A Taxonomic Survey of the Genus Montagnea (Gasteromycetes) with Special Reference to South Africa
S.Afr.l.Bot.,1991,57(3) 161 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 X 9.0 - 14.0 (-16.0) J.l.m, and that these were the product of A collection of Montagnea Fr. was made by one of us predominantly 2 - 3-spored basidia. We have proposed (D.A.R.) in the Roodeplaat Nature Reserve, near Pretoria, naming this last-mentioned taxon, as yet unrecorded from Northern Transvaal. Attempts to name this woody, agaric South Africa, the Large Spored Namaqua Black Cap (M. like, coprinoid Gasteromycete in Bottomley's (1948) mono arenaria var. macrospora Reid & Eicker, var. nov.). graph of the Gasteromycetes of South Africa failed since she considered it to represent a true agaric genus and so Descriptions excluded it from consideration. Montagnea Fries, Genera Hymenomycetum 7, 1836. However, the genus Montagnea became familiar, if not Montagnites Fr., Epicrisis 240, 1838. by name, to a fairly wide public in South Africa with the publication of the Afrikaans edition of the mushroom field Sporophores with a dried-up wizened, coprinoid aspect, and guide of Levin et al. (1987), Veldgids tot die sampioene van with a tough woody stipe. Pileus l.5 - 6 cm diam., conico Suid-Afrika, where it was very well illustrated under the truncate, truncate-campanulate, or convex, often with a rather seductive popular name of 'Namaqua Black Cap'. slight central umbo, pallid, but with exceedingly thin flesh However, its true identity remained a mystery and no Latin which is stretched over the gills and soon ruptures radially name was cited. Indeed, the authors commented that the between them so that the major part of the surface appears family name and scientific name was as then unknown to silvery grey to blackish where the gills are partially to fully them. Attempts to rectify this situation led to a revision of exposed from above. Lamellae crowded, black and radially the genus Montagnea. orientated, partially to fully exposed by rupture of the very It soon became obvious that two species of Montagnea thin overlying flesh, often said to be unbranched, but we occurred in South Africa, viz. the Small Spored Namaqua suspect some forking and anastomosis may occur, although Black Cap (M. haussknechtii Rab.), with small elliptic this needs confirmation from living material. When exposed spores, 6.0 - 8.0 (-10.0) X 3.0 - 5.0 (-6.0) J.l.m, appearing in this manner the attachment of the lamellae to the central diamond-shaped in optical section, and the Medium Spored disc appears very slight and precarious, and to involve Namaqua Black Cap [M. arenaria (DC.) Zeller] with much merely their extremity nearest the stipe. They, therefore, larger elliptic to broadly elliptic spores, (11.5-) 13.0 - 16.0 appear to radiate from the central disc much as would the (-17.0) X 7.5 - 10.5 J.l.m, appearing broadly ovoid to spokes of an umbrella when the covering material has been pyriform or pip-shaped in optical section. In both species the tom away from all but the central area. This led early spores are the product of 4-spored basidia. However, it also workers to postulate erroneous theories of lamellar develop became obvious that in certain parts of the world, notably ment. Stipe 2.5 - 30.0 cm tall, 0.3 - 2.0 cm wide, tough and Australia, an even larger spored variant of M. arenaria woody, longitudinally wrinkled, surface smooth to fibrillose, occurred with spores measuring (14.0-) 16.0 - 23.5 (-27.0) often with adpressed scales, pallid, with a fringed papery 162 S.-Afr.Tydskr.Plantk., 1991 , 57(3)
volva. Volva of young specimens said to be gelatinous Montagnites candollei var. coprinoides P. Henn., Hed (Savulescu, 1935). Annulus lacking. Hymenium comprising wigia 40, (98), 1901. basidia, or basidia separated by pseudoparaphyses. Cystidia Montagnites candollei var. somala Bacc., Eumycet. have been reported by some authorities but their presence Somalia 191, 1916. needs confirmation; they have not been observed in this Montagnites spegazzinii Sacco & Trott., SyU. Fung. 23, study. Basidia difficult to observe in dried material as they 327,1925. collapse to an agglutinated honeycomb-like structure in which individual elements will not separate, but either Sporophores coprinoid, centrally stipitate with a hard woody (1)2-3-4(5)-spored. Spore mass black. Spores sessile, dark stipe. Pileus 1.5 - 3.5 (-5.5) cm diam., usually truncate brown to black-brown, elliptic with a distinct wall, and a campanulate or truncate-hemispherical, occasionally trun hyaline germ-pore, which is often conspicuous and very cate-cylindIic, more rarely becoming convex or applanate, eccentric. In M. haussknechtii 6.0 - 8.0 (-10.0) X 3.0 - 5.0 sometimes with a capping of sand particles. Flesh of pileus (-6.0) J.1m, elliptic in side view but angular-shaped in reduced to a delicate membrane stretched over the lamellae, optical section; in M. arena ria (11.5-) 13.0 - 16.0 (-17.0) but thicker at the centre, and at an early stage splitting X 7.5 - 10.0 J.1m, elliptic to broadly elliptic in side view, radially to the vicinity of the disc, such that a very thin and ovoid to pyriform in optical section; in M. arenaria var. very narrow strip of tissue adheres to the dorsal surface of macrospora borne on (1-)2-3(-4)-spored basidia, and each gill, but sometimes sloughing off completely toward the margin of the cap. The gills may then appear free and to measuring (14.0-) 16.0 - 23.5 (-27.0) X 9.0 - 14.0 (-16.0) hang vertically downward attached to the disc only by their J.1m. In the latter taxon 3-spored basidia dominate in some width (see Levin et al. 1987). Colour ranging from dirty fruitbodies. Clamp connexions are reported by Singer (1976), but we have been unable to demonstrate their pres ence. Habitat: in dry, open, sandy situations such as coastal o\) dunes, steppes, savannah and deserts. o () a (jOO o 0AOo Q, (j () Discussion
