Linum Flos-Carmini (Linaceae), a New Species from Northern Morocco Author(S): José Ruiz-Martín, Pedro Jiménez-Mejías, Juan M
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Linum flos-carmini (Linaceae), a New Species from Northern Morocco Author(s): José Ruiz-Martín, Pedro Jiménez-Mejías, Juan M. Martínez-Labarga & Rocio Pérez-Barrales Source: Annales Botanici Fennici, 52(5–6):383-395. Published By: Finnish Zoological and Botanical Publishing Board DOI: http://dx.doi.org/10.5735/085.052.0521 URL: http://www.bioone.org/doi/full/10.5735/085.052.0521 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. Ann. Bot. Fennici 52: 383–395 ISSN 0003-3847 (print) ISSN 1797-2442 (online) Helsinki 8 October 2015 © Finnish Zoological and Botanical Publishing Board 2015 Linum flos-carmini (Linaceae), a new species from northern Morocco José Ruiz-Martín1, Pedro Jiménez-Mejías2,3, Juan M. Martínez-Labarga2,4 & Rocio Pérez-Barrales1,5,* 1) Departamento de Biología Vegetal y Ecología, Universidad de Sevilla, ES-41012 Sevilla, Spain 2) Departamento de Biodiversidad y Conservación, Real Jardín Botánico, CSIC, ES-28014 Madrid, Spain 3) School of Biological Sciences, Washington State University, P.O. Box 644236, Pullman, WA 99164-4236, USA 4) Departamento de Sistema y Recursos Naturales, Escuela de Ingeniería Forestal y del Medio Natural, Universidad Politécnica de Madrid, ES-28040 Madrid, Spain 5) School of Biological Sciences, University of Portsmouth, Portsmouth PO1 2DY, UK (*corresponding author’s e-mail: [email protected]) Received 9 Jan 2015, final version received 28 Aug. 2015, accepted 1 Sep. 2015 Ruiz-Martín J., Jiménez-Mejías P., Martínez-Labarga J.M. & Pérez-Barrales R. 2015: Linum flos- carmini (Linaceae), a new species from northern Morocco. — Ann. Bot. Fennici 52: 383–395. Linum bicolor is a problematic North African taxon associated with L. setaceum. A consensus on the taxonomic status of a putative form (L. setaceum var. bicolor f. robusta) has not been achieved yet. We conducted a morphological study based on herbarium and field collections, combined with nuclear (ITS) and plastid (ndhF5-8 and trnL-F) phylogenies to clarify its status. The phylogenetic analysis did not reveal molecular divergence, but a comparative morphological study revealed substantial dif- ferences in traits previously used to distinguish the two taxa (stems, leaves, corolla and calyx size). In addition, we found that the indumentum of sepals, petal colour, and the arrangement of anthers and stigmas differed so clearly between the form robusta and L. setaceum s. stricto, that recognizing the former taxon as an independent species was justified. We provide an identification key to the yellow- and white-flowered Linum species in NW Africa. We also revised the unplaced name L. bicolor Schoubs. ex DC. and lectotypified it to avoid nomenclatural problems. Introduction showed that some sections were not monophyl- etic. This illustrates how crucial it is to investi- Linaceae is a cosmopolitan family of 14 genera, gate the taxonomy and systematics of the Linum of which Linum is the most diverse (ca. 250 species and to disentangle the infrageneric clas- species). It has a wide geographic distribution, sification. from temperate to tropical regions (McDill et al. The taxonomy of the Linum species in North 2009, 2011). The most recent taxonomic review Africa is particularly unclear. Linaceae was not of Linum recognized five sections (McDill et al. included in the extensive (although incomplete) 2009), although the molecular data in that study revision by Maire (1952–1987). His notes at the 384 Ruiz-Martín et al. • ANN. BOT. FENNICI Vol. 52 genus level were recorded in the periodical fas- NW Africa, and revise the nomenclatural and cicles of the Contributions à l’étude de la Flore taxonomic status of, as well as lectotypify, the de l’Afrique du Nord (cf. Lebrun & Stork 1978). name L. bicolor Schoubs. ex DC. More recent treatments and checklists adopted a conservative taxonomical and nomenclatural approach (cf. Quézel & Santa 1963, Greuter et Material and methods al. 1989, Valdés et al. 2002, Fennane et al. 2007) and did neither review the genus profoundly nor Morphological study include new combinations or synonyms. Linum bicolor illustrates well the taxonomic Morphological characters of nine specimens confusion. De Candolle (1824: 428) described from eight collections of L. setaceum var. bicolor the species from near Tanger (Morocco) as “Spe- f. robusta, as well as of 25 specimens and collec- cies non satis notae” (“species not well known”), tions of the Ibero-North African L. setaceum with a brief diagnosis in which he implicitly indi- s. stricto were measured and compared (Appen- cated that it