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DOI: 10.3852/mycologia.98.4.650
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Mycologia, 98(4), 2006, pp. 650-658.
© 2006 by The Mycological Society of America

A re-evaluation of the Cortinarius scobinaceus and Cortinarius impolitus complex in the Mediterranean area


Antonio Ortega 1

     Departamento de Botánica, Facultad de Ciencias, Universidad de Granada, C/Severo Ochoa s/no. E-18071 Granada, España

Fernando Esteve-Raventós 2

     Departamento Biología Vegetal, Facultad de Farmacia, Universidad de Alcalá de Henaresm E-28071 Alcalá de Henares, Madrid, España

Francisco Bruno Navarro 3

     Grupo de Sistemas Forestales, Área de Recursos Naturales, Centro de Investigación y Formación Agraria (IFAPA, CICE, Junta de Andalucía), Camino de Purchil s/no, Aptdo. 2027, E-18080 Granada, España


   ABSTRACT
TOP
 ABSTRACT
INTRODUCTION
 MATERIAL AND METHODS
 RESULTS
 KEY TO TAXA
 DISCUSSION
 LITERATURE CITED
 

We have conducted a taxonomic study of Cortinarius belleri, C. cistohelvelloides, C. impolitus and C. scobinaceus from material collected in Spain, France, Italy, Portugal and Morocco. Based on anatomy, morphology, ecology and distribution (geographical data) we recognize two species, C. impolitus and C. scobinaceus, and the new combination C. scobinaceus var. cistohelvelloides is made. Cortinarius belleri, C. cistohelvelloides, C. impolitus and C. scobinaceus have been typed, and a key for identification of these taxa is included.

Key words: Cistus, Cortinarius, Mediterranean fungi, Spain, taxonomy


   INTRODUCTION
TOP
 ABSTRACT
 INTRODUCTION
MATERIAL AND METHODS
 RESULTS
 KEY TO TAXA
 DISCUSSION
 LITERATURE CITED
 
Malençon and Bertault (1970)Go reported Cortinarius scobinaceus from North Africa: a small species of Telamonia sect. Incrustati Melot, growing mainly in the spring, under various trees, Quercus suber, Eucalyptus spp. and Pinus pinea, or in maquis-shrub communities with Q. suber. The diagnostic characters of this species are (i) the small basidiomata (pileus 1.5–2 cm diam, stipe 1.8–2.2 x 0.15–0.25 cm); (ii) pileus surface covered by tiny woolly ochraceous squamules; (iii) stipe showing an abundant veil, forming bands or bracelets; (iv) amygdaliform-cylindrical spores, measuring 10–14 x 5–6 µm; and (v) the presence on the lamellar edge of septate cheilocystidia, with a claviform, subglobose or pyriform apical cell.

Beller (1982)Go described from France C. longisporus, another small Telamonia species, growing on sandy soils under Cistus salviifolius and in open areas of sandy pine forests C. longisporus is characterized by its ellipsoid-fusiform spores, (10–)12–13.5(–15) x 4.5–5.5 µm, and septate cheilocystidia and a fibrillose white veil on the pileus surface but not forming tiny woolly scales.

Moser (1983)Go redescribed Beller’s species and renamed it C. belleri M.M. Moser, owing to the existence of the prior valid binomial C. longisporus Peck. He argued that the presence of cheilocystidia was not a constant character and suggested that C. belleri was related closely to C. incisus (Pers.) Fr. sensu Moser and C. semivestitus M.M. Moser. C. belleri subsequently has been reported from different Mediterranean areas (e.g. Italy [Contu and Lavorato 1986Go, Ballero et al 1992Go, Quadraccia in press], Spain [Moreno et al 1990Go] and France [Moser 1983Go; Bon 1986Go, 1995Go]) always growing under Cistus, indicating a typical Mediterranean distribution.

Cortinarius cistohelvelloides, a taxon similar to C. belleri, also occur under Cistus. It was described by Bon (1992)Go and distinguished from C. scobinaceus by the purplish tinge of the lamellae and smaller spores [8.5–10.5(–11.5) x 5–5.5(–6) µm]. Ortega (1995)Go, after the study of a large number of collections of C. belleri from several Spanish and Italian localities and a re-evaluation of type material of C. scobinaceus, concluded that the taxa were conspecific. Campo (2004)Go concurs.

Moser and Ammirati (1996)Go discussed the taxa in sect. Incrustati, which includes some of the above taxa. They concluded that C. impolitus Kauffman (= C. incisus sensu Moser 1983Go) is closely related to C. belleri and that the latter might be an ecological variant growing under Cistus. However they considered C. scobinaceus a distinct species, related to C. angelesianus A.H. Sm. or C. psammocephalus (Bull.) Fr. Moser (personal communication) considered C. belleri and C. scobinaceus separate taxa based on these characters.

