A taxonomic survey of the genus Montagnea (Gasteromycetes) with special reference to South Africa

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A taxonomic survey of the genus Montagnea (Gasteromycetes) with special reference to South Africa Derek A. Reid and Albert Eicker* Department of Botany, University of Pretoria, Pretoria 0002, Republic of South Africa Accepted 4 March 1991

This paper is a taxonomic survey of the genus Montagnea Fr. Three taxa are recognized, viz. Montagnea arenaria, M. arenaria var. macrospora var. nov. and M. haussknechtii, based largely on spore size and the number of spores produced per basidium. For each of these taxa full descriptions are provided, and their world-wide geographical distribution indicated, based on examination of herbarium material in Kew and reports in the literature. Special emphasis is given to the occurrence of these taxa in South Africa where they are collectively known under the popular name of Namaqua Black Caps. Hierdie artikel is 'n taksonomiese oorsig van die genus Montagnea Fr. Drie taksons word erken, nl. Montagnea arenaria, M. arenaria var. macrospora var. nov. and M. haussknechtii, wat hoofsaaklik gebaseer word op spoorgrootte en die getal spore wat per basidium voortgebring word. Volledige beskrywings word vir elkeen van hierdie taksons voorsien en hul wereldwye geografiese verspreiding, gebaseer op die bestudering van herbariummateriaal in Kew en verslae in die literatuur, word voorsien. Besondere aandag word gegee aan die voorkoms van hierdie taksons in Suid-Afrika, waar hulle gesamentlik onder die populere naam Namakwa swartmusse bekend staan. Keywords: Gasteromycetes, Montagnea arenaria, Montagnea arenaria var. macrospora, Montagnea haussknechtii, Namaqua Black Caps 'To whom correspondence should be addressed.

Introduction

A collection of Montagnea Fr. was made by one of us (D.A.R.) in the Roodeplaat Nature Reserve, near Pretoria, Northern Transvaal. Attempts to name this woody, agariclike, coprinoid Gasteromycete in Bottomley's (1948) monograph of the Gasteromycetes of South Africa failed since she considered it to represent a true agaric genus and so excluded it from consideration. However, the genus Montagnea became familiar, if not by name, to a fairly wide public in South Africa with the publication of the Afrikaans edition of the mushroom field guide of Levin et al. (1987), Veldgids tot die sampioene van Suid-Afrika, where it was very well illustrated under the rather seductive popular name of 'Namaqua Black Cap'. However, its true identity remained a mystery and no Latin name was cited. Indeed, the authors commented that the family name and scientific name was as then unknown to them. Attempts to rectify this situation led to a revision of the genus Montagnea. It soon became obvious that two species of Montagnea occurred in South Africa, viz. the Small Spored Namaqua Black Cap (M. haussknechtii Rab.), with small elliptic spores, 6.0 - 8.0 (-10.0) X 3.0 - 5.0 (-6.0) J.l.m, appearing diamond-shaped in optical section, and the Medium Spored Namaqua Black Cap [M. arenaria (DC.) Zeller] with much larger elliptic to broadly elliptic spores, (11.5-) 13.0 - 16.0 (-17.0) X 7.5 - 10.5 J.l.m, appearing broadly ovoid to pyriform or pip-shaped in optical section. In both species the spores are the product of 4-spored basidia. However, it also became obvious that in certain parts of the world, notably Australia, an even larger spored variant of M. arenaria occurred with spores measuring (14.0-) 16.0 - 23.5 (-27.0)

X 9.0 - 14.0 (-16.0) J.l.m, and that these were the product of predominantly 2 - 3-spored basidia. We have proposed naming this last-mentioned taxon, as yet unrecorded from South Africa, the Large Spored Namaqua Black Cap (M. arenaria var. macrospora Reid & Eicker, var. nov.).

Descriptions

Montagnea Fries, Genera Hymenomycetum 7, 1836. Montagnites Fr., Epicrisis 240, 1838. Sporophores with a dried-up wizened, coprinoid aspect, and with a tough woody stipe. Pileus l.5 - 6 cm diam., conicotruncate, truncate-campanulate, or convex, often with a slight central umbo, pallid, but with exceedingly thin flesh which is stretched over the gills and soon ruptures radially between them so that the major part of the surface appears silvery grey to blackish where the gills are partially to fully exposed from above. Lamellae crowded, black and radially orientated, partially to fully exposed by rupture of the very thin overlying flesh, often said to be unbranched, but we suspect some forking and anastomosis may occur, although this needs confirmation from living material. When exposed in this manner the attachment of the lamellae to the central disc appears very slight and precarious, and to involve merely their extremity nearest the stipe. They, therefore, appear to radiate from the central disc much as would the spokes of an umbrella when the covering material has been tom away from all but the central area. This led early workers to postulate erroneous theories of lamellar development. Stipe 2.5 - 30.0 cm tall, 0.3 - 2.0 cm wide, tough and woody, longitudinally wrinkled, surface smooth to fibrillose, often with adpressed scales, pallid, with a fringed papery

