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Departamento de Biología Vegetal, Facultad de Biología, Universidad de Alcalá, E-28871 Alcalá de Henares, Madrid, Spain
Víctor J. Rico 2
Departamento de Biología Vegetal II, Facultad de Farmacia, Universidad Complutense, E-28040 Madrid, Spain
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ABSTRACT |
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 TOP ABSTRACT INTRODUCTIONMATERIALS AND METHODSTAXONOMYLITERATURE CITED |
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A taxonomic study of the species with nutant, pleurotoid and cyphelloid basidiomata of the genus Arrhenia in the Iberian Peninsula is presented. This study is based on the examination of recent specimens collected in the field and from dried herbarium collections. Five species and one variety of this genus are recognized on the basis of morphological and ecological features: Arrhenia acerosa var. acerosa, A. acerosa var. tenella, A. auriscalpium, A. lobata, A. retiruga and A. spathulata. Arrhenia auriscalpium is new to Spain, where it is not restricted to alpine zones of the Eurosiberian region and is more common in the Mediterranean region than previously reported. In contrast, A. acerosa var. tenella currently is known from the Eurosiberian subalpine belt and represents the first report to the Iberian Peninsula, and A. retiruga is known only from the mesomediterranean to supramediterranean belts of the Mediterranean region. Arrhenia acerosa var. acerosa, A. lobata and A. spathulata are widespread in both the Eurosiberian and the Mediterranean regions. Non-Iberian materials of A. acerosa var. tenella and A. spathulata also were studied for comparison. Cyphella cochlearis var. subsessilis, Dictyolus lagunae, Leptoglossum muscigenum var. azonum and Pleurotellus acerosus var. tenellus are lectotypified. Cyphella cochlearis var. subssesilis, Dictyolus lagunae and Leptoglossum muscigenum var. azonum are synonymized with Arrhenia spathulata, and Cyphella cochlearis var. auriformis is placed in synonymy with A. auriscalpium. These taxa are illustrated, described and discussed, based on Iberian material with emphasis on features of the basidioma, pigment of the pileipellis, presence or absence of clamps and shape and size of the basidiospores. A diagnostic key also is given.
Key words: Agaricales, Arrhenia, Basidiomycota, nomenclature, taxonomy, Tricholomataceae
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSTAXONOMYLITERATURE CITED |
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, Redhead et al 2002). In this sense, the genus Arrhenia constitutes a monophyletic bryophilous group that includes: (a) forms with nutant, pleurotoid or cyphelloid basidiomata (Arrhenia s. str.) and (b) the more typical mesopodal lamellate omphalinoid forms with grayish basidiomata formerly treated under the genus Omphalina (cf. Norvell et al 1994) but excluding those reddish-brown species related to O. pyxidata (Bull. : Fr.) Quél., the conserved lectotype of Omphalina (Moncalvo et al 2000, 2002, Redhead et al 2002). Micromorphological features, such as nonamyloid spores, subregular to irregular hymenophoral trama and pileipellis with incrusting pigment, are shared by all members of the genus Arrhenia.
Five species currently included in Arrhenia have been reported previously from the Iberian Peninsula, but these accounts are scattered in rare and general catalogues of fungi and the taxa often are listed as synonyms of A. acerosa (Fr.) Kühner, A. auriscalpium (Fr.) Fr., A. lobata (Pers. : Fr.) Kühner and Lamoure ex Redhead, A. retiruga (Bull. : Fr.) Redhead and A. spathulata (Fr.) Redhead. Arrhenia acerosa (see for example: Ortega et al 1997, Coutinho 1932 sub Agaricus acerosus) and A. lobata (see for example: Ballarà 1996, Ortega et al 1997, Esteve-Raventós et al 1997, Barrio et al 1985 sub Leptoglossum lobatum) have been reported primarily from wet sites in the mountains of the Mediterranean region. Arrhenia auriscalpium, usually considered an arctic-alpine species (Høiland 1976, Gulden 1988a), has been reported from the Iberian Peninsula in the alpine belt of the Pyrenees (Andorra, Vila et al 1996) and from extra-alpine areas of the Mediterranean region (Beira Baixa in Portugal, Redhead 1984 sub Cyphella cochlearis). Arrhenia retiruga has been found scarcely in lowlands of the Mediterranean region (Barcelona in Spain, Rocabruna 1984 sub Leptoglossum retirugum), but A. spathulata is widespread and has been cited frequently (see for example: Malençon and Bertault 1976 sub Leptoglossum muscigenum, Ballarà 1996, Gómez et al 1995, Campoamor and Molina 2001).
The results presented here and in Barrasa and Esteve-Raventós (2000) and Barrasa and Rico (2001) are part of a wider investigation of the species of Arrhenia, Lichenomphalia Redhead, Lutzoni, Moncalvo & Vilgalys (the lichenized Omphalina fide Redhead et al 2002) and Omphalina in Europe that focus on the poorly known mycobiota of the Mediterranean region. This paper deals with five species and one variety of Arrhenia from the Iberian Peninsula (Andorra, Portugal and Spain, including the Balearic Islands) on the basis of study of fresh and dried material, precise descriptions of their ecology and geographical distribution, an exhaustive review of the literature and the re-evaluation of the nomenclature of the Iberian taxa.
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MATERIALS AND METHODS |
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and Redhead et al (2002). For more details on the distribution and range of some of the species studied, see Redhead (1984, 1989).
Macroscopic photographs were shot with an Olympus SC-35 camera coupled to an Olympus SZ-PT stereo microscope with a Highlight 2001 Illuminator base. Light microscopy photographs were shot with an Olympus BX 50 with bright field and phase contrast. Drawings were made with a camera lucida device. For mycological concepts and terminology we followed Kirk et al (2001), except for terms such as pleurotoid and amygdaliform in which we followed Vellinga (1988). The term cyphelloid refers to cup-like basidiomata. Basidiomata colors were determined according to Munsell (1994). Phytoclimatic terminology is according to Rivas-Martínez (1987). Spore size was based on 30 measurements of at least one basidioma of each studied collection. Comparative spore shape and size between taxa was established following the Q method of Heinemann and Rammeloo (1985). Extreme values have been noted between brackets when they represented no more than 10% of the measurements.
