Watling, R.; Taylor, G.M. 1987: Observations on the Bolbitiaceae: 27. Preliminary account of the Bolbitiaceae of New Zealand. Bibliotheca Mycologica. 117.
Details
Descriptions
Segedin 1631 (Mt Eden, Auckland, legit Hasnain, 19.IX.1979) had similar basidiospores and cheilocystidia and some pleurocystidia with digitate apices similar both to those figured by Overholts (1927) for A. acericola, and seen in collections by one of us (RW) of the same species from Michigan, USA.
This fungus was originally described as Agaricus (Pholiota) acericolus Peck from wood and trunks of maple (Acer) covered with moss but it is not infrequent in N. America on rotten Acer spp., old trunks of Fagus grandifolia Ehrh. and on piles of sawdust and chips from felling activities in frondose woodland or used as mulching in shrubberies. The Auckland material was on wood-chips and could conceivably have been introduced into New Zealand.
This was listed by Berkeley in Hooker's New Zealand Flora (1855) and Massee (1898) as Agaricus (Pholiota) erebius collected by Colenso from Ahuriri; the record should be accepted with some reservation, although it may indicate that this taxon is present in the New Zealand flora. Cunningham is purported to have collected this taxon at Weraroa (material in PDD) but this is a quite different fungus, and as much as we know about the collection is tabulated below. A. erebia should be looked out for under shrubs in gardens and parks, and if present is probably introduced.
Another closely related species but in too poor condition to determine occurred in a garden at Weraroa.
Basidiospores smooth, generally with thick wall and prominent germ-pore, basidia usually 4-spored, in a few species 2-spored; gill trama regular becoming less regular with age. Pileipellis a distinct palisadoderm at first with or without dermatocystidia but may soon disorganize and become mixed with filamentous units; 'scalp' cellular. Cheilocystidia always present, vesiculose to lageniform or cylindric; pleurocystidia present or absent, either simple or prominently differentiated. Stipitipellis of cylindric, hyaline to slightly brownish hyphae; caulocystidia usually present especially at stipe-apex.
Development where known paravelangiocarpic, except in one species (A. cylindrica) where it is bivelangiocarpic.
On the ground in woods, pastures, heaths etc.; on dung, plant remains, refuse and wood; often in gardens, farmyards and greenhouses; saprophytes, or in the case of A. parasitica a weak parasite.
As presently described by Singer (1978) this is a rather broadly conceived taxon and probably will be divided into at least two taxa in the future. As the New Zealand material does not allow detailed examination the above epithet is utilised meantime; the New Zealand material was said to have a mild not a bitter taste, usually a character of A. howeana. It is found not only in North America, from where Peck described it, but also S. America and Central Africa, where it was probably introduced. It should be noted that Peck firstly referred this species to Stropharia because of the purplish tinge to the immature gills; comparison should be made with Stropharia coronilla. A second collection in PDD (solitary, amongst grass on ground, Weraroa, Wellington, 2 X 1919, Cunningham 680) is hesitantly placed here; the pleurocystidia are not as frequent. Some of Cunningham's information is utilised in the description above but because the species is so poorly documented in New Zealand a description of N. American material is offered for comparison:-
Pileus 53-100 mm, convex then expanding, becoming wavy at margin in broad undulations, rich tawny (pale)-straw colour paling to straw yellow at concentrically wrinkled margin, darker but often only slightly so towards disc or in larger basidiomata tawny orange at disc and similar although pale outwards, humid, smooth or fairly rugulose. (One Basidioma with pileus tinged distinct olivaceous buff 'throughout) Stipe 40-68 x 8-13 mm, narrow and relatively thin for size of pileus, swollen slightly at stipe-base to form small bulb with white rhizoidal strands, annulate, pale buff to pallid, irregularly fibrillose-striate below ring, striate at white apex, hollow; ring apical, thin, membranous, soon collapsing onto stipe and then almost disappearing or leaving in age a thin skin from collapsing or after abrasion. Gills buff-pallid with a purple-brown tinge especially when young then becoming snuff-brown. Flesh white in pileus, pinkish brown tinge in stipe downwards, within stipe around 'mid' brown. Taste bitter astringent not mealy (or at most only a hint which is masked and not persistent); smell not mealy, unpleasant but not persistent or penetrating, very earthy, of vegetables immediately on cutting.
In clump; under collapsed Manzanita ( Arctostaphylos sp.) by trackside, Selyer, near Willow Creek, N. California 11 iii 1984, Wat. 17222 in E.Pileus <52 mm plano-convexus vel subumbonatus, brunneo-olivaceus, centraliter ochraceus et rugosus. Stipes 150 x 6 mm (13 mm ad basem) bubalinus, ad apicem attenuatus et gradatim pallescens, squamulis ochraceis subcingulatis irregulariter basim versus instructus, tandem deorsum fuscans, rhizomorphis albis crassis longisque praeditus. Lamellae porphyro brunneae, subliberae. Carp sapore miti.
Basidiosporae cumulatae brunneae, 11-12.5 x 7-8 µm ellipsoideae, laeves, tunica crassa poro germinativo conspicuo instructae. Basidia 4 -sporigera, 34-40 x 10-11 µm, clavata.
Cystidia aciei lamellarum vesiculosa, catenata, fibulata, 20-30 x 15-18 µm. Cystidia faciei lamellarum clavata apice dilatato 46-62 x 23-28 µm. Cystidia stipitis lageniformia. Ad terram inter ramos mortuos, Novazelandia. Typus Horak ZT 69/88.
This species is common throughout New Zealand and probably recognised by Colenso (1890) and Massee (1898) as Agaricus (Pholiota) pudicus. Indeed Colenso's record of Pholiota heteroclita (Fr.) Quel. may refer to the same species but the material (b 624) in K lacks basidiospores. It is an excellent edible mushroom very closely related to A. cylindrica ( DC.: Fr.) Maire, indeed at the start of the present study following Singer (1969) the present authors included it within their circumscription of A. cylindrica.