white at the disc, where the surface may disrupt into small almost certainly belongs here). Africa West: Canary adpressed fibrillose spot-like scales, or remain intact, or Islands: Las Palmas (Spegazzini, 1915; Saccardo & Trotter, rupture radially to give an almost cog-wheel effect, passing 1925 - type of Montagnites spegazzinii, described with through a silvery-grey zone where the flesh has been spores 8.0 - 10.0 X 4.0 - 4.5 11m, almost certainly belongs stretched or partially ruptured, and finally to the broad here). Angola: in sandy dry ground amongst grasses, blackish outer region where the lamellae are exposed. Maionga, Loanda, J. Gosseweiler (289), 1921 (in K); Africa Lamellae black, very crowded, radially orientated, probably North: Algeria: near Biskra (Hennings, 1901 - two col forking but this needs confmnation from a study of fresh lections forming the basis of M. candollei var. coprinoides material. Stipe woody, 6 - 9 (15) cm high, equal, or more Henn. are somewhat dubious on account of the slightly usually markedly tapered downward, from 0.3 - 0.5 (1.0) larger spores, quoted as 7 - 11 x 4 - 6 11m). Tunisia: in cm at the apex to 0.5 - 1.5 cm at the base, where there is a sand, Gares (patouillard. 1894 - type of M. tenuis). Egypt: small, rather inconspicuous, fibrillose volva, with a fringed amongst sand, Nile Valley, Scott-Elliott (type of M. elliottil) margin, which is often left behind in the soil unless the Massee 1892, in K); Wadi RicMd. pro Hekian, 19 Feb. specimen is carefully excavated; whitish, longitudinally 1900; 11 March 1900 (Hennings, 1901 type of M. candollei wrinkled, with a smooth or fibrillose surface, which may n.var. minor). Africa East-Central: Northern Sudan: occasionally disrupt to form coarse squarrose scales toward solitary in sand, sand-dunes west of massif, Jebel Tageru, K. the base, or bear small, scattered patches of tissue else Neumann (230a), 28 Jan. 1964 (in K). Eritrea: on rocky where. Hymenium in dried material comprising a uniform ground. Asciorum Pass, 2800 ft. alt., P.O. Bally (B. 6753), brown honeycomb-like surface of collapsed and strongly 23 March 1949 (in K). Middle East: Israel: (Reichert & agglutinated. 4-spored basidia, which are impossible to Avizohar-Herschenzon, 1959; Dring & Rayss 1963). Trans separate individually in microscope squash preparations. jordan: Kalirrhoe, Prof. T. Rayss, 10 March 1936 (in K). Spores 6.0 - 8.0 (-10.0) X 3.0 - 5.0 (-6.0) 11m, black in the Jordan: in sand, Rum, 100 m. alt., D. Poore & J. Gillet mass, but brown under the microscope, elliptic in side-view, (5993/A), 29 April 1963 (in K). Bahrain: in sandy gully, but appearing angular and diamond-shaped or ovoid in D.A. Bellamy (6), April 1981. Iran: in sandy soil, Enzeli, optical section, with a slightly thickened wall, and a germ south-west coast of Caspian Sea, Prof. Haussknecht (Raben pore which is often placed very obliquely, sessile, apiculus horst, 1870 - type of M. haussknechtii). Aden: (Hollos virtually lacking. 1904). Oman: on sandy bank of small dry water-course, amongst rocks, near Masafi (25°19'N, 56°08'E), 1500 ft, Distribution J. George (32), 9 Feb. 1970 (in K). Western Asia: Russia: In open dry sandy places, in grassland or savannah country, Asiatic: Aral Sea (Sorokin, 1884). West Pakinstan: in sandy etc. South African specimens in K: Cape Province: in grassy soil, common, Sangla Hill, Panjab, Sept. 1938 (in K). places, on banks of river soil. Van Wyksfontein, Kimberley Kashmir: (Hongo, 1965). Asia: China: Nei Monggol, District, J.P. Acocks (328), 22 May 1936; in sandy soil, Xinjiang (Liu 1984). Australia: (Cleland 1934 - it is Kalahari Gemsbokpark, J.V. van Greuning, 14 May 1988. possible that the collection cited from Waitpinga, Encounter Eastern Transvaal: from open bare sandy soil in savannah, Bay, with spores elliptical to almost triangular 7.5 - 5.5 Roodeplaat Nature Reserve, near Pretoria, D.A. Reid, 11m, may belong here but we have not studied the material). Veldie and Martmari van Greuning, 26 Feb. 1989. South America: Argentina: (Spegazzini 1912 - a number Outside of South Africa M. haussknechtii has a wide geo of forms are discussed under Montagnites argentina of graphical distribution extending throughout Africa including which form 'a' with spores elliptic, 8 - 10 x 3 - 4 11m from the Canary Islands, the Middle East, Western Asia, and Cacheuta, prope Mendoza, Mat. 1910, could represent M. possibly also Australia and South America as follows: haussknechtii, but we have not examined the material). Africa East: Somalia: (Baccarini, 1916 - type of Montag nites candollei var. somala, with spores 7.2 X 4.5 11m, Illustrations Coloured illustrations are to be found: Baccarini (1916), Tab. 5; Rabenhorst (1870), Tab. 3, Figure 1; and black-and white illustrations: Dring & Rayss (1963), PI. lA - holo type of M. elliottii; PI. IB upper figure only; Hongo (1965), Figures 4 - 5.