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TAXONOMY |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSTAXONOMYLITERATURE CITED |
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Agaricus acerosus Fr., Syst. mycol. 1: 191. 1821 (basionym).
: 869–870.
Arrhenia glauca (Batsch) Bon & Courtec., Doc. Mycol. 18(69): 37. 1987.
Arrhenia glauca (Batsch) Høiland in L. Hansen & Knudsen eds., Nordic macromycetes 2: 99. 1992; superfl. comb. and basionym or reference not indicated.
Arrhenia latispora (J. Favre) Bon & Courtec., Doc. Mycol. 18(69): 37. 1987. Basidiomata pleurotoid, 1–2.7 x 1–3 cm. Pileus 1–3 cm wide, flabellate to spathulate or reniform, flattened, margin lobed and undulate, glabrous, longitudinally slightly zonate, hygrophanous, gray-brown to gray (10YR 5/1, 10YR 6/1 to 6/2) or light gray to whiteish (10YR 7/1 to 8/1) when moist, gray dark to fuscous or very pale brown on drying (10YR 4/1 to 4/2, 10YR 3/1 to 3/2, 10YR 8/3 to 8/4). Pileipellis a cutis of parallel hyphae, 4–10.5 µm diam, usually with parietal and ± fine encrusting pigment, irregularly distributed on walls. Hymenophore concolorous with pileus, lamellate. Lamellae thin (L = 7–25), moderately spaced, somewhat decurrent to adnate, with irregular lamellulae (l = 1–2). Stipe absent or very short, 0.3–0.6 x 0.2–0.4 cm, lateral to eccentric, whitish, tomentose, solid. Cystidia absent. Basidia 33–42 x 6–8 µm, cylindrical to clavate, 4-spored, sterigmata usually straight and up to 3 µm long. Spores 7.5–10(–11) x (4–)5–6.5 µm, Q = 1.5–1.7, ellipsoidal to broadly ellipsoidal, oval or lacrimiform, smooth, hyaline, nonamyloid. Clamps present in all tissues.
Specimens examined. ANDORRA. ORDINO: Coll d’Ordino, 31TCH8112, 1820 m, on mosses in a Pinus uncinata Ramond ex DC. in Lam. & DC. and Abies alba Miller forest, 16-X-2002, E. Sarrionandía (AH 30614). Ordino-Alcalis, Basses del Port de Rat, 31TCH7520, 2380 m, on plant and mosses debris in a Nardus stricta L. prairie, 16-X-2002, P. P. Daniels, N. Rodríguez, C. Lado, V. J. Rico and J. M. Barrasa (AH 30615). SPAIN. GRANADA: Barranco de San Juan, Sierra Nevada, 30SVG6704, 2500 m, in bare soil in wet grasslands on acidic soil, 5-IX-1996, L. Alcoba, F. Esteve-Raventós, A. Ortega, M. Villarreal and E. Horak (AH 20885 sub Arrhenia acerosa var. acerosa). GUIPÚZCOA: Tolosa, Barrio de Santa Lucía, in a prairie with mosses, 29-XI-1995, P. Pasaban (ARAN s.n.) MADRID: puerto de la Morcuera, 1800 m, among mosses, 28-V-2000, F. Esteve-Ravent;aaos & M. Villarreal (AH 3/502 duplo). NAVARRA: Leitza, Leitzalarrea, 30TWN8974, 600 m, on fallen leaves near a stream, 20-VII-2002, J. M. Lekuona (ARAN s.n.). SEGOVIA: Riofrío de Riaza, on plant debris, mosses and humid soil among mosses, 27-X-2000, F. Esteve-Raventós and V. J. Rico (AH 26853 sub Arrhenia latispora). SWITZERLAND. GRISSONS: val Sesvenna, bord de la Sesvenna, à l’amont de Marangun, sur brindilles mortes de mousse, 2400 m, 24-VIII-1943, J. Favre Z.A.21 (HOLOTYPUS G K13896 of Pleurotellus acerosus f. latisporus J. Favre: two portions, probably belonging to a single basidioma).
Arrhenia acerosa var. acerosa is a variable taxon in some macro and microscopic features and in habitat. Different taxonomic levels (species, forms and varieties) have been used to accommodate every set of the morphological features observed. Specimens with reduced lamellae to vein-like wrinkles have been found and described mainly from Nordic countries (Blytt 1905, Egeland 1911, Høiland 1976, 1992).
The spore width of Arrhenia acerosa var. acerosa is of 3–6 µm (Favre 1955, Kühner and Lamoure 1972, Høiland 1976, Redhead 1984, Watling and Gregory 1989, Kuyper 1995b) but collections with somewhat broader spores (6.5–7.5 µm) have been described from alpine zones and a new form, Pleurotellus acerosus f. latisporus, was proposed based on this criterion (Favre 1955). The separation of these taxa was followed by some authors (Kühner and Lamoure 1972, Høiland 1976), although the recognition of Arrhenia latispora as a different species also has been considered (Bon and Courtecuisse 1987, Watling and Gregory 1989). In contrast, and based on the extremely variable spore size of the specimens examined, Redhead (1984) and Kuyper (1995b) accepted the unique taxon A. acerosa and did not recognize forms or varieties of this species. Pleurotellus acerosus f. latisporus differs only from the typical form of A. acerosa in the spore width (Favre 1955). However, we observed a large range in spore width in the type material (5–7.5 µm) with some overlap with the spore size of A. acerosa var. acerosa. Variation in spore size also was observed in the Iberian collections; AH 20885 and AH 26853 possessed spores 4.5–6.5 µm wide, and no differences were observed among the spores of A. acerosa var. acerosa and A. latispora. Arrhenia acerosa is known from the oromediterranean and supramediterranean belts of that region, in southern (Granada, Ortega et al 1997) and central Spain (Segovia), respectively, and from the coline to Eurosiberian alpine belts of Andorra and northern Spain (Guipúzcoa, Navarra).
