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Tortella Mediterranea (Pottiaceae), a New Species from Southern Europe, Its Molecular Affinities, and Taxonomic Notes on T

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Tortella Mediterranea (Pottiaceae), a New Species from Southern Europe, Its Molecular Affinities, and Taxonomic Notes on T Tortella mediterranea (Pottiaceae), a new species from southern Europe, its molecular affinities, and taxonomic notes on T. nitida Author(s): Heribert Köckinger, Michael Lüth, Olaf Werner and Rosa M. Ros Source: The Bryologist, 121(4):560-570. Published By: The American Bryological and Lichenological Society, Inc. https://doi.org/10.1639/0007-2745-121.4.560 URL: http://www.bioone.org/doi/full/10.1639/0007-2745-121.4.560 BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Tortella mediterranea (Pottiaceae), a new species from southern Europe, its molecular affinities, and taxonomic notes on T. nitida Heribert Kockinger¨ 1,4, Michael Luth¨ 2, Olaf Werner3 and Rosa M. Ros3 1 Roseggergasse 12, A-8741 Weisskirchen, Austria; 2 Emmendinger Strasse 32, D-79106 Freiburg, Germany; 3 Departamento de Biolog´ıa Vegetal, Facultad de Biolog´ıa, Universidad de Murcia, Campus de Espinardo, 30100 Murcia, Spain ABSTRACT. Tortella mediterranea is described as a new species from the Mediterranean Basin of southern Europe. It is mainly characterized by (1) absence of a stem central strand, (2) lanceolate leaf shape, (3) rather gradual limb-sheath transition, (4) partial presence of elongate marginal cells in the upper half of leaf and (5) ventral costal stereid band usually stronger than the dorsal. Chloroplast rps4 and nuclear ITS sequences were invariable within the studied specimens of the species, suggesting a low genetic variability, and the phylogenetic position is rather isolated with only obscure affinities to the group of closely related taxa around T. tortuosa and to T. fasciculata. The new species, seemingly a rather rare plant, was collected from dry and sunny to shaded limestone rocks. In addition, new molecular data for T. nitida s.str. and putative var. irrigata are provided and taxonomic problems within this variable species are discussed. KEYWORDS. Balearic Islands, Croatia, DNA sequencing, Greece, nuclear ITS marker, rps4 chloroplast marker, phylogeny, Spain, taxonomy. ^^^ For several years, the circumscription of the genus ITS sequence (Werner et al. 2005) but confirma- Tortella and the number of species included within tion of its status from an additional marker seems it have been in a state of flux. The European desirable. Werner et al. (2014) showed that the checklist of Hill et al. (2006) cites 12 species and former Trichostomum arcticum Kaal. molecularly several varieties, one of which, T. inclinata var. represents an unambiguous member of Tortella, densa (Lorentz & Molendo) Limpr., is better and therefore must be named T. spitsbergensis treated at species rank (Crundwell & Nyholm (Bizot & Ther.)´ O.Werner, Kockinger¨ & Ros. In 1962; Hedenas¨ 2015; Nebel & Philippi 2000; Smith Europe,itisknownonlyfromSvalbard.Theplant 2004). On the other hand, T. limosella (Stirt.) reported as Trichostomum cuspidatissimum Cardot P.W.Richards & E.C.Wallace, known only from the &Ther.´ (in the sense of T. arcticum)fromnorthern type material, probably represents only an ‘‘aber- Sweden (Engelmark 1974) proved to be a hybrid rant form or short-lived genetic variant of another between T. spitsbergensis and T. arctica (Arnell) species’’ (Blockeel et al. 2014). Recent papers using Crundwell & Nyholm, and should be named molecular tools (Grundmann et al. 2006; Werner et Tortella 3cuspidatissima (Cardot & Ther.)´ O.Wern- al. 2005) demonstrated that Pleurochaete Lindb. is er, Kockinger¨ & Ros. Since both parents seemingly deeply nested within Tortella, suggesting its inclu- do not occur within its European range, an sion in that genus. Alternatively, however, it may alternative approach is to treat it as an indepen- be treated as a putatively young genus which dent, hybridogenous species analogous to T. rigens evolved from the interior of an older one. Tortella Alberts., both reproducing only vegetatively. The inflexa (Bruch) Broth. was presumed to be report of one of its parents, T. arctica,fromSweden excluded from the genus on the basis of a single by L. Hedenas¨ and P. Eckel (in Ellis et al. 2011) has 4 Corresponding author’s e-mail: proved to be erroneous. Nuclear and chloroplast [email protected] sequences of the Swedish material showed that it DOI: 10.1639/0007-2745-121.4.560 belonged to T. tortuosa (Hedw.) Limpr. s.l. (see The Bryologist 121(4), pp. 560–570 Published online: December 10 2018 0007-2745/18/$1.25/0 Copyright Ó2018 by The American Bryological and Lichenological Society, Inc. Kockinger¨ et al.: A new Tortella species from southern