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volva. Volva of young specimens said to be gelatinous (Savulescu, 1935). Annulus lacking. Hymenium comprising basidia, or basidia separated by pseudoparaphyses. Cystidia have been reported by some authorities but their presence needs confirmation; they have not been observed in this study. Basidia difficult to observe in dried material as they collapse to an agglutinated honeycomb-like structure in which individual elements will not separate, but either (1)2-3-4(5)-spored. Spore mass black. Spores sessile, dark brown to black-brown, elliptic with a distinct wall, and a hyaline germ-pore, which is often conspicuous and very eccentric. In M. haussknechtii 6.0 - 8.0 (-10.0) X 3.0 - 5.0 (-6.0) J.1m, elliptic in side view but angular-shaped in optical section; in M. arena ria (11.5-) 13.0 - 16.0 (-17.0) X 7.5 - 10.0 J.1m, elliptic to broadly elliptic in side view, ovoid to pyriform in optical section; in M. arenaria var. macrospora borne on (1-)2-3(-4)-spored basidia, and measuring (14.0-) 16.0 - 23.5 (-27.0) X 9.0 - 14.0 (-16.0) J.1m. In the latter taxon 3-spored basidia dominate in some fruitbodies. Clamp connexions are reported by Singer (1976), but we have been unable to demonstrate their presence. Habitat : in dry, open, sandy situations such as coastal dunes, steppes, savannah and deserts. Discussion When Fries (1836) first conceived the genus he named it Montagnea Fr. in honour of the French mycologist C. Montagne, who had initially sent him the dried material which formed the basis of his generic description. Later Fries (1838) changed the epithet to Montagnites Fr. to avoid confusion with the generic name Montafioa Lallav. & Lex. (Compositae), which De Candolle (1836) emended to Montagnaea DC. in order to more correctly express, when latinized, the name of the gentleman to whom the genus was dedicated - Dr Lud. Montano. However, the publication of the emended name Montagnaea DC. did not appear until September 1836, whereas Montagnea Fries was published in April of the same year and therefore has clear priority. Despite this the name Montagnites received widespread acceptance in mycological circles until relatively recently. Montagnea Fr. has been made the type genus of the Montagneaceae Sing. (1976), along with two other genera Panaeolopsis Sing. and Polyplocium Berk. However, Singer (I.c.) observed that the affinities of his new family Montagneaceae lie with Coprinaceae in the Agaricales. The same author, Singer (1986), wrote to Montagnea 'Many authors consider this genus as belonging about the Agaricales, somewhere near Coprinus. There is no doubt in the author's mind but that Montagnea is, indirectly, related with the Coprinaceae. However, if there is such a thing as Gasteromycete, Montagnea is one of them.'

Montagnea haussknechtii Rab., Sitzungsber. natuTW. Ges. 'Isis' Dresden 8, 1870. (Figures 1 A-F; 2.) Montagnites elliottii Mass., Grevillea 21, 1, 1892. Montagnites tenuis Pat., fl. Bot. 8, 219, 1894. Montagnea tenuis (pat.) Teng. Fungi of China 762, 1964. Montagnites candollei var. minor P. Henn., Hedwigia 40, (98), 1901.

Montagnites candollei var. coprinoides P. Henn., Hedwigia 40, (98), 1901. Montagnites candollei var. somala Bacc., Eumycet. Somalia 191, 1916. Montagnites spegazzinii Sacco & Trott., SyU. Fung. 23, 327,1925. Sporophores coprinoid, centrally stipitate with a hard woody stipe. Pileus 1.5 - 3.5 (-5.5) cm diam., usually truncate campanulate or truncate-hemispherical, occasionally truncate-cylindIic, more rarely becoming convex or applanate, sometimes with a capping of sand particles. Flesh of pileus reduced to a delicate membrane stretched over the lamellae, but thicker at the centre, and at an early stage splitting radially to the vicinity of the disc, such that a very thin and very narrow strip of tissue adheres to the dorsal surface of each gill, but sometimes sloughing off completely toward the margin of the cap. The gills may then appear free and to hang vertically downward attached to the disc only by their width (see Levin et al. 1987). Colour ranging from dirty o\)

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Figure 1

A-F. Monlagnea haussknechlii. A. Spores. B. Spores 740. C. Hymenium of collapsed and agglutinated basidia. All from Van Wyksfontein, Acocks (328). D. Spores. E. Spores X 740. Both from Roodeplaat specimen. F. Spores X 740, from holotype of M. elUol/ii. G-I. Monlagnea arenaria. G. Spores X 740. H. Spores. I. Hymenium of collapsed basidia and pseudoparaphyses. All from Andriesfontein, Acocks (408). J-K. Monlagnea arenaria var. macrospora. I. Spores X 740. K. Spores. Both from holotype. L. Monlagnea sp . Sinclair, New Zealand. All X 340 except where otherwise indicated. X

S.AfrJ.Bot., 1991,57(3) white at the disc, where the surface may disrupt into small adpressed fibrillose spot-like scales, or remain intact, or rupture radially to give an almost cog-wheel effect, passing through a silvery-grey zone where the flesh has been stretched or partially ruptured, and finally to the broad blackish outer region where the lamellae are exposed. Lamellae black, very crowded, radially orientated, probably forking but this needs confmnation from a study of fresh material. Stipe woody, 6 - 9 (15) cm high, equal, or more usually markedly tapered downward, from 0.3 - 0.5 (1.0) cm at the apex to 0.5 - 1.5 cm at the base, where there is a small, rather inconspicuous, fibrillose volva, with a fringed margin, which is often left behind in the soil unless the specimen is carefully excavated; whitish, longitudinally wrinkled, with a smooth or fibrillose surface, which may occasionally disrupt to form coarse squarrose scales toward the base, or bear small, scattered patches of tissue elsewhere. Hymenium in dried material comprising a uniform brown honeycomb-like surface of collapsed and strongly agglutinated. 4-spored basidia, which are impossible to separate individually in microscope squash preparations. Spores 6.0 - 8.0 (-10.0) X 3.0 - 5.0 (-6.0) 11m, black in the mass, but brown under the microscope, elliptic in side-view, but appearing angular and diamond-shaped or ovoid in optical section, with a slightly thickened wall, and a germpore which is often placed very obliquely, sessile, apiculus virtually lacking. Distribution

