Species diversity of Genea (Ascomycota, Pezizales) in Europe Pablo ALVARADO Julio CABERO Gabriel MORENO Zoltán BRATEK Nicolas VAN VOOREN Vasileios KAOUNAS Giorgos KONSTANTINIDIS Carlo AGNELLO Zsolt MERÉNYI Matthew E. SMITH Summary: Here we revisit the species diversity within the genus Genea in Europe. Nine new species are proposed: Genea brunneocarpa, G. compressa, G. dentata, G. fageticola, G. oxygala, G. pinicola, G. pseudobalsleyi, G. pseudoverrucosa, and G. tuberculata to accommodate new collections that do not match any previously described taxa. The new species are described with accurate morphological and ecological data. Genea lobulata is also proposed as a new combination to elevate the previously recognized variety, Genea sphaerica f. lobulata, to the species level. Useful macroscopic images and spore drawings of all ten new taxa are included to help discriminate them from closely related species. Keywords: hypogeous fungi, trules, Europe, phylogeny, taxonomy, Pyronemataceae. Ascomycete.org, 6 (3) : 41-51. Juillet 2014 Mise en ligne le 01/07/2014 Introduction The genus Genea Vittad. was published by Carlo VITTADINI (1831), who dedicated it to zoologist Dr. Joseph Gené. The genus is characterized by its hypogeous ascomata, that are sometimes folded and usually have a basal tuft of hyphae. Genea ascomata typically have an apical oriice, but this can be diicult to locate because of the many lobes that sometimes surround it. The ascomata develop an organized hymenium (ptycothecium) that is protected by an epithecium formed by coalescent paraphyses. Asci contain eight ornamented spores and are inamyloid. Traditionally, a limited number of Genea species are recognized in Europe. VITTADINI (1831) irst proposed the blackish G. verrucosa Vittad. and the reddish G. papillosa Vittad. Soon after, DIETRICH & KLOTZSCH (1839) published a description of some specimens of G. verrucosa, which they considered conspeciic with the taxon Hydnocaryon fragrans Wallr. (WALLROTH, 1833). BERKELEY & BROOME (1846) proposed that these specimens to be accommodated in the new independent species Genea klotzschii Berk. & Broome. However SACCARDO (1889) correctly identiied the appropriate epithet as Genea fragrans (Wallr.) Paoletti. This species has bigger spores than those of G. verrucosa (BERKELEY & BROOME, 1846), although diferent measures have been reported: 35–38 μm (CORDA, 1854), 31–45 × 21– 32 μm (FISCHER, 1897), or 25–34–45 × 20–27–32 μm (HAWKER, 1954). Genea fragrans also has ornamentation formed by large, truncated, block-like warts that are distinct from the small conical warts of G. verrucosa (MATTIROLO, 1900a). BERKELEY & BROOME (1846) also published a description of some G. papillosa specimens from the United Kingdom that were quite diferent from those originally described by VITTADINI (1831). These were later recognized as a new species by TULASNE & TULASNE (1851) under the name Genea hispidula Berk. ex Tul. & C. Tul. This species has small brown ascomata and a peridium coated with pyramidal warts and densely covered by brown hairs. It has a single inner chamber covered with a black, warted epithecium. Its spores are larger than those of G. verrucosa (38–42 × 32 μm) and covered with coarse warts that are sometimes bicuspidate (with the appearance of a molar tooth). A diferent type of spore ornamentation was recorded by TULASNE & TULASNE (1851) in some French specimens, which were accommodated in the taxon Genea sphaerica Tul. & C. Tul. This species produces perfectly sphaerical black ascomata, with a labyrinth-like inner chamber that has characteristic broad white tramal plates. The spores of G. sphaerica are ornamented with obtuse roundish warts. CORDA (1854) proposed many new GENEA species, although most of them have been rarely collected since they were described. First, he published the easily recognizable species G. lespiaultii Corda, which has spores ornamented by lat warts and has been recorded frequently. He also proposed a second species with hairy peridium, Genea pulchra Corda, which difered from G. hispidula because of its smaller spores (20–28 × 15–24 μm), spore ornamentation composed of minute warts, and a pseudoparenchymatic peridium with scattered lacunae (locules inside the ascomata that are lined with a palisade of hymenium). Finally, he erected two species closely resembling G. verrucosa, namely the brownish G. kunzeana Zobel, and the more tuberculate G. perlata Corda. However, G. verrucosa was thought to vary in color from brown to dark, and more rarely black by later authors such as MATTIROLO (1900a). In fact, MATTIROLO (1900a) regarded G. papillosa and G. kunzeana as morphological variations of G. verrucosa and incorrectly considered them as synonyms of his own G. verrucosa var. badia Mattir. In addition, he added a new European species, G. vagans Mattir., which is easily distinguished because of its large spores (35 × 27 μm on average) that are ornamented with large conical warts tending to coalesce at their bases. This taxon has also been recorded in Russia (BUCHOLTZ, 1901; ŁAWRYNOWICZ, 1990). In the 20th century, VELENOVSKÝ (1922) proposed Genea neuwirthii Velen., which has a smooth brownish peridium. It has a single inner chamber and spores that are 35–40 μm long and ornamented with prominent thick warts with blunt rounded edges. Scattered reports about additional specimens of the most commonly accepted European species can be found in the following years. VACEK (1951) presents interesting descriptions of G. pulchra (1947, 1949), and the original form of G. sphaerica (1951) from the Czech Republic. ECKBLAD (1954) suggested that G. pulchra and G. hispidula could represent a single species based on a wide range of spore sizes and ornamentation variability, which they considered developmental stages of this fungus. Later, major reviews of hypogeous fungi did not recognize any additional new Genea species in the European mycota. For example, the work on British hypogeous fungi by HAWKER (1954) recognized only four species (G. verrucosa, G. fragrans, G. sphaerica, G. hispidula) and the publication on Danish hypogeous macromycetes by LANGE (1956) recognized only three species (G. sphaerica was not included). CERUTI (1960) described and illustrated six European species (those mentioned above as well as G. lespiaultii and G. vagans). In Sweden, KERS described new reports of G. verrucosa (1979) and G. hispidula (1990), while PEGLER et al. (1993) mentions 41 that G. hispidula, G. verrucosa, G. sphaerica and G. fragrans as present in Britain. In Spain, published reports of Genea began with CALONGE (1985) and later VIDAL et al. (1991, 1997a). Recently, Genea sphaerica f. lobulata Mor.