Discussion 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 evi dence 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 Figure 2 Montagnea haussknechtii. Dried fruitbodies, Gems spores measuring 12.0 - 19.0 X 6.0 - 11.2 11m. Dring & bokpark 14 May 1988, PRUM 2475. 65% of natural size. Rayss (I.c.) merely remarked, 'We are not convinced that it 164 S.-Afr.Tydskr.Plantk .•1991.57(3)
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 study of collections available to them, and a survey of the the moment, consider this unduly significant, as a number of world literature, were able to distinguish under the compre the small-spored collections we have studied show a tend hensive 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 This seems a convenient starting, although as more data pileus truncate-cylindric, but again we do not consider this 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 overall spore picture to which group a particular sporophore angularly diamond-shaped than is perhaps usual. However, 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 col Mag. 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 different fruitbodies, one comprising a pileus and the stipe (98), 1901; see also Hollos, Gasterom. vonotk. helyesb. tapering downward but lacking a base; the other with a base Termeszetr. Fuzet. 25, 96-98, 1902. but no pileus. ? Montagnea radiosa (Pallas) Sebek, Ceska Myk. 8, 144, When Massee (l.c.) described the species he published an 1954. illustration, which not only figures a mature sporophore, but ? Montagnites pallasii Fr., Epicrisis 241 , 1838. also showed sections through the developing fungus. Judged from these illustrations the young fruitbody develops within Sporophores with a dried-up wizened appearance, from a membranous 'egg', but these figures appear very schem short and squat to tall and erect, coprinoid, with a central atic, and there are no immature specimens in the Kew woody stipe. Pileus 2 - 4 (-6) cm diam., conico-truncate or collection which could have formed the basis for Massee's truncate-campanulate to convex, often with a slight umbo, figures. However, the holotype was evidently divided, with and exceedingly thin flesh, which is stretched over the gills a part being retained at K and part going to NY, for Zeller and soon ruptures radially between them. At maturity the (1943) comments on the spores of the NY material, "spores cap colour passes from whitish at the intact or somewhat in the type of M. elliottii (in NY Bot. Gard. Herb.) ovate scaly disc, through silvery grey where the underlying gill and 6.0 - 7.75 x 4.5 - 5.0!lm instead of '12 x 7' as colour is apparent, to a broad greyish-black zone extending described by Massee". Zeller (l.c.) makes no mention of to the margin in which the dorsal surface of the gills is fully any immature developmental stages being represented in exposed. Lamellae numerous, black, radially orientated, NY. However, his observations on the spore size of the probably somewhat forked, or anastomosing. Stipe 2.5 - 24 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) 165
adpressed scales, and at the base an inconspicuous fringed *Jaczewski 1893; Hennings, 1901; Hollos 1904). Tunisia: papery volva, which in the young specimens is gelatinous (patouillard 1908). Libia: (Baroni 1892; El-Buni & Rattan (Savulescu 1935). Hymenium comprising 4-spored basidia, 1981). Middle East: Egypt: (Hollos 1904; Reichert 1921). which appear to be more or less regularly disposed over the Israel: (*Reichert & Avizohar-Her~enzon 1959; Dring & gill surface, separated by pseudoparaphyses, as illustrated by Rayss 1963). Iraq: (Eckblad 1970). Iran: (petrak, 1939; Hollos (1904). The basidia do not collapse into a honey *1949; Esfandiari 1951; Ershad 1977; Watling & Gregory comb-like pattern as in M. haussknechtii, but appear more 1977; Leonard 1981; *Saber 1986). Aden: (Hollis 1904). discrete. Spore powder black. Spores (11.5-) 13.0 - 16.0 Europe: Spain (Cuatrecasas 1926; Bellot 1942; Rivas (-17.0) X 7.5 - 1O.5/lm, elliptic to broadly elliptic in side Martinez & Losa-Quintana 1969; *Calonge 1975a, b; view but ovoid to pyriform or pip-shaped in optical section, *Honrubia, Calonge, Demoulin, Moreno, & Llimona 1982; sessile, dark brown, with a conspicuous, hyaline, very *Moreno, Manjon & Zugasa 1986; *Rolland 1986; Esteban oblique germ-pore. 1988; Ortega & Buendea 1989; lllana, Heykoop, Esteve Raventos & Moreno 1989). South of France: (*De Candolle Distribution 1816 - type of M. arenaria; *Corda 1854; De Seynes In dry, open sandy localities, in warmer parts of the world, 1862; 1863; Konrad & Maublanc 1924 - 1927; Malencon including coastal dunes, steppes, deserts etc. South African 1982). Italy: Sardinia: (Cavara 1901; *Belli 1908 - type of specimens in K: Cape Province: in red sand on top of M. radiosus var. isosporus. Italy: Central and Southern: Langeberge, Andriesfontein, Barkly West, J.P.H. Acocks (*Bergamasco 1909; *Pacioni & Lalli 1981). Italy: Sicily: (408), 18 June 1936 (No. 28643). [Illustrated in Dring & (Cavara 1902). Jugoslavia: (Babos 1980). Greece: Attica Rayss 1963, PI. 