spores, SEM, x 10000. (a) Micropsailiota cfr globocystis (Little Flower 419); (b) M . pudica; (c) M. subalba; (d) M. alba; (e) M. pleuro~ystidiata;V) M. cfr cinnamomeopallida (Leelavathy F38); (8)M. albosericea; (h) M. cymbispora; (i) M. malabarensis; 6)M. avellanea. All holotypus except a and f.
Micropsalliota subalba Heinem. & Little Flower, Bull. lard. Bot. Nat. Befg. 53: 83 (1983).
(Fig. 6c)
SEM: Sports nearly smooth o r finely vermcose with dense, roundish, irregular, less than 0.05 wrn large verrucae. Specimens examined: Kerala: Calicut, 4 Oct. 1980, Little Flower 596 (BR holotypus, CUBH, isotypus); id, 24 Oct. 1980, Little Flower 618 (CUBH, BR).
Key to Micropsalliota species from Kerala Sfafe, India In the following key, t h e slenderness of fruit bodies is indicated b y a n index: IG (Indice de gracilite', Heinemann, 1 9 8 3 ) which is calculated as follows: St
IG
= ----,
D.d
where St = length of stipe; D = diameter of pileus; d = diameter of stipe.
1. Pileus white or whitish sometimes with the centre becoming brownish: . . M. pulverulenta 2. Pileal cuticle containing globose cells; spore 4.5-5.2 x 3.5-4.1 pm . 2. Pileal cuticle devoid of globose cells: 3. Spores 5.2-6.6 X 3.3-4.1 pm: 4. Fruitbody slender (IG > 30); pileus white: 5. Context yellowing and reddening; exsiccatum dark coloured; cheilocystidia 40-50 x 5-10.5 pm; spores slightly verrucose . M . pudica in SEM . . . . . M.alba , , 5. Context not discolouring: exsiccatum pale; cheil~c~stidia 26-37 x 4.0-7.5 Dm; spores nearly smooth in SEM. 4. Fruitbody squat (IG < 30); cheilocystidia with capitellurn 7-10 Um diarn; context not discolouring; pileus white then . . . M. sltbalba brownish . 3 . Spores 4.0-5.5 x 2.7-3.5 pm; fruitbody very slender: 6. Cheilocystidia truly capitate, sometimes intermixed with subcapitate ones: 7. Pileus white, whitish or pale greyish; exsiccatum dark coloured . . . M. arginea 7. Pileus pale with the centre cinnamon; exsiccatum greyish brown with the centre dark coloured . M. cinnamomeopallida . 6. Cheilcystidia subcapitate, sometimes intermixed with club-shaped ones:
Micrapsalliota
in Kerala
346
8. Spores ellipsoid, vermcose in SEM; pileus squamulose . . . . . .M. plumaria 8. Sporesf cymbifom, at least a number of them with the adaxial side concave or straight, nearly smooth in SEM; ~ i l e u ssilky
M.
albosericea
1. Pileus rosy, purplish or brown, or with coloured scales on a pale background: 9. Spores 44-6.1 x 2.8-3.8 prn: 10. Sporesf cymbiform, at least a number of them with the adaxial side concave or straight; no pleurocystidia: 11. Cuticular hyphae with encrusted pigmentation: 12. Fruitbody rather squat (IG < 30); cheilocystidia capitate, with capitellum 5.5-7.5 pm diarn; pileus brown with erect squamules
.
.
.
.
M.