Arrhenia acerosa var. tenella (Kühner) Aronsen, Persoonia 14: 427. 1992. Figs. 7, 8
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Pleurotellus acerosus var. tenellus Kühner, Bull. Soc. Naturalistes d' Oyonnax 8: 76, 82. 1954 (basionym). Basidiomata pleurotoid, 0.4–1.7 x 0.4–2 cm. Pileus 0.4–2 cm wide, spathulate to reniform, margin lobed and undulate, glabrous, longitudinally slightly zonate, hygrophanous, gray-brown to pale gray when moist (10YR 3.5/3 to 2.5, Y 6.5/3, 7/3), gray dark to fuscous on drying (10YR 4/1 to 4/2, 10YR 3/1 to 3/2). Pileipellis a cutis of parallel hyphae, 4–9 µm diam, usually with parietal and ± fine encrusting pigment, irregularly distributed on walls. Hymenophore concolorous with pileus, lamellate. Lamellae thin, moderately spaced, somewhat decurrent to adnate, with irregular lamellulae. Stipe very short, 0.1–0.3 x 0.1–0.4 cm, lateral to eccentric, whitish, tomentose, solid. Cystidia absent. Basidia 23–32 x 5–9 µm, cylindrical to clavate, 2-spored or rarely 1-, 3- and 4-spored, sterigmata usually straight and up to 8 µm long. Spores 10–13.5 x 5–6 µm, Q = 2.0, elongate, smooth, hyaline, nonamyloid. Clamps present in all tissues.
Specimens examined. ALGERIA. ALGER: Villa des Bois, Sur la terre, 30-II-1932, R. Kühner II Alg 61 (LECTOTYPUS G in herb. R. Kühner s.n. of Pleurotellus acerosus var. tenellus Kühner, here designated: several portions, probably belonging to a single basidioma). FRANCE. SAVOIE: Parc Nat. de la Vanoise, Pralognan, environs de Pralognan, Cirque du Génépy, 2250 m, 24-VIII-1963, R. Kühner K-63–185 (G in herb. R. Kühner sub P. acerosus var. tenellus); id., 2400 m N.W., 8-IX-1971, R. Kühner K71–115 (G in herb. R. Kühner sub P. acerosus var. tenellus). HAUTE-SAVOIE: Préalpes du Faucigny, du Cirque des Fonts au Col d"Anterne, dans la mégaphorbiacée à Cicerbita alpina de l'aunaie, 26-VIII-1959, R. Kühner Sa59–4 (G in herb. R. Kühner sub P. acerosus var. tenellus). Sine loc., 17-IX-1959, R. Kühner (G in herb. R. Kühner s.n. sub P. acerosus var. tenellus). SPAIN. GERONA: Ripollés, Queralbs, Vall de Núria, Santuari de Núria, 31T4304694, 2220 m, on moss debris, 16-VIII-1996, J. Vila (JVG 960816–1 sub "Arrhenia glauca").
Arrhenia acerosa var. tenella is characterized by its bisporic basidia, larger spores (type material: 10–14.5 x 6–8 µm, Q = 1.8) and tendency to grow in wetlands. It has been reported from wetlands of mountainous and alpine areas in France (Kühner and Romagnesi 1954, Kühner and Lamoure 1972), Germany (Schmid-Heckel 1985) and Switzerland (Gulden 1988c) and also in wetlands of boreal areas of southern Norway (Aronsen 1992), as well as in dunes in The Netherlands (Kuyper 1995b). The only collection of A. acerosa var. tenella in the Iberian Peninsula (Figs. 7 and 8) was reported from wet sites of the subalpine belt of the Spanish Pyrenees (Gerona) within the Eurosiberian region (Vila et al 1997 sub "Arrhenia glauca").
Specimens examined for this study produced bisporic basidia, but monosporic, trisporic and tetrasporic basidia occasionally were observed. Monosporic and trisporic basidia have been described in this taxon by Gulden (1988c) and Kühner and Lamoure (1972), respectively. Although the micromorphological and ecological characters distinguishing this variety could be used to recognize this taxon at the species level, we have elected to examine additional collections to confirm the findings of our initial survey.
Arrhenia auriscalpium (Fr.) Fr., Summa veg. Scand., Sectio posterior: 312. 1849. Figs. 9–14
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Cantharellus auriscalpium Fr., Elench. fung. 1: 54. 1828 (basionym).
Cyphella cochlearis var. auriformis Bres., Brotéria 2: 88. 1903. Basidiomata nutant becoming pleurotoid, 0.4–1.8 x 0.35–1.5 cm. Pileus 0.35–1.5 cm wide, becoming flabellate or spathulate, margin undulate, glabrous, irregularly slightly zonate, hygrophanous, opaque to obscurely translucent, gray to dark gray or dark grayish-brown when moist (10YR 6/1, 2.5Y 4/1, 2.5Y 3/1 to 3/2), grayish-brown to brown or black when drying (10YR 5/2 to 5/3, 2.5Y, 2.5/1). Pileipellis a cutis of parallel hyphae, 3–11 µm diam, usually with parietal and dense encrusting pigment, irregularly distributed on walls or somewhat zebra-like. Hymenophore concolorous with pileus, smooth or venose to intervenose, somewhat anastomosing, consisting of thick veins, typically delimited, especially from the stipe, by a sterile margin continuous with the pileus. Stipe 0.2–0.8 x 0.08–0.2 cm, lateral, ascending with pileus, concolorous with pileus but sometimes whitish toward the base, glabrous to slightly pubescent, solid. Cystidia absent. Basidia 28–39 x 4–8 µm, cylindrical to clavate, 4-spored, sterigmata straight to slightly curved and 4–6 µm long. Spores 7.5–10.5(–11) x 4.5–7 µm, Q = 1.4–1.7, ellipsoidal to oval or amygdaliform, smooth, hyaline, nonamyloid. Clamps present in all tissues.