-Arr., J. Gómez & Calonge (MORENO-ARROYO et al., 1998a) was erected. This variety is similar to G. sphaerica because of its labyrinthlike inner chamber with broad white tramal plates, but its spore ornamentation does not seem to match the original description. Another species putatively related to G. verrucosa was described from southern Spain as Genea subbaetica Mor.-Arr., J. Gómez & Calonge (MORENO-ARROYO et al., 1998b). The spores are similar to those of G. verrucosa (26–30 × 22–26 μm) and ornamented with 2–3 μm high warts that are conical to cylindrical. However, G. subbaetica also has smaller cells in the outer peridial layer (15–30 μm in diam.) and bigger asci (240–280 × 25–30 μm in G. subbaetica vs. 200–220 × 26– 30 μm in G. verrucosa). Furthermore, this species is collected in early winter, while G. verrucosa is usually found in spring or early summer. In Italy, the major works by MONTECCHI & LAZZARI (1993), MONTECCHI & SARASINI (2000) and GORI (2005) contributed with very useful macro and microscopic images to illustrate the species that had already been described by the classical authors. It is also worth mentioning the work on Sicilian hypogeous fungi by VENTURELLA et al. (2004). The genus Genea has been reported also from Hungary (HOLLÓS, 1911; SZEMERE, 1965; BRATEK et al., 2013), Greece (DIAMANDIS & PERLEROU, 2008, KONSTANTINIDIS, 2009), and Turkey (TÜRKOĞLU & CASTELLANO, 2014). For the present work we have studied more than 160 new collections and herbarium specimens from across Europe, with particular attention to brownish European specimens and those with strikingly diferent spore ornamentation. In a few cases, a combination of morphological characters and ecological traits were also employed to deine species. We have also obtained molecular data from all species and these have been compared with described taxa from Europe and North America to support the assertion that the new taxa presented here do not match any of the previously established species. Material and methods Morphological study. —All collections are preserved in the herbarium of the Universidad de Alcalá (AH) and Hungarian Natural History Museum (BP) at Budapest, although the codes referring to the origin of each sample used during the study are also provided: AH (Alcalá de Henares Herbarium), AVM (Asociación Vallisoletana de Micología), BM (Baldomero Moreno-Arroyo), CA (Carlo Agnello), GH (Gunnar Hensel), GK (Giorgos Konstantinidis), JC (Julio Cabero), NV (Nicolas Van Vooren), VK (Vasileios Kaounas), and ZB (Zoltán Bratek, from mycotheca of the First Hungarian Trule Society, EMSzE). Microscopic study was performed in distilled water, lactophenol cotton blue, Trypan blue or Congo red-loxine. KOH 5% and Melzer’s reagent were not used for spore measurements because these mounting media have been shown to dissolve spore ornaments in some hypogeous Pezizales (SMITH et al., 2006). Spores measurements are provided on 50 items outside asci. X represents the average of spore dimensions. Qm represents the mean ratio of the spore length divided by the width. Spore dimensions are given without ornamentation. The width of ornamentation corresponds to the width at the base of warts. Phylogenetic analysis. — The molecular identity of the new taxa proposed in the present work was studied and compared with other described Genea species from Europe and North America. All of the new species described below represent independent monophyletic lineages that are distinct from any of the described taxa. A detailed phylogenetic analysis will be presented as part of an upcoming paper. 42 Taxonomic results Genea brunneocarpa G. Moreno, J. Cabero & V. Kaounas, sp. nov. — MB 809071 Etymology: brunneo- comes from the latin brunneo, meaning brown, and –carpa is derived from latin carpus, fruit, the name meaning Genea with brownish ascomata. Diagnosis: Ascomata hypogeous, reddish to brownish in color, with a single inner chamber with peridium wall projections, lined with a light brown epithecium. Ascospores measuring less than 32 μm, excluding spore ornamentation which consists of truncatedconic warts that are sometimes cubic in appearance. This species fruits under Quercus ilex with or without Pinus halepensis. Holotype here designated: SPAIN: Valladolid, Tiedra, Montes Torozos, 784 m asl, under Quercus ilex, leg. J. Cabero, 03-IV-2011, JC6, AH44112. Description Ascomata hypogeous, subglobose or slightly lobed, measuring up to 20 mm, reddish to brown in color, covered with small, irregular, lat warts with the apices somewhat darkened; apical oriice present, and also a basal tuft of hyphae that often adheres to soil particles. Odor weak and unremarkable. Peridium bilayered measuring 250– 340 μm, composed by an external pseudoparenchymatous layer 160–190 μm thick, formed by hyaline subglobose or isodiametric cells measuring 33.5–55.0 × 21.5–35.0 μm, whose walls look thicker, more angular and more pigmented closer to the surface of the peridium, and an inner layer 130–180 μm thick, also pseudoparenchymatous but turning prosenchymatous in the vicinity of the hymenium. Inner chamber usually not divided, but sinuose due to wall projections, lined with a light brown epithecium with scattered minute warts with darkened apices, overall appearance and pseudoparenchymatic structure similar to that of the peridium. Hymenium arranged as a palisade under the epithecium, composed of cylindrical pedicelate asci, mixed with long paraphyses measuring 3–8 (–10) μm thick, and only slightly exceeding asci in length, not inlated at the top. Asci 234–288 × 28–32 μm, containing eight uniseriate or slightly disorganized spores. Ascospores ellipsoid or occasionally globose, (26.5–) 28.0–31.5 (–32.5) × (18.5 ) 19.5–24.0 (–25.0) μm, X = 29.5 × 21.5 μm, Qm= 1.35, ornamented by truncated-conic warts that sometimes look almost cubic, only rarely with secondary warts protruding from the apex; truncated-conic warts measure up to 2.5 × 3.5 μm, while cubic warts up to 3.8–5.5 μm in height; microwarts present and numerous. Ecology: Fruiting in winter and spring (Feb.