C. scobinaceus: (i) Pileus and stipe surface covered by ochraceous, lanose squamulae; (ii) cheilocystidia always present, broader than basidia; (iii) habitat always under Cistus spp., often only with this host present.

C. belleri: (i) Pileus and stipe surface covered by ephemeral lanose squamules, these always whitish and constituted by hyaline hyphae; (ii) cheilocystidia present or absent and always similar to basidia; (iii) habitat always under Mediterranean Pinus spp. and therefore collections listed under Cistus ssp. (in Pinus forests plant communities) in reality form mycorrhizas with Pinus species.

Vila and Llimona (2002)Go described C. cistohelvelloides var. phyllophlebophorus from Catalonia (Spain). It was separated from var. cistohelvelloides by the larger size and intervenose lamellae and was from Cistus stands where the typical variety also has been found.

In the present paper we have studied and evaluated a large number of collections, including type collections, under the names C. belleri, C. cistohelvelloides, C. scobinaceus and C. impolitus, either from the Iberian Peninsula or from other western Mediterranean countries. A comparison of basidiospore shape and ornamentation was made with scanning electron microscope (SEM). On the basis of all basidiomata characteristic and ecological data we conclude that only two species can be recognized in this complex, C. scobinaceus and C. impolitus, and consider C. cistohelvelloides a variety of C. scobinaceus.


   MATERIAL AND METHODS
TOP
 ABSTRACT
 INTRODUCTION
 MATERIAL AND METHODS
RESULTS
 KEY TO TAXA
 DISCUSSION
 LITERATURE CITED
 
The materials studied are conserved in these herbaria: AH (Univ. Alcalá de Henares, Madrid, Spain), BCC-SCM (Univ. Barcelona, Herbarium of the Societat Catalana de Micologia, Spain), GDAC (Univ. Granada, Spain), JVG (personal herbarium of J. Vila, Barcelona, Spain), MES (personal herbarium of R. Mahiques, Quatretonda, Valencia, Spain), ROHB (Univ. Roma, Italy), CAG (Cagliari, Italy), MICH (Michigan, USA), MPU (Univ. Montpellier, France), IB (Univ. Innsbruck, Austria) and Herb. M. BON (personal herbarium of M. Bon, St. Valérysur-Somme, France).

These characteristics are included (TABLE IGo): (i) basidiospore measurements, with values of maximum length (LM), minimum length (Lm), mean length (Ld), maximum width (AM), minimum width (Am), mean width (Ad), maximum length/width ratio (QM), minimum length/width ratio (Qm) and mean length/width ratio (Qd); (ii) habitat (Hb) (a) under Cistus spp., (b) in maquis shrubs with presence of Cistus and Quercus or Pinus, (c) under Quercus suber, (d) in mixed broadleaf forests, or (e) under Pinus; (iii) cheilocystidia (Cis) (a) absence, (b) unicellular and (c) pluricellularseptate; (iv) bioclimatic belt (according to Rivas Martínez et al [1997]Go and Rivas Martínez and Loidi [1999]Go) (Pb) (a) supramediterranean, (b) thermo- or mesomediterranean; and (v) phenology (Fe) (a) spring, (b) autumn and (c) winter.


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  		TABLE I. Data used in correspondence analysisa

 
From these data only the mean values of basidiospore measurements have been used. A correspondence analysis (CA) with Statistica 6.0 to establish possible relations among populations of the species is provided. The number in parenthesis, after the name of each locality, refers to the material studied (TABLE IGo) for basidiospore measurements (n = 30 basidiospores x 3 basidiomata), was used to calculate the average of each parameter and the ratio length/width (Q: L/l). These values enabled us to calculate the average basidiospore size of each collection. The Spearman rank correlation was used to relate the mean length and width of the basidiospores (metric variables), as well as the length of the basidiospores and the absence or presence of one-celled or multicelled cystidia (nominal variable).

A one-way ANOVA was performed to determine the dependence of the basidiospore length with respect to the absence of cheilocystidia and types. Bartlett’s test was used to test the homogeneity of the variance, and Turkey’s multiple-comparison test was used to establish the differences between the mean values. Samples used for SEM of basidiospores were submitted to critical point drying, according to the method of Moreno et al (1995)Go.