In open dry sandy places, in grassland or savannah country, etc. South African specimens in K: Cape Province: in grassy places, on banks of river soil. Van Wyksfontein, Kimberley District, J.P. Acocks (328), 22 May 1936; in sandy soil, Kalahari Gemsbokpark, J.V. van Greuning, 14 May 1988. Eastern Transvaal: from open bare sandy soil in savannah, Roodeplaat Nature Reserve, near Pretoria, D.A. Reid, Veldie and Martmari van Greuning, 26 Feb. 1989. Outside of South Africa M. haussknechtii has a wide geographical distribution extending throughout Africa including the Canary Islands, the Middle East, Western Asia, and possibly also Australia and South America as follows: Africa East: Somalia: (Baccarini, 1916 - type of Montagnites candollei var. somala, with spores 7.2 X 4.5 11m,

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almost certainly belongs here). Africa West: Canary Islands: Las Palmas (Spegazzini, 1915; Saccardo & Trotter, 1925 - type of Montagnites spegazzinii, described with spores 8.0 - 10.0 X 4.0 - 4.5 11m, almost certainly belongs here). Angola: in sandy dry ground amongst grasses, Maionga, Loanda, J. Gosseweiler (289), 1921 (in K); Africa North: Algeria: near Biskra (Hennings, 1901 - two collections forming the basis of M. candollei var. coprinoides Henn. are somewhat dubious on account of the slightly larger spores, quoted as 7 - 11 x 4 - 6 11m). Tunisia: in sand, Gares (patouillard. 1894 - type of M. tenuis). Egypt: amongst sand, Nile Valley, Scott-Elliott (type of M. elliottil) Massee 1892, in K); Wadi RicMd. pro Hekian, 19 Feb. 1900; 11 March 1900 (Hennings, 1901 type of M. candollei n.var. minor). Africa East-Central: Northern Sudan: solitary in sand, sand-dunes west of massif, Jebel Tageru, K. Neumann (230a), 28 Jan. 1964 (in K). Eritrea: on rocky ground. Asciorum Pass, 2800 ft. alt., P.O. Bally (B. 6753), 23 March 1949 (in K). Middle East: Israel: (Reichert & Avizohar-Herschenzon, 1959; Dring & Rayss 1963). Transjordan: Kalirrhoe, Prof. T. Rayss, 10 March 1936 (in K). Jordan: in sand, Rum, 100 m. alt., D. Poore & J. Gillet (5993/A), 29 April 1963 (in K). Bahrain: in sandy gully, D.A. Bellamy (6), April 1981. Iran: in sandy soil, Enzeli, south-west coast of Caspian Sea, Prof. Haussknecht (Rabenhorst, 1870 - type of M. haussknechtii). Aden: (Hollos 1904). Oman: on sandy bank of small dry water-course, amongst rocks, near Masafi (25°19'N, 56°08'E), 1500 ft, J. George (32), 9 Feb. 1970 (in K). Western Asia: Russia: Asiatic: Aral Sea (Sorokin, 1884). West Pakinstan: in sandy soil, common, Sangla Hill, Panjab, Sept. 1938 (in K). Kashmir: (Hongo, 1965). Asia: China: Nei Monggol, Xinjiang (Liu 1984). Australia: (Cleland 1934 - it is possible that the collection cited from Waitpinga, Encounter Bay, with spores elliptical to almost triangular 7.5 - 5.5 11m, may belong here but we have not studied the material). South America: Argentina: (Spegazzini 1912 - a number of forms are discussed under Montagnites argentina of which form 'a' with spores elliptic, 8 - 10 x 3 - 4 11m from Cacheuta, prope Mendoza, Mat. 1910, could represent M. haussknechtii, but we have not examined the material). Illustrations

Coloured illustrations are to be found: Baccarini (1916), Tab. 5; Rabenhorst (1870), Tab. 3, Figure 1; and black-andwhite illustrations: Dring & Rayss (1963), PI. lA - holotype of M. elliottii; PI. IB upper figure only; Hongo (1965), Figures 4 - 5. Discussion

Figure 2 Montagnea haussknechtii. Dried fruitbodies, Gemsbokpark 14 May 1988, PRUM 2475. 65% of natural size.

Hitherto there has been considerable reluctance to accept more than one species in the genus Montagnea Fr., and even as recently as 1963 Dring & Rayss (1963), despite the evidence of collections with highly divergent spore sizes, were unwilling to follow Zeller (1943) and accept that the type material of Montagnites elliottii Massee, with irregularly ovoid spores, 6.0 - 8.0 x 4.5 - 5.5 11m, represented a species distinct and apart from M. arenarius which latter taxon Zeller interpreted as having large ellipsoid to ovoid spores measuring 12.0 - 19.0 X 6.0 - 11.2 11m. Dring & Rayss (I.c.) merely remarked, 'We are not convinced that it