-Apr.) in calcareous soils, under Quercus ilex, with or without Pinus halepensis. Distribution: known thus far only from Mediterranean Europe (Spain and Greece). Notes Genea brunneocarpa can look microscopically similar to G. fragrans, but both the spores and spore ornaments are smaller in G. brunneocarpa. It also has a brownish peridium and epithecium and the inner chamber is illed with projections from the wall but these internal epithecium projects are not folded as they are in G. fragrans. The North American taxon G. compacta is also brownish, has a single irregularly cavernous inner chamber, and spores 32–34 × 24–28 μm with spore ornamentation of large irregular truncate or conical warts (HARKNESS, 1899; GILKEY, 1939). The European specimens have not been accommodated in G. compacta because of the dubious status of this exceedingly rare species. Genea compacta has only been found a handful of times and this species is morphologically similar to G. arenaria. Genea compacta also overlaps with the distribution of the highly morphologically variable G. arenaria, suggesting that it may actually be a synonym (SMITH et al., 2006). In addition, the original description of G. compacta suggests that it is PLATE 1 A. Genea compressa JC28 (J. Cabero); B. G. lobulata VK122 (V. Kaounas); C. G. sphaerica GH20100724 (G. Hensel); D. G. lespiaultii RM2140 (R. Martínez); E. G. fageticola GK4129 (G. Konstantinidis); F. G. fragrans AH39100 (M.A. Sanz); G. G. pseudoverrucosa AH39104 (M.A. Sanz); H. Genea cf. anthracina JC3 (J. Cabero); I. G. dentata NV-R (P. Ribollet); J. G. verrucosa PSS3703 (F. Sáinz); K. G. pseudobalsleyi JC29 (J. Cabero); L. G. vagans JC17 (J. Cabero); M. G. oxygala JC14 (J. Cabero); N. G. tuberculata JC32 (J. Cabero); O. G. brunneocarpa JC6 (J. Cabero); P. G. pinicola JC12 (J. Cabero); Q. G. hispidula JC23 (J. Cabero); R. G. arenaria JC7 (J. Cabero); S. G. thaxteri JC16 (J. Cabero); T. Genea cf. subbaetica VK3291 (V. Kaounas). 43 slightly smaller (1 cm), stellate in shape, light brown in color, has a coarsely verrucose peridium, and spore ornamentation that is more crowded than in G. brunneocarpa. Genea papillosa, G. kunzeana and G. verrucosa f. badia have brownish ascomata, but spores are ornamented with small pointed warts, similar to those of G. verrucosa. Genea neuwirthii is light brownish, has larger spores (35–40 μm on average), and was collected in central Europe (Czech Republic). Studied collections: GREECE: Attica, Katsimidi, preference for Quercus ilex in a mixed stand with Pinus halepensis on calcareous soil, leg. V. Kaounas, 24-II-2013, VK2922, AH44113. SPAIN: Guadalajara, Castilblanco de Henares, under Q. ilex, calcareous soil, leg. M.A. Sánz, 13-III-2011, AH42934. Valladolid, Tiedra, Montes Torozos, 784 m asl, under Q. ilex, leg. J. Cabero, 03-IV-2011 JC6, AH44112 HOLOTYPE. Valladolid, Urueña, Montes Torozos, under Q. ilex, calcareous soil, leg. J. Cabero, 20-IV-2014, JC35, AH44111. Genea compressa Z. Merényi, J. Cabero & G. Moreno, sp. nov. — MB 809072 Etymology: compressa refers to the compressed peridium wall projections that extend into the inner chamber. Diagnosis: Ascomata hypogeous, subglobose, lobate, with a black peridium covered with small irregular warts. Inner chamber illed with numerous compressed wall projections. Ascospores measuring on average 25 μm, ornamented with pointed conical warts approximately 3 μm in height. Found under Quercus spp. in alpine forests of Mediterranean Europe and hills of central Europe in autumn. Holotype here designated: SPAIN: Zamora, Parque Natural de Sanabria, Llanes, hypogeous under Quercus pyrenaica, 994 m asl, leg. J. Cabero, 04-III-2013, JC28, AH44118. Description Ascomata hypogeous, subglobose, lobate, measuring 8–14 mm. in diam.; black peridium covered with small irregular to polygonal low black warts. An apical oriice covered with warts similar to those on the peridium is present, and the fungus is attached to the substrate by a basal tuft of hyphae. Odor weak and unremarkable. Peridium total width of 350–490 μm, bilayered, with an external layer measuring 230–250 μm, composed of cells arranged as a hyaline pseudoparenchyma, appearing subglobose or slightly angular, and pigmented close to the surface with individual cells 38–47.5 × 29– 36 μm. The internal layer is 120–240 μm thick, and formed of loose hyphae perpendicularly arranged that can look as difuse small elements when sectioned. Inner chamber containing numerous compressed coarse wall projections, coated with an epithecium that is and more or less similar to the external peridium surface but sometimes more brownish. The epithecium measures 90–130 μm and is pseudoparenchymatic in structure, most cells angular but a few subglobose. Hymenium formed by asci and cylindrical, septate, somewhat tortuous paraphyses arranged in a palisade measuring 310–380 × 3–5 μm, forming an epithecium above the hymenium. Asci cylindrical, sometimes inlated, indehiscent, 200–280 × 25–38 (–42) μm, with a short, eccentric sinuose or even coiled peduncle. Ascospores uniseriate, subglobose to ellipsoid, (23.5–) 24.0–26.5 (– 30.5) × (18.0–) 19.0–21.0 (–25.5) μm, X= 25.0 × 21.5 μm, Qm = 1.26, ornamented by conical warts, often digitate at the top, measuring (2.3–) 3.3 (–5.3) × (2.0–) 3.3 (–5.8) μm. Ecology: Autumn (Sept.–Nov.), under a variety of host plants, including Quercus pyrenaica, Q. cerris, and Quercus sp., in the alpine belt of the Mediterranean basin and in the hill country of central Europe. Distribution: known from Spain, Hungary, and Morocco; potentially widely distributed across Europe. 44 Notes This species is similar to G. vagans in several features: it has black ascomata covered with irregular to polygonal warts, a chambered inner cavity, and spores ornamented by conical warts. However, Genea compressa has smaller spore ornaments — typically 2–5 μm high — than those of G. vagans — typically 5–8 μm high (MATTIROLO, 1900a, 1900b; CERUTI, 1960; VIDAL, 1997). Genea compressa sometimes resembles other blackish species, such as G. fragrans or G. verrucosa, but these difer from G. compressa due to their spore sizes or the morphology of their spore ornaments. Finally, G. echinospora Gilkey produces spores with similar ornamentation, but is described as mummy-brown in color and hispid, has conspicuously larger spores (36–40 μm), and was collected at lower elevation sites in North America (GILKEY, 1939, 1954). Studied collections: HUNGARY: Borsod-Abaúj-Zemplén county, Bogács, Bükk Mountains, 200-250 m asl, under Quercus sp., Quercus cerris and Carpinus betulus, 4-X-1999, ZB1790, BP104836. KomáromEsztergom county, Szárliget, 250-300 m asl, under Quercus sp. and Carpinus betulus, 16-IX-2006, ZB3332, BP104837. MOROCCO: Ifrane, road from Azrou to Ain Leuh, under Cedrus atlantica with scattered Quercus sp., leg. M.A. Sánz, P. Alvarado, J.L. Manjón & J. Álvarez, 18XI-2010, AH42944. SPAIN: Zamora, Parque Natural de Sanabria, Llanes, under Quercus pyrenaica, 994 m asl, leg. J. Cabero, 04-III-2013, JC28, AH44118 HOLOTYPE. Genea dentata Van Vooren, J. Cabero & Hensel, sp. nov. — MB 809074 Etymology: dentata from latin dens (tooth), refers to the teethlike appearance of the spore ornamentation (either fang-like or molar-like). Diagnosis: Hypogeous ascomata of reduced dimensions (2– 5 mm), with a black peridium topped by prominent warts. Ascospores measure less than 32 μm without ornamentation, which is composed of long (5–6 μm) and thin fang-like or molar-like