   RESULTS
TOP
 ABSTRACT
 INTRODUCTION
 MATERIAL AND METHODS
 RESULTS
KEY TO TAXA
 DISCUSSION
 LITERATURE CITED
 
Cortinarius impolitus Kauffman, Publications Mich. Geol. Biol. Surv., Biol. Ser. 5(26):419, 1918. FIG. 1Go = Cortinarius incisus Fr. sensu M.M. Moser FIG. 2Go


Figure 1
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  		FIGS. 1–6. 1. Cortinarius impolitus (holotype). Basidiospores. 2. Cortinarius incisus (IB 70-297). Basidiospores. 3. Cortinarius longisporus (holotype). Basidiospores. 4. Cortinarius scobinaceus (GDAC 44206). Basidiospores. 5. Cortinarius scobinaceus (GDAC 44206): Cheilocystidia. 6. Cortinarius cistohelvelloides (holotype). Basidiospores.

 
Material examined. – USA: New Richmond (1), on sandy soi1 among mosses in low moist places in white pine and beech woods, 19-9-1912, leg. Kauffman, MICH 10366 (Holotype). SWEDEN: Femsjö (26), Småland, Schnoden, under Pinus and Picea, 15-9-1990, leg. M.M. Moser, IB 70/241. DENMARK: Keby, Hedeskov, Fühen (27), under Picea, 24-9-1970, leg. M. Lange & M.M. Moser, IB 70/297. SPAIN: Ávila, Peguerinos (46), under Pinus sylvestris, 13-4-1990, leg. F. Esteve-Raventós, AH 30963. Guadalajara, Ciruelos del Pinar (50), under Pinus pinaster, 24-5-1993, leg. F. Esteve-Raventós, AH 30970. Madrid, El Paular, mirador de los Robledos (47), under Pinus sylvestris, 27-4-1988, leg. F. Esteve-Raventós, AH 30964. Ibidem (48), 26-6-1992, AH 30965. Madrid, Navacerrada, pradera de las Cortes (49), under Pinus sylvestris, 22-5-1993, leg. F. Esteve-Raventós, AH 30969. Segovia, Sepúlveda, Parque Natural del río Duratón (45), under Pinus pinaster, 30-10-1994, leg. F. Esteve-Raventós, AH 18992.
Sel. Descr. Kauffman, Cortinarius Fr. North American Flore 10:336, 1932; Moser, Mycol Helvet 1(1):3, 1983 (as Cortinarius incisus); Moser and Ammirati, Mycotaxon 58:400–404, 1996. Sel. Icon. Bresadola, Iconograph Mycol XIII: tab. 656, 1927 (as Cortinarius incisus); Moser, Mycol Helvet 1(1): FIGS. 1 and 3CGo, 1983 (as Cortinarius incisus).

Notes. – Diagnostic characters of this species are (i) habitat diverse, either in coniferous (Pinus, Abies) or mixed forests with the presence of Fagus (Moser and Ammirati 1996Go); (ii) it is a mesophilous or montane species; (iii) the pileus is covered by an arachnoid-fibrillose veil, fugacious and remaining only on the pileus margin; (iv) basidiospores (FIGS. 1, 2Go) ellipsoid-subamygdaliform to cylindrical and relatively narrow (7.8–) 8–11(–12) ((3.8–)4–5.5 µm, Xm = 8.9–10.3 (4.2–5 µm, Q: L/l= 1.6–2.5, Qm = 1.8–2.3; (v) cheilocystidia absent; and (vi) stipe covered by a fugacious whitish to cream-colored veil, which generally forms an annular band toward the lower half.

Moser and Ammirati (loc. cit. 402) conclude that C. impolitus y C. incisus sensu Moser are conspecific. We have studied the type material of C. impolitus and two collections typical of C. incisus (IB 70/241, IB 70/297) and we can support this hypothesis, given the similarity of characters between these collections (basidiomata, basidiospores, habitat, etc.). For this reason we consider the two taxa synonymous.

Cortinarius scobinaceus Malençon & Bertault, Fl. Champ. Supér. Maroc 1:541, 1970. We recognize two varieties:

Cortinarius scobinaceus Malençon & Bertault, Fl. Champ. Supér. Maroc 1:541, 1970 var. scobinaceus = Cortinarius belleri M.M. Moser, Mycol Helvet I (1):4, 1983.

