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Ribosomal ITS Diversity Among the European Species of the Genus Hydnum (Hydnaceae)

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Ribosomal ITS Diversity Among the European Species of the Genus Hydnum (Hydnaceae) hydnum:11-Hydnum 10/12/2009 13:27 Página 121 Anales del Jardín Botánico de Madrid Vol. 66S1: 121-132, 2009 ISSN: 0211-1322 doi: 10.3989/ajbm.2221 Ribosomal ITS diversity among the European species of the genus Hydnum (Hydnaceae) by Tine Grebenc1, María P. Martín2 & Hojka Kraigher1 1 Slovenian Forestry Institute, Večna pot 2, SI-1000 Ljubljana, Slovenia. [email protected]; [email protected] 2 Departamento de Micología, Real Jardín Botánico, CSIC, Plaza de Murillo 2, E-28014 Madrid, Spain. [email protected] Abstract Resumen Grebenc, T., Martín, M.P. & Kraigher, H. 2009. Ribosomal ITS di- Grebenc, T., Martín, M.P. & Kraigher, H. 2009. Diversidad de las versity in the European species of the genus Hydnum (Hyd- secuencias ITS del ADN ribosómico nuclear en las especies del naceae). Anales Jard. Bot. Madrid 66S1: 121-132. género Hydnum (Hydnaceae) en Europa. Anales Jard. Bot. Madrid 66S1: 121-132 (en inglés). Several morphological species of the genus Hydnum L. are En Europa, sobre la base de la morfología se han identificado known to occur in Europe, but little molecular evidence exists to distintas especies en el género Hydnum L.; sin embargo, no se confirm the exact number and delimitation of the species. The tenían datos moleculares para confirmar el número exacto de present study seeks to investigate the genus Hydnum through táxones y las relaciones entre los mismos. Este trabajo se basa sequence analysis of the nuclear ribosomal ITS regions and en los análisis filogenéticos de las secuencias ITS del nrDNA, through morphological studies. The DNA sequences phyloge- que se comparan con los estudios morfológicos y los análisis es- netic analysis revealed high diversity among the ITS region se- tadísticos. Los análisis filogenéticos revelan una alta diversidad quences in H. repandum (two clades) and H. rufescens (six en las secuencias de las regions ITS en H. repandum (dos cla- clades) while the specimens of H. albidum, H. umbilicatum and dos) y en H. rufescens (seis clados), mientras que las muestras H. ellipsosporum formed one and clearly separated clade per de H. albidum, H. umbilicatum e H. ellipsosporum se agrupan morphological species. Phylogenetic distances among the en clados únicos, que coinciden con especies tradicionales recognised species and the obtained morphologically unsup- basadas en caracteres morfológicos. Los análisis morfológicos y ported clades are comparable and support the idea of several filogenéticos son similares y apoyan la idea de que en este new, yet undescribed species. The intraspecific variability in the género existen todavía un número de especies no descritas. En sequence data among phylogenetic species is generally low. De- las posibles especies filogenéticas, la variabilidad intraespecífi- tailed morphological analysis of putative informative morpho- ca de las secuencias es baja. Por otro lado, el resultado del de- logical characteristics could not support any of the observed tallado análisis morfológico no apoya ninguno de los clados de non-monophyletic DNA-sequences clades within H. repandum H. repandum o H. rufescens, por lo que todavía no queda claro or H. rufescens, and the proper use of names is not yet clear. el táxon al que designan estos nombres. Una variabilidad in- Similar intraspecific variation has also been observed in many traespecífica similar se ha observado en otros géneros de hon- other ectomycorrhizal genera and could be explained by inten- gos ectomicorrícicos y podría explicarse por especiación inten- sive speciation within variable groups under the influence of var- siva bajo la influencia de diversos factores (efecto de nicho, se- ious factors (niche effect, ectomycorrhizal partner selection). lección del hospedante ectomicorrícico). Keywords: Hydnum repandum, H. rufescens, DNA-sequences Palabras clave: Hydnum repandum, H. rufescens, relaciones phylogenetic relationships, morphological traits, nrDNA, in- filogenéticas, caracteres morfológicos, ADN ribosómico nuclear, traspecific variability. variabilidad intraespecífica. Introduction tive geotropic spines, ranging from small granular warts to clear individual spines (Ainsworth & al., 1973). Members of the family Hydnaceae Chevallier Recent literature cites six valid genera in the family. (1826) are primarily identified by the presence of posi- To our knowledge, five of them were never included hydnum:11-Hydnum 10/12/2009 13:27 Página 122 122 T. Grebenc & al. in any molecular analyses: Corallofungus Kobayasi, rDNA. Additionaly the variability of the restriction Dentinum Gray, Gloeomucro R.H. Petersen, Nigro- pattern within H. rufescens was observed after diges- hydnum Ryvarden, and Phaeoradulum