In addition Stevenson (1982 a & b) describes the same species on Alectryon excelsum Gaertn., Carpodetus serratus Forst., Corynocarpus laevigatus Forst., Dysoxylum spectabile Blume, Hoheria sp., Laurelia novaezelandiae A. Cunn., Nothofagus solandri Oerst. and Plagianthus betulinus A. Cunn. To this list can now be added Erythrina caffra. Leptospermum ericoides. Macropiper excelsum. Magnolia grandiflora Linn. (Univ. of Auckland), Metrosideros excelsa. Populus italica (Duroi) Muench. (Wynyard Car Park, Univ. of Auckland).
Thus A. parasitica has now been found on both native and introduced trees and would appear to be far commoner than A. cylindrica is in Europe which, although it has been recorded on several hosts, is much more restricted in its host preference. One record exists in New Zealand for A. parasitica on Ulmus glabra Huds., a host on which A. cylindrica is found in Europe. Illustrations of this species occur in Taylor (1981; 1983).
A. parasitica (as A. cylindrica) from Australia is illustrated by Macdonald and Westerman 1979 and Fuhrer 1985. Dr Fuhrer kindly sent a collection to Edinburgh for study and it confirmed our identification. Material examined: on Atherospermum moschatum. Talbot Drive, near Marysville, Vie., Australia, v 1985, B. Fuhrer B236. This may be the first record of A. parasitica in Australia, where it appears to be not uncommon in Victoria at least, and is probably widespread.
Singer (1969) synonymised Agaricus crassivelus Spegazzini (in Ann. Soc. Cienc. Argent. 9: 279, 1880), Pholiota formosa Spegazzini (in Bot. Acad. Cienc. Cordoba 28; 311, 1926) and Pholiota impudica Spegazzini (in Bol. Acad. Nac. Cienc. Cordoba ii» 414, 1889) with A. cylindrica (as A. aegerita) and recognised considerable variation within this taxon. He suggested that several taxa may ultimately be described in the complex. A. parasitica is just one such taxon; unfortunately during the present study Spegazzini's types were not examined and so it has not been possible to compare the microscopic characters demonstrated in our material with those published for the S. American taxa. However, collections from Brazil examined by one of us (RW) demonstrate that quite a different fungus is at least present there. Agaricus phylicigenus Berkeley from Tristan da Cunha is also claimed to be A. cylindrica (Singer, 1969), but the type in K is so badly preserved that it does not help to clarify the situation.
We have compared our material with the following collections of A. cylindrica from Europe (all in E) - The Netherlands; on Populus. Italy: on Ficus, Genova, Erbar Crittogam. Ital., Ser. II. British Isles: Petersham, Surrey, 27 v 1908, legit Hartley Smith; on Populus, Hereford, Ross-on-Wye. 29 x 1969, Orton 3503; on Acer campestre L., Archer Wood, Coppingford, Huntingdonshire, 5 v 1973, legit Sheila S Wells; on Populus, Offord, Huntingdonshire, 7 vii 1974, legit S. Wells; on Sambucus, Castor Henglands NNR, Cambridgeshire, 3 x 1976, legit S. Wells. Other material from the British Isles was examined during the preparation of the British Fungus Flora (Watling, 1982) and the data obtained there from has been used in this study.
In the Dutch and some English material on Populus the pleurocystidia are very rare, and the cheilocystidia vesiculose to ventricose and never with a long appendage. This is in fact parallel to material collected in Kashmir (on Populus alba L., Gulmarg, 28 ix 1978, Wat. 13067 in E & Abraham - 6RRL 7748 in E); see Watling & Gregory, 1980 and Watling & Abraham (ined.). In those British collections with abundant pleurocystidia these are always clavate, although often with a long pedicellate base, and subhymenial or even tramal in origin. In this way they resemble collections from Afghanistan (in garden, Kabul, 12 v 1951 legit Gehli; on Populus pyramidalis Rozier (= P. italica (Duroi) Muench) Kabul, 21 v 1951; both det. R. Singer in MICH), although the basidiospores of these agree more with A. parasitica (11-12 x 6-7 µm).
Thus whereas the European collections appear to be heterogeneous, those from New Zealand are uniform and characterised by the following cluster of characters;- relatively large basidiospores with small, although quite distinct germ-pore, elongate cheilocystidia and abundant development of pleurocystidia with long apical appendage.
Taylor 867 is notable in showing an excellent development of hyphal material from its bivelangiocarpic growth in parallel to the condition described for A. cylindrica by Reijnders (1979); however, this collection consists of immature basidiomes, judging from lack of spore-development; the basidiospores available are in fact comparatively small.
Some variation is found in the detailed structure of the pileipellis in the collections of A. parasitica but this simply reflects the state of maturity. At first the pileus is covered in hyphal strands under which a palisadoderm is formed; as the pileus expands these hyphae become intermixed with ellipsoid-pedicellate to spheropedunculate cells and at maturity are even joined by hyphae derived from the pileus-trama. This may explain the variation found in A. cylindrica where rugulose and smooth-capped forms occur but more critical notes are required on individual collections to ascertain the significance of such a character and whether it is correlated with other micro-data.
The variation in basidia, cystidial morphology and their abundance in Brazilian collections examined by one of us (RW) indicates that almost totally 2-spored forms are common there, 2- and 4-spored basidia were found on many of the New Zealand collections, e.g. Taylor 867, and in European collections from the Netherlands and from England (Orton 3503). Singer (1969) indicates that similar variation is found in the S. American collections he has examined, he also indicates that within these same collections of A. cylindrica (as A. aegerita) he found populations in which the basidiospores are "distinctly truncate whereas in others they are not, populations in which the pleurocystidia are acute and others in which the pleurocystidia are obtuse ventricose-ampullaceous, and populations which are exclusively 2-spored and others exclusively 4-spored.