Specimens examined. ANDORRA. ORDINO: El Serrat, Llacs Tristaina, 31TCH7919, 2260 m, on soil among mosses and lichens, 15-VII-1993, J. Vila (BCC-SCM 2246). PORTUGAL BEIRA BAIXA: Castelo Branco, São Fiel, ad terram inter muscos minores, XII-1902, C. Torrend (LECTOTYPUS S F21076 of Cyphella cochlearis Bres. var. cochlearis: from left to right, the third basidioma of the upper and longer row; cf. Redhead 1984:870). SPAIN. SEGOVIA: Riofrío de Riaza, puerto de La Quesera, 30TVL6264, 1700 m, on pastures among mosses, 25-X-2002, J. M. Barrasa (AH 30618). MADRID: Colmenarejo, among mosses, 20-I-2001, F. Prieto (AH 19643). TOLEDO: La Iglesuela, on mosses, 16-I-1996, M. Heykoop, J. L. Aguirre and M. Villarreal (AH 30501). SWEDEN. OSTROGOTHIA: H. v. Post (UPS in herb. E. Fries s.n. sub Cantharellus glaucus (Batsch) Fr.).
Arrhenia auriscalpium is the type species of the genus Arrhenia (Donk 1957), and it is described as a typical member of the arctic and alpine mycota. This species frequently is found in North America (Redhead 1984, 1989), northern Europe, Greenland and Iceland (Lange 1957, Ohenoja 1971, Høiland 1976, Hallgrímsson 1981, Gulden 1988a) and also in the Alps (Favre 1955). Arrhenia auriscalpium grows among leafy gametophytes (and presumably on protonemata) on naked, sandy or burnt soils in the early stages of colonization by bryophytes and lichens (Høiland 1976, Redhead 1984). Within the Iberian Peninsula, it represents an uncommon species that has been found in the alpine belt of the Eurosiberian region (Andorra, Vila et al 1996) but also in the supramediterranean (Segovia in Spain; Beira Baixa in Portugal, Bresadola 1903 sub Cyphella cochlearis var. cochlearis) and mesomediterranean (Madrid and Toledo in Spain) belts of the Mediterranean region. The collections from Madrid, Segovia and Toledo represent the first reports of this species in Spain. Specimens collected in the Mediterranean region were morphologically variable in comparison with material from the Eurosiberian region and possessed smaller basidiomata and scarcely venose or even completely smooth hymenophores (AH 19643, 30501, 30618).
When Bresadola (1903) described Cyphella cochlearis, he also described two new varieties for the species, C. cochlearis var. subsessilis Bres. and C. cochlearis var. auriformis Bres., to which it is necessary to add the corresponding autonym (C. cochlearis var. cochlearis). As pointed out by Redhead (1984:871), the type collection consist of two species: Arrhenia auriscalpium and Arrhenia spathulata. Redhead (1984:871) lectotypified only the species name and autonym by selecting the elements corresponding to A. auriscalpium. We select here the portion referred to A. spathulata as the lectotype of C. cochlearis var. subsessilis and synonymize this name with A. spathulata. According to Redhead (pers comm), the fact that the two species were mixed reflects the fact that Bresadola (1903) ascribed only two varieties to his species and did not name A. cochlearis var. cochlearis. Cyphella cochlearis var. auriscalpium is considered here to be a homotypic synonym of C. cochlearis var. cochlearis because we think it probable Bresadola intended to describe the autonym under this name.
Arrhenia lobata (Pers. : Fr.) Kühner & Lamoure ex Redhead, Canadian J. Bot. 62: 871. 1984. Figs. 15–18
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Merulius lobatus Pers., Syn. meth. fung. 2: 494. 1801 (basionym). Sanctioning author: Fr., Syst. mycol. 2(2): 323. 1823 sub Cantharellus lobatus (Pers.) Fr. TYPUS: M. Vahl, Fl. dan., vol. 6, fasc. 18, pl. 1077, fig. 1 (upper figures). 1793 sub Helvella membranacea Dicks. (LECTOTYPUS!, cf. Redhead 1984:873).
Leptoglossum lobatum (Pers. : Fr.) Ricken, Blätterpilze Deutschland Fasc. 1: 6. 1910. Basidiomata pleurotoid, 1–4 cm wide. Pileus 1–4 cm wide, flabellate, dorsally attached, margin lobate and undulate, glabrous, transversely slightly zonate, hygrophanous, translucent, dark-grayish to gray-brown when moist or drying (10YR 4/2 to 4/4, 10YR 3/3 to 3/4). Pileipellis a cutis of radial to parallel hyphae, 4–12 µm diam, smooth to usually with parietal and moderately encrusting pigment, irregularly distributed on walls or sometimes restricted to specific hyphal portions. Hymenophore concolorous with pileus, intervenose, more or less reticulate, veins anastomosing. Stipe absent. Cystidia absent. Basidia 25–38 x 6–9 µm, clavate, 4-spored, sterigmata usually curved or sometimes straight and up to 4 µm long. Spores 7.5–10(–12) x 5.5–8(–9) µm, Q = 1.3, broadly ellipsoidal, lacrimiform, smooth, hyaline, nonamyloid. Clamps present in all tissues.