= C. longisporus Beller non Peck, Doc Mycol 12(46):32, 1982. FIG. 3Go

Material examined of Cortinarius scobinaceus var. scobinaceus. – ITALY: Roma, Sandalo-Nettuno (12), under Pinus pinea, 17-11-1986, leg. L. Quadraccia, ROHB No. 1051 LQ.- Sardegna, Maracalagonis (21), under Cistus, 6-10-1988, leg. M.Contu, CAG s.n. Sardegna, prov. Sassari, Tempio P., S. Bachisio, Baldo (56), among mosses near Quercus suber, 23-5-1999, leg. M. Contu, CAG 5/8 61B. Ibidem (57), 2-4-1999, CAG 5/8 61C. Ibidem (58), 22-4-1998, CAG 5/8 61D. MOROCCO: Rabat, Fôret de Mamora (3), on sandy soil under Quercus suber, 4-1-1936, leg. G. Malençon, Herb. G. Malençon No. 214 (MPU) (Holotype). PORTUGAL: Extremadura, Marinha Grande, S. Pedro de Moel, Pedras Negras (54), in coastal dunes under Cistus and Pinus pinaster, 7-XI-2000, leg. F. Esteve-Raventós, AH 29848. Trasos-Montes, Rabal (51), under Cistus ladaniferus, 21-X-1999, leg. F. Esteve-Raventós, AH 29799. SPAIN: Girona, Puig Margall, Vilajuïga, Alt Empordà (31), alt. 170 m, under Cistus monspeliensis, 18-11-1997, leg. J. Vila, JVG971118-8. Girona, Castle of Quermançó, Vilajuïga, Alt Empordà (34), alt. 95 m, under C. monspeliensis, 25-1-1999, leg. J. Vila & X. Llimona, JVG990125-5. Girona, Cala Portaló, Cadaqués, Alt Empordà (43), alt. 35 m, under C. monspeliensis, 8-1-1999, leg. J. Vila and X. Llimona, JVG90108-12. Granada, La Alcaicería, Prados del Pinar (14), under C. laurifolius, 8-11-1994, leg. A. Ortega and L. Alcoba, GDAC 44207. Ibidem (16), 14-11-1988, leg. A.Ortega, GDAC 30560. Granada, Casa forestal de Bolones, Natural Park of Sierra de Huétor (18), under Pinus halepensis, Cedrus atlantica, Quercus ilex subsp. ballota and Cistus albidus, 16-5-1993, leg. A.Ortega, GDAC 44206. Jaén, El Centenillo (22), under Cistus laurifolius and C. ladaniferus, 23-11-1988, leg. A.Ortega, GDAC 30785. Madrid, Miraflores de la Sierra (55), under Cistus laurifolius, 21-10-1990, leg. F. Esteve-Raventós and V. González, AH 30962. Segovia, Fresno de Cantespino, Prado Pinilla (52), under Cistus laurifolius and Quercus pyrenaica, 11-3-1995, leg. J.M. Barrasa and F. Esteve-Raventós, AH 16933. Sevilla, Aznalcázar, under Cistus monspeliensis, 2-12-2000, leg. L. Alcoba and A. Ortega, GDA 47374. Valencia, Els Surars, Pinet, Vall d’ Albaida (17), under Cistus crispus and C. salvifolius, 3-12-1994, leg. R. Mahiques, MES 2363. Ibidem (19), under C. crispus, 12-10-1996, leg. R. Mahiques, MES 2926. Ibidem (23), 19-11-1994, MES 2332.

Material examined of Cortinarius belleri. – FRANCE: Moléts (2), on sandy soils under Cistus salviifolius, in open pine woods, 27-2-1972, leg. J. Beller, Herb. M. BON No. 824-2402.1, fiche Bon: 7202 2801 (Holotype of Cortinarius longisporus Beller). L’Oustaou de Dieu, Porquerolles (25), under Cistus, 13-11-1980, leg. C. Furrer, IB 80/621. SPAIN: Cáceres, Finca de las Cansinas, Natural Park of Monfragüe (15), under C. ladaniferus, 5-11-1987, leg. G. Moreno and F. Esteve-Raventós, AH 10658. Granada, La Alcaicería, Prados del Pinar (11), under Cistus laurifolius, 12-1-1990, leg. A. Ortega, GDAC 31395. Ibidem (20), 14-11-1991, GDAC 36771. Valencia, Pla de Suros, Barx, Safor (9), under C. crispus and C. salviifolius, 12-10-1995, leg. R. Mahiques, MES 2524.

Sel. Descr. Malençon and Bertault, Flore des Champignons Supérieurs du Maroc 1:537–541, 1970; Moser, Mycol Helvet 1(1):4–5, 1983 (as Cortinarius belleri); Contú and Lavorato, Gruppo Micologico G. Bresadola 29(1–2): 94-95, 1986 (as Cortinarius belleri).

Sel. Icon.: Contú and Lavorato, Gruppo Micologico G. Bresadola 29(1–2):96, 1986 (as Cortinarius belleri); Vila and LLimona, Rev Catalan Micol 21:136, 1998; Campo, J Journé europ Cortinaire 7(6):64, 2004.