As the basidiospores of members of the A. cylindrica group germinate so readily and the axenic cultures prepared fruit readily these agarics offer ideal subjects for the study of variation and interfertility. Esser and his colleagues (Esser, Semerdzieva & Stahl, 1974; Esser & Meinhardt, 1977; Meinhardt & Esser, 1981) have demonstrated the ways by which variability in European isolates can be studied, indeed they have analysed some 2-spored forms. Thus the conspecificity or autonomous nature of the various isolates in the A. cylindrica complex could be determined in parallel to the recent techniques used for analysing ranges of Armillaria isolates. A. cylindrica is in fact cultivated commercially in Europe; see Ferri, 1973.
In Taylor 1263 the basidiospores were slightly larger especially in breadth, 12.5-15(-16) x 8-10(-11) µm, and may represent an independent but closely related taxon but without field data it is impossible to decide, especially as only two collections are available and each may represent the two extremes of a range of variation. In Segedin 1220 the basidiospores had a slightly excentric germ-pore.
In the literature 'Naucoria' pediades is recorded in Colenso (1886) and Massee (1898) but this taxon has undoubtedly been confused in the past. Colenso b 113 is a mixed collection of which the smaller element refers to A. pediades; the other part of b 113 refers to A. semiorbicularis q.v.
Pileus (20-) 33-80 mm, convex with inrolled margin, purplish brown when very young soon yellow ochre to yellowish, browning with age, smooth, texture of kid-glove; margin smooth with narrow greyish border where flesh is thin, when immature persistent purplish brown, often with velar remnants. Stipe 60-80 x 6-8 mm, equal or tapered upwards from swollen or bulbous base, annulate, whitish and striate at apex, yellowish below ring but browning on handling, solid then hollow, attached to conspicuous white mycelial cords at base; ring membranous, superior, whitish, darkening with age. Gills adnate-sinuate, crowded, pale cream-colour at first becoming pinkish brown-grey finally darkening. Flesh relatively thick, pale cream-colour with a grey line above gills, creamy white, fibrous in stipe. Smell sour.
Basidiospores deep brown in mass, 8-9.5(-10) x (4.5-)5-6 µm, ellipsoid, smooth, thick-walled, relatively strongly coloured in water and aqueous alkali solutions; germ-pore central, distinct. Basidia 4-spored, 20-25 x 6-7.5 µm. Cheilocystidia a mixture of clavate to vesiculose cells (30-45 x 18-20 µm) with a few ventricose cells 15-20 x 5-10 µm; pleurocystidia frequent, prominent, mucronate, lageniform or even ampulliform, 45-55 x 18-25 µm, sometimes with minute granulations at apex. Hymenophoral trama parallel with a few oleiferous hyphae. Pileipellis a palisadoderm of spheropedunculate cells 17-38 µm diam. Clamp-connections present.
Colenso b 283 cannot be added to this list meantime although Horak (1971b) states 'all found characters correspond well with those of the type' i.e., A. praecox.
A widespread agaric of gardens and fields in western Europe from late spring until summer even into late August in more northerly regions; probably introduced into New Zealand. Material has been compared with that from Europe (Scotland; Orton 2101-3 inci & Wat. 11745 in E). This is a world wide species of cultivated and disturbed areas: it is a rather variable species, or more likely a complex of taxa; see Walling, 1985. Cultural studies would be valuable. This species has been recorded by Colenso (1886) and figured by Massee (1898) The description is taken from Taylor's collections although the material in PDD accompanied by short descriptions agree in all major details.
A collection in PDD (on sawdust, Mt. Albert, Auckland City, ix 1969, legit K. R. W. Hammett, PDD 29208) thought to be A. praecox has processes on the pleurocystidia similar to, although less distinct, than those in A. arvalis. This type of pleurocystidia is also found in A. acericola (Peck) Singers. Overholts and the Hammett collection may represent this widespread N. American taxon. See above.
A collection from Otago (amongst Ammophila in dunes, near Brighton, 28 xii 1969, Austwick 1921) may also represent a slender A. praecox but in the absence of field-data, other than an uninformative pencil sketch, this cannot be confirmed; in such areas in Britain A. sphaleromorpha (Bull.: Fr.) Fayod might be expected. Vesiculose cheilocystidia could not be located although typical ampulliform pleurocystidia clearly approached the gill-edge; some basidiospores in any one mount were also rather large, e.g. 11-12 x 7-8 µm. Equally, filamentous cheilocystidia taken as characteristic of A. cf. puiggarii were not found; see below.
Leucoagaricus holosericeus (Fries) Moser apud Gams
As Agaricus praecox, in field, Napier, Colenso, b 283 and retained there by Horak (197ib) but we differ in our interpretation of the material. The basidiospores are metachromatic, dextrinoid (9.5-10.5 x 6 µm) and the cheilocystidia characterise a member of the L. naucina group. Parallel species are known from New Zealand and they may well have been introduced; they are generally more frequent in countries with warmer climates. A collection has been located in PDD (40317) filed under Armillaria.
This species was originally described, as Pholiota based on a collection from grassy fields at Apiahy, Brazil. Singer (1950) recognised it as an Agrocybe and placed it close to A. praecox and A. molesta (Lasch) Singer (= A. dura (Bolt.: Fr.) Singer).