'B' lower figure.] (Hollos 1904). East Germany: (Rauschert 1964; 1965; Sebek Outside of South Africa M . arenaria has a very wide 1966; Dorfelt 1974; Kreisel 1987). Poland: (Skirgiello geographical distribution throughout the tropical and warm 1977). Czechoslovakia: (Simr 1935; *Smarda 1952; *Sebek temperate regions of the world along beaches or in desert or 1958; 1961; 1963). Austria: (Lohwag 1928). Hungary: steppe conditions. There are specimens in K from Algeria: (Hollos 1904; Babos 1980). Romania: (Alexandri 1932; ex. Herb. Berkeley, no further data; in desert, 50 paces from Eliade 1965). Turkey: Manavgat (Watling & Gregory 1977). an artificial waterhole near cypress trees, Sahara Edjeleh, Russia: European: (Vasil'kov 1955); Elevationis Volgensis: Mrs. Dean Netscher 1972. Southern France: Monspelii (Davydkina, Ivanov & Komirnaja 1989); Stavropol, Cauca (specimen from Montagne); ad mare in sabulosis, circa sus (Schwarzman & Filimonova 1970). (*Sosin 1973). Monspelium, Petit detit, Martio 1824, Herb. J. Gay; Gall. Ukraine: (Zerova 1956; *1974; Wasser 1973; Wasser & Soldatova 1977; *Zerov Peresipkin 1979. Asia: Russia: merid., Prof. Mertens, 1806, Herb. Dawson Turner. Egypt: & Shakrat el Delem, G.W. Murray (3692), 1925 - 1926 (Herb. (Vasil'kov 1955). Kasakhstan (Svarcman 1959; Byzova 1964; Pisareva 1964; Kalymbetov 1969; *Schwarzman & D.N. Simpson) near Mersa Shat, G.W. Murray (3748), 1925 Filimonova 1970). Tadzhikistan (Pilat 1965). Turkmenistan: - 1926 (Herb. D.N. Simpson). Crete: in pure sand on beach, (Kalymbetov 1956; Schwarzman & Filimonova 1970; Mallia, D.A., D.G. & P.M. Reid, 2 June 1966. Arabia: in Koshkelova, Frolov, Orlov, Bezukhova, Baryrova 1983; hard damp sand near granite outcrop 5 km north of road, BatYrova 1985; Teterevnikova-Babayan 1987). Uzbekistan; 3000 ft. alt., J.S. Collenette (5729). 11 March 1986. (panfilova & Gaponenko 1963; Schwarzman & Filimonova Romania: bord de la chaussee, terrains sablonneux, 1970). Kirghizskaya SSR: (Domacova 1958; Eljchibayev Dobrogea, dist. Constanta - intre Babadag si Ienisala, Tr. 1969). Middle Asia: Irtin: (pallas 1773 - type of Agaricus Savulescu & AI. V. Alexandri (Herbarium Mycologicum radiosus Pallas and M. pallasii Fr.), (*Sosin 1973). Siberia: Romanicum, Fasc. X, No. 481). West Pakistan: Karachi, S. (*Sosin 1973). Eastern Asia: China: Nei Monggol, Gunsu, Ahmad (254), 18 Aug. 1967; in sandy soil, Ledhar (Shei Xinjiang (Liu 1984). Afghanistan: (Lange 1953; Eckblad khupura), G.A. (2716), 15 Aug. 1948; in sandy soil, Lahore, 1970). West Pakistan: (*Ahmad 1952). North America: G.R. (15687), 25 Oct. 1960. Tibet: Kambajong, Major D. USA: (McKnight & McKnight 1987; & Zeller 1943); on the Prain, Sept. 1903. United States: Seattle, bank of Columbia plains of eastern Oregon, on California deserts, in New River, a few miles from Ginkgo Museum, Miss M. Holden, Mexico, in the region of San Diego and on the peninsula of 5 Sept. 1969; Texas, C. Wright (as M. candollei var. California (Morse 1948). Central America: Mexico: texensis B. & C.); New Mexico, C. Wright ex Curtis (as M . (*Guzman & Herrera 1969; 1973). Cuba: (Kreisel 1971). candollei B texensis ex Ravenel's Herbarium - recd 1891). South America: Argentina: [*Spegazzini 1899 - type of Additional South African collections are listed by Doidge M. argentina with spores quoted as 14 - 20 x 7 - 10 /lm, (1950) under Montagnites candollei mostly from the Barkly but type specimen annotated by Spegazzini as having spores West, Kimberley, Griqualand, Kalahari area, but since M. 14 x 9 /lm, fide Horak, 1967, who found them to have a haussknechtii also occurs in this region it is impossible, range of 11.5 - 14.0 X 8.5 - 9.5 /lm, which is within the without spore data, to know which of the two taxa are in range of M. arenaria var. arenaria. Another collection - volved. Chubut, Feb. 1899, was said by Spegazzini, on the packet, Outside of South Africa there are reports from North to have spores 12 - 20 X 8 - 12 /lm; Horak I.c., found Africa: Canary Islands (Wildpret 1977; Beltran Tejera & them to measure 22.5 - 24.5 X 16.4 - 17.5 (13.2 - 14.8) Santos 1972; Eckblad 1962; 1975; Beltran Tejera & Wild Ilm, which suggests M. arenaria var. macrospora. Later pret 1977; Beltran Tejera 1980). Mauritania: Sahel: (Hariot Spegazzini (1912) when amplifying his account of M. & Patouillard 1910). Morocco: (Sebek 1958; Malen~on & argentina gave details of five unnamed forms 'a-e'. Of Bertault 1960). Algeria: (Fries 1838; 1874; Montagne 1843; these, 'a' is possibly to be referred to M. haussknechtii, 166 S.-Afr.Tydskr.Plantk., 1991, 57(3)