Notes. – Diagnostic characters of this taxon are (i) its habitat in typically Mediterranean plant communities, with Pinus halepensis, P. pinea, Quercus ilex subsp. ballota, Q. suber, mostly with the presence of different Cistus species (C. albidus, C. crispus, C. ladaniferus, C. laurifolius, C. monspeliensis and C. salviifolius), in thermo- and mesomediterranean belts; (ii) pileus surface covered when young with an abundant fibrillose, whitish to cream-ochraceous veil, often forming lanose and normally persistent squamules, at least toward the center, this veil is made of hyphae with an incrusting parietal pigment, variable in thickness and density; (iii) lamellae reddish-brown to ochraceous, with a whitish edge, although they may show a purplish tinge in some collections (e.g. JVG990108-12); (iv) basidiospores (FIG. 4Go) ellipsoid-amygdaliform to cylindrical-fusiform, 9–13.5 x 4.5–6.5 µm, Xm = 10.2–11.8 x 4.6–5.8 µm, Q: L/l = 1.6–2.6; Qm = 1.9–2.4; (5) cheilocystidia present in most collections, pluricellular-articulate (FIG. 5Go), with claviform to subglobose apical cell, 8.5–22 µm wide, seldom unicellular.

C. scobinaceus var. scobinaceus differs from C. impolitus by the habitat, more developed and persisting veil and bigger basidiospores, and from C. scobinaceus var. cistohelvelloides, by the differently shaped and somewhat larger basidiospores.

Based on a study of the holotype and several collections labeled or determined as C. belleri, we have observed in some collections only slight quantitative differences related to the amount of veil that covers the pileus and stipe surface; this character, variable and influenced by environmental conditions, is not taxonomically significant. Basidiospore (FIG. 3Go) shape and measurements of C. belleri (9–13.7 x 4.5–6.3 µm, Xm = 10.2–11.8 x 4.9–5.5 µ, Q: L/l = 1.7–2.5; Qm = 2.1–2.2), fit within the range of variation for C. scobinaceus var. scobinaceus. The cystidia of the holotype (IB 80/621) show articulate cheilocystidia, with claviform to subglobose apical cell, 10–24 µm diam. However in other collections they are not well differentiated or look similar to basidioles (GDAC 31395, 36771; MES 2524); the presence/absence of cheilocystidia was pointed out by Malençon (Malençon and Bertault 1970Go), suggesting that the arrangement of cheilocystidia along the lamellar edge is not uniform and that some lamellae portions away from the pileus margin could be devoid of these cells. We also assume that this character is tied to age and developmental conditions, and what we call "cystidia" are terminal elements of the hymenophoral trama, similar to those of Inocybe subgenus Mallocybe Kühner.

C. belleri would require the presence of Pinus to develop, according to Moser (personal communication), but this opinion has not been confirmed by this study. For example IB 80/621, GDAC 31395, GDAC 36771 and MES 2524 grow in pure Cistus stands. Also collection IB 80/621 (from Porquerolles, France) was gathered from the same locality and habitat as C. cistohelvelloides collection (33). On the basis of these morphological and ecological data (habitat, basidiospore features and cheilocystidia) we consider C. belleri as conspecific with C. scobinaceus (FIG. 5Go.) Quantitative differences in veil amount cannot justify even a treatment at the varietal level.

Cortinarius scobinaceus Malençon & Bertault var. cistohelvelloides (Bon) A.Ortega & Esteve-Rav., comb. Nov.

Basionym: Cortinarius cistohelvelloides Bon, Doc Mycol 85:52, 1992.

= Cortinarius cistohelvelloides var. phyllophlebophorus Vila & Llimona, Rev Catalan Micol 24:91, 2002.