There is evidence that this species may be more widespread than the literature suggests and that it may occur in New Zealand. Three collections with characteristic flexuous filamentous to lageniform cheilocystidia and mucronate-vesiculose pleurocystidia were located amongst the material in PDD. Unfortunately little field data accompanies the collections so the identifications are meantime preliminary. It, however, indicates that yet another member of Agrocybe sg. Agrocybe is to be found in New Zealand and fresh collections to substantiate the record are urgently required. The New Zealand material had the following characters:
a. Broadly ellipsoid to ovoid basidiospores 10.5-14 x 7-8(-10) µm,
b. Basidia mostly 2-spored,
c. Absence of pleurocystidia and
d. Lageniform cheilocystidia with ± subcapitate head.
It is widespread on lawns and in grasslands. Details of the New Zealand collection are as follows;-
Basidiospores 14-16 x (8-)8.5-ll µm, ellipsoid-ovoid, medium ochraceous in water, darker in aqueous alkali, smooth; germ-pore distinct. Basidia 2-spored, a few 3- and 4-spored basidia seen, 25-30 x 7.5-10 µm. Cheilocystidia 20-30 x 5-7.5 µm, ampulliform with slightly swollen obtuse apex (3-5 µm broad); pleurocystidia absent. Caulocystidia subcapitate to slightly swollen at apex, 20-30 x 6-8 µm, apex 4-5 µm.
A collection in K labelled Agaricus erebius (H1257/80) is undoubtedly a member of sg. Agrocybe possessing spores (7-) 7.5-8.5(-10) x 5.5-6(-6.5) µm with a distinct truncate germ-pore. In the A. erebia group the spores are fusiform to boletoid and lack a germ-pore (i.e. sg. Aporus sect. Velatae). Horak (1971b) also infers that this is a member of the genus Agrocybe. The dark coloured pileus suggested from the original naming indicated that this is very different from anything we know. It is too young to extract any further information.
The K material of seven basidiomata and one isolated pileus possesses the following characters (compounded from all specimens);
Basidiospores (7-) 7.5-10 x 5.5-6(-6.5) µm, ovoid, broadly elliptic in face-view, slightly flattened in side-view. Basidia 4-spored, c. 25 x 7.5 µm, clavate. Cheilocystidia gill-margin grazed; pleurocystidia ventricose to shortly lageniform, 25-35 x 9-15 µm, apex 7.5-8 µm broad. Pileipellis damaged, a mixture of filamentous and some vesicular cells.
Another specimen in PDD, 682 also labelled Pholiota near erebia (on ground, Weraroa, 3 x 1919, G H Cunningham), is heavily moulded but might represent the same species; both lageniform cheilo- and pleurocystidia were located. It has similarly shaped spores of parallel size and although the basidiomata are poorly preserved similar pleurocystidia to those on the K material were found (27-38 x 12.5-14 µm, apex 5.5-8.5 µm); cheilocystidia are present, vesiculose to mucronate with obtuse apex, 20-25 x 13-14 µm, apex 4 µm.
A collection also in K simply labelled 'Agaricus strophosus Fries, New Zealand' is not a species of Hebeloma as would be assumed from the use of Fries' name but a species of Agrocybe. We are grateful to Egon Horak for drawing this Colenso collection, which appears to be from Wairarapa (Massee, 1898), to our attention. The collection consists of a partially expanded basidioma with distinct velar remains on both the pileus-margin and stipe. The gill-margin is badly damaged but the basidiospores are characteristically bolbitiaceous, with thick-wall and large, truncate germ-pore; the basidia are 4-spored.
It can only be assumed that the basidioma when fresh was relatively pale-coloured from the epithet chosen but it is impossible without field data to trace the relationships of this collection. The following additional data are offered:
Basidiospores 10-12 x 7-8.5 µm, ellipsoid, slightly flattened in side-view, smooth, thick-walled, ochraceous brown in water, darker in aqueous alkali solutions; germ-pore distinct, broad. Cystidia not found.
This species is recorded in both Colenso (1890), and Massee (1898).
A. temulenta is sporadically recorded in Europe in grassy places, often in previously disturbed areas and one of us (RW) has seen a recent collection from North America (New York State). It is easily recognised when young by the uniformly bright ochraceous yellow pileus and stipe, which fade and become flushed with saffron, and the stuffed stipe with tough cortex. The basidiospores, (10.5-)11-13(-14.5) x (6.5-)7.5-8.5(-9) µm, are ellipsoid and basidia 2-or 4-spored.
Basidiospores smooth, relatively thick-walled, with distinct, often very prominent germ-pore, rust-colour in mass. Basidia 4-spored, very rarely 2-spored, and in one New Zealand species more than 4-spored, usually separated by distinct, inflated brachycystidia. Gill-trama regular becoming irregular to alveplate with age. Cheilocystidia distinct, vesiculose to inflated, sometimes with differentiated neck; pleurocystidia if present usually similar to cheilocystidia. Pileipellis a distinct palisadoderm covered with a gelatinous pellicle; 'scalp' cellular. Stipitipellis of hyaline, cylindric, parallel cells covered in part or throughout in caulocystidia similar to those on gill-margin.
Development paravelangiocarpic.
On the ground in woods, pastures, heaths etc., on dung, plant remains and refuse, and on wood; saprophytes. Those species on wood have often been placed in the separate genus Pluteolus.
One species B. muscicola appears to be endemic and widespread in mixed forest on very soft wood in which it produces a stringy white rot; the other common species is B. vitellinus. Two further collections are at the moment unplaced.
Basidiospores orange-tawny in mass, 7.5-8.5(-9.5) x 4.5-5.5(-6) µm ellipsoid, very slightly amygdaliform in side-view, smooth, relatively thick-walled, truncate from broad germ-pore. Basidia 4-spored, clavate-pedicellate, hyaline, in 'pavement' with shorter
brachycystidia. Cheilocystidia lageniform with short to elongate neck, 17.5-35 x 6-10.5(-12) µm, apex slightly swollen 2.5-5(-7) µm; pleurocystidia absent. Pileipellis a mixture of chains of inflated cells, 14-22.5 µm long, and vesiculose to spheropedunculate cells, 29-40 x 15-26 µm, some end-cells filled with pale greyish to lilaceous brown vacuolar sap, long, flexuous, septate hyphae also present pushing through gelatinous pellicle.