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is sufficiently different from M. arenaria to merit specific Another aspect of the holotype of this species is that the rank, but we think that it is better considered merely as form surface of the stipe is silky fibrillose and without scales. of M. arenaria'. This feature was the subject of the comment by both Zeller Earlier Reichert and Avizohar-Hershenzon (1959), after a (l.c.) and Dring & Rayss (1963). However, we do not, for the moment, consider this unduly significant, as a number of study of collections available to them, and a survey of the world literature, were able to distinguish under the comprethe small-spored collections we have studied show a tendhensive name M. candollei [= arenaria] , three types of ency towards a fibrillose surface to the stem, especially in fungus based on spore data: with small spores 5 - 8 x 4 - 5 the lower region. !lm, with medium spores 12 - 16 x 5 - 7 !lm, and with Judged from the published description of Patouillard large spores 20 - 26 x 9 - 15 !lm. (1894), Montagnites tenuis Pat. is unusual in having the pileus truncate-cylindric, but again we do not consider this This seems a convenient starting, although as more data become available from different geographical areas it may feature significant. The specimen from northern Sudan cited require modification. Also it should be noted that in some above also appears to have had a similarly shaped pileus. fruitbodies belonging to the first group, occasional larger The only comment which needs to be made in relation to spores may be present which, if incorporated into the overall the South African collections of M. haussknechtii is that the spore range, would blur the distinction with the other sporophores from the Kalahari and the Roodeplaat Nature groups. However, it is usually obvious from a study of the Reserve show spores which, in optical section, appear more angularly diamond-shaped than is perhaps usual. However, overall spore picture to which group a particular sporophore should be assigned. sporophores of Montagnea are rather persistent and it is The oldest available name for the small-spored taxon is possible that prolonged weathering may have had an effect Montagnea haussknechtii Rab. based on material collected on spore shape. from the south-western shore of the Caspian Sea on the Iranian side. We consider the following to be synonyms: Montagnea arenaria (DC.) Zeller, Mycologia 35, Montagnites elliottii Massee from Egypt, M. tenuis Pat. 418, 1943. (Figure 1 G-l). from Tunisia; M. spegazzinii Sacco & Trott. from the Canary Agaricus arenarius DC., Flore Franc. 6,45, 1815. Islands, together with the following, each published as a Montagnites arenarius (DC.) Morse, Mycologia 40, 256, variety of M. candollei: M.c. minor P. Henn. from Egypt, 1948. M.c. eoprinoides P. Henn. from Algeria (this taxon has Montagnites candollei Fr., Epicrisis 241, 1838. somewhat larger spores than is usual, i.e. 7 - 11 x 4 - 6 Montagnea candollei (Fr.) Corda, leones Fung. 6, 85, !lm, and on that account its relegation to synonymy under 1854. M. haussknechtiiis slightly dubious) and M.c . somala Bacc. Montagnea candollei var. texensis Berk. & Curt. Ann. Montagnites elliottii was described from specimens colMag. Nat. Hist. Series 2, 12,422, 1853. lected by Scott-Elliott in the Nile Valley, but the locality Montagnites argentina Speg., An Mus. Nac. Buenos Aires was later erroneously cited by Saccardo (1895) and others as 6, 160, 1899. New Zealand. Sporophores of the holotype are taller than Montagnites radiosus var. isosporus Belli, Ann. Bot. 6, the other specimens available for study: Massee (1892) 526, 1908. described them as 10 - 15 cm tall. At a casual glance the ? Agaricus radiosus Pallas, Reise Russischen Reichs 2 Kew material appears to represent a single sporophore with (2), 744, 1773 (see epithet on Tab. 'W' Figure 3). the stipe broken in two and the respective portions mounted ? Montagnites radiosus (pallas) P. Henn., Hedwigia 40, side by side. However, in fact the two portions are from (98), 1901; see also Hollos, Gasterom. vonotk. helyesb. different fruitbodies, one comprising a pileus and the stipe Termeszetr. Fuzet. 25, 96-98, 1902. tapering downward but lacking a base; the other with a base ? Montagnea radiosa (Pallas) Sebek, Ceska Myk. 8, 144, but no pileus. 1954. When Massee (l.c.) described the species he published an ? Montagnites pallasii Fr., Epicrisis 241 , 1838. illustration, which not only figures a mature sporophore, but also showed sections through the developing fungus. Judged Sporophores with a dried-up wizened appearance, from from these illustrations the young fruitbody develops within short and squat to tall and erect, coprinoid, with a central a membranous 'egg', but these figures appear very schemwoody stipe. Pileus 2 - 4 (-6) cm diam., conico-truncate or atic, and there are no immature specimens in the Kew truncate-campanulate to convex, often with a slight umbo, collection which could have formed the basis for Massee's and exceedingly thin flesh, which is stretched over the gills figures. However, the holotype was evidently divided, with and soon ruptures radially between them. At maturity the a part being retained at K and part going to NY, for Zeller cap colour passes from whitish at the intact or somewhat (1943) comments on the spores of the NY material, "spores scaly disc, through silvery grey where the underlying gill in the type of M. elliottii (in NY Bot. Gard. Herb.) ovate colour is apparent, to a broad greyish-black zone extending and 6.0 - 7.75 x 4.5 - 5.0!lm instead of '12 x 7' as in which the dorsal surface of the gills is fully to the margin described by Massee". Zeller (l.c.) makes no mention of exposed. Lamellae numerous, black, radially orientated, any immature developmental stages being represented in probably somewhat forked, or anastomosing. Stipe 2.5 - 24 NY. However, his observations on the spore size of the material in NY apply also to the spores present on the K cm tall (even taller in some American specimens), 0.4 - 2.0 portion of the holotype. Clearly Massee (l.c.) was in error cm wide, equal, woody, pallid, longitudinally wrinkled, and when he cited the spore size of M. elliottii as 12 x 7 !lm. either smooth to somewhat fibrillose, often bearing large flat