while fonns 'b-d' most probably represent M. arenaria var. Discussion arenaria. Fonn 'e' has spores 15 - 18 x 8 - 10 Ilm which There has been a strong tendency amongst mycologists, are somewhat intennediate in size between those of M. are while recognizing the great variability in spore size between naria var. arenaria and M. arenaria var. macrospora and different collections, to interpret Montagnea arenaria as a one would have to examine the material microscopically species with large spores. Examination of three collections before attempting to assign it to one of the taxa recognized in K from the topotype locality (vicinity of Montpellier in in this paper. Morse, 1948; Wright, 1949, Singer 1968]. southern France) has revealed fructifications with unifonnly Galapagos Islands: (Morse 1948). Australasia: Australia: large spores with a range of (11.5-) 13.0 - 16.0 x (8.0-) (*Cleland 1934 - it is just possible that some of the several 8.5 - 10.5 Ilm. It therefore seems reasonable to follow collections cited may represent M. arenaria var. arenaria, tradition and interpret M. arenaria as a large-spored taxon. while others refer to M. arenaria var. macrospora, and yet The problem arises as to which epithet should be used for others may possibly represent M. haussknechtii. Gatherings this species. Some authorities have accepted the epithet with spherical or irregularly spherical spores, 5.0 - 6.5 Ilm, based on Agaricus radiosus Pallas (1773), which is without may represent an undescribed species, but we have seen no doubt the oldest epithet available. However, Pallas based his material with spores of this shape and size). Victoria (Willis taxon on materia! collected in Middle Asiatic Russia (lrtin) 1957). and in this region there is a strong probability of an overlap It should be noted that when compiling the above distrib in distribution with M. haussknechtii Rab., originally from utional data for Montagnea arenaria, many records were Iran, at Engeli, on the south-west coast of the Caspian Sea. culled from regional lists of species, from ecological studies It is, therefore, in our opinion, premature to reject the well or from papers concerned with distribution which lacked established and correctly applied name M. arenaria, in microscope data. Since most of the authors involved did not favour of that based on Agaricus radiosus Pallas, which is attempt to distinguish between M. arenaria vaT. arenaria, of far less certain application, and may eventually even M. arena ria var. macrospora and M. haussknechtii it prove to be an earlier name for M. haussknechtii Rab. follows that such records are somewhat ambiguous. How Much has been written on the variability of the spore size ever, where spore data are provided and substantiate the in Montagnea arenaria, and even after the segregation of M. record such references are marked with an asterisk. 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 large Illustrations spored taxon in which there is wide variation in spore size, Since the taxa of Montagnea are currently impossible to in the overall range of (11.5-) 12.0 - 27.0 x (7 .5-) 8.5 - distinguish macroscopically, published illustrations said to 14.0 (-16.0) Ilm. Microscopic examination of collections of represent this species are cited below without any attempt at this large-spored taxon in K has shown that this variation is due to the variable number of spores produced per basidium, critical evaluation. ranging from 1 - 4(-5). Coloured illustrations may be consulted in: Belli (1908), Australian collections, from central Australia and south Tab. V (excellent); Bergamasco (1909), Tab. 7 A (excellent); west Queensland in particular, are often 2-3-spored, or Calonge (1975a), Figure 117 (excellent); Hollos (1904), Tab occasionally the 2-3-spored basidia are intermixed with l I, Figures 16 - 23; Tab II Figures 1 - 4 (excellent); Jaczew and/or 4-spored basidia (one 5-spored basidium was ob ski (1893); McKnight & McKnight (1987), pI. 34; Malen~on served), resulting in a combined spore range of (14.0-) 16.0 (1982), Atlas 229 (excellent); Moreno, Manjon & Zugaza - 23.5 (-27.0) x 9.0 - 14.0 (-16.0) Ilm. It seems to us de (1986), Figure 417; Rolland (1986), Figure 294 (poor); sirable to segregate this predominantly 2-3-spored taxon at Zerova (1974), Tab. 166 Figure 1. varietal level, as