malabarensis
12. Fruitbody slender (lG > 40); cheilocystidia capitate, with capitellum 3.7-4.5 prn diam; pileus rosy or vinaceous brown with . . . . M. cymbispora appressed squamules . 11. Cuticular hyphae with vacuolar pigmentation; cheilocystidia, with capitellum 4.5-5.5 urn diarn; pileus reddish brown, slightly squamulose . . . . . M. avellanea 10. Spores ellipsoid; pleurocystidia present; fruitbody rather large (D = 3-5 cm) . . . . M. pleurocystidiata 9. Spores 62-7.1 x 3-7-4.2 vm; cheilocystidia subcapitate with large capitellum; fruitbody slender; pileus purplish brown, somewhat squamulose, on a white background towards the margin . . M. globocystis
REFERENCES
Heinemann, P. (1983).Cle de dCtermination de Micropsalliota (Agaricaceat) et description de deux especes nouvelles. Bulletin du lardin Botanique National de Belgique 53, 85-95. Heinemann, P. & Little Flower (1983).Micropsalliota de Kerala (Inde). Bulletin du Jardin Botanique National de Belgique 53, 75-84. Heinernann, P. & Rammeloo, J. (1985). De la mesure des spores et de son expression. Agarica 6, 366-380. Hoehnel, F. von (1909). Fragmente zur Mykologie 6. Sitzungberichte der Ahdernie der Wissenschaften, Wien, Mathematik-Nafumissenschaften Klasse 118,275-452 (Received for publication 26 January 1990)
Hoehnel, F. von (1914). Fragmente zur Mykologie 16. Sitzungberichte der Ahdemie der Wissenschaften, Wien, Mathematik-NatumissenschaftenKlasse 123,49-155. Pegler, D. N. (1977). A ~reliminar~ Flora of East Africa. Kew Bulletin, Additional Series 6 , 615 p. Petch, T. (1917). A Revision of Ceylon Fungi (part V). Annals of the Royal Botanic Garden, Peradeniya 6 , 307-355.
341
Printed in Greai Brifain
The genus Micropsalliota (Agaricaceae) in Kerala State, India
P. HEINEMANN Faculth des Sciences Agronomiques i7 B. 5030 Gembloux, Belgium and lardin Botanique national de Belgique 2 B. 1860 Meise, Belgium K. M. LEELAVATHY Mycology Research Laboratory, Department of Bofany, University of Calicuf, Kerala 673 635, India
Thirteen species of Micropsalliota, three of which are new, are considered. Full descriptions and illustrations are given for the new species, namely M . albosericea, M . pulverulenta and M . ptldica as well as for two collections attributed tentatively to M . cinnamomeopallida and to M . globocysfis, the others benefiting from complementary notes on the spores examined in SEM. A key to the species treated is given as an appendix.
The genus Micropsalliota v. Hoehn. emend Pegler is widely distributed in the tropical zones. It seems more represented in Asia, less in Africa and America but this is perhaps due to incomplete investigation. Wherever the small basidiomes are systematically investigated, several species are discovered, and nearly fifty species are now known. From Kerala, Heinemann & Little Flower (1983) described six new species, not mentioning some species known from other countries. Since then, new collections have been made by one of us (K.M.L.) and as a result thirteen species are known from Kerala. They all have been collected from the campus of the Calicut University which lies in the Malabar coast of southern India. The species are treated alphabetically. In our treatment, we give descriptions to the new species and two collections tentatively named, viz. M. cinnamomeopallida and M. globecystis. However, for the other species, existing descriptions are completed with results obtained from SEM examination of the spores. All data are taken from types unless otherwise stated. The spores measurements follow the procedure proposed by Heinemann & Rammeloo (1985).
Micropsalliota alba Heinem. & Little Flower, Bull. lard. Bot. Nat. Belg. 5 3 : 75, fig. 1 (1983). SEM: Spores nearly
(Fig. 6d)
smooth with very fine asperities
( < 0.1 pm) especially towards both apex and base ( x 20 000).
Specimen exnmined: Kerala: Calicut, 19 Aug. 1980, Little Flower 567 (BR holotypus, Calicut University Botany Herbarium (CUBH) isotypus).
Micropsalliota albosericea Heinem. & Leelavathy, sp. nov. (Figs I, 6s) Minusculus. Pileus albus, sericeus. Stipes graciIis, albus, tomentosus, annulo praeditus. Lamellae albidae deinde roseo-brunneae. Sporae
Fig. 1. Micropsalliofa albosericea (holotypus). (a) Cheilocystidia, x 1000; (b) spores, x 2000; (c) ca~loc~stidia, x 1000. 4.0-5.0 x 2.8-3.2 pm, plus minusve cymbifomes. Cheilocysfidia clavata, piriformia vel subcapitata apice usque 10 pm lato.