Specimens examined. SPAIN. GRANADA: Barranco de San Juan, Sierra Nevada, 30SVG6704, 2500 m, on bryophytic communities near a stream on acidic soil, 5-IX-1996, F. Esteve-Raventós, A. Ortega, M. Villarreal, L. Alcoba and E. Horak (AH 21649, 21650); id., on the moss Cratoneuron commutatum (Hedw.) G. Roth, 19-XI-1996, L. Alcoba and A. Ortega (AH 21657). HUESCA: Balneario de Panticosa, ibón Azul Superior, 30TYN2541, 2450 m, among mosses, 16-VII-1997, F. Arenal, F. Esteve-Raventós, M. Villarreal and V. González (AH 23653); id., ibón de Bachimaña Superior (AH 23658). Puerto de El Portalet, El Formigal, among bryophytes, 18-VII-1997, F. Arenal, F. Esteve-Raventós, M. Villarreal and V. González (AH 23657). Chisagüés, Sierra de Liena, Peña de Hércules, among bryophytes, 18-VIII-1997, F. Arenal, F. Esteve-Raventós, M. Villarreal and V. González (AH 23654). Benasque, ibón de Paderna, among bryophytes, 22-VIII-1997, F. Arenal, F. Esteve-Raventós, M. Villarreal and V. González (AH 23655). Liri, collado de la Corva, among bryophytes, 21-VIII-1997, F. Arenal, F. Esteve-Raventós, M. Villarreal and V. González (AH 23656). MADRID: Puerto de los Cotos, on Sphagnum sp., 11-IV-1976, J. M. Barrasa (AH 12367 sub Leptoglossum lobatum). Canencia, Puerto de Canencia, on mosses in humid sites, 24-IV-1999, F. Esteve-Raventós (AH 25000); id., arroyo de Canencia, 30TVL3527, 1240 m, on Aulacomnium palustre (Hedw.) Schwaegr., 5-IV-1982, L. Barrio (AH 3216, 3217 sub L. lobatum); id., on Calliergonella cuspidata (Hedw.) Loeske, L. Barrio (AH 3210, 3211, 3212, 3213, 3214, 3215 sub L. lobatum); id., on Sphagnum auriculatum Scimp., L. Barrio (AH 3219 sub L. lobatum); id., on Sphagnum teres (Schimp.) Ångstr., L. Barrio (AH 3218 sub L. lobatum); id., on Sphagnum subsecundum Nees, L. Barrio (AH 3220 sub L. lobatum). Laguna Grande de Peñalara, 30TVL1921, 2050 m, on Drepanocladus uncinatus (Hedw.) Warnst., 12-V-1982, L. Barrio (AH 3228 sub L. lobatum); id., on Sphagnum teres, L. Barrio (AH 3224, 3225, 3229 sub L. lobatum); id., on Rhizomnium punctatum (Hedw.) T. Kop., L. Barrio (AH 3222, 3223 sub L. lobatum); id., on Sphagnum sp., 8-XI-1982, L. Barrio (AH 2712 sub L. lobatum); id., on Calliergonella cuspidata, L. Barrio (AH 3230 sub L. lobatum); id., on Bryum caespiticium Hedw., 6-VII-1981, L. Barrio (AH 3221 sub L. lobatum); id., on Calliergon stramineum (Brid.) Kindb., 20-VII-1983, L. Barrio (AH 3226 sub L. lobatum); id., on Drepanocladus aduncus (Hedw.) Warnst., L. Barrio (AH 3227 sub L. lobatum). Pico del Nevero, 30TVL2937, 2150 m, on Calliergonella cuspidata, 6-XI-1982, L. Barrio (AH 3209 sub L. lobatum). Puerto de la Morcuera, on mosses, 24-IV-1999, F. Esteve-Raventós (AH 24999).
This species frequently has been reported from arctic and alpine areas and is mainly circumpolar in distribution (Kallio and Kankainen 1964, Gulden and Lange 1971, Høiland 1976, Redhead 1984, 1989), but it also is known from central Europe in the Alps, the Carpathian and the Ilgaz-Dagh mountains (Favre 1955, Ohenoja 1971, Kühner and Lamoure 1972). Arrhenia lobata is associated consistently with mosses and a parasitic relationship with moss gametophytes has been suggested (Savile and Parmelee 1964, Savile in Horak 1982).
Arrhenia lobata is a common species in mountainous areas of the Iberian Peninsula. This species appears to be restricted to the supramediterranean and oromediterranean belts of the Mediterranean (Madrid in Spain, Barrio et al 1985; Granada in Spain, Ortega et al 1997) and to the Eurosiberian montane, subalpine and alpine belts (Huesca in Spain, Esteve-Raventós et al 1997). In addition, it frequently is found on peaty sites (usually above 1200 m in the Iberian Peninsula) fruiting on mosses belonging to the genera Aulacomnium Schwaegr., Calliergonella Loeske, Drepanocladus (C. Muell.) G. Roth and Sphagnum L. (Barrio et al 1985, Ortega et al 1997). In contrast, this species never has been found on Sphagnum in northern Europe (Høiland 1976, Hallgrímsson 1981, Gulden 1988b). Although several specimens of this fungus were found fruiting in autumn (see specimens examined), most of the material studied from the Iberian Peninsula was collected between April and August (Barrio et al 1985, Esteve-Raventós et al 1997, Ortega et al 1997). Like Kuyper (1995a), we believe that A. lobata could represent a vernal species. Apart from the features of the basidioma, A. lobata can be distinguished easily from other nonomphalinoid species of Arrhenia by its broadly ellipsoidal spores (Q = 1.3).
Arrhenia retiruga (Bull. : Fr.) Redhead, Canadian J. Bot. 62: 873. 1984. Figs. 19–21
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Helvella retiruga Bull., Herb. France 11: 289, pl. 498, fig. 1. 1791 (basionym). Sanctioning author: Fr., Syst. mycol. 2(2): 324. 1823 sub Cantharellus retirugus. TYPUS: Bull. op. cit., pl. 498, fig. 1 (LECTOTYPUS!, cf. Redhead 1984: 873, 875).