S.Afr.J.Bot., 1991, 57(3) adpressed scales, and at the base an inconspicuous fringed papery volva, which in the young specimens is gelatinous (Savulescu 1935). Hymenium comprising 4-spored basidia, which appear to be more or less regularly disposed over the gill surface, separated by pseudoparaphyses, as illustrated by Hollos (1904). The basidia do not collapse into a honeycomb-like pattern as in M. haussknechtii, but appear more discrete. Spore powder black. Spores (11.5-) 13.0 - 16.0 (-17.0) X 7.5 - 1O.5/lm, elliptic to broadly elliptic in side view but ovoid to pyriform or pip-shaped in optical section, sessile, dark brown, with a conspicuous, hyaline, very oblique germ-pore. Distribution In dry, open sandy localities, in warmer parts of the world, including coastal dunes, steppes, deserts etc. South African specimens in K: Cape Province: in red sand on top of Langeberge, Andriesfontein, Barkly West, J.P.H. Acocks (408), 18 June 1936 (No. 28643). [Illustrated in Dring & Rayss 1963, PI. 'B' lower figure.] Outside of South Africa M . arenaria has a very wide geographical distribution throughout the tropical and warm temperate regions of the world along beaches or in desert or steppe conditions. There are specimens in K from Algeria: ex. Herb. Berkeley, no further data; in desert, 50 paces from an artificial waterhole near cypress trees, Sahara Edjeleh, Mrs. Dean Netscher 1972. Southern France: Monspelii (specimen from Montagne); ad mare in sabulosis, circa Monspelium, Petit detit, Martio 1824, Herb. J. Gay; Gall. merid., Prof. Mertens, 1806, Herb. Dawson Turner. Egypt: Shakrat el Delem, G.W. Murray (3692), 1925 - 1926 (Herb. D.N. Simpson) near Mersa Shat, G.W. Murray (3748), 1925 - 1926 (Herb. D.N. Simpson). Crete: in pure sand on beach, Mallia, D.A., D.G. & P.M. Reid, 2 June 1966. Arabia: in hard damp sand near granite outcrop 5 km north of road, 3000 ft. alt., J.S. Collenette (5729). 11 March 1986. Romania: bord de la chaussee, terrains sablonneux, Dobrogea, dist. Constanta - intre Babadag si Ienisala, Tr. Savulescu & AI. V. Alexandri (Herbarium Mycologicum Romanicum, Fasc. X, No. 481). West Pakistan: Karachi, S. Ahmad (254), 18 Aug. 1967; in sandy soil, Ledhar (Sheikhupura), G.A. (2716), 15 Aug. 1948; in sandy soil, Lahore, G.R. (15687), 25 Oct. 1960. Tibet: Kambajong, Major D. Prain, Sept. 1903. United States: Seattle, bank of Columbia River, a few miles from Ginkgo Museum, Miss M. Holden, 5 Sept. 1969; Texas, C. Wright (as M . candollei var. texensis B. & C.); New Mexico, C. Wright ex Curtis (as M . candollei B texensis ex Ravenel's Herbarium - recd 1891). Additional South African collections are listed by Doidge (1950) under Montagnites candollei mostly from the Barkly West, Kimberley, Griqualand, Kalahari area, but since M. haussknechtii also occurs in this region it is impossible, without spore data, to know which of the two taxa are involved. Outside of South Africa there are reports from North Africa: Canary Islands (Wildpret 1977; Beltran Tejera & Santos 1972; Eckblad 1962; 1975; Beltran Tejera & Wildpret 1977; Beltran Tejera 1980). Mauritania: Sahel: (Hariot & Patouillard 1910). Morocco: (Sebek 1958; Malen~on & Bertault 1960). Algeria: (Fries 1838; 1874; Montagne 1843;

165 *Jaczewski 1893; Hennings, 1901; Hollos 1904). Tunisia: (patouillard 1908). Libia: (Baroni 1892; El-Buni & Rattan 1981). Middle East: Egypt: (Hollos 1904; Reichert 1921). Israel: (*Reichert & Avizohar-Her~enzon 1959; Dring & Rayss 1963). Iraq: (Eckblad 1970). Iran: (petrak, 1939; *1949; Esfandiari 1951; Ershad 1977; Watling & Gregory 1977; Leonard 1981; *Saber 1986). Aden: (Hollis 1904). Europe: Spain (Cuatrecasas 1926; Bellot 1942; RivasMartinez & Losa-Quintana 1969; *Calonge 1975a, b; *Honrubia, Calonge, Demoulin, Moreno, & Llimona 1982; *Moreno, Manjon & Zugasa 1986; *Rolland 1986; Esteban 1988; Ortega & Buendea 1989; lllana, Heykoop, EsteveRaventos & Moreno 1989). South of France: (*De Candolle 1816 - type of M. arenaria; *Corda 1854; De Seynes 1862; 1863; Konrad & Maublanc 1924 - 1927; Malencon 1982). Italy: Sardinia: (Cavara 1901; *Belli 1908 - type of M. radiosus var. isosporus. Italy: Central and Southern: (*Bergamasco 1909; *Pacioni & Lalli 1981). Italy: Sicily: (Cavara 1902). Jugoslavia: (Babos 1980). Greece: Attica (Hollos 1904). East Germany: (Rauschert 1964; 1965; Sebek 1966; Dorfelt 1974; Kreisel 1987). Poland: (Skirgiello 1977). Czechoslovakia: (Simr 1935; *Smarda 1952; *Sebek 1958; 1961; 1963). Austria: (Lohwag 1928). Hungary: (Hollos 1904; Babos 1980). Romania: (Alexandri 1932; Eliade 1965). Turkey: Manavgat (Watling & Gregory 1977). Russia: European: (Vasil'kov 1955); Elevationis Volgensis: (Davydkina, Ivanov & Komirnaja 1989); Stavropol, Caucasus (Schwarzman & Filimonova 1970). (*Sosin 1973). Ukraine: (Zerova 1956; *1974; Wasser 1973; Wasser & Soldatova 1977; *Zerov & Peresipkin 1979. Asia: Russia: (Vasil'kov 1955). Kasakhstan (Svarcman 1959; Byzova 1964; Pisareva 1964; Kalymbetov 1969; *Schwarzman & Filimonova 1970). Tadzhikistan (Pilat 1965). Turkmenistan: (Kalymbetov 1956; Schwarzman & Filimonova 1970; Koshkelova, Frolov, Orlov, Bezukhova, Baryrova 1983; BatYrova 1985; Teterevnikova-Babayan 1987). Uzbekistan; (panfilova & Gaponenko 1963; Schwarzman & Filimonova 1970). Kirghizskaya SSR: (Domacova 1958; Eljchibayev 1969). Middle Asia: Irtin: (pallas 1773 - type of Agaricus radiosus Pallas and M. pallasii Fr.), (*Sosin 1973). Siberia: (*Sosin 1973). Eastern Asia: China: Nei Monggol, Gunsu, Xinjiang (Liu 1984). Afghanistan: (Lange 1953; Eckblad 1970). West Pakistan: (*Ahmad 1952). North America: USA: (McKnight & McKnight 1987; & Zeller 1943); on the plains of eastern Oregon, on California deserts, in New Mexico, in the region of San Diego and on the peninsula of California (Morse 1948). Central America: Mexico: (*Guzman & Herrera 1969; 1973). Cuba: (Kreisel 1971). South America: Argentina: [*Spegazzini 1899 - type of M. argentina with spores quoted as 14 - 20 x 7 - 10 /lm, but type specimen annotated by Spegazzini as having spores 14 x 9 /lm, fide Horak, 1967, who found them to have a range of 11.5 - 14.0 X 8.5 - 9.5 /lm, which is within the range of M. arenaria var. arenaria. Another collection Chubut, Feb. 1899, was said by Spegazzini, on the packet, to have spores 12 - 20 X 8 - 12 /lm; Horak I.c., found them to measure 22.5 - 24.5 X 16.4 - 17.5 (13.2 - 14.8) Ilm, which suggests M. arenaria var. macrospora. Later Spegazzini (1912) when amplifying his account of M. argentina gave details of five unnamed forms 'a-e'. Of these, 'a' is possibly to be referred to M. haussknechtii,