warts that resemble teeth. It can be found under broadleaved trees in temperate Europe. Holotype here selected: FRANCE: Loire-Atlantique, Orvault, border of stream “Cravatte”, under Corylus avellana, Prunus laurocerasus, Ilex aquifolium and Ruscus sp., leg. P. Ribollet, 31-V-2009, NV-R, AH44121 HOLOTYPE. Description Ascomata hypogeous, globose to subglobose, minute, measuring 2–5 mm in diam. Black peridium covered with prominent pyramidal warts. Basal tuft composed of brownish-reddish hyphae. Apical oriice present. Single inner chamber lined with brownish lat warts. Peridium pseudoparenchymatic measuring 350–500 μm, bilayered; external layer 200–400 μm thick, composed of angular dark brown cells, more or less stretched, e.g. 42 × 23 μm, or triangular, e.g. 35 × 30 μm, with thick wall (4–7 μm); internal layer about 100–150 μm thick, comprised of brownish cells, angular or almost spherical, measuring 7–45 × 7–25 μm, with thin wall. Asci cylindrical, 260–300 × 33–40 μm, containing eight uniseriate spores. Paraphyses hyaline and septate, forming an epithecium above the palisade of asci. Ascospores hyaline, generally uniguttulate, broadly ellipsoid to ellipsoid or even subcylindrical, (26.5–) 27.0–32.0 (–33.0) × (19.0–) 20.0–22.0 (–23.0) μm, X = 29.5 × 21.0 μm, Q=1.3–1.6, ornamented with long warts that are curved and spiny (similar to the fangs of a cat) or sometimes truncated and digitate at the top (like typical human molars), measuring 5.0–6.0 × 1.5–2.5 μm. Ecology: Under deciduous trees, in siliceous soil of temperate Europe. Spring (May-Jun.). Distribution: known from France, Germany and putatively from Spain. Notes This species is proposed to accommodate two small black specimens with long, spiny spore ornaments that vaguely resemble teeth under the light microscope. A third collection (JC19, AH44120 from Spain) seems somewhat deviant, both genetically and morphologically, but is nevertheless considered conspeciic for now. It has slightly larger and more ellipsoid spores that are ornamented with long, scattered warts with more rounded tips. GH20060610 (AH44119) was immature, although the sparsely ornamented spores were morphologically similar to those of the holotype specimen (PR0970 NVR, AH44121). The morphological features reported here should be conirmed with new collections matching the genetic proile of the holotype collection. Studied collections: FRANCE: Loire-Atlantique, Orvault, border of stream “Cravatte”, under Corylus avellana, Prunus laurocerasus, Ilex aquifolium and Ruscus sp., leg. P. Ribollet, 31-V-2009, NVR, AH44121 HOLOTYPE, Ribollet (personal herbarium) PR0970 ISOTYPE. GERMANY: Nickelsdorf, Zeitzer Forst, MTB5038/1, leg. G. Hensel, under Quercus sp., in siliceous soil, 10-VI-2006, GH20060610, AH44119. SPAIN: Asturias, Pola de Somiedo, under Fagus sylvatica, calcareous soil, leg. F. García, 12-VI-2013, JC19, AH44120. Genea fageticola Konstant., J. Cabero & F. García, sp. nov. — MB 809075 Etymology: fageticola refers to the apparently strict association with Fagus sylvatica. Diagnosis: Hypogeous blackish ascomata covered with irregular warts, with an abundant basal tuft of hyphae, a single inner chamber lacking wall projections lined with a brownish epithecium. Ascospores larger than 32 μm without ornamentation, which consists of prominent truncated or rounded warts approximately 4 μm in height. It is apparently a strict symbiotic associate of Fagus sylvatica. Holotype here designated: GREECE: Drosopigi Florina, under Fagus sylvatica, along with Hydnotrya tulasnei (Berk.) Berk. & Broome, 1150 m asl, leg. D. Klisiari (dog Sisi), 27-IX-2009, GK4129, AH44122. Description Ascomata hypogeous, subglobose, slightly depressed, measuring 7–15 mm in diam.; black, covered with small, irregular, lat or roundish warts; with an apical oriice and a proliic basal tuft of reddish or brownish hyphae. Peridium 200–400 μm thick, bilayered; external layer measuring 150–320 μm, pseudoparenchymatous, composed of hyaline inlated subglobose or angular cells measuring 25.0–60.0 × 20.0–45.0 μm, becoming more pigmented and thickwalled toward the peridium surface. The inner layer is 50–70 μm thick, composed of yellowish inlated subglobose or angular cells measuring 14–40 × 9–25 μm. Single inner chamber with some small lobes (but lacking wall projections) lined with a light brown epithecium formed by minute papillae, which is also pseudoparenchymatous in structure and composed mostly of angular cells. Hymenium arranged as a palisade, formed by asci measuring 305–370 × 25–26 μm, 8-spored, cylindrical, with a slightly lateral and short, somewhat sinuous peduncle measuring 23.4 × 6.5 μm on average. Interestingly, some asci are much bigger than the space illed by the spores (these inlated asci measure 305–370 × 25–26 μm), even when the ascospores are completely mature and have reached their full size. Paraphyses septate, cylindrical, measuring 19.5–24.5 × 3.5– 4.9 μm in diam., forming an epithecium over the palisade of asci. Ascospores ellipsoid, (28.0–) 29.0–34.5 (–36.5) × (18.0–) 20.5–24.5 (–26.0) μm, X = 31.5 × 22.5 μm, Qm=1.41, thick walled, sometimes with a blackish-lilac oil droplet, ornamented by crowded cylindrical, truncated or rounded warts, measuring 2.1–6.5 × 2.0–6.0 μm. Ecology: under Fagus sylvatica, sometimes with scattered Quercus petraea trees. Late summer and winter (Sept.-Feb.). Distribution: known from Greece and Spain. Notes Genea fageticola macroscopically resembles other blackish Genea species. The more or less globose ascomata distinguish it from G. verrucosa, which has more irregular ascomata, smaller spores, different spore ornamentation, and is more commonly found with Quercus ilex. Genea fageticola can be confused with G. fragrans, which can also be found under Fagus and is supposed to have similarly large spores ornamented with prominent truncated warts (BERKELEY & BROOME, 1846; TULASNE & TULASNE, 1851; CORDA, 1854; HAWKER, 1954). However, G. fragrans usually has folded ascomata and a labyrinthine inner chamber when mature, a small inconspicuous basal tuft of hyphae, a black epithecium covered with warts similar to those of the external peridium, and a more prominent and scattered spore ornamentation, sometimes with tiny secondary warts among the primary warts. Spore ornamentation of G. fageticola can also resemble that of G. hispidula, another taxon that occasionally fruits under Fagus sylvatica. However, these two species can be easily discriminated because G. hispidula has a brownish peridium with copious peridial hairs. Genea vagans can also be present in the same habitats and has spores that are similar in size to G. fageticola (35 × 27 μm on average), but G. vagans can be separated due to its conical spore ornamentation and chambered inner cavity. Genea fageticola difers from G. compressa by its single inner chamber without obvious peridium wall projections, its more conspicuous basal tuft of brownish hyphae, its bigger spores ornamented with truncated or rounded warts, and a host preference for Fagus sylvatica. In contrast, G. compressa has an inner chamber illed with packed wall projections, an inconspicuous or absent basal tuft of hyphae, smaller ascospores ornamented with conical pointed warts, and is commonly found under Quercus spp without F. sylvatica. Studied collections: GREECE: Drosopigi Florina, under Fagus sylvatica, along with Hydnotrya tulasnei (Berk.) Berk. & Broome, 1150 m asl, leg. D. Klisiari (dog Sisi), 27-IX-2009, GK4129, AH44122 HOLOTYPE. SPAIN: León, Soto de Sajambre, bosque de Vegabaño, under F. sylvatica and several Quercus petraea, humid climate, 1432 m asl, calcareous soil, leg. F. García, 16-II-2008 H-139, AH44123. Genea lobulata (Mor.