Material examined. – FRANCE: Biot (Alpes Maritimes) (32), under Cistus and Quercus suber, 7-11-1991, leg. M. Bon, Herb. M. BON No. 91244 (Holotype). Porquerolles (33), under Cistus salviifolius and C. monspeliensis, 9-11-1991, leg. M. Bon, Herb. M. BON No. 91244 (Isotype). SPAIN: Cádiz, Grazalema, cortijo de Chusco, under Cistus albidus, 6-1-2004, leg. F. Prieto, AH 30902. Girona, Mas dels Rabassers de Baix, Cadaqués, Alt Empordà (36), alt. 105 m, under C. monspeliensis and C. salviifolius, 25-1-1999, leg. J. Vila and X. Llimona, JVG990125-1. Ibidem, JVG990125-32. Ibidem (40), JVG990125-9. Ibidem (44) (as Cortinarius cistohelvelloides var. phyllophlebophorus), BCC SCM B-4060 (Holotype). Girona, Sant Pele de Rodes, El Port de la Selva, Alt Empordà, (38), alt. 470 m, under C. monspeliensis and C. albidus, 8-1-1999, leg. J. Vila and X. Llimona, JVG990108-38. Girona, Coll de la Perafita, Roses, Alt Empordà (39), alt. 250 m, under C. monspeliensis, 8-1-1999, leg. J. Vila and X. Llimona, JVG990108-6. Granada, La Alcaicería, Prados del Pinar (35), under C. laurifolius, 14-12-1988, leg. A.Ortega, GDAC 44205. Ibidem (6), under Cistus ladaniferus, 5-11-1990, leg. A.Ortega, GDAC 36768. Ibidem (41), 5-11-1999, leg. L. Alcoba and A. Ortega, GDAC 44365. Ibidem (42), GDAC 44366. Huelva, Sierra de Gimón Peréz, Natural Park of Sierra de Aracena and Picos de Aroche (7), under C. laurifolius, 21-11-1990, leg. A. Ortega. GDAC 36769. Jaén, El Centenillo (4), under C. ladaniferus, 13-11-1990, leg. A. Ortega, GDAC 36770. Segovia, Fresno de Cantespino, Prado Pinilla (53), under Cistus laurifolius, leg. J.M. Barrasa and F. Esteve-Raventós, 13-5-1995, AH 16951. Valencia, Els Surars, Pinet, Vall d’ Albaida (29), under C. crispus, 7-11-1992, leg. R. Mahiques, MES 1812. Valencia, Pla de Suros, Barx, Safor (30), under C. crispus, 17-11-1993, leg. R. Mahiques, MES 2112.

Sel. Descr. Bon, Doc Mycol 22 85):52–53, 1992, (as Cortinarius cistohelvelloides).

Sel. Icon. Bon, Doc Mycol 22(85): FIG. A, 1992, (as Cortinarius cistohelvelloides); Mahiques, Butlletí Soc Micol Valènc 6:129, 2001 (as Cortinarius cistohelvelloides); Vila and LLimona, Rev Catalan Micol 24:119, 2002, (as Cortinarius cistohelvelloides var. phyllophlebophorus).

Notes. – Diagnostics characters of this taxon are (i) found always under Cistus (C. albidus, C. crispus, C. ladaniferus, C. laurifolius, C. monspeliensis, C. salviifolius), in xeric plant communities, in thermo- and mesomediterranean belts; (ii) pileus mostly covered with copious whitish to cream-ochraceous veil, forming lanose upturned scales, normally persisting, and also present in herbarium specimens; (iii) lamellae, in most collections, brown-purplish (chocolate), with a whitish or pale edge; (iv) stipe covered by an abundant whitish veil, sometimes forming a peronate sheath, sometimes showing squamulose bands; (v) basidiospores (FIG. 6Go) ellipsoid to ellipsoid-subamygdaliform (never fusiform), 8–12.5 x 4–6(–6.5) µm, Xm= 8.8–10.5 x 4.3–5.3 µm, Q: L/l = 1.6–2.4; Qm = 1.8–2.1; and (vi) cheilocystidia similar to basidioles or more frequently unicellular claviform, pyriform or sometimes pluricellular multiseptate, like those of var. scobinaceus near the pileus margin or with 1–2 septa, the apical cell being cylindrical to claviform, 10–13(–18) µm wide.

The cistophilous habitat and the larger basidiospores, with a more ellipsoidal-subamygdaliform shape (Qm: L/l = 1.8–2.1) distinguish this taxon from C. impolitus. From var. scobinaceus it differs in most collections by purplish tinged lamellae and the differently shaped basidiospores. The basidiospores also are smaller, especially in length, with a slightly lower ratio length/width (1.8–2.1 versus 1.9–2.4), although some overlapping between both taxa occurs, placing C. cistohelvelloides at the varietal level based on these results. Smaller basidiospores were reported for C. scobinaceus by Malençon and Bertault (1970Go:538).