166

while fonns 'b-d' most probably represent M. arenaria var. arenaria. Fonn 'e' has spores 15 - 18 x 8 - 10 Ilm which are somewhat intennediate in size between those of M. arenaria var. arenaria and M. arenaria var. macrospora and one would have to examine the material microscopically before attempting to assign it to one of the taxa recognized in this paper. Morse, 1948; Wright, 1949, Singer 1968]. Galapagos Islands: (Morse 1948). Australasia: Australia: (*Cleland 1934 - it is just possible that some of the several collections cited may represent M. arenaria var. arenaria, while others refer to M. arenaria var. macrospora, and yet others may possibly represent M. haussknechtii. Gatherings with spherical or irregularly spherical spores, 5.0 - 6.5 Ilm, may represent an undescribed species, but we have seen no material with spores of this shape and size). Victoria (Willis 1957). It should be noted that when compiling the above distributional data for Montagnea arenaria, many records were culled from regional lists of species, from ecological studies or from papers concerned with distribution which lacked microscope data. Since most of the authors involved did not attempt to distinguish between M. arenaria vaT. arenaria, M. arena ria var. macrospora and M. haussknechtii it follows that such records are somewhat ambiguous. However, where spore data are provided and substantiate the record such references are marked with an asterisk.

S.-Afr.Tydskr.Plantk., 1991, 57(3)

Discussion There has been a strong tendency amongst mycologists, while recognizing the great variability in spore size between different collections, to interpret Montagnea arenaria as a species with large spores. Examination of three collections in K from the topotype locality (vicinity of Montpellier in southern France) has revealed fructifications with unifonnly large spores with a range of (11.5-) 13.0 - 16.0 x (8.0-) 8.5 - 10.5 Ilm. It therefore seems reasonable to follow tradition and interpret M. arenaria as a large-spored taxon. The problem arises as to which epithet should be used for this species. Some authorities have accepted the epithet based on Agaricus radiosus Pallas (1773), which is without doubt the oldest epithet available. However, Pallas based his taxon on materia! collected in Middle Asiatic Russia (lrtin) and in this region there is a strong probability of an overlap in distribution with M. haussknechtii Rab., originally from Iran, at Engeli, on the south-west coast of the Caspian Sea. It is, therefore, in our opinion, premature to reject the wellestablished and correctly applied name M. arenaria, in favour of that based on Agaricus radiosus Pallas, which is of far less certain application, and may eventually even prove to be an earlier name for M. haussknechtii Rab. Much has been written on the variability of the spore size in Montagnea arenaria, and even after the segregation of M. haussknechtii, with small spores in the range of 6.0 - 8.0 (-10.0) x 3.0 - 5.0 (-6.0) Ilm, one is still left with a largespored taxon in which there is wide variation in spore size, in the overall range of (11.5-) 12.0 - 27.0 x (7.5-) 8.5 14.0 (-16.0) Ilm. Microscopic examination of collections of this large-spored taxon in K has shown that this variation is due to the variable number of spores produced per basidium, ranging from 1 - 4(-5). Australian collections, from central Australia and southwest Queensland in particular, are often 2-3-spored, or occasionally the 2-3-spored basidia are intermixed with land/or 4-spored basidia (one 5-spored basidium was observed), resulting in a combined spore range of (14.0-) 16.0 - 23.5 (-27.0) x 9.0 - 14.0 (-16.0) Ilm. It seems to us desirable to segregate this predominantly 2-3-spored taxon at varietal level, as Montagnea arenaria var. macrospora var. nov.

Illustrations Since the taxa of Montagnea are currently impossible to distinguish macroscopically, published illustrations said to represent this species are cited below without any attempt at critical evaluation. Coloured illustrations may be consulted in: Belli (1908), Tab. V (excellent); Bergamasco (1909), Tab. 7 A (excellent); Calonge (1975a), Figure 117 (excellent); Hollos (1904), Tab I, Figures 16 - 23; Tab II Figures 1 - 4 (excellent); Jaczewski (1893); McKnight & McKnight (1987), pI. 34; Malen~on (1982), Atlas 229 (excellent); Moreno, Manjon & Zugaza (1986), Figure 417; Rolland (1986), Figure 294 (poor); Zerova (1974), Tab. 166 Figure 1. Black-and-white photographs or line drawings are to be found in: Ahmad (1952); Bellot (1942), Figure 1; Bertram & Wildpret (1977), pI. 2C; Cleland (1934), Figure 31; Corda (1854), Tab. 20, Figure 416; Dring & Rayss (1963), PI. IB, lower figure is of South African specimen from Andriesfontein, Acocks (408); Guzman & Herrera (1969), Figures 67, 68; Leonard (1981), Figure IB; Levin, Branch, Rappoport, Mitchell (1987), p. 109 (excellent); Liu (1984), Figure 46; Melik-Khatrayan & Martirosyan (1971), Figure 1 (poor); Morse (1948), Figures 1 - 9; Pacioni & Lalli (1981), Figure 1.8; Pallas (1773), Tab. 'W'; Pilat (1965), very poor photographs; Rauchert (1964), Figure 2; Rauchert (1965), Figures 1 - 3; Reichert & Avizohar-Hershenzon (1959), pI. iV, A; Saber (1986), Figure 17 (poor photograph); Schwarzman & Filimonova (1970), Figures 14 - 15; Sebek (1958), Figures 55, 57; Sebek (1963), Figure 40; Smarda (1952), Figures 13,