Fruitbody very slender and fragile. Pileus less than 0.5 cm diam, convexo-campanulate to plano-convex with a small umbo; surface pure white, silky smooth. Stipe 0.8-1.2 x ca 0.05 cm, white, hairy; annulus superior, membranous. Lamellae free, white to pinkish brown. Flesh white, not changing colour on bruising. Exsiccaturn ochre brownish. Spores 4.0452-5.0 x (2.6) 2.8-2.96-3.2 um (Q: (1.36) 1.39-1-53-1.66 (1.73); n = 30), brownish, shortly ovoid or f cymbiform, the apex obscurely conical; wall fairly thin; endosporium slightly thickened at the apex. Basidia 15-20 x 5-7.5 pm, clavate, hyaline, with 4 sterigmata. Cheilocystidia (10) 20-25 x 6-5-9-0 (12.5) pm, clavate-pyriform, rarely ventricose with a capitate head up to 10 Vm diam. Trama subregular; hyphae with swollen cells, up to 15 pm diam. Cuticle of radial hyphae, 5-10 pm diam, suberect in places with the endsf capitate. Stipe with tufts of caulocystidia (8) 15-28 x 5-8.5 um, clavate or f capitate. Clamp-connexions none. SEM: Spores nearly smooth to lowly verrucose; verrucae irregular, less than 0.05 um diam.
Micropsalliota in Kerala Specimen examined: Kerala: Calicut, University Campus, on the ground, I2 Nov. 1983, Leelavathy F 142 (BR holotypus, CUBH, isotypus). This species differs principally from M . plumaria (Berk. & Br.) v. Hoehn. by its cymbiform spores; other species have similar spores but their pileus is not white.
Micropsalliota arginea (Berk. & Br.) Pegler & Rayner,
IC!72
Fig. 2. Micropsalliota cinnamomeopallida, spores, x ZOO0 (a) Leelavathy F38; (b) holotypus.
Bull. 2 3 : 367 (1969). Agaricus argineus Berk. & Br., lour. Linn. Soc., Bot. 1 1 : 551 (1871). Stropharia arginea (Berk. & Br.) Sacc., Syll. Fung. 5 : 1026 (1887). Psalliofa arginea (Berk. & Br.) v. Hoehn., Akad. Wiss. Wien Math.-Nafumiss. Klasse 1 1 8 : 12, fig. 3 (1909). SEM: Spores with roundish verrucae, 0.084.5 pm large, distant, smaller at the apex ( x 20 000). Kerala: Calicut, University Campus, on the ground among litter, 30 Oct. 1982, Leelavathy F43 and F44 (CUBH, BR). The species was described from Sri Lanka and further defined by Petch (1917) from the same country and by v. Hoehnel(1909) from Java. It has also been recorded by Pegler (1977) from East Africa but the published sketch does not fit the habit of the species. There is also some doubt about the the two collections from Kerala State, for they show slight differences in spore form.
Micropsalliota avellanea Heinem. & Little Flower, Bull. lard. Bof. Nat. Belg. 5 3 : 77 (1983).
(Fig. 6j)
SEM: Spores verrucose, with roundish vermcae, 0.05-0.15 pm, rather spaced on a rugose background. Specimens examined: Kerala: Calicut, 8 Jun. 1980, Little Flower 483 (BR holotypus, CUBH isotypus); id., 23 Oct. 1980, Little Flower 617 (CUBH, BR).
SEM: Spores irregularly verrucose, with roundish 0.054.15 pm large verrucae (rarely and locally much larger), progressively smaller towards the apex which is sometimes nearly smooth ( x 20 000). Specimens examined: Kerala: Calicut University Campus, 29 Oct. 1982, Leelavathy F38 (CUBH, BR). The above description is exclusively based on the Kerala collection. We have seen the holotypus of M. cinnamomeopallida and found the following discrepancies with the Kerala collection: (a) spores slightly larger: 4.8-5.30-5.7x 2.9-3.10-3.3 pm (Q: 1.59-1.71-1.84; n = 31). In SEM the ornamentation pattern is similar but perhaps more regular; (b) cheilocystidia are more of a 'classic' form though they are intermixed with clavate, broad elements not signalized in the original description; (c) the pigmentation of the pileal surface is clearly membranal but we suspect that both types of pigmentation may occur on the same fruitbody, with variable intensity according to the place examined. A new species is not proposed on such slight differences but the simple assimilation might be hazardous. Micropsalliota species, though pantropical, seem to exhibit a certain endemism, at least at a continental level. M. cinnamome~~allida was described from central America (Costa Rica).
Micropsalliota cymbispora Heinem. & Little Flower, Bull. Micropsalliota cfr cinnamomeopallida Singer apud Singer & Gomez, Brenesia 19-20: 37 (1982).