   KEY TO TAXA
TOP
 ABSTRACT
 INTRODUCTION
 MATERIAL AND METHODS
 RESULTS
 KEY TO TAXA
DISCUSSION
 LITERATURE CITED
 

1. Basidiospores large, Xm = 10.2–11.8 x 4.6–5.8 µm, (Qm = 1.9–2.4); habitat diverse, under Cistus, Eucalyptus spp., Pinus pinea, Quercus suber, etc., in thermomediterranean and mesomediterranean biogeographic belts; cheilocystidia mostly articulate Cortinarius scobinaceus var. scobinaceus
1'. Basidiospores smaller, Xm = 8.8–10.5 x 4.2–5.3 µm. 2
2. Basidiospores Xm = 8.8–10.5 x 4.3–5.3 µm, (Qm = 1.8–2.1); habitat strictly under Cistus spp. in thermomediterranean and mesomediterranean belts; cheilocystidia mostly a single cell C. scobinaceus var. cistohelvelloides (=C. cistohelvelloides)
2'. Basidiospores Xm = 8.9–10.3 x 4.2–5 µm, (Qm = 1.8–2.3); habitat diverse, under coniferous and broadleaf trees in mesophilous forests (supramediterranean belt), never in xerophilous habitats under Cistus spp.; without cheilocystidia C. impolitus


   DISCUSSION
TOP
 ABSTRACT
 INTRODUCTION
 MATERIAL AND METHODS
 RESULTS
 KEY TO TAXA
 DISCUSSION
LITERATURE CITED
 
Most of the collections can be placed in one of two groups based on correspondence analysis of data (TABLE IGo, FIG. 7Go).


Figure 2
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  		FIG. 7. Correspondence analysis of data (TABLE IGo). Cb: Cortinarius belleri; Ci: Cortinarius impolitus; Cc: Cortinarius cistohelvelloides; Cs: Cortinarius scobinaceus. (T): holotype; (IT): Isotype. Number with each taxon belongs to sample number. Ba: Samples without cheilocystidia. Bb: Samples with unicellular-cheilocystidia. Bc: Samples with articulate-pluricellular-cheilocystidia.

 
Group A: This includes C. impolitus collections (1), (26), (27) and (45), which produce basidiomata in autumn, and also collections (46), (47), (48), (49) and (50), which produce them in spring. All collections share these characters: (i) occurrence in the supramediterranean belt or area, (ii) habitat in Pinus forests, (iii) absence of cheilocystidia and (iv) relatively small basidiospores (Xm = 9.5 x 4.7 µm).

Group (B): This is heterogeneous and represents those samples collected under Cistus ssp. in areas within the thermo- and mesomediterranean bioclimatic belts. In this group we can differentiate three subgroups: (Ba) including those samples devoid of cheilocystidia, such as C. belleri collections (9), (11) and (20), and C. cistohelvelloides collections (29) and (37). Subgroup (Bb) includes samples with unicellular cheilocystidia, such as C. cistohelvelloides collections (32), (33), (35), (36), (38), (39) and (40), and C. scobinaceus collections (15), (31) and (51). A third subgroup (Bc) includes samples with articulate-pluricellular cheilocystidia, such as C. belleri collections (2) and (25), C. cistohelvelloides collections (4), (6), (7), (30), (41), (42) and (44) and C. scobinaceus collections (14), (15), (16), (17), (19), (21), (22), (34) and (43).

Spore measurements (FIG. 8Go) also have helped to establish some good comparison criteria, as is shown in the taxonomic key.


Figure 3
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  		FIG. 8. Basidiospore size variation (mean values) of the different taxa. Cb: Cortinarius belleri; Ci: Cortinarius impolitus; Cc: Cortinarius cistohelvelloides; Cs: Cortinarius scobinaceus. (T): holotype; (IT): Isotype. Number with each taxon belongs to sample number.

 
As cited by Moser (1983)Go we confirm that the presence/absence of cheilocystidia and cystidial morphology are respectively inconsistent and variable characters and cannot be used as discriminatory taxonomic features. Articulate-pluricellular cheilocystidia are more usual than unicellular ones in C. scobinaceus (=C. belleri), which together with the larger basidiospore size (10.2–11.8 x 4.6–5.8 µm), help differentiate it from C. cistohelvelloides (with or without cheilocystidia and smaller spore size, 8.8–10.5 3 4.3–5.3 µm).

There was a significant difference between the mean length of the basidiospores in different populations in relation to absence: (i), types of cystidia (2 to 3) (one-way ANOVA results: F = 8.74, df = 2, P-value = 0.0006; FIG. 9Go). This indicates that basidiospore mean length can be used as a diagnostic character of those acystidiate (1) species as compared to those with pluricellular cystidia (3).


Figure 4
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  		FIG. 9. One-way Anova (P-value = 0.0006) for mean length ± SE of the basidiospores in acystidiate species (1), species with unicellular cystidia (2) and species with pluricellular cystidia (3). Tukey test was applied as post hoc test in multiple comparisons. Different letters show significant differences at a 95% significance level.

 
The mean length of the basidiospores increased positively from the noncystidiate species to those with unicellular cystidia, or pluricellular cystidia (Spearman correlation coefficient r = 0.52, P-value = 0.0001). However a clear overlapping in spore measurements in the studied collections with unicellular (2) and pluricellular cystidia (3), which may correspond both to C. cistohelvelloides and C. scobinaceus, has been observed, suggesting that it would be more reasonable to treat them as varieties of the same species.