Europe 561 Figure 1. ITS phylogram obtained by Bayesian analysis. Tortella mediterranea is situated on a well-supported clade (pp¼1) close to T. arctica, T. densa, T. fragilis, T. inclinata, T. pseudofragilis and T. tortuosa. Kockinger¨ & Hedenas¨ 2017, Fig. 1). The most During morphological studies on the variability recent change in the European Tortella flora was of European Tortella tortuosa, the first author’s presented by Kockinger¨ & Hedenas¨ (2017). They attention was attracted by photos with this name discovered that T. bambergeri (Schimp.) Broth. in from Mallorca in Luth’s¨ ‘‘Pictures of Bryophytes’’ its original sense belongs to T. tortuosa s.l., whereas (Luth¨ 2014), seemingly representing something the plants understood as T. bambergeri during outside the known variation of T. tortuosa. The loan recent decades belong to two distinct taxa, T. of this plant and of additional Mediterranean fasciculata (Culm.) Culm and T. pseudofragilis collections of T. tortuosa from Luth’s¨ private (Ther.)´ Kockinger¨ & Hedenas.¨ herbarium revealed a further specimen from Greece 562 The Bryologist 121(4): 2018 almost identical to the Mallorcan plant. In addition, genetic variability, four samples of T. fasciculata were HK detected a third similar collection from Croatia included. In the case of T. tortuosa we deliberately in his own herbarium. chose four genetically very different plants whose The main aim of the present study is to establish identity is known to us. Trichostomum crispulum the correct rank and systematic position of this Bruch was used to root the trees. neglected plant. For this purpose, OW agreed to The chloroplast rps4 region was chosen to undertake a molecular investigation. We also represent the chloroplast DNA because together with checked the synonymy of similar species for any the atpB-rbcL spacer it is the region with the highest name that might be available for our plant. The number of available Tortella sequences at GenBank overall appearance of this plant is 6 intermediate and the number of base differences between sequences between Tortella nitida (Lindb.) Broth. and T. are in the same range for these two regions. The fact tortuosa. Since the first author, in collaboration with that rps4 is a protein coding gene reduces possible L. Hedenas,¨ is already engaged in a thorough ambiguities in the alignment. For rps4region,the investigation of the T. tortuosa complex, where this same three deviating Tortella specimens, two of T. issue was taken into account, we concentrated our nitida s.str. and two of T. nitida cf. var. irrigata were search on T. nitida. A detailed discussion of the newly sequenced. The sequences of eleven Tortella general variation within this mainly Mediterranean species were added from public databases. Three of species was provided by Eckel (1998). She also them belonged to T. fasciculata, four to T. tortuosa. studied and illustrated the Lindberg type. We needed Trichostomum crispulum was again used to root the to pay particular attention to the younger synonym trees. GenBank accession numbers of all sequences T. cirrifolia (Mitt.) Broth. and to T. nitida var. used in this study are given in Table 1. irrigata (H.Winter) R.H.Zander, since the descrip- DNA isolation and amplification of nrITS and tions of both these Macaronesian taxa come closer to cprps4. Total DNA was extracted from dry material our neglected plant than usual expressions of T. using the NaOH extraction method as explained in nitida. Werner et al. (2002). The nrITS region was amplified with the primers AB 101F and AB 102R (Douzery et METHODS al. 1999). In the case of the rps4 region, the primers Morphological study. The description is based used were rps5 (Nadot et al. 1994) and trnas (Buck on all three known collections. To detect further et al. 2000). Two lL of stock DNA were used as material of the deviating plant many samples template in 25 lL reaction volume. To this were identified as Tortella nitida and T. tortuosa from added 200 lM of dNTP, 2 mM MgCl2, 2 units Taq MUB herbarium were revised by RMR and a series of polymerase (DreamTaq, ThermoFisher), one lL doubtful plants also checked by HK. Tom Blockeel BLOTTO (10% skimmed milk powder and 0.2% was searching for similar plants among his Greek NaN3 in water) and the buffer (provided by the collections and provided specimens. David Holyoak enzyme supplier). BLOTTO attenuates the PCR sent some T. nitida collections. inhibition caused by plant compounds (De Boer et The type material of Tortella cirrifolia from NY al. 1995). The amplification conditions were as and MO and Tortella nitida var. irrigata (H.Winter) follows: 3 min at 948C, 35 cycles with 30 sec at 948C, R.H.Zander from JE was borrowed and studied by 30 sec at 508C(rps4) or 558C (ITS) and 1 min at RMR.
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