(Figs 2, 6f)
Pileus 0.5-1.2 cm diam, campanulate with broad umbo or convex, white with the disk pinkish brown owing to minute light brown scales, at first with a white floccose covering. Sfipe 1.5-2.5 x 0.15-0.2 cm, cylindrical, white, nearly smooth; annulus white, cottony, sometimes fugacious. Lamellae free, white to dull pinkish brown, crowded, with lamellulae; edge serrate. Flesh white, not changing colour on bruising. Exisccafum: pileus ochre brownish, darker in the centre, stipe and lamellae light brown. Spores 4.1-462-5.1 x 2.5-2.87-3.1 ym ( Q : (1.46) 1.49-1.62-1.74 (1.78); n = 31), brownish, ellipsoid with the adaxial surface more applanate than the abaxial; wall rather thin, slightly thickened at the apex. Basidia 4-spored, 10-15 x 5-6 ym. Cheilocysfidia polymorphous, lanceolate, generally capitate, 20-30 x 5-7 pm, with the capitellum 4-6 ym diam, sometimes thickened or with digitate projections at the apex; some light brownish. Cuticle formed by cylindrical sometimes branched hyphae, 3-6 (10) ym diam; pigment probably vacuolar, brownish, $-soluble in NH,OH in which it discolours olivaceous. Surface of sfipe with parallel cylindrical hyphae, some with capitate apices, e.g. 4.7 pm diam.
lard. Bof. Naf. Belg. 5 3 : 78 (1983).
(Fig. 6 h)
SEM: Spores verrucose, with roundish, rather irregular verrucae, 0.05-0.3 pm in diam. Specimens examined: Kerala: Calicut, 5 Aug. 1980, Little Flower 555 (BR holotypus, CUBH isotypus); id., I0 Jun. 1980, Leelavathy in Little Flower 428 (CUBH, BR).
Micropsalliota cfr globocystis Heinem., Bull. lard. Bof. Nat. Belg. 5 0 : 5 7 (1980).
(Figs 3, 6a)
Pileus 1.5-2.4 cm diam, conical or convex then plane, umbonate; surface brown or light brown with the umbo dark purplish brown, fibrillose - tomentose, shaggy at the centre; margin becoming striate. Sfipe 3.5-4.6 x 0.1-0.15 cm, cylindric, probably solid then hollow, &cartilaginous, concolorous to the pileus. Lamellae free, 0.14.2 cm wide, moderately crowded, with lamellulae, date brown; edge serrate; annulus membranous, concolorous to the stipe, not persistent. Flesh thin, dull white then greyish brown. Spore-print brown. Exsiccatum deep brown sometimes with a violet hue; stipe twisted. Spores 5.9-6.74-7.8 (8.2) x 3.5-3.98-4.3 pm (Q: 1.48-1.69-2.00; n = 48), brownish, ellipsoid or feebly amygdaliform; wall fairly thick; with a light apical thickening.
P. Heinemann and K. M. Leelavathy
Fig. 3. Micropsalfiota cfr globocystis. (a) and (c) Cheilocystidia, x 1000; (b) and (a-c) Little Flower 419; (d-e) Little Flower 420).
(4 spores, x 2000; (e) hyphae of pileipellis, x 500.