Pat. (Kirk & al. tion of the amplified PCR product with HinfI en- 2001); while Hydnum as the type genus, was the only donuclease. The observed additional differences indi- genus represented in phylogenetic studies. At higher cate possible variability of collections from different taxonomic rank Hydnum was placed in Cantharellales sites (Agerer & al., 1996). Ostrow & Beenken (2004) first by Kreisel (1969) and later confirmed with molec- found a good correlation for selected morphological ular data by Pine & al. (1999) and subsequent papers. and molecular characters for four European species In European taxonomic reviews and determination with only few samples sequenced for each species. books the following species and varieties/forms have They reported no sequence diversity within H. ru - been mentioned in the genus: Hydnum albidum Peck, fescens, although only for H. repandum and H. ellip- H. repandum L.:Fr. and H. rufescens Pers. (Maas sosporum was the absence of any such intraspecific Geesteranus, 1975; Jülich, 1984; Courtecuisse & variability clearly stated. Duhem, 1995), H. repandum var. rufescens (Fr.) Barla Comparison of rDNA ITS sequences is a valuable and H. repandum f. rufescens (Pers.) Nikol. (synonym tool in phylogenetic studies, and to provide more ac- of H. rufescens Pers.) (Marchand, 1973; Cetto, 1976; curate species delimitation (Taylor & al., 2000). Cur- Gerhardt, 1997). In Slovenia two more taxa have been rently there is a poor overlap between morphological cited: H. umbilicatum Peck (Petkovšek & Vrščaj, and molecular species concept based on the variabili- 1998) [the species generally known from North ty of the rDNA ITS sequences in studied Hydnum America (Hall & Stuntz, 1971) and Asia (Maas collections. To support the results obtained at the Geesteranus, 1971)] and H. repandum f. amarum molecular level, selected morphological characters in- Vrščaj (Stropnik & al., 1988); however, the second dicative of taxonomic affiliation in the genus Hydnum taxon was never published with a comprehensive de- were measured and correlated to the clades retrieved scription, thus according to Art. 36.1, CABI Bio- in the DNA-sequences phylogenetic analyses. Multi- science Databases (Kirk & al., 2003) the name H. variate statistics were employed for these analyses. repandum f. amarum Vrščaj is treated as nom. inval. Hydnum rufescens and H. repandum are distributed Materials and methods over an exceptionally wide area and are even recog- DNA analyses were undertaken in the laboratories nised in the Far East (Asia) although several syno - in Slovenia (SFI) and in Spain (RJB). The different nyms from different areas and for local populations protocols were standardized at both sites, such that were published and many local names were con- the final results obtained from the same sample were specific with European species (Maas Geesteranus, equal. Thin layer cromatography (TLC) analysis was 1971). Hydnum elatum Massee and two more un- carried out in RJB in Spain. named Hydnum species were recognised for Asia and Australia in addition to H. umbilicatum (Maas Fungal material Geesteranus, 1971). The latter was described in North America by Peck (1902). Despite it was com- Specimens from the genus Hydnum included in the monly found on several continents (Hall & Stuntz, study (Table 1) were either collected from various lo- 1971) its presence in Europe was only confirmed for calities in the years 1999-2002 and stored in the herba- Finland (Huhtinen & Ruotsalainen, 2006). rium at Slovenian Forestry Institute (LJU) or ob- Clear delimitation of species cited in Europe is not tained from herbarium MA-Fungi (Madrid, Spain). always easy. Morphological characters can vary with We have tried to locate the type or representative ma- the developmental stage of pileus and environmental terial for European species in different institutional conditions during the growth period (Hall & Stuntz, herbaria (UPS, MSB, and PC; Holmgren & al., 1998). 1971; Maas Geesteranus, 1975). Spore size and shape However, according to the curators, the material ei- can well separate H. albidum and H. ellipsosporum ther does not exist or was not possible to locate. Even from the others (Ostrow & Beenken, 2004) while taxo - though H. umbilicatum has been cited in Slovenia, no nomical position of H. rufescens within the genus is reference material was available from the area; the two confusing, not only after classical identification but collections included in the study were kindly sent by also after molecular data have become available. Mole- Lorelei L. Norvell from the Pacific Northwest Mycolo - cular identification of H. repandum and H. rufescens gy Service. ectomycorrhizae on Norway spruce showed distinct Specimens with fully developed basidiomata and restriction patterns of amplified ITS region in genomic spores were used for examination of macro- and
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