Montagnea arenaria var. macrospora Black-and-white photographs or line drawings are to be var. nov. found in: Ahmad (1952); Bellot (1942), Figure 1; Bertram & Wildpret (1977), pI. 2C; Cleland (1934), Figure 31; Corda Montagnea arenaria var. macrospora Reid & (1854), Tab. 20, Figure 416; Dring & Rayss (1963), PI. IB, Eicker, vaT. nov. (Figure 1 J-K). lower figure is of South African specimen from Andries fontein, Acocks (408); Guzman & Herrera (1969), Figures ? Coprinus psamathonophilus Speg., An. Mus. Nac. 67, 68; Leonard (1981), Figure IB; Levin, Branch, Rappo Buenos Aires 6, 156, 1899. ? Montagnites psamathonophilus (Speg.) Horak. Darwin port, Mitchell (1987), p. 109 (excellent); Liu (1984), Figure ian a 14, 372, 1967. 46; Melik-Khatrayan & Martirosyan (1971), Figure 1 (poor); Morse (1948), Figures 1 - 9; Pacioni & Lalli (1981), Figure A varietate typica basidiis bisporis vel triporis dominantibus sed 1.8; Pallas (1773), Tab. 'W'; Pilat (1965), very poor photo interdum basidiis monosporis vel quadrisporis intermixtis, sporis graphs; Rauchert (1964), Figure 2; Rauchert (1965), Figures (14.0-) 16.0 - 23.5 (-27.0) X 9.0 - 14.0 (-16.0) Il-m maxime 1 - 3; Reichert & Avizohar-Hershenzon (1959), pI. iV, A; variabilibus, plerumque ellipticis poro-germinativo hyaIino et Saber (1986), Figure 17 (poor photograph); Schwarzman & vaIde excentrico differt. Filimonova (1970), Figures 14 - 15; Sebek (1958), Figures Holotypus.- Ernabella, Musgrave Range, Central Australia, J.B. 55, 57; Sebek (1963), Figure 40; Smarda (1952), Figures 13, 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 Distribution (1956), Figure 3. The collection from Ernabella has been selected as holotype S.Afr.J.Bot., 1991, 57(3) 167
of the variety macrospora, since Cleland (1934) has var. arenaria. Accordingly, we have listed M. argentina published a very detailed account of the gathering, including Speg. in synonymy under M. arenaria var. arenaria. fresh material, and the fruitbodies show predominantly tri However, another collection in LPS - Chubut, II 1899, was sporous basidia, along with many which are 2-spored. The said by Spegazzini, on the packet, to have spores 12 - 20 x spore range of this material is 14 - 22 x 9 - 13 f..lm . Other 8 - 12 f..lm . Horak found them to measure 22.5 - 24.5 x Australian collections of var. macrospora in K are from: 16.4 - 17.5 (13.2 - 14.8) f..lm, which is strongly suggestive Central Australia: George Gunn Road, Kathleen Spring, of M. arenaria var. macrospora. Spegazzini also described A.C. Beauglehole (K74), 7 July 1964 (spores 16.0 - 20.0 x Coprinus psamathonophilus Speg. with spores 18 - 21 x 9 9.75 - 13.0 f..lm, basidia mostly 2-spored); side of dry water - 11 f..lm while Horak (loc. cit.) found them to measure 17.0 course 300 miles west of Woomera, F.L. Hill, December - 19.0 (19.5) x 12.5 - 13.5 (10.6 - 11.5) f..lm . Again these 1953 (spores 15.0 - 23.5 x 10.0 - 14.0 f..lm, basidia mostly data suggest the occurrence of M. arenaria var. macrospora 2-spored). South West Queensland: foot of a sandhill, in Argentina, but unfortunately no observations as to the Arrabury, E.N. Marks, 29 Aug. 1967 (spores 17.5 - 26.0 x number of spores produced per basidium are given. Should 11.0 - 16.0 f..lm, basidia 1-2-spored). it ultimately prove that our suggested synonymy is correct, Cleland (1934) also reported additional large-spored Aus then, should one wish to recognize this very large-spored tralian collections under the name Montagnites candollei taxon at specific rank, the epithet psamathonophilus would which almost certainly represent var. macrospora: Western take priority, but would need to be transferred to Mon Australia: Kurrawang. South Australia: Miller Creek near tagnea. Alternatively, should one agree with our concept of Mount Eba; Ooldea. Central Australia: north of Charlotte the taxon with (1-)2-3(-4)-spored basidia as representing a Waters; Woodforde Creek. New South Wales: Forbed. variety of the 4-spored M. arenaria var. arenaria, the Other collections cited by Cleland (1.c.) either appear correct epithet would be M. arenaria var. macrospora. Later typical of M. arenaria, or may represent M. haussknechtii, Spegazzini (1912) described a number of unnamed forms of while yet others with spherical or irregularly spherical his M. argentina in which for the most part the spore size spores may ultimately prove to belong to a new undescribed was more typical of M. arenaria var. arenaria. It seems, species. therefore, that both 2- and 4-spored varieties of M. arenaria occur in Argentina. Discussion Saber (1986) quoted the spore size of Iranian material as The occurrence of occasional basidia with fewer than the 13 - 22 