Holotypus.- Ernabella, Musgrave Range, Central Australia, J.B. Cleland, Aug . 1933, Ex Waite Agricultural Research Institute, No.

15; Sorokin (1884), Tab. 14, Figures 53 - 57; Sosin (1973),

7003 (K).

Figure 100; Svarcman (1959), Figure 18; Vasil'kov (1955), Figure 18; Zerov & Perisipkin (1979), Figure 183; Zerova (1956), Figure 3.

Distribution

Montagnea arenaria var. macrospora Reid & Eicker, vaT. nov. (Figure 1 J-K).

? Coprinus psamathonophilus Speg., An. Mus. Nac. Buenos Aires 6, 156, 1899. ? Montagnites psamathonophilus (Speg.) Horak. Darwinian a 14, 372, 1967. A varietate typica basidiis bisporis vel triporis dominantibus sed interdum basidiis monosporis vel quadrisporis intermixtis, sporis (14.0-) 16.0 - 23.5 (-27.0) X 9.0 - 14.0 (-16.0) Il-m maxime variabilibus, plerumque ellipticis poro-germinativo hyaIino et vaIde excentrico differt.

The collection from Ernabella has been selected as holotype

S.Afr.J.Bot., 1991, 57(3)

of the variety macrospora, since Cleland (1934) has published a very detailed account of the gathering, including fresh material, and the fruitbodies show predominantly trisporous basidia, along with many which are 2-spored. The spore range of this material is 14 - 22 x 9 - 13 f..lm. Other Australian collections of var. macrospora in K are from: Central Australia: George Gunn Road, Kathleen Spring, A.C. Beauglehole (K74), 7 July 1964 (spores 16.0 - 20.0 x 9.75 - 13.0 f..lm, basidia mostly 2-spored); side of dry water course 300 miles west of Woomera, F.L. Hill, December 1953 (spores 15.0 - 23.5 x 10.0 - 14.0 f..lm, basidia mostly 2-spored). South West Queensland: foot of a sandhill, Arrabury, E.N. Marks, 29 Aug. 1967 (spores 17.5 - 26.0 x 11.0 - 16.0 f..lm, basidia 1-2-spored). Cleland (1934) also reported additional large-spored Australian collections under the name Montagnites candollei which almost certainly represent var. macrospora: Western Australia: Kurrawang. South Australia: Miller Creek near Mount Eba; Ooldea. Central Australia: north of Charlotte Waters; Woodforde Creek. New South Wales: Forbed. Other collections cited by Cleland (1.c.) either appear typical of M. arenaria, or may represent M. haussknechtii, while yet others with spherical or irregularly spherical spores may ultimately prove to belong to a new undescribed species. Discussion The occurrence of occasional basidia with fewer than the normal 4 sterigmata is always a chance possibility, resulting in the production of a few giant spores in any microscope preparation, but such abnormal basidia would be expected to be in the vast minority in any fruitbodies to be assigned to var. arenaria. If these giant spores were to be included, without comment, in the accepted spore range for a given collection this would confuse the situation, and make for difficulty when trying to assign it to either var. arenaria or var. macrospora. Such is frequently the situation encountered in the literature, where very large spores are reported, and it is only after microscopic study of the actual material that confirmation of the occurrence of var. macrospora is possible. When Berkeley & Curtis (1853) published their variety texensis as Montagnites candollei var. texensis, they merely indicated that it was smaller, and had slightly larger spores than in the typical European and Algerian form. Our study of the holotype of var. texensis in K (Texas, C. Wright, No. 3917) failed to confirm these observations. It shows the spores to measure 13 - 16 x 8 - 9 f..lm and to originate from 4-spored basidia, which is typical of Montagnea arenaria var. arenaria. Furthermore, the fruitbodies are not particularly small when considered on a worldwide basis, being 8 - 11 cm tall. The taxon is therefore relegated to synonymy of Montagnea arenaria. However, Zeller (1943) cited spores from North American material as measuring 12.0 - 19.0 x 6.0 - 11.2 f..lm, which might indicate the occurrence of var. macrospora in the United States. When Spegazzini described his Montagnites argentina Speg., he described the spores as 14 - 20 x 7 - 10 f..lm, although he annotated the holotype as having spores 14 x 9 f..lm. Horak (1967) found the spores on the latter to measure U.5 - 14.0 x 8.5 - 9.5 f..lm, which is typical of M. arenaria

