Evaluation of the taxonomic status of the genus Aliella (Compositae, Gnaphalieae): a recircumscription of the genus Phagnalon

NOEMÍ MONTES-MORENO; NÚRIA GARCIA-JACAS; CARLES BENEDÍ; LLORENÇ SÁEZ

A taxonomic evaluation of the genus Aliella, endemic to the Moroccan Atlas Mountains, is presented. We evaluate the taxonomic status of Aliella using a morphologic and molecular approach. Firstly, we discuss the variability and usefulness of its morphological diagnostic characters. Secondly, we analyse nuclear ETS and ITS, and chloroplast ycf3-trnS and trnT-trnL spacers. Phylogenetic analyses of the nrDNA and cpDNA spacers suggest the paraphyly of Aliella and Phagnalon. Two species of Aliella, A. ballii and A. embergeri, form a strongly supported clade. In contrast, relationships of A. platyphylla to A. ballii and A. embergeri are only weakly supported, and A. iminouakensis do not form a group with the other species and shows two different haplotypes. The morphological and diagnostic characters of Aliella are described and compared with an extensive sampling of the closely related genus Phagnalon. Our results strongly suggest that Aliella should be merged into Phagnalon. For each accepted taxon, taxonomical, chorological, and ecological data are provided. Six taxa are recognized, three species and three subspecies. Three lectotypifications of specific names and three new combinations are proposed. New descriptions and distribution maps of the recognized taxa are given.

-Phytotaxa 148 (1): 1–31 (2013) www.mapress.com / phytotaxa / Copyright © 2013 Magnolia Press ISSN 1179-3155 (print edition) Article PHYTOTAXA ISSN 1179-3163 (online edition) http://dx.doi.org/10.11646/phytotaxa.148.1.1 Evaluation of the taxonomic status of the genus Aliella (Compositae, Gnaphalieae): a recircumscription of the genus Phagnalon NOEMÍ MONTES-MORENO1*, NÚRIA GARCIA-JACAS1, CARLES BENEDÍ2 & LLORENÇ SÁEZ3 1 Botanic Institute of Barcelona (IBB-CSIC-ICUB), Passeig del Migdia s.n., ES-08038 Barcelona, Spain. E-mail: n.montesmoreno@gmail.com (corresponding author), ngarciajacas@ibb.csic.es 2 Departament de Productes Naturals, Biologia Vegetal i Edafologia, Unitat de Botànica, Facultat de Farmàcia, Universitat de Barcelona, Avda. Joan XXIII s.n., ES-08028 Barcelona, Spain. E-mail: cbenedi@ub.edu 3 Departament de Biologia Animal, Biologia Vegetal i Ecologia, Unitat de Botànica, Facultat de Biociències, Universitat Autònoma de Barcelona, ES-08193 Bellaterra, Barcelona, Spain. E-mail: llorens.saez@uab.es Abstract A taxonomic evaluation of the genus Aliella, endemic to the Moroccan Atlas Mountains, is presented. We evaluate the taxonomic status of Aliella using a morphologic and molecular approach. Firstly, we discuss the variability and usefulness of its morphological diagnostic characters. Secondly, we analyse nuclear ETS and ITS, and chloroplast ycf3trnS and trnT-trnL spacers. Phylogenetic analyses of the nrDNA and cpDNA spacers suggest the paraphyly of Aliella and Phagnalon. Two species of Aliella, A. ballii and A. embergeri, form a strongly supported clade. In contrast, relationships of A. platyphylla to A. ballii and A. embergeri are only weakly supported, and A. iminouakensis do not form a group with the other species and shows two different haplotypes. The morphological and diagnostic characters of Aliella are described and compared with an extensive sampling of the closely related genus Phagnalon. Our results strongly suggest that Aliella should be merged into Phagnalon. For each accepted taxon, taxonomical, chorological, and ecological data are provided. Six taxa are recognized, three species and three subspecies. Three lectotypifications of specific names and three new combinations are proposed. New descriptions and distribution maps of the recognized taxa are given. Key words: Aliella, endemism, ETS, Gnaphalieae, infraspecific variation, ITS, Mediterranean region, Phagnalon, trnTtrnL, ycf3-trnS Introduction The genus Aliella Qaiser & Lack (1986: 487) (Compositae, Gnaphalieae) was described as a segregate from Phagnalon Cassini (1819: 173) on the basis of vegetative and reproductive characters (Qaiser & Lack 1986). It is currently accepted as an independent genus (Anderberg 1991, Dobignard 1997, Bayer et al. 2007, Ward et al. 2009). Aliella comprises four species and two subspecific taxa: A. ballii (Klatt 1896: 836) Greuter (2003: 241); A. ballii subsp. ballii; A. ballii