Finally it bears mentioning that the phenology is not a discriminatory taxonomic character. C. impolitus (Group A, FIG. 7Go) produces basidiomata in spring and autumm, while C. scobinaceus sensu lato (Group B), does so from spring to winter.

A large number of taxa in sect. Incrustati are represented in Europe, most of them growing in montane or boreal woodlands. C. semivestitus M.M. Moser appears to be close to C. impolitus in habitat and spore characters; it can be differentiated by the yellow-ochraceous veil, formed by hyphae with yellowish walls (x 5–11 µm), whereas C. impolitus has a whitish to pale ochraceous veil with hyaline walls (x 3–7 µm) according to Moser (1983)Go and Moser and Ammirati (1996)Go. Another species with a yellow veil is C. luxnymphae Melot (= C. incisus sensu auct.), but the spores are much smaller, measuring 6.5–8 x 3–3–5 µm (Brandrud et al. 1994Go).

Some species of this section are characterized by the long and "boletoid" spores; this is the case of C. arcanus G. Moreno, Heykoop & E. Horak from Mediterranean areas, characterized by its stout habit, a distinct brown annular zone at the stipe base and narrow spores (11–15 x 3.5–5 µm) (Moreno et al 2004Go). C. arcanus and C. heterosporus Bres. share similar spore shapes although they are smaller in the latter (7.5–10.5 x 2.3–3.5 µm), according to Moser (2000), or 8.2–10.5 x 2.4–3 µm, according to Moreno et al (loc. cit.). Some similarity, especially in veil development, also be can observed among C. scobinaceus, C. angelesianus A.H. Sm. and C. psammoce-phalus (Bull.) Fr.; these two species show smaller spores than C. scobinaceus, measuring 7–8.5 x 4.5–5.5 µm (Brandrud et al 1994Go) and 7.5–9 x 4.5–6 µm (Brandrud et al 1998Go) respectively.


   ACKNOWLEDGMENTS
 
We thank Dr M.M. Moser for his valuable taxonomical opinions and Prof. M. Bon (St Valéry-sur-Somme, France) for his comments on C. cistohelvelloides and the loan of the holotype; the curators at MICH (Michigan, USA), MPU (Montpellier, France) and lB (lnnsbruck, Austria), Dr M. Contú (Olbia, Italia), Prof. X. Llimona (Universitat de Barcelona), R. Mahiques (Valencia) and J. Vila (Barcelona) for the loan of types and other collections that have made this contribution much easier. Finally we thank Drs J.D. Bueno and A. González (Servicio de Apoyo Técnico a la Investigación, Universidad de Granada), for their useful help in SEM techniques and study of the spores.


   FOOTNOTES
 
Accepted for publication May 5, 2006.

2 E-mail: fernando.esteve{at}uah.es Back

3 E-mail: francisco.bruno.ext{at}juntadeandalucia.es Back

1 Corresponding author. E-mail: aortegad{at}ugr.es


   LITERATURE CITED
TOP
 ABSTRACT
 INTRODUCTION
 MATERIAL AND METHODS
 RESULTS
 KEY TO TAXA
 DISCUSSION
 LITERATURE CITED
 
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———. 1992. Agaricomycétes mediterraneéns ou meridionaux. Doc Mycol 85:51–62.

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———, Esteve-Raventós F, Illana C. 1990. Estudios micológicos en el parque natural de Monfragüe y otras zonas de Extremadura (España), IV. Agaricales. Boletín Soc Micol Madrid 14:115–142.

———, Heykoop M, Horak E. 2004. Un nouveau Cortinaire à spores fusoïdes décrit d’Espagne. Bull Soc mycol France 120(1–4):157–168.

Moser MM. 1983. Notizen zu einigen Cortinarien aus der untergattung Telamonia. Mycol Helvetica I(1):1–16.

———. 1997. On the ocurrente of two Cortinarius species described by Bresadola in North America. Bollettino del Gruppo Micologico G. Bresadola-Nuova serie 40(2–3):337–342.

———, Ammirati JF. 1996. Studies in North American Cortinarii II. Interesting and new species collected in the north Cascade Mountains, Washington. Mycotaxon 58:387–412.

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———, Loidi J. 1999. Bioclimatology of the Iberian peninsula. Itinera Geobotan 13:41–47.

Vila J, Llimona X. 2002. Novedades sobre el component fúngic de les comunitats de Cistus de Catalunya. Rev Catalan Micol 24:75–121.





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