Basidia (2) 4-spored, clavate, hyaline, 12.5-16.5 x 6-7.5 pm. Micropsalliota plumaria (Berk. & Br.) v. Hoehn., Akad. Wiss. Wien. Math.-Nafumiss, Klasse 1 2 3 : 32 (1914), auct. pl. Cheilocysfidia (20) 30-37.5 x 7-10 pm, capitate, the capitellurn (5) 9-1 1 prn diam. Pleurocysfidia probably none. ~ ~ m e n o ~ h o r a l non v. Hoehn. franta regular to subregular, the hyphae 4.5-12 pm diam; Agaricus plumarius Berk. & Br., Journ. Linn. Soc. Bot. 1 1 : 552 subhymenium narrow. Cuticle of ferect hyphae, 7-16.5 (1871). pm diam, somewhat constricted at the septa; wall somewhat SEM: Spores nearly smooth to lowly verrucose; verrucae thickened, with yellow brown incrustations. Sfipe surface with irregular, less than 0.05 pm diarn., smaller towards the apex cylindric hyphae, 5-9 pm diam. (Kerala collection). SEM: Spores finely verrucose with verrucae, 0.050.1 ym diam., smaller or absent at the apex ( x 20000; both Specimen examined: Kerala: Calicut, University Campus, on the soil, among litter, 9 Nov. 1982, Leelavathy F. 62 (CUBH, BR). collections). The Kerala collection corresponds fairly well with the Specimens examined: Kerala: Calicut 9 Sept. 1980, Little Flower 419 and 420 (CUBH, BR). species described by Berkeley & Broome but not with its interpretation by v. Hoehnel, whose fungus is said to have a The description is based upon the two Kerala collections finely squamulose pileus becoming wine red and brown and which are probably represented by fairly old fruitbodies. The spores 6-7 x 3 ym. The latter is evidently too large when comassimilation with M. globocysfis remains dubious some features pared with the spores of the type. The type (K. Thwaites 191) of it having not been seen on the Kerala material, especially has spores regularly ellipsoid with sometimes a slight apical a yellow and red discolouration of the context on bruising, but thickening, measuring: 4.5-497-5.5 (5.7) x 3.1-3.44-3.7 pm this may be the result of ageing. Furthermore, the dimensions (Q: (1.22) 1.26-1.45-1.63 (1.66); n = 31); SEM: spores with of the microscopical elements are somewhat smaller in the distant, roundish, 0.05-0.15 pm large verrucae, progressively Kerala collections. smaller towards the apex which is nearly smooth. The Kerala collection fits well this pattern. Micropsalliota malabarensis Heinem. & Little Flower, Bull. (Fig. 6i) lard. Bof. Naf. Belg. 53: 8 1 (1983). SEM: Spores verrucose with irregular, roundish verrucae, 005-0.15 ym diam. ( x 20000). Specimen examined: Kerala: Calicut, 4 Sept 1980, Little Flower 571 (BR holotypus, CUBH isotypus). Micropsalliota pleurocystidiata Heinern. & Little Flower, (Fig. 6e) Bull. lard. Bot. Naf. Belg. 5 3 : 81 (1983). SEM: Spores verrucose, with dense, roundish, 0.1-0.2 ym large verrucae which are often confluent especially near the apiculus. Specimens examined: Kerala: Calicut, 13 Jun. 1980, Leelavathy in Little Flower 449 (BR holotypus, CUBH isotypus).
Micropsalliota pudica Heinem & Leelavathy sp, nov. (Figs 4, 6 b) Carpophorw parvus, gracilis, albus, lutescens et mbescens. Pileus convexo-campanulatus, laevis, margo appendiculatus. Stipes elongatus, laevis; annulus absens. Larnellae liberae, albidae deinde brunneae. Sporae 5.2-6.4 x 3.4-4.1 pm, amygdalifomes, apice incrassatae et tmncatae, microscopio electronico leviter vermcosae. Cheilocystidia capitata, 40-50 x 5-10.5 pm, capitulo 4-65 pm diam. Fmifbody white, becoming yellow and orange red on bruising. Pileus up to 1.5 cm &am, convexo-campanulate; white, silky smooth; margin appendiculate. Stipe 3-3-5 x 0.15-0-2 cm, cylindrical, hollow; surface white,
Micropsalliota in Kerala
Fig. 4. Micropsalliota pudica (holotypus). (a) Cheil~c~stidia, x 1000; (b) spores, x 2000; (c) basidiurn and basidiole, x 1000; (d) ~ossibly a pleurocystidium, x 1000.
smooth; annulus absent. Lamellae free, up to 0.2 cm broad, dull white then brown; edge serrate. Flesh up to 0.15 cm thick near centre, white. Exsiccatum greyish brown. Spores 5-2-5.786.4 x 3.4-3.77-4.1 pm (Q: 1.41-1.54-1.68; n = 32), pinkish brown, amygdaliform with the apex obscurely truncate; wall thick with an apical thickening. Basidia &pored, ca 20-24 x 6 4 5 pm. Ckeilocystidia abundant, capitate with ventricose base, hyaline, (26) 40-50 (54) x 5-10.5 (12.5) ym, the head 4.5-6.5 (8.5) ym diam; wall smooth or somewhat incrusted. Pleurocystidia none, except perhaps near the lamella. of cylindrical hyphae, edge (one seen 40 x 12 ~ m ) Cuticle 3.5-7 ym diam, hyaline, without incrustations. Clamp-connexions not detected.