Leptoglossum retirugum (Bull. : Fr.) Ricken, Blätterpilze Deutschland Fasc. 1: 6. 1910. Basidiomata cupulate to discoid, 0.35–1 x 0.25–1 cm. Pileus 0.25–1 cm wide, dorsal to ± central attached or sometimes fixed by one point, margin lobate, slightly hairy or glabrous, nonzonate, hygrophanous, yellowish gray or ochraceous to pale-brown when moist (10YR 6/4 to 6/6, 10YR 7/4 to 7/6), pale brown to yellowish when drying (10YR 8/4 to 8/6, 10YR 7/4 to 7/6). Pileipellis a cutis of parallel hyphae, 4–10 µm diam, usually with parietal and fine encrusting pigment, irregularly distributed on walls. Hymenophore brown to concolorous with pileus, smooth or very shallow to slightly venose, fully delimited by a sterile margin. Stipe absent. Cystidia absent. Basidia 25–32 x 4–8 µm, clavate, 4-spored, sterigmata usually straight and 3–4.5 µm long. Spores 7–11 x 4–6 µm, Q = 1.6, ellipsoidal to oval, sometimes lacrimiform, smooth, hyaline, nonamyloid. Clamps absent in all tissues.
Specimens examined. SPAIN. BARCELONA: Dosrius, 31TDG5004, 160 m, on Pseudoscleropodium purum (Hedw.) Fleisch. in Broth., 1-II-1984, A. Rocabruna and M. Tabarés (BCC-SCM 184 sub Leptoglossum retirugum). ZAMORA: Galende, 29TPG9264, 1000 m, on mosses in banks with Alnus glutinosa, 19-X-1999, F. Esteve-Raventós and M. Villarreal (AH 29774).
Few noteworthy microscopic differences separate Arrhenia retiruga and A. spathulata, and these species have been suggested to be ecological variants of the same taxon (Reid 1964). However, the hymenophore of A. retiruga is delimited by a sterile margin that is absent in A. spathulata (Redhead 1984). Specimens of A. retiruga collected in the Iberian Peninsula also can be distinguished from A. spathulata by the fine encrusting pigment of the pileipellis and spore shape. Only two collections of A. retiruga were studied, and more specimens should be examined to confirm these microscopic differences.
Arrhenia retiruga and A. spathulata can be distinguished by macroscopic features that might be an ecological adaptation. The cupulate habit of A. retiruga, implicating dorsal attachment and fixation by one point to the substrate, was described as an adaptation to growth on pleurocarpic mosses, where only small sites or cavities are available for the development of the hymenium (Horak 1982, Redhead 1984). The stipe of A. spathulata also is usually well developed and might represent an adaptation to growth on small compact carpets of acrocarpous mosses that serve to elevate the hymenium.
In the Iberian Peninsula A. retiruga is known only from the Mediterranean region, lowlands of the mesomediterranean belt (Barcelona in Spain, Rocabruna 1984, Rocabruna et al 1985, Tabarés and Rocabruna 1985, 1987) and riversides of the supramediterranean belt (Zamora in Spain).
Arrhenia spathulata (Fr.) Redhead, Canadian J. Bot. 62: 876. 1984. Figs. 22–25
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Cantharellus spathulatus Fr., Elench. fung. 1: 53. 1828 (basionym).
Thelephora muscigena Pers. 1801. Sanctioning author: Fr., Syst. mycol. 2(2): 323. 1823 sub Cantharellus muscigenum. TYPUS: Bull. op. cit., pl. 288 (LECTOTYPUS!, cf. Redhead 1984:876–877).
Leptoglossum muscigenum (Bull. : Fr.) P. Karst., Bidrag Kännedom Finlands Natur Folk 32: 242. 1879.
Arrhenia muscigena (Bull. : Fr.) Honrubia & Folgado in Folgado, Honrubia & M. J. Costa [sub "muscigenum"], Int. J. Mycol. Lichenol. 1: 357. 1984, nom. illegit., Art. 53.1.
Cantharellus lagunae (Lázaro Ibiza) Sacc. & D. Sacc., Syll. fung. 17: 37. 1905.
Basidiomata pleurotoid, 2.2–2.8 x 0.5–2.2 cm. Pileus 0.5–2.2 cm wide, spathulate to somewhat flabellate, margin lobate, glabrous, transversely slightly zonate, sometimes azonate, hygrophanous, gray, gray-brown to pale brown on moist or drying (2.5Y 6/1 to 6/2, 2.5Y 5/1, 2.5Y 4/1). Pileipellis a cutis of parallel hyphae, 3–7 µm diam, usually with parietal and ± dense encrusting pigment, zebra-like. Hymenophore concolorous with pileus, smooth or venose to intervenose, when venose often dichotomous or somewhat anastomosing, nondelimited by a sterile margin and continuous with the pileus. Stipe 0.3–0.5 x 0.1–0.3 cm, lateral, whitish to concolorous with pileus, glabrous to minutely hairy, solid. Cystidia absent. Basidia 28–37 x 4–8 µm, cylindrical to clavate, 4-spored, sterigmata usually curved and 4–6 µm long. Spores 7–10 x 4–5.5(–6) µm, Q = 1.7, elongate, sometimes ellipsoidal or lacrimiform, smooth, hyaline, nonamyloid. Clamps absent in all tissues.