subsp. nitida (Emberger 1935: 224) Qaiser & Lack (1986: 493); A. embergeri Humbert & Maire in Maire (1928: 52) Qaiser & Lack (1986: 493); A. iminouakensis (Emberger 1932: 189) Dobignard & Jeanmonod in Dobignard (1997: 143) and A. platyphylla (Maire 1924: 85) Qaiser & Lack (1986: 490). Aliella helichrysoides (Ball 1873: 364) Qaiser & Lack (1986: 492) is a synonym of Aliella ballii. The generic diagnosis was based on the presence of bracts on the peduncle similar in shape and size to the involucral ones, the presence of waxy cushions on the corolla lobes, tubular female florets, caudate anthers, and pappus setae barbellate from the base to apex. The species of Aliella are chamaephytes which grow in calcareous or siliceous rock crevices in the Atlas Mountains of Morocco, at altitudes of 1800 m to 3600 m. Accepted by Alexander Sennikov: 4 Oct. 2013; published: 28 Nov. 2013 1 Previous classifications by other authors (Ball 1873, Klatt 1896) originally included some species of Aliella in other genera: Gnaphalium Linnaeus (1753: 850) or Helichrysum Miller (1754: 462). Subsequent taxonomic difficulties sometimes led to different and somewhat confusing infrageneric classifications and taxonomic treatments (Maire 1928, Emberger 1932, 1935, Quézel 1951, Qaiser & Lack 1986, Dobignard 1997, Greuter 2008), and these are shown in Table 1. TABLE 1. Major treatments of taxa (at species or subspecies rank) included in Aliella and Phagnalon sect. Gnaphaliopsis. Maire (1928) Emberger (1932, 1935) Quézel (1951) Qaiser & Lack (1986) Anderberg (1991) Dobignard (1997) Greuter (2008) Phagnalon helichrysoides Phagnalon helichrysoides Phagnalon helichrysoides Aliella helichrysoides Aliella bracteata Aliella helichrysoides Aliella ballii Phagnalon helichrysoides var. nitidum – Aliella helichrysoides subsp. nitida Aliella helichrysoides subsp. nitida Aliella ballii subsp. nitida Phagnalon platyphyllum Phagnalon platyphyllum Phagnalon platyphyllum Aliella platyphylla Aliella platyphylla Aliella platyphylla Aliella platyphylla Phagnalon embergeri Phagnalon embergeri Phagnalon embergeri Aliella embergeri Aliella embergeri Aliella embergeri Phagnalon lepineyi – – – – Phagnalon iminouakense – – – Aliella iminouakensis – – Aliella iminouakensis The generic delineation of Aliella and Phagnalon has been discussed by several authors. Maire (1928) pointed out a close relationship between Ph. platyphyllum Maire (1924: 85) (= Aliella platyphylla) and Ph. latifolium Maire (1929: 134) based on general appearance and geographic distribution: both taxa, together with A. ballii, were found in the same locality (see additional specimens examined and Appendix 1). According to Anderberg (1991), Phagnalon was nested within a clade with Anisothrix Hoffmann (1898: 129), whereas Aliella was more closely related to other taxa of Gnaphalieae. Finally, a molecular phylogenetic study of Aliella, Phagnalon and other early branching genera of the tribe Gnaphalieae (the “Relhania clade” sensu Ward et al. 2009) revealed that the monophyly of Aliella and Phagnalon s. str. was not supported (MontesMoreno et al. 2010). Regarding chromosome numbers, all the reports for Aliella show 2n = 18 (Quézel 1957, Humphries et al. 1978, Galland & Favarger 1985), although one conflicting report shows 2n = 14 for A. ballii (Humphries et al. 1978). All reports on the chromosome number of Phagnalon show 2n = 18. Pollen studies revealed that the morphology of the pollen grains of Aliella and Phagnalon is very homogeneous, both at specific and generic levels (El Ghazaly & Anderberg 1995). However, differences in the exine, which was thicker in Aliella, and in the internal foramina, which were less visible and vestigial in Aliella, were observed (El Ghazaly & Anderberg 1995). The results confirmed that these two genera belong to the tribe Gnaphalieae, as they showed an irregularly interlaced inner exine layer, which is characteristic of the Gnaphalieae (Anderberg 1991). Otherwise, the subtribal affinities of Aliella were poorly resolved. Qaiser & Lack (1986) suggested that this genus occupies an intermediate position between the subtribes Athrixiinae and Gnaphaliinae of Merxmüller et al. (1977). This subtribal affinity was suggested because of its tubular to infundibuliform florets and sweeping hairs distributed apically, which characterise the Athrixiinae, and also because of its caudate anthers and parallel stigmatic rows, which define the Gnaphaliinae. Based on a cladistic analysis of morphological data, Anderberg (1988) included Aliella within the “basal group taxa”, which was an informal group within the Gnaphalieae unassigned to any subtribe. On the basis of molecular data, Aliella is currently nested in the “Relhania clade”, which is mainly distributed throughout South and Tropical Africa and Madagascar (Montes-Moreno et al. 2010) with several representatives in Macaronesia and the Mediterranean, North Africa, Arabia, the Middle East and the Irano-Turanian regions (Anderberg 1991). 