x 9 - 13 f..lm (av. 17.25 x 10.7 f..lm) in a collection normal 4 sterigmata is always a chance possibility, resulting from Rafsanjan (illustrated Figure 17), which is strongly in the production of a few giant spores in any microscope suggestive of var. macrospora, but only 9 - 15 x 6 - 10 preparation, but such abnormal basidia would be expected to f..lm (av. 11.0 x 7.2 f..lm) in a collection from Dashte Kavir, be in the vast minority in any fruitbodies to be assigned to which is nearer the range of var. arenaria. Ahmad (1952) var. arenaria. If these giant spores were to be included, quoted 13 - 20 x 8 - 21 f..lm as the range for spores from without comment, in the accepted spore range for a given material in West Pakistan, and again there is the suggestion collection this would confuse the situation, and make for that var. macrospora may be involved. difficulty when trying to assign it to either var. arenaria or In Europe Corda (1854) noticed the presence of bisporous var. macrospora. Such is frequently the situation and trisporous basidia in Montagnea candollei [= arenaria], encountered in the literature, where very large spores are and in his characterization of the genus Montagnea he reported, and it is only after microscopic study of the actual described the basidia as '2 - 3 spora (forsan pleiospora?)', material that confirmation of the occurrence of var. macro but quoted an equivocal spore length of 0.000 58 p.p.p. spora is possible. [= 16.1 f..lm] for material from Montpellier. Rolland (1986), When Berkeley & Curtis (1853) published their variety writing on Spanish fungi, indicated a spore range of 10 - 25 texensis as Montagnites candollei var. texensis, they merely x 5 - 9 f..lm, which could include var. macrospora. 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. Montagnites schuppii Rick, Egatea 13,434, 1928. 3917) failed to confirm these observations. It shows the The original description of this species is as follows: spores to measure 13 - 16 x 8 - 9 f..lm and to originate from 4-spored basidia, which is typical of Montagnea 'Pileo 1 dm. lato, 1 cm. crasso, saepe volvae restis tecto, parum arenaria var. arenaria. Furthermore, the fruitbodies are not depresso, albido, adpresse brunneo-squarroso, squamis fibrosis, particularly small when considered on a worldwide basis, orbiculariter dispositis, lamellis densis 8 mm latis, aequalibus a being 8 - 11 cm tall. The taxon is therefore relegated to stipite 8 mm remotis: stipite 1 dm. alto, 1 cm. crasso, albo, synonymy of Montagnea arenaria. However, Zeller (1943) squamulosa: volva 4 - 6 cm. alta, inferius bulbosa, superius nigra cited spores from North American material as measuring inferius albida: basidiis 15.5 X 8 - 9 11m, sterigmata 3 11m longa: 12.0 - 19.0 x 6.0 - 11.2 f..lm, which might indicate the sporis badio-atris, laevibus, globoso ovoideis 5 - 6.5 X 4.5 - 5 - occurrence of var. macrospora in the United States. 5.5 11m. In terra arenosa. When Spegazzini described his Montagnites argentina Pulcherrima species praec1aro P. Ambrosio Schupp jam de Speg., he described the spores as 14 - 20 x 7 - 10 f..lm, functo, qui earn 1906 legit, dedicata'. although he annotated the holotype as having spores 14 x 9 We have not been able to study the above material which f..lm . Horak (1967) found the spores on the latter to measure seems to have been a very large squat fungus, with cap far U.5 - 14.0 x 8.5 - 9.5 f..lm, which is typical of M. arenaria 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, 11. Lahore, Panjab University Press. bearing circularly disposed, adpressed, brown squarrose ALEXANDRI, A.V. 1932. Monlagniles radiosus (Pall.) Holl. in fibrous scales; also in having a very prominent volva, 4 - 6 der Dobrogea und im Siiden von Bessarabien. Bullelin de la cm high; and globose to ovoid spores 5.0 - 6.5 x 4.5 - 5.0 Seclion scientifique de l'Academie roumaine 15 (7-8): 1-3. - 5.5 Jlm. A revision of the holotype is needed in order to BABOS, M. 1980. Seltene Pilzarten der Sandgebiete Ungams III. Siudia Bolanica Hungarica 14: 55 - 61. determine whether this species belongs in Montagnea. BACCARINI, P. 1916. Eumyceles Somalia in Missione Siefanini Paoli nella Somalia Italiana. 189 - 200; 228. Firenze. Galletti 'Montagnea sp' (Figure 1 L). e Cocci. BARONI, E. 1892. Sopra alcune crittogame Africane raccolte In K there is a specimen (with coloured sketch) from New presso Tripoli di Barberia dal Prof. Rafaello Spigai. Bullelino Zealand, collected by Sinclair, 9 February 1856, filed in the della Sociela Bolanica Italiana 1: 239 - 243. Montagnea folder, but merely labelled 'Montagnites?'