SEM : Spores nearly smooth to finely and irregular vermcose; verrucae 0.05 Vm or less diam, frequently smaller or even missing near the apex, the latter obviously truncate ( x 20000). Specimen examined: Kerala: Calicut, University Campus, in soil, isolated or two together, 4 Nov. 1983, Leelavathy F126 (BR holotypus, CUBH isotypus). This species is characterized, among the slender white species, by the apically truncated spores. An orange red colour oozes out into water when (fresh) material is crushed or cut.
(1.46); n = 231, yellowish brown, subcymbifonn; wall fairly thick, with an apical thickening. Basidia 4-spored. Cheilocysfidia cylindrical, some with ventricose base, capitate or subcapitate, 34.5-57.5 x 3.5-10 pm, the capitellurn ca 5 ym; content hyaline or brown near the apex. Cuticle with globose cells, 11-25 ym diam, slightly thickened wall, interspersed with pileocystidia; pigmentation membranal, Pileo- and caulocysfidia analogous to cheilocystidia. Clamp-connexions none.
SEM: Spores verrucose with roundish or irregular verrucae, 0.06-0.25 pm diam. ( x 20 000). Specimen examined: Kerala; Calicut, on the ground, 31 Oct. 1982, Leelavathy F39 A (CUBH, BR); id., 19 Sep. 1984, Leelavathy F271 (holotypus BR, isotypus CUBH). The description is based only on the holotypus, but a clear understanding of the relationship between the spherocyts and the pileal context was not discernible. The other collection shows abnormal variability of the spores. This is a puzzling species: the pileal globose cells would lead to Hymenagaricus Heinem. subgen. Xnnfhagaricw but the spores and cheilocystidiae retain it in Micropsalliota.
Micropsalliota pulverulenta Heinem. & Leelavathy, sp. nov. (Fig. 5) Minutus. Pileus albus; pulverulentus, mbescens. Stipes breviusculus, tornentosus, albus, ~bescens,annulo praeditus. Lameiiae liberae, ventricosae, albae deinde roseobmnneae. Sporae 4.5-5.2 x 3.5apici incrassatae. Cheil~c~stidia, pileocystidia 4.1 prn, s~bc~rnbiformes, et caulocystidia capitata vel subcapitata, 34.5-575 x 3.5-10 pm. Cutis pilei cellulis globosis praedita.
Pileu 0.3-0.6 cm diam, convex to plane, with a slight umbo; surface pruinose, pure white becoming dull vinaceous on drying, cracking up in some. Sfipe 0 - 4 4 . 7 x 0.034.05 cm, tomentose, white becoming vinaceous; annulus superior, white, evanescent. Lamellae free, up to 0.1 cm broad, ventricose, white becoming pinkish brown, moderately spaced, some forked near the margin, with lamellulae; edge fimbriate. Spores 4.5-4.82-5.2 x 3.5-3.83-4.1 Wm ( Q = 1.13-1.26-1.40
Fig. 5. Micropsalliota pulverulenta (holotypus). (a) Spherocyts of piieipellis, x 1000; (b) spores, x 2000; (c) cheilocystidia, x 1000.
P. Heinemann a n d K. M. Leelavathy
345
Fig. 6. Spores, SEM, x 10000. (a) Micropsailiota cfr globocystis (Little Flower 419); (b) M . pudica; (c) M. subalba; (d) M. alba; (e) M. pleuro~ystidiata;V) M. cfr cinnamomeopallida (Leelavathy F38); (8)M. albosericea; (h) M. cymbispora; (i) M. malabarensis; 6)M. avellanea. All holotypus except a and f.
Micropsalliota subalba Heinem. & Little Flower, Bull. lard. Bot. Nat. Befg. 53: 83 (1983).
(Fig. 6c)
SEM: Sports nearly smooth o r finely vermcose with dense, roundish, irregular, less than 0.05 wrn large verrucae. Specimens examined: Kerala: Calicut, 4 Oct. 1980, Little Flower 596 (BR holotypus, CUBH, isotypus); id, 24 Oct. 1980, Little Flower 618 (CUBH, BR).
Key to Micropsalliota species from Kerala Sfafe, India In the following key, t h e slenderness of fruit bodies is indicated b y a n index: IG (Indice de gracilite', Heinemann, 1 9 8 3 ) which is calculated as follows: St
IG
= ----,
D.d
where St = length of stipe; D = diameter of pileus; d = diameter of stipe.