-Arr., J. Gómez & Calonge) P. Alvarado & Mor.-Arr., comb. nov. — MB MB809076 Basionym: Genea sphaerica f. lobulata Mor.-Arr., J. Gómez & Calonge, in Moreno-Arroyo et al., Boll. Gruppo Micol. ‘G. Bresadola’ (Trento), 41(3): 207 (1998). Description Ascomata hypogeous, subglobose to lobed, measuring 15–40 mm in diam. An apical oriice and a basal tuft of hyphae reddish brown in color are present. Peridium black, covered with polygonal lat warts, lacking peridial hairs, bilayered, with an external pseudoparenchymatic layer measuring 250–300 μm wide, composed of angular cells measuring 30–70 × 22–50 μm just below the surface, and an inner pseudoparechymatic layer 150–200 μm wide formed of globose cells that turn prosenchymatic near the hymenium. Inner chamber highly folded due to projections of the hymenium, brainlike appearance when cut. Hymenium covered by a dark brown, warted pseudoparenchymatous epithecium that is commonly black, but sometimes brownish when young. Asci cylindrical, 200– 225 × 26–28 μm, containing eight uniseriate spores. Paraphyses 210–325 × 6–10 μm. Ascospores subglobose to ellipsoidal, 24–30 × 20–26 μm, ornamented with spiny warts, more rarely conical with roundish top, measuring 1–2.5 μm high × 1–2 μm wide, with minute warts among them. 45 Ecology: This species is found fruiting under Quercus ilex in winter, spring and summer (Jan.-Aug.). Distribution: known from Greece, Spain and Cyprus. Holotype here designated: SPAIN: Zamora, Toro, Bosque de Montelarreina, under Pinus pinea and P. pinaster, 16-II-2008, JC14, AH44177. Notes This species is similar to Genea sphaerica due to its black and minutely warted peridium, and labyrinthine inner chamber with thick sterile white tramal plates (TULASNE & TULASNE, 1851; FISCHER, 1897; VACEK, 1951). However, several authors observed diferences with this taxon, which lead to the proposition of an independent variety (MORENO-ARROYO et al., 1998a), which is here elevated to the species level as a result of molecular studies (data not shown). Three main features can help discriminate G. lobulata from G. sphaerica. First, G. sphaerica is smaller (10–20 mm), almost perfectly round, more or less radially symmetrical, and the projections of the inner cavity are not delimited into closed chambers. In contrast, G. lobulata is bigger, often irregularly lobed, and rarely has any obvious symmetry when sectioned. Furthermore, the peridium wall projections produce closed chambers when the ascomata are sectioned. Second, ascospores in G. sphaerica are ornamented with conspicuous warts that are very round, while those of G. lobulata are ornamented with spiny-tuberculate warts (MATTIROLO, 1903; MORENO-ARROYO et al., 1998a). Finally, G. sphaerica seems to occur in alpine Mediterranean habitats as well as central and northern Europe lowlands during summer and autumn under deciduous trees such as Carpinus, Corylus, Fagus or TILIA (HESSE, 1894; FISCHER, 1897; VACEK, 1951; ŁAWRYNOWICZ, 1990). In contrast G. lobulata is commonly found in winter, spring and summer in lowland Mediterranean Europe under Quercus ilex or Q. faginea (MATTIROLO, 1903; VIDAL, 1997; MORENO-ARROYO et al., 1998a; MONTECCHI & SARASINI, 2000). The only other Genea species putatively resembling G. lobulata and G. sphaerica is G. lespiaultii, which can be easily separated because of its distinctive spore ornamentation of lat warts. Description Ascomata hypogeous, subglobose, highly lobed or tuberculate, 10–25 (–32) mm in diam. Surface brown to dark brown with blackish areas when mature, covered with minute irregular warts or papillae, with a basal tuft of dark brownish mycelium, and with an apical opening, sometimes diicult to detect because of the highly folded fruiting body. Odor pleasant, similar to yogurt. Peridium comprised of one layer, measuring around 320–350 μm thick. Microscopically, it has a pseudoparenchymatic structure composed of hyaline inlated subglobose or isodiametric cells that are angular and pigmented with the cells walls becoming thicker near the surface of the peridium. Peridium cells variable in size, 19–45 × 12–27 μm. Inner cavity divided into a variable number of irregular chambers, lined with a light brown-colored verrucose epithecium. Hymenium arranged in a regular palisade, composed of asci and septate paraphyses, which are somewhat more inlated and longer than asci (29–35 μm between each septa, exceeding slightly the length of asci × 3.5–5.0 μm thick), forming an epithecium about 170–200 μm thick, with a pseudoparenchymatic structure composed of angular cells up to 46 × 38 μm. Asci irregularly cylindric, 220–290 × 29–35 μm, pedicelate (peduncule measuring approximately 57 × 9.5 μm), and indehiscent, containing eight uniseriate ascospores. Ascospores ellipsoid to ovoid, with rounded ends, (24–) 24.5–28 (–29) × (18.2–) 19.0–21.8 (–22.4) μm, X = 26 × 20.5 μm, Qm= 1.28, ornamented by widely separated, low and hemisphaerical warts, measuring 1.5–4 (–5) μm high and 2.5–7.5 μm wide. Ecology: associated with pine trees such as Pinus pinea or P. pinaster, in siliceous sandy soil. Fruiting in winter or spring in Mediterranean climates. Distribution: known only from Spain. Studied collections: CYPRUS: unknown location, under deciduous trees, leg. Th. Alexandridis, 01-VII-2010, GK5078, AH44141. Unknown location, under deciduous trees, leg. S. Mavros, 15-VI2011, GK5770, AH44143. GREECE: Attica, Parnitha, under Quercus ilex and Q. ithaburensis subsp. macrolepis, argillaceous soil, 500 m asl, leg. V. Kaounas, 30-III-2011, VK2122, AH44149. Same locality, 30V-2011, VK2213, AH44150. Same locality, 17-VI-2011, VK2238, AH44151. Same locality, VK2239, AH44152. Korestia Kastoria, under deciduous trees, leg. G. Setkos, 07-XII-2010, GK5384, AH44142. SPAIN: Guadalajara, Valdearenas, under Q. ilex