Specimens examined. ANDORRA. LA MASSANA: Sispony, Cortals de Sispony, 30TCH7509, 1700 m, on Tortula ruralis, 17-X-2002, J. Campoamor (AH 30616). CZECH REPUBLIC. BOHEMIAE: Bohemia Centralis, Rèporyje, loco stepposo-saxa sectilia, in Festucetis, matrix: Syntrichia ("Tortula sp.") ruralis, 9-XI-1952, Z. Pouzar (NEOTYPUS PRM 171770 of Cantharellus spathulatus Fr. sub Leptotus spathulatus sensu Velen.; cf. Redhead 1984:876). Karlík, inter muscos ("Tortula sp.") ad tectum domi, 13-IX-1955, F. Kotlaba and Z. Pouzar (PRM 516633 sub Leptoglossum spathulatum (Fr.) Velen.). Locus "Teyssleruv mlyn" prope Radotín, loco stepposo, matrix: Syntrichium (Tortula) rurale, solo calcareo!, 9-XI-1952, Z. Pouzar (PRM 171771 sub Leptotus spathulatus sensu Velen.). Chynice prope Radotín, loco stepposo cum Stipa capillata, Festuca valesiaca, matrix: Syntrichium ("Tortula sp.") rurale, solo calcareo!, 9-XI-1952, Z. Pouzar (PRM 171772 sub L. spathulatus sensu Velen.). ITALY. VERCELLIS: sero autumno, in clivo sabuloso ad muscos, socio Agarico atrorufo, "on Rhacomitrium canescens", Cesati [Rabenh. Klotzschii Herb. vivum Mycol. 1-n° 1605] (LECTOTYPUS PRM 515156 of Leptotus queletii Pilát & Svrcek sub Cantharellus glaucus (Batsch) Fr.; cf. Redhead 1984:876). PORTUGAL. BEIRA BAIXA: Castelo Branco, São Fiel, ad terram inter muscos minores, XII-1902, C. Torrend (LECTOTYPUS S F21076 of Cyphella cochlearis var. subsessilis Bres., here designated: from left to right, the fifth basidioma of the upper and longer row). ESTREMADURA: Ribeira de Caparide, Alto do Moínho, Sôbre os grandes Musgos, XII-1929, P. Coutinho (LECTOTYPE LISU P48864-upper-sheet-material of Leptoglossum muscigenum var. azonum Cout., here designated: in the upper row, the first basidioma from left to right); id., II-1925 (LISU P48863 sub L. muscigenum var. azonum); id., I-1928 (LISU P48863 sub L. muscigenum var. azonum); id., XII-1928 (LISU P48864 sub L. muscigenum var. azonum); id., I-1929 (LISU P48864 sub L. muscigenum var. azonum). Marinha Grande, São Pedro de Moel, 29SME9801, 25 m, on mosses in dunes, 7-XI-2000, V. J. Rico and J. M. Barrasa (AH 29159). SPAIN. ALBACETE: Mesones, on mosses, 19-XI-1983, P. P. Moreno (MA-Fungi 43982 sub Leptoglossum muscigenum). BARCELONA: Dosrius, El Maresme, among mosses, 30-I-1994, A. Rocabruna (BCC-SCM 2403 sub Phaeotellus rickenii (Hora) Bon); id.: 13-I-1996, A. Rocabruna (BCC-SCM 2818); id.: on mosses among Cistus sp., 22-III-1991, A. Rocabruna (BCC-SCM 1716). Sant Cugat del Vallés, on mosses, 13-XII-1986, A. Rocabruna and M. Tabarés (BCC-SCM 168 sub Leptoglossum muscigenum). Santa Agnes de Malanyanes, on mosses under Cistus laurifolius L., 8-XII-1998, A. Rocabruna (BCC-SCM 3497). BURGOS: Río de Losa, Valle de Losa, 30TVN7755, 650 m, on Tortula ruralis, 8-XI-2002, E. Sarrionandía, I. Salcedo & I. Olariaga (AH 30621). GRANADA: Pantano de Cubillas, 7-XII-2000, among mosses, under Pinus halepensis Miller, F. D. Calonge (MA-Fungi 48132 sub L. muscigenum). GUADALAJARA: Molina de Aragón, on mosses, 17-XI-1975, C. Casas and J. Girbal (AH 751 sub L. muscigenum). Tamajón, on mosses, 25-X-1990, C. Ochoa, G. Moreno and M. Heykoop (AH 12808 sub L. muscigenum). Codes, on mosses, 13-X-1994, F. Esteve-Reventós and M. Trasviña (AH 25364). Huertahernando, 30TVL6562, 760 m, on mosses on calcareous soil with Pinus nigra Arnold, 26-X-2001, F. Esteve-Raventós and J. M. Barrasa (AH 29272). GUIPÚZCOA: Zumaia, Santioko hondartza, 30TWN6094, 10 m, on mosses in dunes, 1-XII-1990, J. M. Lekuona (ARAN A03981). MADRID: San Lorenzo de El Escorial, Herreria, en el suelo entre el musgo, I-1874, M. Laguna (LECTOTYPUS MAF-Fungi 1 of Dictyolus lagunae Lázaro Ibiza sub Cantharellus- laevis? Fries, here designated); id., Silla de Felipe II, 30TVL0291, 1100 m, on mosses on granitic, sandy soil in a forest of Quercus pyrenaica Willd., 23-XI-2002, V. J. Rico (AH 30619); id., on mosses on granitic rock in a forest of Quercus pyrenaica, 23-XI-2002, V. J. Rico (AH 30620). Guadalíx de la Sierra, 30TVL4315, 840 m, on mosses on calcareous rocks in a forest of Quercus faginea Lam. and Quercus rotundifolia Lam., 23-XI-2002, V. J. Rico (AH 30622). El Pardo, among mosses, 28-I-1975, G. Moreno (AH 11858 sub Leptoglossum muscigenum). Puerto de la Cruz Verde, among mosses with vegetation of Rubus sp. and Quercus sp., 1-XII-1975, T. Saenz, A. Egana and J. Salcedo (AH 11952 sub L. muscigenum). El Escorial, on Tortula ruralis (Besch.) Grout with vegetation of Quercus ilex L., 8-I-1976, J. Gómez, C. Gómez, J. A. Barbera and G. Moreno (AH 752 sub L. muscigenum). Cercedilla, on soil with mosses and Quercus pyrenaica, 7-XI-1976, F. D. Calonge (MA-Fungi 14747 sub L. muscigenum). Móstoles, Parque de Coimbra, among mosses, 22-I-1984, F. D. Calonge (MA-Fungi 5943 sub L. muscigenum). Carretera Madrid-Toledo, km 43, among living mosses, 26-II-1978, E. Álvarez (MA-Fungi 2786 sub L. muscigenum). San Martín de Valdeiglesias, río Cofio, 