The spotting, pattern of raised lines and scrobiculae of the pileus and their colouring is very variable, and probably depends on age of the basidioma and environmental conditions, although the microscopic characters are relatively constant. However, a collection on Nothofagus fusca (Horak ZT 67/192) differed in the cheilocystidia being slightly forked at their apex, the basidiospores slightly paler in overall pigmentation and slightly smaller, and the pileipellis apparently lacking numerous elongate hyphae. The collection, however, comes well within the variation accepted for B. muscicola.
Horak (1971b) agrees with our disposition of Stevenson's taxon although he prefers to use the generic name Pluteolus. Stevenson's material differs only in that a few 2-spored basidia were found; this undoubtedly explains the slightly larger spore-size for the type material; also see Stevenson, 1982a. ZT 69/46 is somewhat different in that although the immature pilei are scrobiculate the old pilei have dark raised lines upon them.
Material collected by Horak (ZT 69/109) from Tophouse Saddle, Nelson may represent a new taxon within the B. vitellinus group or an extreme form.
The persistently yellow stipe is significant, and microscopically this collection can be characterised by the innumerable, inflated vesiculose hyaline cheilocystidia and polymorphic frequently septate caulocystidia. B. vitellinus generally has a cream straw or yellowish buff stipe; B. variecolor Atk. has a deeper yellow stipe and the pileus is olivaceous.
Several collections examined possess the delicate facies of an agaric which in Europe has been interpreted as B. titubans (Watling, 1982). Without more extensive field data, however, the majority of records cannot be substantiated. A collection from Piha Valley (1 iv 1974, legit J.C. Segedin, Segedin 1171) and one from above Huka Falls (on mown grass-straw, beside Waikato River, 27 x 1982, Taylor 1260) possessed the broader basidiospores associated with B. titubans. Taylor 190 (with grass, roadside bank, Holmes St., Oamaru, 26 iv 1964, legit L R Taylor) is immature but may represent the same taxon. in the Huka Falls collection some development of biporate spores was observed, something which was found to an even greater degree in Segedin 1161 (in grass outside entrance to Kauri Glen, Auckland, 28 iv 1974). Unfortunately the little field data available only indicates that the agaric is coloured much the same as members of the B. vitellinus group.
The large number of bi-porate basidiospores resemble those of many members of the strophariaceous gastromycetoid series; similar spores have been found in collections of Agrocybe from Kashmir and Poona, India (Watling & Abraham, in press). This fungus needs to be refound in New Zealand. In some ways the collection resembles Gastrocybe lateritia Watl. and G. incarnata (Peck) Baroni, although it is a typical agaric. This is yet another example of a bolbitiaceous fungus which reduces the hiatus between the true agarics and the so-called gastromycetes (secotioid). The Segedin collection lacks the colour typical of G. lateritia and G. incarnata but resembles the fungus named (ad interim) as Bolbitius rogersii (Heim, 1968); see Walling, Quadraccia & Tabares (ined).
The reader is referred to the discussion under B. vitellinus of which B. titubans has variously been considered a synonym, variety or subspecies (Watling & Gregory, 1981); Bell's record (1983) of B. vitellinus may refer here.
Pileus conical (15-30 mm), rapidly becoming plano-convex or plane (15-60 mm), with or without an umbo, bright yellow (yellow - chrome) darker at the disc, paler towards the margin where it becomes cinnamon-brown, shining, glutinous, finally fawn throughout except for yellow centre; margin striate, splitting and partially collapsing. Stipe 70-110 x 2-3 mm, tapering upwards, sometimes somewhat wider at the base, whitish or extremely pale yellow, pruinose or farinaceous throughout becoming white-silky, stuffed. Gills free or adnexed, pale yellowish with paler watery margin, becoming fawn, finally red-brown, fairly crowded, very thin and papery. Flesh very thin, distinction between pileus and stipe obvious, whitish or pale yellowish.
Basidiospores 11.5-14 x 7-8.5 µm, ellipsoid, thick-walled, sienna in water more strongly pigmented in aqueous alkali solutions, smooth; germ-pore large, central. Basidia 4-spored, a few 2-spored, clavate-pedicellate, 21-24 x 9.5-13 µm, hyaline, intermixed with sterile brachycystidia (pavement cells) 14-17.5 µm broad. Cheilocystidia irregularly lageniform with short or long and then flexuous neck, some simply vesiculose, 23-36(-48) x 9.5-12(-19) µm, hyaline; pleurocystidia infrequent, utriform or shortly lageniform,
23.5-28.5 x 7-10 µm. Pileipellis of spheropedunculate cells. Stipitipellis of hyaline, cylindric hyphae with clusters of irregular clavate, lageniform to vesiculose cells, especially at apex.
B vitellinus is widespread in temperate Europe and N America and along with B. titubans could well have been introduced into New Zealand with stock, feedstuffs or allied material; see Stevenson, 1982a and Bell. 1983. It grows naturally on a whole range of substrates and so could easily survive in a new area. The increased frequency of biporate basidiospores in collections of B. titubans and the appearance of 5-spored basidia in Taylor 246 may indicate some genetic in balance in certain New Zealand collections.
B vitellinus is a very variable fungus for which several varieties and subspecies have been recognised depending on size and growth pattern, although in many cases these are probably simply an expression of the food status of the substrate. After further field work Bolbitius sp. 1 (ZT 69/109), described below may be found to be included within the concept of this taxon. At present the persistently yellow stipe is significant.