167

var. arenaria. Accordingly, we have listed M. argentina Speg. in synonymy under M. arenaria var. arenaria. However, another collection in LPS - Chubut, II 1899, was said by Spegazzini, on the packet, to have spores 12 - 20 x 8 - 12 f..lm. Horak found them to measure 22.5 - 24.5 x 16.4 - 17.5 (13.2 - 14.8) f..lm, which is strongly suggestive of M. arenaria var. macrospora. Spegazzini also described Coprinus psamathonophilus Speg. with spores 18 - 21 x 9 - 11 f..lm while Horak (loc. cit.) found them to measure 17.0 - 19.0 (19.5) x 12.5 - 13.5 (10.6 - 11.5) f..lm. Again these data suggest the occurrence of M. arenaria var. macrospora in Argentina, but unfortunately no observations as to the number of spores produced per basidium are given. Should it ultimately prove that our suggested synonymy is correct, then, should one wish to recognize this very large-spored taxon at specific rank, the epithet psamathonophilus would take priority, but would need to be transferred to Montagnea . Alternatively, should one agree with our concept of the taxon with (1-)2-3(-4)-spored basidia as representing a variety of the 4-spored M . arenaria var. arenaria, the correct epithet would be M. arenaria var. macrospora. Later Spegazzini (1912) described a number of unnamed forms of his M. argentina in which for the most part the spore size was more typical of M. arenaria var. arenaria. It seems, therefore, that both 2- and 4-spored varieties of M. arenaria occur in Argentina. Saber (1986) quoted the spore size of Iranian material as 13 - 22 x 9 - 13 f..lm (av. 17.25 x 10.7 f..lm) in a collection from Rafsanjan (illustrated Figure 17), which is strongly suggestive of var. macrospora, but only 9 - 15 x 6 - 10 f..lm (av. 11.0 x 7.2 f..lm) in a collection from Dashte Kavir, which is nearer the range of var. arenaria. Ahmad (1952) quoted 13 - 20 x 8 - 21 f..lm as the range for spores from material in West Pakistan, and again there is the suggestion that var. macrospora may be involved. In Europe Corda (1854) noticed the presence of bisporous and trisporous basidia in Montagnea candollei [= arenaria], and in his characterization of the genus Montagnea he described the basidia as '2 - 3 spora (forsan pleiospora?)', but quoted an equivocal spore length of 0.000 58 p.p.p. [= 16.1 f..lm] for material from Montpellier. Rolland (1986), writing on Spanish fungi, indicated a spore range of 10 - 25 x 5 - 9 f..lm, which could include var. macrospora. Montagnites schuppii Rick, Egatea 13,434, 1928. The original description of this species is as follows: 'Pileo 1 dm. lato, 1 cm. crasso, saepe volvae restis tecto, parum depresso, albido, adpresse brunneo-squarroso, squamis fibrosis, orbiculariter dispositis, lamellis densis 8 mm latis, aequalibus a stipite 8 mm remotis: stipite 1 dm. alto, 1 cm. crasso, albo, squamulosa: volva 4 - 6 cm. alta, inferius bulbosa, superius nigra inferius albida: basidiis 15.5 X 8 - 9 11m, sterigmata 3 11m longa: sporis badio-atris, laevibus, globoso ovoideis 5 - 6.5 X 4.5 - 5 5.5 11m. In terra arenosa. Pulcherrima species praec1aro P. Ambrosio Schupp jam defuncto, qui earn 1906 legit, dedicata'.

We have not been able to study the above material which seems to have been a very large squat fungus, with cap far larger than in the other taxa belonging in Montagnea. It

168

S.-Afr.Tydskr.Plantk., 1991, 57(3)

would also be unique in the genus in having a whitish cap, bearing circularly disposed, adpressed, brown squarrose fibrous scales; also in having a very prominent volva, 4 - 6 cm high; and globose to ovoid spores 5.0 - 6.5 x 4.5 - 5.0 - 5.5 Jlm. A revision of the holotype is needed in order to determine whether this species belongs in Montagnea. 'Montagnea sp'

(Figure 1 L).

In K there is a specimen (with coloured sketch) from New Zealand, collected by Sinclair, 9 February 1856, filed in the Montagnea folder, but merely labelled 'Montagnites?'. The unusual features of this specimen were briefly discussed by Dring & Rayss (1963) in their generic characterization of Montagnea, and they obviously considered it to belong in this genus; we would prefer to reserve judgement until we have seen more material, but we are led to doubt this disposition. A description follows in which details of the fresh plant are taken from the collectors' notes: Sporophore tall, delicate, coprinoid. Pileus 3.5 cm diam., shallowly convex and slightly umbonate, with a densely radially sulcate blackish-brown zone extending from the irregularly fimbriate margin to the small central fleshcoloured disc, although the illustration shows the central area as yellowish-buff without pink tones. The dried specimen is unusual in that the surface of the cap is entirely covered with a distinct, loose tangle of hyphae which appears almost pulverulent under a lens and does not conform to the illustration, which shows the sulcate area of the cap as seemingly glabrous. Such a cap surface, if really a part of the fungus, would be unique in Montagnea. Lamellae dark blackish-brown, numerous and not exposed by radial fissuring of the overlying cap surface as in the other species. Stipe tall and elegant, apparently not woody, 12 cm high, 0.25 cm wide, expanding gradually to 0.5 cm at the white base, elsewhere pale buffy-brown with flesh-coloured apex but again the illustration shows the apex as yellowish buff without pink tints; no obvious annulus or vol va and neither is shown on the illustration. Microscopic structure difficult to interpret. Spores in mass blackish; there is an accompanying 'spore-print' which has a stained watersoaked appearance suggesting it was taken under very moist conditions - possibly with the gills having started to deliquesce - again strongly suggestive of a coprinoid fungus rather than a Montagnea . Spores 8.0 - 10.0 x 6.0 - 8.0 Jlm, brown and translucent, varying in shape from ovate or somewhat amygdaliform to broadly elliptic, with a complex, slightly thickened wall, an inconspicuous germ-pore, and a small oblique apiculus. [Dring & Rayss (1963) noted them as 'ellipsoid 8 - 12 X 6 .- 8 Jlm, dark brown, asymmetrically apiculate after the manner of the ballistospore of an agaric'] .

Acknowledgments The authors gratefully acknowledge the financial support of the University of Pretoria and the South African Foundation for Research Development.

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