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Hebeloma Vesterholtii, a New Species in Section Theobromina

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Hebeloma Vesterholtii, a New Species in Section Theobromina Mycol Progress (2010) 9:215–223 DOI 10.1007/s11557-009-0627-z ORIGINAL ARTICLE Hebeloma vesterholtii, a new species in section Theobromina Ursula Eberhardt & Henry J. Beker Received: 20 July 2009 /Revised: 4 September 2009 /Accepted: 24 September 2009 /Published online: 15 October 2009 # The Author(s) 2009. This article is published with open access at Springerlink.com Abstract The species Hebeloma vesterholtii spec. nov. is other species of sect. Theobromina mentioned above and described. It is morphologically and molecularly closely H. plesiocistum which shares a number of characters with related to Hebeloma theobrominum from which it can be the members of sect. Theobromina. distinguished using either morphological or molecular characters. Both of these species belong to section Theobromina. The new species is known from 13 collec- Materials and methods tions from six European countries. ITS sequence data were obtained from all known col- Keywords Hebeloma theobrominum . Ectomycorrhizal lections of Hebeloma vesterholtii and all collections of fungi . Agaricales . Basidiomycota H. theobrominum examined in the context of this study. Sequences were produced by direct sequencing following the protocols detailed in Eberhardt et al. (2009)and Introduction references therein. In some of the tracefiles evidence was found for length variation within the template PCR products. Hebeloma section Theobromina was proposed by Vesterholt These were taken into account by including both length (2005) and includes H. theobrominum (the section type), variants as separate sequences into GenBank and into the H. erumpens and potentially another taxon that cannot be phylogenetic analyses. Collection details and GenBank clearly differentiated from H. erumpens (Eberhardt et al. accession numbers of new accessions are listed in Table 1. 2009). In this paper we describe a new species, H. Additional sequences considered in the molecular analyses vesterholtii spec. nov., that fits both morphologically in were EU570163—EU570189, EU693013, EU881922, and the sect. Theobromina and molecularly in the clade / EU887517 (GenBank acc. numbers) for which collection theobromina. We also provide molecular evidence using details were published by Eberhardt et al. (2009). To avoid ITS sequence data that H. vesterholtii is a taxon distinct from confusion stemming from the application of different the other members of its section. In addition, we include a species concepts we did not include sequences in the morphological key to separate this species from the two analysis for which the corresponding collections have not been morphologically examined within this project. Se- quence alignment was carried out using Se-al (vs. 2.0a11; * U. Eberhardt ( ) Rambaut 2002). Maximum likelihood analyses were carried CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, out as a heuristic search (100 replicates) in RAxML (vs. NL-3584 CT Utrecht, Netherlands 7.0.3; Stamatakis 2006) under the GTRMIX model includ- e-mail: [email protected] ing bootstrap analyses (500 replicates). FigTree (vs. 1.1.2; Rambaut 2008) was used to visualize the results, relying on H. J. Beker Rue Père de Deken, 19, njplot (vs. 2.2; Perrière and Gouy 1996) for the correct 1040 Bruxelles, Belgium placement of the bootstrap values. 216 Mycol Progress (2010) 9:215–223 Table 1 Collection details and GenBank accession no. of new Hebeloma spp. ITS sequences included in the phylogenetic analyses Taxon Collection codeaand GenBank acc. no. Origin Host plants herbarium number H. erumpens C, 41584, double HJB, FJ816618 ITALY: Prov Foggia, Mattinata, Tratturita Cistus monspeliensis; HJB12301 Quercus coccifera; Q. ilex H. theobrominum HJB, HJB10467 FJ816619 ENGLAND: Derby, Coombsdale Helianthemum sp. H. theobrominum HJB, HJB10679 FJ816620 ENGLAND: East Sussex, St Dunstan’s broadleaves H. theobrominum C, JV00-435, double HJB, FJ816621 DENMARK: LFM, Hamborg Skov, Fagus sp. HJB10909 Lolland: Fuglsang Storskov UTM PF 812,741 TBU 36. H. theobrominum C, JV00-314, double HJB, FJ816622 NORWAY, Leikanger, Sanden by Klyvi broadleaves HJB10908 H. theobrominum HJB, HJB10063 FJ816623 BELGIUM: Luxembourg, Han sur Lesse broadleaves H. theobrominum HJB, HJB11382 FJ816624 SPAIN: Catalunya, Brunyola Quercus sp. H. theobrominum C, JV06-1095, double HJB, FJ816625 ITALY: Grosseto, Cala Violina NW of Quercus cerris; Q. ilex; HJB11848 Grosseto Q. suber H. vesterholtiib BR, BR-MYCO 166528-76, FJ816626, FJ816627 BELGIUM: Luxembourg, Han sur Lesse Carpinus sp.;Corylus sp.; double, HJB, HJB1000133, Fagus sp.;Quercus sp. double C H. vesterholtii HJB, HJB10117 FJ816628 BELGIUM: Luxembourg, Han sur Lesse Carpinus sp.;Corylus sp.; Fagus sp.;Quercus