The only subgenus not as yet found in New Zealand is Galerella but it should be looked for in grassland, on lawns, and close to human habitation. Subgenus Conocybe sect. Singerella has been recorded from New Guinea and Malaysia; and sect. Nodulososporae from Japan and Malaysia.
The genus is probably widespread but easily overlooked as 'little brown agarics'. The species are defined not only by the fresh colours of the pileus and stipe but also by the shape and distribution of the cheilo- and caulocystidia. Fourteen distinct entities have been recognised, of which ten have been named. Nine are placed in sg. Conocybe; three are new.
Pileus 10.5-11 mm, hemisphericus deinde convexus, ochraceo-brunneus, submicaceus, hygrophanus, in siccitate vel aetate pallescens, ad discam subrugosus, ad marginem substriatus. Stipes 33 x 1 (-1.5) mm, cylindricus, ad basem subbulbosus, pileo concolore, apice minute pruinoso, deorsum fibrillosus, siccus, fragilis; annulo striato subpersistente. Lamellae adnexo-adnatae, ventricosae, ochraceo-brunneae, marginibus concoloribus vel albidis. Caro inodora et insipide.
Basidiosporae 8.5-10 x 5-6 µm, ellipsoideae. Basidia 4-sporigera. Cystidia aciei lamellarum lageniformia, 30-40 x 8-12 µm ad apicem 2.5-3 µm. Cellulae cuticulae pilei pyriformes vel spheropedunculatae 18-24µm diam. Cystidia stipitis similia.Ad terram.
Pileus ≤20 mm (-25 mm when expanded) conical, collapsing on expansion, white with faint central ochre flush in age, striate-grooved in outer half. Stipe 60-70 x 1-2 mm, tapered upwards, abruptly bulbous at the base, white satiny with traces of mealy granules, twisted, hollow, very fragile. Gills subfree, crowded, very thin in substance, sienna. Flesh thin and translucent in pileus and stipe. Spore-print sienna.
Basidiospores (12-) 12.5-14 x (7-) 8.5-9.5 µm, broadly ellipsoid to almost ovoid, thick-walled, strongly pigmented in water and aqueous alkali solutions; germ-pore prominent, large. Basidia 4-spored, hyaline, clavate with distinct pedicel, separated by distinct brachycystidia. Cheilocystidia lecythiform, 21-27 x (7-) 9-11 µm, capitulum 3.5-4.5 µm diam.; pleurocystidia absent. Pileipellis a palisadoderm of spheropedunculate cells 16-19 µm broad x <38 µm long
Pileus 6-20 mm, convex to campanuiate becoming expanded, argillaceous when wet, to paler honey-brown on drying, hygrophanous, dry, pruinose under hand-lens; margin conspicuously striate when young. Stipe 22.5-42 x 1-1.5 mm, base 2.5-3 mm, cylindric with subbulbous base, pale honey-brown, hollow, ± smooth but pruinose above and appressed fibrillose towards base. Gills adnexed, ventricose, whitish or pale becoming ochraceous rust-colour, with whitish, fimbriate edge. Flesh concolorous. Taste and smell not distinctive.
Basidiospores (7.5-) 8.5-9(-9.5) x 4-5 µm, elliptic in face-view, slightly flattened in side-view, relatively thick-walled, ochraceous brown in both water and aqueous alkali solutions, darker in the latter, non-amyloid, smooth; germ-pore prominent and broad. Basidia 4-spored, hyaline, clavate with very short pedicel, 18-22.5 x 7-9 µm. Cheilocystidia lecythiform, 15-21.5 x 7.5-12.5 (-15) µm, capitulum 3-4.5 µm diameter, hyaline; pleurocystidia absent. Pileipellis a palisadoderm of spheropedunculate to ellipsoid cells, 18-40 x 7.5-12 µm with slightly darkened pedicels and tibiiform dermatocystidia; pilocystidia 25-40 x 4.5-10 µm, capitulum 3-4 µm. Stipitipellis of filamentous, parallel hyphae covered in lecythiform caulocystidia, 20-25 x 10-12.5 µm, capitulum 3-4 µm. Clamp-connections present.
Pileus <25 mm conicus vel campanulatus, ochraceo-flavidus, ad discum fulvus, siccus, impolitus. Stipes 50-55 mm, cylindricus sed diameter variabilis, tortilis, fibrosus, politus, ad apicem pallidus deorsum fuscans, farctus, facile ab pileo secedens. Lamellae brunneo-ochraceae, subliberae; acie alba serrataque. Caro pilei straminea tenuisque; stipitis concolora.
Basidiosporae ellipsoideo-amygdaliformes, 9.5-10.5 x 5.6 µm. Basidia 4-sporigera. Cystidia aciei lamellarum irregulariter lageniformia 29-54 x 7-10.5 µm, ad apicem 3.5-6 um. Cellulae cuticulae pilei pyriformes vel spheropedunculatae vel lageniformes 23.5-36 x 6-9.5 µm. Cystidia stipitis similia.Ad terram in pascuo ovino.
Pileus <35 mm, convex-umbonate to plano-convex but retaining umbo, yellow-brown or tawny brown, drying paler (yellowish), disc rather darker, moist, matt, radially wrinkled about umbo; margin radially striate, splitting. Stipe 35-40 x 3 mm, equal but for abruptly bulbous base, concolorous or paler than pileus, hollow. Gills adnexed, yellow-brown, crowded. Flesh extremely thin in pileus, whitish, distinct from the easily removable stipe, fibrous, very brittle in stipe.