in basic soil, leg. M.A. Sanz, 30-V-2009, AH42940. Navarra, Salinas de Oro, under Q. ilex, leg. P.M. Pasaban & F. Sáinz, 01-IV-2011, PSS3501, AH44147. Same locality, 07-IV-2011, PSS3502, AH44148. Valladolid, Santa Espina, Montes Torozos, under Q. ilex, calcareous soil, leg. J. Cabero, 30-V-2013, JC2, AH44144. Same locality, 26-III-2013, JC4, AH44145. Valladolid, Urueña, Montes Torozos, under Q. ilex, calcareous argillaceous soil, 06-VI-2012, JC9, AH44146. Córdoba, Doña Mencía, under Q. ilex, leg. B. Moreno-Arroyo, 26-V-2013, BM1043, AH44140. Genea oxygala J. Cabero & F. García, sp. nov. — MB 809077 Etymology: oxygala is derived from the ancient greek words oxy(acid) and –gala (milk), which Galen describes as a fermented milk product. It is still a word applied to yogurt among some greek-speakers formerly inhabiting Turkey (G. Konstantinidis, personal observation). Diagnosis: Large brownish ascomata, highly lobed, surface covered with minute irregular warts, intense pleasing odor that is similar to yogurt, ascospores not exceeding 32 μm, ornamented with scattered wide hemisphaerical warts measuring 1.5–4 × 2.5–7.8 μm. Strictly associated with Pinus spp. in Mediterranean habitats. 46 Notes This species was previously reported as G. thaxteri Gilkey by VIDAL et al. (1997) and MONTECCHI & SARASINI (2000). Genea thaxteri was irst collected by Roland Thaxter from Maine and Tennessee (USA) and published by GILKEY (1939). VIDAL et al. (1997) and MONTECCHI & SARASINI (2000) reported Genea thaxteri from Europe, despite the fact that the European material did not exactly match the original description of this North American species. Genea thaxteri is smaller than the European taxon; while G. oxygala ranges from 10 to 25 mm in diameter, GILKEY (1939) reported G. thaxteri as only 7 mm in diameter. The color of G. thaxteri was described as “Sudan brown” (GILKEY, 1954), while that of G. oxygala is darker, reported by VIDAL et al. (1997) as “burnt umber”. The ascomata of G. thaxteri are only slightly or not at all lobed whereas G. oxygala is highly lobed to tuberculate. Finally, ascospores of G. thaxteri are reportedly ornamented with crowded, rounded to pointed papillae, while those of G. oxygala have widely spaced low hemisphaerical warts that are not pointed. The very rare European species G. neuwirthii has smooth, lobed light brownish ascomata with a single inner chamber, and its large ascospores (35–40 μm) are ornamented with prominent thick warts that have rounded edges. These spore ornaments difer dramatically from the low wide hemispherical warts of G. oxygala. Studied collections: SPAIN: Segovia, Cuellar, sandy soil under Pinus pinea, leg. A. García Blanco & J.M. Sanz Carazo, 14-II-2004, AVM1745, AH44175. Same locality, Pinar de Pociage, under Pinus pinaster, leg. F. García, 19-I-2013 (H-201). Zamora, Toro, under P. pinea, leg. A. García Blanco, M. Sanz Carazo, J. Cabero, 18-I-2004, AVM1842, AH44176. Zamora, Toro, Bosque de Montelarreina, under P. pinea and P. pinaster, 16-II-2008, JC14, AH44177 HOLOTYPE. Genea pinicola V. Kaounas, J. Cabero & F. García, sp. nov. — MB 809078 Genea pseudobalsleyi Agnello, Bratek & J. Cabero, sp. nov. — MB 809079 Etymology: pinicola refers to the association of this species with Pinus spp. Etymology: pseudobalsleyi refers to the close morphological and phylogenetic relationship with the North American species G. balsleyi. Diagnosis: Hypogeous brownish ascomata with a single inner chamber lacking peridium wall projections, ascospores measure less than 32 μm and are ornamented with cylindrical warts 1–3 μm. It is apparently associated with Pinus sp., with or without other tree species. Holotype here designated: SPAIN: Valladolid, Rábano, hypogeous under Pinus pinea, 795 m asl, leg. J. Cabero, 04-III-2013, JC12, AH44153. Diagnosis: Subglobose to irregularly lobed hypogeous ascomata with black warted peridium, morphologically similar to the epithecium which lines the peridium wall projections within the inner chamber. Ascospores measure less than 32 μm and are ornamented with truncated or conic warts about 2 μm high. Holotype here designated: ITALY: Brindisi, Bosco Compare, under Quercus ilex on calcareous soil, 30 m asl, leg. C. Agnello, 29-V-2010, CA01, AH44156. Description Ascomata hypogeous, subglobose to depressed with few lobules, measuring 7–15 mm in diam., yellowish brown, brown or reddish brown in color. Peridium covered with minute irregular low warts giving a rough appearance. A large apical opening is present, as well as a basal tuft of adherent hyphae. Odor unremarkable. Peridium two-layered, 210–260 μm thick, formed by an external pseudoparenchymatic layer, 120–160 μm thick, composed of hyaline subglobose or angular cells, measuring 20–40 × 20–34 μm, the internal layer a prosenchyma, 80–90 μm thick, with scattered inlated cells. Single inner chamber lacking conspicuous wall projections, warted, and similar in color to the peridium. Hymenium formed by asci arranged in palisade, with interspersed septate, iliform paraphyses measuring 260–400 × 1–4 μm, and forming a pseudoparenchymatous epithecium above asci. Asci cylindrical, 220–340 × 23–27 μm, containing eight uniseriate spores. Ascospores ellipsoid, 26.0–29.5 × (16.0–) 17.0–20.5 μm, X = 27.5 × 18.5 μm, Qm=1.49, ornamented by crowded truncated-conic or cylindric warts measuring 1–3 μm high and 1–3 μm wide, with or without digitations at their apices. Ecology: in calcareous soil, under Pinus halepensis, P. pinea and probably P. sylvestris, with or without Quercus ilex. In winter and probably spring (March). Distribution: known from Spain and Greece. Notes This taxon appears to preferentially or exclusively associate with species of Pinus. DNA from this species has been detected on the ectomycorrhizal roots of P. sylvestris in Iran (BAHRAM et al., 2013) and was likely introduced from the Mediterranean region. Due to its yellowish or brownish color, G. pinicola can be confused with young G. arenaria or G. thaxteri, but G. arenaria usually has peridial hairs while the latter has more lobed ascomata when mature. In addition, these two species are not typically collected under Pinus spp. Genea brunneocarpa is brownish and single-chambered, but is not commonly found with Pinus spp., rarely has ascomata that are depressed, has more pointed peridial warts, and more prominent and scattered spore ornamentation. The rare G. neuwirthii is described with smooth, lobed, light brown ascomata with a single inner chamber, but this species has bigger ascospores (35–40 μm) that are ornamented with prominent thick warts that have rounded edges. Studied collections: GREECE: Katsimidi Attica, under Quercus ilex and Pinus halepensis on calcareous soil, about 650 m asl, leg. V. Kaounas, 