sp. H. vesterholtii HJB, HJB10339 FJ816629, FJ816630 ITALY: Carpegna Castanea sp.;Quercus sp. H. vesterholtii HJB, HJB11206 FJ816631 BELGIUM: Luxembourg, Han sur Lesse Carpinus sp.;Fagus sp.; Quercus sp. H. vesterholtii HJB, HJB11199 FJ816632 BELGIUM: Luxembourg, Han sur Lesse Carpinus sp.;Fagus sp.; Quercus sp. H. vesterholtii HJB, HJB11195 FJ816633, FJ816634 BELGIUM: Luxembourg, Han sur Lesse Carpinus sp.;Fagus sp.; Quercus sp. H. vesterholtii HJB, HJB11377 FJ816635 SPAIN: Pais Vasco, Urdiain Quercus rubra H. vesterholtii C, JV08-132, double HJB, FJ816636, FJ816637 DENMARK, Eastern-Jylland: Elbæk Skov Fagus sylvatica; HJB12321 Elbæk Skov E of Horsens, N of Horsens Quercus robur Fjord H. vesterholtii C, JV08-133, double HJB, FJ816638, FJ816639 DENMARK, Eastern-Jylland: Elbæk Skov Fagus sylvatica; HJB12331 E of Horsens, N of Horsens Fjord Quercus robur H. vesterholtii C, DB8207, double HJB, FJ816640, FJ816641 BULGARIA, Sarantsi Quercus cerris; Q. HJB12344 frainetto;Q. sp. H. vesterholtii HJB, HJB12696 FJ816642, FJ816643 ENGLAND, Calke Abbey Fagus sylvatica, Quercus sp. H. vesterholtii HJB, HJB10674 FJ943237, FJ943238 ENGLAND, East Sussex, St Dunstan’s broadleaves H. vesterholtii HJB, HJB11869 FJ943239, FJ943240 ITALY, Sardinia, Mount Limbara not specified a Collection codes: C, Copenhagen; HJB, personal herbarium H. J. Beker; BR Meise, Belgium. b type of H. vesterholtii Details of the morphological analyses are given in (Vesterholt 2005). It is also important, when determining the Eberhardt et al. (2009). For each collection at least 50 spores average apex width, not to be selective with regard to the were measured in Melzer’s reagent, excluding the apiculus. cystidia chosen for measurement. To determine the average The maximum length and width of each spore was measured width at the apex about 100 cheilocystidia were measured and its Q value (ratio of length to width) calculated. Average on the lamellae edge. For other measurements, at least 20 length, width and Q value were calculated and recorded cheilocystidia, separated from the lamella edge, were mea- alongside the median, standard deviation and 5% and sured from each collection. Because of the complex shapes 95% percentiles. The assessment and coding of spore of the cheilocystidia four measurements were made: length, characters follows Vesterholt (2005). The average width width at apex, width at narrowest point in central region and of the cheilocystidium apex appears to be an important maximum width in lower half. The measurements are given character in the separation of species within Hebeloma in this order and an average value was calculated for Mycol Progress (2010) 9:215–223 217 each of these measurements. The measurements were Results and discussion made in 5% KOH. Unless mentioned otherwise, cited specimens are presently located in the personal collection The sequences of the currently known collections belong- of H.J. Beker (HJB). ing to the species described below as Hebeloma vesterholtii Fig. 1 Best of 100 ML tree of DB8207 12344 FJ816640 BULGARIA from a heuristic search with 100 JV08-132 FJ816637 DENMARK replicates based on ITS data; JV08-133 FJ816639 DENMARK bootstrap support was calculated BR-MYCO 166528-76 FJ816627 BELGIUM type with 500 replicates. For speci- HJB11206 FJ816631 BELGIUM mens showing length variation BR-MYCO 166528-76 FJ816626 BELGIUM type in the ITS, both variants were HJB10339 FJ816629 ITALY included in the tree and indicat- ed by the same shade font. HJB10674 FJ943237 ENGLAND Single sequences are in black DB8207 12344 FJ816641 BULGARIA HJB10674 FJ943238 ENGLAND HJB12696 FJ816643 ENGLAND Hebeloma vesterholtii HJB10117 FJ816628 BELGIUM HJB11199 FJ816632 BELGIUM HJB11195 FJ816633 BELGIUM HJB12696 FJ816642 ENGLAND 97 HJB11195 FJ816634 BELGIUM T HJB11869 FJ943240 ITALY h HJB11377 FJ816635 SPAIN e HJB11869 FJ943239 ITALY o JV08-133 FJ816638 DENMARK b r JV08-132 FJ816636 DENMARK o HJB10339 FJ816630 ITALY m JV06-1095 FJ816625 ITALY i 74 n HJB10009 EU570181 ESTONIA a 74 HJB10063 FJ816623 BELGIUM JV02-782 EU570182 DENMARK HJB9293 EU570180 ENGLAND Hebeloma theobrominum HJB11382 FJ816624 SPAIN HJB10467 FJ816619 ENGLAND JV00-314 FJ816622 NORWAY JV00-435 FJ816621 DENMARK 51 HJB10679 FJ816620 ENGLAND JV05-746 EU570186 SPAIN C41584 FJ816618 ITALY 92 Hebeloma erumpens JV05-740 EU570185 SPAIN JVG1071029-8 EU570187 SPAIN 51 H. cf. erumpens JVG991224 EU570184 SPAIN H. cf. erumpens JVG991224 EU570188 SPAIN 62 Hebeloma sp. HJB9381 EU570179 ENGLAND Hebeloma sp. HJB10897 EU570183 GERMANY 100 HJB11868 EU570172 ITALY JVG1021214 EU570170 SPAIN Hebeloma plesiocistum JVG1000201-6 EU570171 SPAIN 99 L960.110-524 EU887517 FRANCE JV05-792 EU570163 SPAIN Hebeloma cavipes 100 CL841104-30 EU693013 ITALY 96 JV95-386 EU570167 DENMARK HJB10370 EU570168 BELGIUM Hebeloma hiemale 77 HJB10472 EU570169 ENGLAND 100 JV04-551 EU570173 ITALY HJB10483 EU570175
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