Plant Syst Evol (2009) 281:209–216 DOI 10.1007/s00606-009-0202-8 ORIGINAL ARTICLE Phylogenetic relationships in the genus Hennediella (Pottiaceae, Bryophyta) inferred from nrITS sequence data Marı´a J. Cano Æ Juan F. Jime´nez Æ M. Teresa Gallego Æ Juan A. Jime´nez Æ Juan Guerra Received: 7 July 2008 / Accepted: 8 June 2009 / Published online: 11 July 2009 Ó Springer-Verlag 2009 Abstract The first phylogenetic study of the genus ledges, taluses, walls, and rocks with accumulated soils Hennediella using nuclear ITS rDNA for 11 of the 15 (Cano 2008). species recognized is presented. Maximum parsimony, The genus as presently defined includes small plants maximum likelihood, and Bayesian analyses were per- characterized mainly by leaves edged with cells that are formed to evaluate the monophyly of the genus and to different in shape, usually rectangular to linear, in 1–4 investigate phylogenetic relationships between the species. strata, less papillose and thicker-walled than the inner cells, The molecular data suggest that the core Hennediella is leaf margins plane, occasionally slightly recurved, mostly monophyletic only including Tortula platyphylla, although denticulate to dentate at least close to the apex, costa the affinities of two species (Hennediella heteroloma and usually with dorsal and ventral surface cells differentiated, H. longipedunculata) are ambiguous. The ITS sequences one band of dorsal stereids usually semicircular, surface of did not fully resolve the internal relationships of Henn- the laminal cells frequently plane, and KOH color reaction ediella while its affinities with Dolotortula, Leptophascum, orange to reddish. The sporophyte varies from long Sagenotortula,orTortula remain ambiguous. The present seta with emergent capsules and peristomes of 32 spirally circumscription of Hennediella is provisional until a mor- teeth with a long basal membrane to short seta with phological revision of genus Tortula is completed. capsules immersed and gymnostomous (Zander 1993; Cano 2008). Keywords Hennediella Á Pottiaceae Á nrITS sequence Á The genus was erected by Brown (1893) and later syn- Phylogeny Á Taxonomy onymyzed with Pottia Ehrh. ex Fu¨rnr (Sainsbury 1952; Wijk et al. 1962). Zander (1989) resurrected the genus Hennediella and revised its circumscription (Zander 1993), on the basis of contributions by Blockeel (1991). The Introduction genus, as defined by Blockeel (1991) and Zander (1989, 1993), comprises 20 species that had previously been Hennediella Paris is a small genus of the Pottiaceae dis- included in Desmatodon Brid., Pottia,orTortula Hedw. tributed throughout much of the world from 81°N in the Cano (2008) monographed the genus, reducing the Arctic to 78°S in the Antarctic, although its highest number of species to 15. Although she established the dif- diversity is found in the Southern Hemisphere. The species ferences between Hennediella and Syntrichia Brid., the of this genus occur from near sea level to 5,000 m in puna boundaries between Tortula and Hennediella remained and paramo formations, growing mainly on soil or on ambiguous. Tortula is distinguished by broader leaves, more convex surface of the laminal cells, absence of elongate marginal cells border, and yellow KOH reaction; M. J. Cano (&) Á J. F. Jime´nez Á M. T. Gallego Á however, some species with markedly recurved margins, J. A. Jime´nez Á J. Guerra for example T. platyphylla Mitt., have differentiated mar- Departamento de Biologı´a Vegetal (Bota´nica), Facultad de Biologı´a, Universidad de Murcia, 30100 Murcia, Spain ginal cells disposed in 1–2 strata and orange to reddish e-mail: [email protected] KOH leaf reaction. Species such as T. solmsii (Schimp.) 123 210 M. J. Cano et al. Limpr. have similar lax areolation and differentiated bor- Mu¨ll. Hal., Aloina Kindb., Chenia (including Leptopha- der, but in this case the leaf KOH reaction is yellow. Cano scum (Mu¨ll. Hal.) J. Guerra & M.J. Cano), Crumia W.B. (2008) noted that most of the species treated in her study Schofield, Dolotortula, Hennediella, Hilpertia, Micro- fulfil the requirements for belonging to Hennediella bryum Schimp., Phascopsis I.G. Stone, Sagenotortula, (H. antarctica (A˚ ngstr.) Ochyra & Matteri, H. arenae (Besch.) Stonea, Streptopogon (Taylor) Wilson ex Mitt., Syntrichia R.H. Zander, H. densifolia (Hook. f. & Wilson) R.H. and Willia Mu¨ll. Hal.) in Syntrichieae. Zander, H. denticulata (Wilson) R.H. Zander, H. heimii To date, there has been no molecular phylogenetic work (Hedw.) R.H. Zander, H. kunzeana (Mu¨ll. Hal.) R.H. on Hennediella. DNA-based analyses of Tortula and rela- Zander, H. longirostris (Hampe ex Mu¨ll. Hal.) R.H. Zander, ted genera have been also very limited (Werner et al. 2002; H. macrophylla R. Br. bis) Paris, H. marginata (Hook. f. Cano et al. 2005). Molecular data using rps4 sequence & Wilson) R.H. Zander, H. polyseta (Mu¨ll. Hal.) R.H. (Werner et al. 2002) suggest that some similar genera, for Zander, H. stanfordensis (Steere) Blockeel and H. steere- example Crossidium, Stegonia, or species of Pterygoneu- ana (R.H. Zander & H.A. Crum) R.H. Zander). However, rum, could also be included in the Tortula concept. In H. bellii (E.B. Bartram) R.H. Zander, H. heteroloma addition, Syntrichia formed a monophyletic clade clearly (Cardot) R.H. Zander, and H. longipedunculata (Mu¨ll. separated from Tortula. Hal.) R.H. Zander, which lack marginal teeth on their In this paper, we present a preliminary phylogenetic leaves, have a combination of characteristics that suggest a analysis based on ITS1, 5.8S, and ITS2 ribosomal DNA close relationship with Tortula. According to Zander and sequence data. The ITS region has been widely used to Eckel (2007), the two genera intergraded somewhat and a resolve phylogenetic relationships in different bryophytes less arbitrary distinction than simple emphasis on KOH groups such as Sphagnum L. (Shaw 2000), Amblystegium color reaction awaits more study. Schimp. (Vanderpoorten et al. 2001), Campylopus Brid. The genus Tortula is one of the most complex and (Stech 2004), Didymodon Hedw. (Werner et al. 2005), or diverse genera in terms of morphological variation in the Tortula (Cano et al. 2005). A review of the applications of Pottiaceae. With the exception of the treatment of the ITS region in bryophyte systematics is provided in Brotherus (1924), the genus Tortula has never been Vanderpoorten et al. (2006). The objectives of this study monographed. Only the taxonomic revision of the genus in are to test the monophyly of the species of Hennediella and South America (Cano and Gallego 2008) and floristic the internal relationship between them based on ITS treatments such as Zander and Eckel (2007) for North sequences; and to explore the phylogenetic relationship of America, Mishler (1994) for Mexico, Smith (2004) for Hennediella with other members of Pottiaceae to which it Great Britain and Ireland, and Cano (2006) for the Iberian is believed to be closely related, for example Dolotortula, Peninsula are outstanding. Zander (1989) recognized gen- Leptophascum, Sagenotortula, and Tortula, on the basis of era such as Chenia R.H. Zander, Dolotortula R.H. Zander, ITS sequences. Hennediella, Hilpertia R.H. Zander, Sagenotortula R.H. Zander, Stonea R.H. Zander or Syntrichia, as segregates of Tortula, but included taxa that traditionally were placed in Materials and methods other genera such as some species of Pottia, Phascum Hedw. (e.g. Phascum cuspidatum Hedw.), and the genus Plant material Desmatodon. In his classification of genera of Pottiaceae, Zander A total of 22 accessions were used from material collected (1993) placed Hennediella close to Dolotortula in the in the field and herbarium material from the following Pottieae. Dolotortula has a reddish orange KOH reaction of herbaria: AAS, ALTA, CAS, CBG, MUB, NY, and per- the leaf, broad and smooth laminal cells, border of elon- sonal herbarium of T.L. Blockeel. Eighteen accessions gated cells, and a costa with dorsal and ventral surface cells belong to species in Hennediella, representing 11 species differentiated and one semicircular dorsal band of stereids of the 15 presently recognized in the genus (Cano 2008). like Hennediella. The genera only differ in the multistra- The specimens included were selected from approximately tose border of stereid cells, undifferentiated dentate leaf 1,100 collections used for a taxonomic revision of the margins and smooth laminal cells of Dolotortula instead of genus (Cano 2008). Material from some species such as the usually unistratose to tristratose border, dentate leaf H. heimii is relatively abundant in the herbaria, however, margins, and, usually, papillose laminal cells of Hennedi- most of the studied species of this genus are very scarce ella. Later, Zander (2006) included Tortula and related and the collections in the herbaria available are often old. genera (Aloinella Cardot, Crossidium Jur., Globulinella When possible, several accessions of the same species were Steere, Pterygoneurum Jur., and Stegonia Vent.) in the extracted and amplified because of the taxonomic problems Pottieae and the rest of the genera of Pottioideae (Acaulon of the group. However, in many cases it was not possible. 123 Phylogeny of Hennediella 211 Hennediella marginata, and H. steereana were not inclu- PAUP 4.0b10 (Swofford 2002). Bayesian analyses were ded in the study because material suitable for sequencing performed using MrBayes v 3.1 (Huelsenbeck and Ron- was not available. Unfortunately, the amplification of quist 2001). Gaps were excluded from all analyses. Both H. macrophylla and H. antarctica extracts was unsuc- MP and ML analyses were conducted with heuristic sear- cessful. The amplifications of some species from the genus ches as described in Harrison and Langdale (2006). For ML Syntrichia, which were used in the study, for example analysis, the choice of the model of sequence evolution S. serrulata (Hook. & Grev.) M.J. Cano, S. robusta Duse´n, was performed using the program Modeltest 3.7 (Posada and S. pseudorobusta (Duse´n) R.H. Zander, also failed. and Crandall 1998) and Modeltest WebServer (http:// Tortula platyphylla was also sampled.