30TUK8774, 590 m, on mosses, 7-X-2002, V. J. Rico and J. M. Barrasa (AH 30617). MÁLAGA: Montejaque, arroyo del Cupil, 30TUF0271, 500 m, 13-XII-2001, A. Castro (MA-Fungi 50728). MURCIA: Teatinos, Sierra de la Fuensanta, on mosses, 20-I-1979, X. Llimona (MA-Fungi 43979 sub L. muscigenum). Puerto de la Cadena, on mosses, 13-II-1982, F. Robledano and F. Dicenta (MA-Fungi 43983 sub L. muscigenum). Rambla de la Bermeja, Sierra de Ricote, on mosses, XI-1980, P. del Olmo and M. Hurtado (MA-Fungi 43980 sub L. muscigenum). Valle del Leiva, Sierra Espuña, on mosses, 14-XI-1982, F. Alcaraz (MA-Fungi 43981 sub L. muscigenum). SEGOVIA: Fresno de Cantespino, 30TVL5576, 1050 m, on mosses on sandy soil of a roadside with Quercus ilex, 30-X-1999, J. M. Barrasa (AH 29262). SORIA: Almazán, dehesa de la Mitiosa, on mosses under Pinus sp., 21-X-1987, A. M. de Azagra (MA-Fungi 21235 sub L. muscigenum). TERUEL: Castillo de Albarracín, on mosses, 26-XI-1994, F. D. Calonge (MA-Fungi 33178 sub Leptoglossum sp.). TOLEDO: Yuncos, on mosses, 31-XII-1978, E. Álvarez (MA-Fungi 5292 sub Leptoglossum muscigenum). La Iglesuela, on mosses, 16-I-1996, M. Heykoop, J. L. Aguirre and M. Villarreal (AH 20741).
Arrhenia spathulata is widespread in the Iberian Peninsula and usually has been cited as Leptoglossum muscigenum (Malençon and Bertault 1976, Barrio et al 1985). In contrast with other species of Arrhenia (A. auriscalpium and A. lobata) known from the medium and upper belts of Eurosiberian and Mediterranean regions, A. spathulata frequently is found from the thermomediterranean (Estremadura in Portugal) to the supramediterranean (Segovia in Spain) belts of the Mediterranean region and more rarely from coline to montane belts of the Eurosiberian region (Andorra, Guipúzcoa in Spain). This species seems to be associated with acrocarpous mosses, and Jahn (1960) has suggested that it is restricted to Tortula ruralis (Hedw.) Gaert. et al (sub Syntrichia ruralis (Hedw.) F. Weber & D. Mohr, Jahn 1960). Arrhenia spathulata frequently is found in autumn and winter in open sites with sandy soils or on coastal dunes throughout the Iberian Peninsula. A similar ecology is reported for this species in central and northern Europe (Jahn 1960, Bon 1970, Kuyper 1995a).
The material from the Iberian Peninsula agrees with the macro and microscopic characters observed in the neotype selected by Redhead (1984) for Cantharellus spathulatus. The pleurotoid stipitate basidiomata, smooth or venose hymenophore, absence of clamps and spore shape are the most noteworthy features differentiating it from the other members of the genus.
Type material of Dictyolus lagunae Lázaro Ibiza (Lázaro e Ibiza 1902) collected in central Spain (MAF) and original material of Leptoglossum muscigenum var. azonum Cout. (Couthino 1932) (LISU) were examined. Lectotypes for these names are designated here and these species are synonymized with A. spathulata. See the commentary for A. auriscalpium for details concerning the new synonym Cyphella cochlearis var. subsessilis.
KEY TO THE TAXA OF ARRHENIA IN THE IBERIAN PENINSULA
1. Species with clamp connections . . . . . 2
1. Species without clamp connections . . . . . 5
2. Basidiomata pleurotoid, with very short or absent stipe, lamellae well developed. Spores broadly ellipsoidal to elongate, Q = 1.5–2.0. . . . . . 3
2. Basidiomata pleurotoid, stipe present or absent, lamellae forked, reduced to vein-like or rib-like wrinkles. Spores broadly ellipsoidal to ellipsoidal, Q = 1.3–1.7 . . . . . 4
3. Basidia tetrasporic. Spores 8–10(–11) x (4–)5–6.5 µm, Q = 1.5–1.7 . . . . . A. acerosa var. acerosa
3. Basidia bisporic, occasionally mono-, tri- or tetrasporic. Spores 10–13.5 x 5–6 µm, Q = [type: 1.8–]2.0 . . . . . A. acerosa var. tenella
4. Basidiomata dorsally or laterally attached, stipe absent. Spores broadly ellipsoidal, Q = 1.3. On mosses in peaty sites . . . . . A. lobata
4. Basidiomata stipitate, hymenophore well delimited from a solid stipe by a sterile lateral margin.. Spores ellipsoidal to slightly elongate, Q = 1.4–1.7. On soil among mosses . . . . . A. auriscalpium
5. Basidiomata cupulate, dorsally attached, stipe absent, hymenophore delimited by a sterile margin. Spores ellipsoidal to oval, Q = 1.6. Usually on pleurocarpous mosses . . . . . A. retiruga
5. Basidiomata pleurotoid, with a lateral and well-developed stipe continuous with the hymenophore. Spores elongate or lacrimiform, Q = 1.7. Usually on acrocarpous mosses . . . . . A. spathulata
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ACKNOWLEDGMENTS |
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FOOTNOTES |
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Accepted for publication February 14, 2003.
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LITERATURE CITED |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSTAXONOMYLITERATURE CITED |
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