Basidiospores brownish yellow-ochre in mass, 5.5-7(8.5) x 3.5-4.5 µm, elongate-elliptic in face-view, perhaps slightly amygdaliform in side-view, smooth, non-amyloid, slightly thick-walled, ochraceous in water; germ-pore absent. Basidia 4-spored, hyaline, clavate-pedicellate. Hymenophoral trama subparallel. Cheilocystidia lecythiform, 16.5-21.5 x 6-7 µm, capitulum 3.5-4 µm, some with brownish plasmatic pigment in venter in KOH. Pileipellis a palisadoderm of pyriform to ellipsoid-pedicellate cells some with slightly brownish encrustations, 10-15 µm broad, some with plasmatic pigment, hair-like cells absent; pilocystidia detached, either long encrusted hyphae with clamp-connections (17.5-32 x (2.5-) 4.5-6 µm or short chains arising from ± vesiculose cells c. 12 µm broad, usually with swollen capitulum at apex and often containing brown plasmatic pigment. Pileus trama of hyphae with yellow contents in KOH. Stipitipellis of parallel to subparallel cells producing lageniform to cylindric caulocystidia intermixed with long flexuous hair-like cells <60 µm. Clamp-connections present in pileus trama, pileipellis and stipitipellis.
The pilocystidia are distinctive and possibly represent remnants of the original blematogen as described for C. rickeniana (Singer) P D Orton and C. mesospora (Kuhner ex) Kuhner & Watling (Watling, 1975). Similar structures have not been found previously in C. piloselloides or C. pilosella (Pers.: Fr.) Kuhner and may indicate the New Zealand material represents an independent yet closely related taxon. Strong developments of pilocystidia are found in C. horakii Walling Taylor (q.v.) and in C. dumetorum (Vel.) Svreck
The fungus is easily recognised by the small basidiospores lacking a germ-pore, strongly developed often-coloured pilocystidia and lageniform to cylindric caulocystidia mixed with hair-like cystidia. For these latter cells Walling (1964 et seq.) has erroneously used the term pilocystidia referring to the dermatocystidia of the pileus as pileocystidia. Patrick Barrows (1979) are correct in pointing out that pilocystidia a term introduced by Buller is derived from Greek where pile- refers to head and not hair (Latin). This has led to some confusion; the hair-like cells are so characteristic we believe they should be recognised as a distinct entity.
Pileus 27 mm, hemispheric, becoming convex to subcampanulate, golden tawny brown, sienna or cinnamon fawn when fresh, strongly hygrophanous, drying out whitish with ochraceous centre, dry, smooth, membranaceous; margin striate. Stipe 30-40 x 2.5 mm (base 5 mm), cylindric, annulate, white then fawn, apex pruinose, below ring longitudinally fibrillose or minutely squamulose, base dark brown, dry, fragile, hollow; ring mobile, whitish, persistent, striate-grooved above, pale woolly below. Gills adnexed to adnate, ventricose, dense, concolorous with fresh pileus or cinnamon to sienna, with whitish, minutely denticulate edge. Flesh white. Smell slight.
Basidiospores 8.5-9.5(-10.5) x 5-5.5 µm, elliptic in face-view slightly flattened in side-view, thick-walled, non-amyloid; germ-pore prominent, broad, central. Basidia 4-spored, clavate with short pedicel, 20-27 x 7 µm. Cheilocystidia lageniform with narrow tapered neck, 27-40 x 8-9 µm and obtuse apex 2-3 µm broad. Caulocystidia similar to those on gill-edge 42-51 x 9-17 µm, with apex 3-3.5 µm, intermixed with vesiculose cells, 23-32 x 9-13 µm. Pileipellis a palisadoderm of spheropedunculate cells 27-36 x 15-17 µm.
On cow dung, near Buller bridge, west of Murchison, Nelson, 25 i 1969, Horak ZT 69/20.
This collection differs from C. vexans in its habitat preference; it is unknown on dung in Europe. Two coprophilous, annulate Conocybe spp. (sg. Pholiotina), occur in North America but they differ in their microscopic characters particularly the smaller basidiospores. C. vexans is the same as Kuhner’s four-spored form of C. blattaria; see Walling & Gregory (1981) and van Waveren (1970).
A second collection, Horak, in ZT 68/440 probably represents the same taxon although the pileus was wrinkled at the centre. We do not place much emphasis on this phenomenon which is not infrequent in many members of the Bolbitiaceae; compare C. rugosa and C. filaris in Walling (1982). This collection, from under Leptospermum ericoides A. Rich., Wharariki, Cape Farewell, Nelson, 13 v 1968, has both slightly narrower, and therefore slightly fusoid spores and narrower cheilocystidia; the following description is offered:
Pileus 14-17 mm, umbonate-convex to campanulate, dark ochre-brown (wet), ochraceous (dry), hygrophanous, micaceous-shiny, at least centre wrinkled-venose, dry, striate; veil remnants absent. Gills adnate to adnexed, ventricose, pale ochre (young), chocolate brown (with rust brown tinge) when old, crowded, with white, fimbriate edge. Stipe 25-34 x 1-1.5 mm (base 2 mm) cylindrical or subclavate, annulate, concolorous with pileus, subpruinose at apex, fibrillose towards base, hollow, dry; ring persistent, striate-grooved, immobile. Smell and taste not distinctive.
Basidiospores 10-12.5 x 5.5-6.5 µm, ellipsoid in face-view, slightly flattened in side-view, thick-walled, ochraceous brown in water, slightly darker in aqueous alkali solutions, nonamyloid; germ-pore central, broad. Basidia 4-spored, 24-25 x 5-9 µm. Cheilocystidia ampulliform to cucurbitiform with distinct elongate neck, 30-37 x 8-11 µm with apex 2 µm broad. Pileipellis a palisadoderm of pedicellate cells, 22-40 x 12-22 µm.