31-III-2009, VK825, AH44155. Same locality, 31-I-2013, VK2865, AH44154. SPAIN: Valladolid, Rábano, hypogeous under Pinus pinea, 795 m asl, leg. J. Cabero, 04-III-2013, JC12, AH44153 HOLOTYPE. Description Ascomata hypogeous, compact, subglobose and moderately lobed, 7–13 mm in diam., homogenously covered with small polygonal black warts that extend through the apical oriice. A small basal tuft of hyphae is present. Odor pleasant, similar to that of Tuber aestivum Vittad. Peridium a single layer of pseudoparenchymatic tissue 160 μm thick, composed of hyaline subglobose cells measuring 20–40 μm, becoming angular and with thicker and darker walls in the outermost layer. Inner cavity sometimes with a single inner chamber but other times with abundant wall projections that create independent sinuous chambers that give the gleba a brain-like appearance. The inner chamber lined with a black or brownish black epithecium with small irregular warts similar to those in the outer peridium surface. Epithecium is 40–70 μm thick, and its structure is also pseudoparenchymatous, composed of angular cells with thick, melanized walls that measure 20–35 μm in diam. Asci arranged in a continuous palisade, irregularly cylindrical, 190–270 × 27–38 μm, containing eight uniseriate spores, and with a short peduncle. Paraphyses cylindrical, occasionally septate, 44– 66 × 3–6 μm, some of them with swollen cells, up to 15 μm in diam. Ascospores (21.7–) 23.4–27.7 (–30.2) × (17.7–) 18.6–22.6 (–26.3) μm, X= 25.1 × 20.9 μm, Q=1.06–1.29 (average Qm=1.21), subglobose to broadly ellipsoidal or ovoid, hyaline or pale yellowish in color, with a conspicuous, irregularly located oil droplet (especially visible when mounted in water), ornamented by truncated warts, rarely pointed, occasionally with small digitations at the top, measuring (1–) 1.5– 2.5 (–4.0) μm high, and 1.5–2 (–4) μm wide. Ecology: under Quercus ilex and Q. pyrenaica, in summer to early winter (Jun.-Dec.). Distribution: known from Hungary, Italy and Spain. Notes This species is morphologically similar to G. balsleyi M.E. Smith (SMITH, 2007) because of its black peridium surface, spore size, and spore ornamentation. However, G. pseudobalsleyi difers in the abundant peridium wall projections that sometimes produce a brain-like, labyrinthine, and chambered inner cavity that is apparently absent in G. balsleyi. So far, G. balsleyi has been recorded only from North America, while G. pseudobalsleyi is present only in Europe. Peridium structure is remarkedly bilayered in G. balsleyi whereas this feature was not observed in G. pseudobalsleyi. Genea pseudobalsleyi difers from G. verrucosa because of its crowded, truncated spore ornamentation, which is more scattered and formed of thinner and more pointed warts in G. verrucosa. Genea compressa can look macroscopically similar, but difers in its conical spore ornamentation. Genea fragrans and G. fageticola have markedly diferent spore sizes and ornamentation. The rare American species G. macrosiphon Gilkey is macroscopically similar and has ascospores similar in size and ornamentation based on the description of GILKEY (1939), but its pseudoparenchymatic peridium cells are conspicuously inlated to 3–4 times the size of spores. 47 Studied collections: HUNGARY: Baranya county, Bóly, 20-XI-2007, ZB3804, BP104851. Baranya county, Ormánság, Marócsa, 29-X-1998, ZB1458, BP104850. Pest county, Gyömrő, sandy soil under Quercus sp., leg. Z. Merényi, É. Bordás, G. Fenyőssy, 18-VII-2010, ZB4192, BP104858. ITALY: Brindisi, Bosco Compare, under Quercus ilex on calcareous soil, 30 m asl, leg. C. Agnello, 29-V-2010, CA01, AH44156 HOLOTYPE. SPAIN: Zamora, Galende, 1017 masl., under Q. pyrenaica, leg. J Cabero, 01-XII-2013, JC29, AH44157. Genea pseudoverrucosa Bratek, Konstant. & Van Vooren, sp. nov. — MB 809080 Etymology: pseudoverrucosa refers to the close morphological ailiation with Genea verrucosa and the fact that this taxon has often been misidentiied as Genea verrucosa, because of the similarity in spore sizes Diagnosis: Black folded ascomata covered with minute inconspicuous warts, with a labyrinthine inner chamber, and ascospores not exceeding 32 μm, ornamented truncated warts approximately 3–4 μm high. Fruiting in summer, autumn and winter, but not spring, under deciduous trees of Mediterranean and central Europe. Holotype here designated: FRANCE: Meurthe-et-Moselle, Sexeyaux-Forges, 250 m asl, under deciduous trees such as Carpinus, Quercus, Corylus, Cornus, calcareous-clayey soil, leg. J.-B. Perez (dog Fii), 05-I-2011, NV-D, AH44160. Description Ascomata hypogeous, subglobose to highly folded, 6–15 mm in diam., with an inconspicuous basal tuft of dark hyphae. Peridium surface black and covered with minute rounded warts, composed by a single layer, pseudoparenchymatic 180–200 μm thick, formed of irregular polygonal cells measuring (22.5–) 27.0 (–34.5) × (33.0–) 37.5 (–45.0) μm. Inner cavity highly folded or labyrinthine but scarcely divided or not divided at all, lined with a black epithecium similar to the peridium. Epithecium structure pseudoparenchymatous, 80–150 μm thick, mostly formed by isodiametric cells. Glebal trama whitish, grayish or pale yellowish. Asci 197–222 × 26–27 μm, containing eight uniseriate spores. Ascospores subglobose to broadly ellipsoid, (23–) 27–30 × 21.5–24 (–24.5) μm, hyaline, ornamented by conical-truncated to irregular warts measuring (2–) 3–4 (–5) μm high and 3–4 (–4.5) μm wide, sometimes with a few interspersed smaller warts. Ecology: in basic soil, associated with temperate deciduous trees (Quercus, Carpinus, Tilia, Corylus), but also in Mediterranean habitats with Quercus ilex or Q. coccifera, from summer to early winter (Jul.Jan.), but apparently not in spring. Distribution: known from France, Greece, Hungary and Romania. Also known in Morocco. Notes This taxon macroscopically resembles Genea fragrans but has been typically misidentiied as G. verrucosa on the basis of ascospore dimensions. Spore ornamentation is formed by truncate warts that are similar to those of G. fragrans, but those of G. pseudoverrucosa are lower, crowded, and are more or less the same size, while those of G. fragrans are usually larger (3–7 μm high) and more scattered, with smaller warts that are frequent interspersed among the larger ones (BERKELEY & BROOME, 1846; TULASNE & TULASNE, 1851; CORDA, 1854; HAWKER, 1954). Spore size and ornamentation of G. pseudoverrucosa can look similar to those of G. brunneocarpa, but some warts are more scattered in G. brunneocarpa, and macroscopic features and ecology help to discriminate these two species. Genea pseudobalsleyi also has a black, warted peridium and has ascospores that are similar to G. pseudoverrucosa, but the spore