1. Pileus white or whitish sometimes with the centre becoming brownish: . . M. pulverulenta 2. Pileal cuticle containing globose cells; spore 4.5-5.2 x 3.5-4.1 pm . 2. Pileal cuticle devoid of globose cells: 3. Spores 5.2-6.6 X 3.3-4.1 pm: 4. Fruitbody slender (IG > 30); pileus white: 5. Context yellowing and reddening; exsiccatum dark coloured; cheilocystidia 40-50 x 5-10.5 pm; spores slightly verrucose . M . pudica in SEM . . . . . M.alba , , 5. Context not discolouring: exsiccatum pale; cheil~c~stidia 26-37 x 4.0-7.5 Dm; spores nearly smooth in SEM. 4. Fruitbody squat (IG < 30); cheilocystidia with capitellurn 7-10 Um diarn; context not discolouring; pileus white then . . . M. sltbalba brownish . 3 . Spores 4.0-5.5 x 2.7-3.5 pm; fruitbody very slender: 6. Cheilocystidia truly capitate, sometimes intermixed with subcapitate ones: 7. Pileus white, whitish or pale greyish; exsiccatum dark coloured . . . M. arginea 7. Pileus pale with the centre cinnamon; exsiccatum greyish brown with the centre dark coloured . M. cinnamomeopallida . 6. Cheilcystidia subcapitate, sometimes intermixed with club-shaped ones:
Micrapsalliota
in Kerala
346
8. Spores ellipsoid, vermcose in SEM; pileus squamulose . . . . . .M. plumaria 8. Sporesf cymbifom, at least a number of them with the adaxial side concave or straight, nearly smooth in SEM; ~ i l e u ssilky
M.
albosericea
1. Pileus rosy, purplish or brown, or with coloured scales on a pale background: 9. Spores 44-6.1 x 2.8-3.8 prn: 10. Sporesf cymbiform, at least a number of them with the adaxial side concave or straight; no pleurocystidia: 11. Cuticular hyphae with encrusted pigmentation: 12. Fruitbody rather squat (IG < 30); cheilocystidia capitate, with capitellum 5.5-7.5 pm diarn; pileus brown with erect squamules
.
.
.
.
M.
malabarensis
12. Fruitbody slender (lG > 40); cheilocystidia capitate, with capitellum 3.7-4.5 prn diam; pileus rosy or vinaceous brown with . . . . M. cymbispora appressed squamules . 11. Cuticular hyphae with vacuolar pigmentation; cheilocystidia, with capitellum 4.5-5.5 urn diarn; pileus reddish brown, slightly squamulose . . . . . M. avellanea 10. Spores ellipsoid; pleurocystidia present; fruitbody rather large (D = 3-5 cm) . . . . M. pleurocystidiata 9. Spores 62-7.1 x 3-7-4.2 vm; cheilocystidia subcapitate with large capitellum; fruitbody slender; pileus purplish brown, somewhat squamulose, on a white background towards the margin . . M. globocystis
REFERENCES
Heinemann, P. (1983).Cle de dCtermination de Micropsalliota (Agaricaceat) et description de deux especes nouvelles. Bulletin du lardin Botanique National de Belgique 53, 85-95. Heinemann, P. & Little Flower (1983).Micropsalliota de Kerala (Inde). Bulletin du Jardin Botanique National de Belgique 53, 75-84. Heinernann, P. & Rammeloo, J. (1985). De la mesure des spores et de son expression. Agarica 6, 366-380. Hoehnel, F. von (1909). Fragmente zur Mykologie 6. Sitzungberichte der Ahdernie der Wissenschaften, Wien, Mathematik-Nafumissenschaften Klasse 118,275-452 (Received for publication 26 January 1990)
Hoehnel, F. von (1914). Fragmente zur Mykologie 16. Sitzungberichte der Ahdemie der Wissenschaften, Wien, Mathematik-NatumissenschaftenKlasse 123,49-155. Pegler, D. N. (1977). A ~reliminar~ Flora of East Africa. Kew Bulletin, Additional Series 6 , 615 p. Petch, T. (1917). A Revision of Ceylon Fungi (part V). Annals of the Royal Botanic Garden, Peradeniya 6 , 307-355.