D. majestaticaHorak, found in South Island with Nothofagus spp., D. phlebophora Horak, also found with Nothofagus but in addition with Leptospermum, and D. gunnii (Berk.) Horak based on Secotium gunnii (Gunn 257 in K) noted by Berkeley (in Massee, 1891) and originally collected at Rotorua. The last species grows with both Leptospermum spp. and Nothofagus spp. and is fairly common in coastal and submontane forests; it is also known from New Guinea (Horak, 1980d). D. gunnii is closely related to the Australian D. recedens (Cooke & Massee) Singer now known to be widespread in Eastern Australia (Horak, 1980d; Watling unpubl. data). Descolea has been recently placed in the Bolbitiaceae by Singer (1972) and Horak (1979b).
As Agaricus (Naucoria) pediades in field, Colenso 269.
This collection belongs in the genus Galerina and agrees with the illustration for Colenso 1039 (Naucoria nasuta), which appears in Horak (197ib); see below.
As Agaricus (Naucoria) pediades, under Kaiwarawarra, T. Kirk (sheet H1257/80 in K).
Resembling in all ways the Galerina figured under N. nasuta Kalchbrenner (Horak, 197ib). The basidiospores are very distinctive in that they possess a roughened calyptrospore; the pleurocystidia usually possess two lobes at the apex.
As Galera tenera, Waitaki, Berggren 60 (K); Berggren material is the subject of a paper by Cooke in Grevillea (1890). Only the following microscopic data was obtainable:-
Chrysocystidia absent. Basidiospores 16.5-18 x 9.5-10.5 µm, mid-brown ochraceous with olivaceous tinge, smooth, thick-walled, sometimes with dark inclusions; germ-pore prominent, Pileipellis filamentous. Neither cheilo- nor pleurocystidia could be located.
We agree with Horak (i971b) in placing this collection in Psilocybe. It is impossible to be any more precise even using Guzman's monograph (1983).
As Agaricus semiorbicularis, Waitaki, Berggren 61 (K). Badly consumed by insects. Only the following microscopic data was obtainable;
Cheilocystidia absent. Basidiospores 11-12 x 6.5-7(-7.5) µm, smooth, thick-walled with truncate germ-pore, sometimes with dark inclusions. Pileipellis filamentous. Neither cheilocystidia nor pleurocystidia could be found.
This belongs to Psilocybe; see comments above. We do not agree with the placement by Horak (197ib).
An illustration in E apparently is a copy by M. C. Cooke of a Berggren collection labelled 'AGARICUS (GALERA), on the ground, New Zealand'. It is impossible to go any further but it is not unlikely that this figure depicts one of the above collections of Psilocybe.
Identification keys
Bolbitiaceae
Agrocybe
Bolbitius
Conocybe
Cited scientific names
- Agaricus heteroclitus sensu Colenso 1887 [1886]
- Agaricus praecox sensu Colenso 1887 [1886]
- Agaricus tasmanicus sensu Colenso 1891 [1890]
- Agrocybe acericola (Peck) Singer 1950
- Agrocybe erebia (Fr.) Kühner 1935
- Agrocybe Fayod 1889
- Agrocybe howeana (Peck) Singer 1951 [1949]
- Agrocybe olivacea Watling & G.M. Taylor 1987
- Agrocybe parasitica G. Stev. 1982
- Agrocybe pediades (Fr.) Fayod 1889
- Agrocybe praecox (Pers.) Fayod 1889
- Agrocybe praecox sensu E. Horak 1971
- Agrocybe puiggarii (Speg.) Singer 1952 [1950]
- Agrocybe semiorbicularis (Bull.) Fayod 1889
- Agrocybe sp. 1 Watling & G.M. Taylor 1987
- Agrocybe sp. 2 Watling & G.M. Taylor 1987
- Agrocybe temulenta (Fr.) Singer 1936
- Agrocybe vervacti (Fr.) Singer 1936
- Bolbitius Fr. 1838
- Bolbitius muscicola (G. Stev.) Watling 1981
- Bolbitius sp. 1 Watling & G.M. Taylor 1987
- Bolbitius sp. 2 Watling & G.M. Taylor 1987
- Bolbitius titubans (Bull.) Fr. 1838
- Bolbitius vitellinus (Pers.) Fr. 1838
- Conocybe Fayod 1889
- Conocybe gracilenta Watling & G.M. Taylor 1987
- Conocybe horakii Watling & G.M. Taylor 1987
- Conocybe huijsmanii Watling 1983
- Conocybe mesospora Kühner ex Kühner & Watling 1980
- Conocybe novae-zelandiae Watling & G.M. Taylor 1987
- Conocybe piloselloides Watling 1983
- Conocybe pubescens (Gillet) Kühner 1935
- Conocybe pubescens sensu A.E. Bell 1983
- Conocybe rickenii (Jul. Schäff.) Kühner 1935
- Conocybe rickenii sensu A.E. Bell 1983
- Conocybe rugosa (Peck) Watling 1981
- Conocybe sp. 1 Watling & G.M. Taylor 1987
- Conocybe sp. 2 Watling & G.M. Taylor 1987
- Conocybe sp. 3 Watling & G.M. Taylor 1987
- Conocybe sp. 4 Watling & G.M. Taylor 1987
- Conocybe vexans P.D. Orton 1960
- Descolea Singer 1950
- Galerina sp. 1 Watling & G.M. Taylor 1987
- Galerina sp. 2 Watling & G.M. Taylor 1987
- Galerina sp. 3 Watling & G.M. Taylor 1987
- Leucoagaricus holosericeus (Fr.) M.M. Moser 1967
- Naucoria temulenta sensu Massee 1899 [1898]
- Psilocybe sp. 1 Watling & G.M. Taylor 1987
- Psilocybe sp. 2 Watling & G.M. Taylor 1987
- Stropharia coronilla (Bull. ex DC.) Quél. 1872