was related to L. tenuifolium but dis- dix). We examined the important characters tinguishable from it by the presence of “annual according to the taxonomic treatment in Europe roots and flower colour”. After that description, and the Mediterranean (i.e. size of sepals, petals L. bicolor was treated by Ball (1878) as a doubt- and stigmata; Quézel & Santa 1963, Ockendon ful taxon. He stated “Quid sit nescio” (literally & Walters 1969, López 1979, Valdés et al. 2002, “I do not know what it is”), and discussed its Fennane et al. 2007), with particular emphasis possible origin as a hybrid between L. setaceum on the characters already reported as distinc- and L. angustifolium (= L. bienne). Later, Maire tive between L. setaceum s. stricto and the form (1933: 206) treated the taxon as L. setaceum var. robusta (habit, leaves, corolla, calyx size; Maire bicolor. Maire (1933: 207) also described a new 1933). Iberian and North African populations of form based on a collection by Pius Font-Quer L. setaceum s. stricto were evaluated separately (L. setaceum var. bicolor f. robusta), presumably to assess possible differences among them. We distinguishable by being larger than the typical also searched for type materials of L. bicolor form bicolor. in eleven of the herbaria listed in Stafleu and The field surveys conducted by the first Cowan (1985) as housing Schousboe’s collec- author, and the herbarium material revised for the tions (B, BM, BR, C, FI, GOET, K, L, S, UPS project Flora Iberica allowed identification of and W). materials ascribable to L. setaceum var. bicolor f. robusta. In this study, we aimed at revisiting this taxon and clarifying its taxonomic status. We Molecular study firstly conducted a morphological study revising Iberian and North African materials ascribable to Sampling L. setaceum s. lato (i.e. L. setaceum s. stricto and L. setaceum var. bicolor f. robusta). We studied Our sampling covered the four sections rec- vouchers from MA and SEV, which house the ognized by McDill et al. (2009) (Table 1). We two main North African collections in Spain, as included 21 samples, 14 of which were taken well as field material, and revised the digital- from McDill et al. (2009) and 7 new samples ized material housed at JSTOR Global Plants from the specimens collected in the field and (plants.jstor.org). We also carried out a phyloge- in the herbarium (Table 2). Two samples of netic analysis based on nuclear (ITS) and plastid L. setaceum (one from the Iberian Peninsula (ndhF5-8 and trnL-F) markers to disentangle the and one from Morocco), and a sample of the phylogenetic relationships. Our results suggest form robusta (hereafter L. flos-carmini) were that plants belonging to the form robusta are studied, representing both known morphological distinguishable from L. setaceum s. stricto at the extremes and distribution of L. setaceum s. lato. species level. We provide a key to identify the Three outgroups were included based on the yellow- and white-flowered Linum species from previous phylogenetic studies (Savolainen et al. ANN. BOT. FENNICI Vol. 52 • Linum flos-carmini, a new species from northern Morocco 385 2000, McDill et al. 2009), namely Sclerolinon were checked manually and poorly aligned posi- digynum, Anisadenia pubescens and Hugonia tions were eliminated using GBlock v0.91b busseana (Table 2). (Castresana 2000), with less restrictive treatment to avoid losing information. For each matrix, we performed maximum DNA extraction, PCR amplification and parsimony (MP) using TNT (Goloboff et al. sequencing 2008), maximum likelihood (ML) using RAxML ver. 7.3.1 (Stamatakis et al. 2006) implemented DNA was extracted from 15–20 mg of silica- in Cipres Science Gateway bioportal (https:// dried leaf tissue using the DNEasy Plant Minikit www.phylo.org), and Bayesian inference using (QIAGEN Inc.) following the protocol of the MrBayes 3.1.2 (Ronquist & Huelsenbeck 2003). manufacturer. We amplified and sequenced the For MP analyses, we set equal weighting of all nuclear ribosomal ITS region, and the plastid characters. The ITS matrix was engaged with ndhF and trnL-F regions. We used the primers 5000 random addition replicates, and the plas- 17SE and 26SE (Sun et al. 1994) for PCR ampli- tidial matrix with 3000 replicates, according to fications of ITS; for the plastid regions, we used computation time. Non-parametric bootstrap the primers described by Jansen (1992) for ndhF analysis under MP criterion took two trees per and by Taberlet et al. (1991) for trnL-F. The replication and TBR (tree bisection-reconnec- three markers were amplified using the follow- tion) swapping to completion on a maximum of ing PCR profile: 94 °C for 5 min; followed by 20 000 minimal trees found.