ornaments in G. pseudobalsleyi are notably thinner (rarely exceeding 1.5–2 μm wide). Finally, the North American species G. balsleyi is similar but can be 48 discriminated because of its bilayered peridium, a more or less regular hollow inner cavity, and mostly conical spore warts (as well as a very diferent molecular proile). Studied collections: FRANCE: Sexey-aux-Forges, Meurthe-et-Moselle, 250 m asl, under deciduous trees such as Carpinus, Quercus, Corylus, Cornus, calcareous-clayey soil, leg. Jean-Baptiste Pérez (dog Fii), 05-I-2011, NV-D, AH44160 HOLOTYPE. GREECE: Anthohori Ioannina, 1040 m asl, under Quercus coccifera and other broadleaf shrubs, leg. George Setkos, Katerina Nola and George Konstantinidis (dog Spina), 31-VII-2010, GK5088, AH44158. Kastoria, leg. George Setkos (dog Cannelle), 28-X-2010, GK5286, AH44159. HUNGARY: Győr-Moson-Sopron county, Fertőrákos, under Quercus cerris, 29-IX2001, ZB2386, BP104854. Heves county, Szilvásvárad, Bükk Mountains, in a Quercetum petreae-cerris phytocoenosis, 11-XI-1998, ZB1473, BP104852. Pest county, Budapest, Budai range, XII-1999, ZB1948, BP104853. MOROCCO: Chefchaouen, Jebel Tissouka, under Quercus sp., leg. M.A. Sánz, J.L. Manjón, P. Alvarado, J. Álvarez, 14-XI2010, AH39104. ROMANIA: Harghita, Cristuru Secuiesc, under Carpinus sp. and Fagus sp., 21-XI-2006, ZB3445, BP104855. Genea tuberculata J. Cabero & P. Juste, sp. nov. — MB 809081 Etymology: tuberculata is derived from the latin root tuberculum, meaning protuberance, regarding the extremely humped and lobed ascomata of this species. Diagnosis: Hypogeous ascomycete, with a large brownish humped and lobed ascoma. The peridium is covered with minute irregular warts. It has a chambered inner cavity, and ascospores measuring less than 30 μm that are ornamented with small irregularly conical warts. It grows in gypsiferous marl soils under Pinus trees in winter. Holotype here designated: SPAIN: Valladolid, Aldealbar, 867 m asl, hypogeous under Pinus pinea and Quercus ilex, leg. J. Cabero, 09-III2014, JC32, AH44178. Description Ascomata hypogeous, subglobose, highly lobed (some specimens with more than 60 lobes), measuring 20–30 mm, brownish to dark brownish in color or occasionally reddish brown, becoming darker with age. Peridium usually covered with minute irregular warts, but sometimes with lattened warts, a feature that could be related to maturity. The peridium is 185–235 μm thick, composed of a single pseudoparenchymatic layer formed of cells 26.0–38.5 × 19.5–22.5 μm which gradually become smaller in the innermost layers. Apical oriice not conspicuous and diicult to locate due to the many lobes of the ascomata. Small basal tuft of hyphae present. Odor intense, similar to yogurt, even in dried specimens. Chambered inner cavity due to the lobes of ascomata, but also due to projections of the inner wall. This is lined with an epithecium formed of small warts similar to those of the external peridium, except that they are more darkened at their tips. The epithecium is 110–150 μm thick, pseudoparenchymatic in structure, and formed of mostly subglobose cells, with some angular elements about 23–32 × 18– 21 μm. The hymenium is arranged as a regular palisade composed of asci and irregularly cylindric but frequently septate paraphyses, 33.0–38.5 μm between each septum, 10.0–16.5 μm thick, more inlated in the areas near the hymenium and epithecium. Paraphyses 310–380 μm in length, considerably exceeding the hymenium. Asci cylindrical, pedicelate, inamyloid and indehiscent, with uniseriate Plate 2 — Ascospores (in Cotton blue). a. Genea brunneocarpa JC6; b. G. compressa JC28; c. G. dentata NVR; d. G. fageticola GK4129; e. G. lobulata BM1043; f. G. pinicola JC12; g. G. pseudobalsleyi CA01; h. G. pseudoverrucosa NV-D; i. G. oxygala JC14; j. G. tuberculata JC32. Scale bars = 10 μm. PLATE 2 49 spore arrangement; many asci are conspicuously larger than the ascospores they contain, reaching up to 252–289 × 28–32 μm, with a central pedicel measuring about 70 × 8 μm. Ascospores subglobose, (24.5–) 26.0–29.0 (–29.5) × (19.5–) 21.5–24.0 (–24.5) μm, X= 26.5 × 22.5 μm, Qm=1.20, ornamented by very small scattered conical or truncated-conical warts, measuring 1.3–1.7 (–2.0) × 0.8–1.5 (– 1.7) μm. Ecology: so far found under Pinus pinea mixed with Quercus ilex, as well as some Q. faginea and Juniperus thurifera in basic soils (pH near 8.0) with gypsum marl of central Spain. In winter (Jan.-Mar.). Distribution: known only from central Spain. Notes The brownish tones, abundant lobes, intense odor, and particular habitat help to easily distinguish G. tuberculata from all other species except for G. oxygala, which is morphologically similar. However, G. oxygala usually has fewer swellings and lobes in its ascomata and has very diferent spore ornamentation. Genea thaxterii is somewhat or not at all lobed and is lighter in color. Studied collections: SPAIN: Valladolid, Aldealbar, 867 m asl, hypogeous under Pinus pinea and Quercus ilex, leg. J. Cabero, 09-III2014, JC32, AH44178 HOLOTYPE. Valladolid, Cogeces del Monte, under P. pinea and Q. ilex, sandy soil of gypsyferous marl, leg. F. García, 12-III-2013, JC34, AH44180. Valladolid, Tudela de Duero, La Manbla, hypogeous under Pinus pinaster, leg. P. Juste, 23-I-2009, JC33 PJ 124-G, AH44179. Acknowledgements We are grateful to Dr. J. Rejos, curator of AH herbarium, for his assistance with the specimens examined in the present study. We also thank P. Chautrand, F. García, A. García Blanco, G. Hensel, P. Juste, M. Loizides, R. Martínez, A. Montecchi, B. Moreno-Arroyo, G. Pacioni, J.B. Pérez, C.M. Pérez del Amo, P. Ribollet, M.A. Sanz, F. Sáinz, and J.M. Vidal for kindly sending us their collections and images. Funding for the participation of M.E. Smith was provided in part by University of Florida’s Institute for Food and Agricultural Sciences (IFAS). 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Pablo Alvarado ALVALAB, La Rochela 47 39012 Santander Spain pablo.alvarado@gmail.com Zoltán Bratek Eötvös Loránd University 1053 Budapest Hungary bratek@caesar.elte.hu Julio Cabero C/ El Sol Nº 6. 49800 Toro (Zamora) Spain fotovideocabero@hotmail.com Gabriel Moreno Universidad de Alcalá 28871 Alcalá de Henares Spain gabriel.moreno@uah.es Nicolas Van Vooren 36 rue de la Garde 69005 Lyon France nicolas@vanvooren.info Vasileios Kaounas Sokratous 40, TK 19016 Artemis Attiki Greece bkaounas@gmail.com George Konstantinidis Agiou Kosma 25, ΤΚ 51100 Grevena Greece manitarock@hotmail.gr Carlo Agnello Via Antonio Gramsci 11 72023 Mesagne Italy agnellocarlo@libero.it Zsolt Merényi Eötvös Loránd University 1053 Budapest Hungary zmerenyi@gmail.com Matthew E. Smith University of Florida FL 32611 Gainesville USA trulesmith@gmail.com 51