. The BATYROVA, G. Sh. 1985. Konspekt Flory Makromitsetov kopet unusual features of this specimen were briefly discussed by daga. Ashkhabad Ylym. Akademia Nauk Turkmenskoi SSR. Dring & Rayss (1963) in their generic characterization of BELLI, S. 1908. Addenda ad Floram Sardoam. Annali di Bolanica Montagnea, and they obviously considered it to belong in 6: 523 - 534. this genus; we would prefer to reserve judgement until we BELLOT, F. 1942. Dos Basidiomicetos Interesantes. Anales de la have seen more material, but we are led to doubt this dispo Real Academia de Farmacia (n.v.). sition. A description follows in which details of the fresh BELTRAN TEJERA, E.B. 1980. Catalogo de los Hongos Sapro fitos presentes en el Archipielago Canario Tenerife. Instituto de plant are taken from the collectors' notes: Estudios Canarios. La Laguna de Tenerife. Sporophore tall, delicate, coprinoid. Pileus 3.5 cm diam., BELTRAN, E. & WILDPRET, W. 1977. Gasteromycetes de las shallowly convex and slightly umbonate, with a densely Islas Canarias. Vieraea 7: 49 - 96. radially sulcate blackish-brown zone extending from the BERGAMASCO, G. 1909. Due novi miceti per la Campania. irregularly fimbriate margin to the small central flesh Nuovo Giornale Bolanico Italiano . Nuova Serie 16: 439 - 443. coloured disc, although the illustration shows the central BERKELEY, M.l. & CURTIS, M.A. 1853. Centuries of North area as yellowish-buff without pink tones. The dried speci American Fungi. Annals and Magazine of Nalural Hislory. men is unusual in that the surface of the cap is entirely Series 1. 12: 417 - 435. covered with a distinct, loose tangle of hyphae which ap BOTIOMLEY, A.M. 1948. Gasteromycetes of South Africa. BOlhalia 4: 473 - 810. pears almost pulverulent under a lens and does not conform BYZOVA, Z.M. 1964. Itohi izucheniya mikoflory Chu-llii'skikh to the illustration, which shows the sulcate area of the cap as gor. Trudy Inslilula BOlaniski Akademii Nauk Kazakhskoy SSR, seemingly glabrous. Such a cap surface, if really a part of 18: 147 - 181. the fungus, would be unique in Montagnea. Lamellae dark CAWNGE, F.D. 1975a. Hongos de Nuestros Campos y Bosques blackish-brown, numerous and not exposed by radial fissur I, Madrid. Ministerio de Agricultura Secretaria General ing of the overlying cap surface as in the other species. Stipe Tecnica. tall and elegant, apparently not woody, 12 cm high, 0.25 cm CAWNGE, F.D. 1975b. Omamentacion de las esporas de wide, expanding gradually to 0.5 cm at the white base, Algunos Gasteromycetes Espanoles. Anales des Inslilulo elsewhere pale buffy-brown with flesh-coloured apex but Bolanico Anlonio Jose Cavanilles 32: 103 - 115. again the illustration shows the apex as yellowish buff DE CANDOLLE, A.P. 1816. Flore Francaise Ed 3, 6 Paris, without pink tints; no obvious annulus or vol va and neither Desray. DE CANDOLLE, A.P. 1836. Prodromus Systematis Naturalis is shown on the illustration. Microscopic structure difficult Regni Vegetabilis V. Paris, Treuttel & Wiirtz. to interpret. Spores in mass blackish; there is an CA VARA, F. 1901. La vegetazione della Sardegna meridionale. accompanying 'spore-print' which has a stained water Nuovo Giornale Bolanico Iialiano 8: 363 - 415. soaked appearance suggesting it was taken under very moist CA V ARA, F. 1902. Di a1cuni miceti nuovi 0 rari della Sicilia conditions - possibly with the gills having started to deli orientale. Bullellino della Sociela Bolanica Iialiana, 11: 186 - quesce - again strongly suggestive of a coprinoid fungus 190. rather than a Montagnea. Spores 8.0 - 10.0 x 6.0 - 8.0 Jlm, CLELAND, l.B. 1934. Toadstools and Mushrooms and other brown and translucent, varying in shape from ovate or larger Fungi of South Australia I, Adelaide. Harrison Weir. somewhat amygdaliform to broadly elliptic, with a complex, CORDA, A.C.l . 1854. Iconum Fungorum 6, Prague. F. Ehrlich. slightly thickened wall, an inconspicuous germ-pore, and a CUATRECASAS, 1. 1926. Monlagniles radios us (Pall.) Ho1l6s. small oblique apiculus. [Dring & Rayss (1963) noted them Bulleli de la Inslilucio calalana d' hisloria natural. Ser. VI. 2: as 'ellipsoid 8 - 12 X 6 .- 8 Jlm, dark brown, 152 - 154 (n.v .). asymmetrically apiculate after the manner of the ballisto DA VYDKINA, T.A., IVANOV, A.1. & KOMIRNAJA, O.N. spore of an agaric'] . 1989. Novitates de Aphyllophoralibus et Gasteromycetibus elevationis Volgensis. Novosli Sislemaliki Nizshikh Raslenit Acknowledgments 26: 60 - 62. DOIDGE, E.M. 1950. The South African Fungi and Lichens to the The authors gratefully acknowledge the financial support of end of 1945. BOlhalia 5: 1 - 1094. the University of Pretoria and the South African Foundation DOMAl:OVA, A.A. 1958. 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