2 • Phytotaxa 148 (1) © 2013 Magnolia Press MONTES-MORENO ET AL. In the present paper, we (1) examine the variability and usefulness of the diagnostic characters of Aliella using a comprehensive sampling of Phagnalon species for comparison of those characters; (2) establish the phylogenetic relationships of the taxa currently included in Aliella; (3) contrast the relationships between sequence data and morphology and evaluate whether Aliella and Phagnalon constitute natural groups or not; and (4) provide a taxonomic revision and nomenclature for the accepted taxa. Material and Methods Phylogenetic study Sampling included one representative from two to three different populations of the five Aliella taxa: A. ballii subsp. ballii, A. ballii subsp. nitida, A. embergeri, A. iminouakensis and A. platyphylla, increasing the sampling of Aliella carried out in Montes-Moreno et al. (2010), and 29 of 36 species of Phagnalon (one population per species). Outgroups were chosen from Anisothrix, Athrixia Ker Gawler (1823: 681) and Pentatrichia Klatt in Schinz (1895: 436) based on the work of Montes-Moreno et al. (2010). The analyses used 172 sequences, including 26 new sequences (46 ETS, 46 ITS, 40 ycf3-trnS, 40 trnT-trnL). Voucher data and GenBank sequence accession numbers are given in the Appendix 2. Total genomic DNA was extracted following the CTAB method of Doyle & Doyle (1987), as modified by Cullings (1992), from silica-gel dried leaves collected in the field, herbarium material, or fresh leaves derived from plants cultivated at the Botanic Institute of Barcelona. DNA was extracted from old herbarium specimens using the Nucleospin Plant Kit (Macherey-Nagel, PA, USA) or the Dneasy Plant Mini Kit (Qiagen, Valencia, CA, USA). Double-stranded DNA of the ITS region was obtained by PCR amplification and sequenced using 17 SE (Sun et al. 1994) and ITS1 (White et al. 1990) as forward primers together with 26 SE (Sun et al. 1994) and ITS4 (White et al. 1990) as reverse primers. The PCR profile used for amplification follows the protocol of Montes-Moreno et al. (2010). The ETS region was amplified by PCR and sequenced using ETS1F (Linder et al. 2000) and AST1F (Markos & Baldwin 2001) as forward primers, and AST1R (Markos & Baldwin 2001), 18S2L (Linder et al. 2000) and 18S-ETS (Baldwin & Markos 1998) as reverse primers. The PCR profile followed the protocol of Montes-Moreno et al. (2010). Double-stranded DNA of the ycf3-trnS spacer was amplified by PCR and sequenced using SP43122F and SP44097R as the forward and reverse primers, respectively (Hershkovitz 2006). Double-stranded DNA of the trnT-trnL spacer was amplified and sequenced using trnT-A2F (Cronn et al. 2002) as the forward primer, and trnL-b as the reverse primer (Taberlet et al. 1991). The PCR profile followed the protocol described by Montes-Moreno et al. (2010). Purification of PCR products was carried out using QIAquick PCR Purification Kit (Qiagen). PCR products of recalcitrant samples were purified using the DNA Clean and Concentrator kit (Zymo Research, CA, USA). Direct sequencing of the amplified DNA segments was performed at the “Serveis Cientificotècnics” of the University of Barcelona using a BigDye Terminator Cycle Sequencing v.3.1 (Applied Biosystems, Forster City, CA, USA), following the manufacturer’s protocol, and analysed on an ABI PRISM 3730 DNA analyser (Applied Biosystems). The nucleotide sequences were edited using Chromas 1.56 (Technelysium Pty, Tewantin, Australia). DNA sequences were aligned visually by sequential pairwise comparison (Swofford & Olsen 1990). Phylogenetic analyses were performed with two different data sets: (1) ETS+ITS, and (2) ycf3-trnS+trnTtrnL. Parsimony analysis involved heuristic searches conducted with PAUP version 4.0b10 (Swofford 2002) using tree bisection reconnection (TBR) branch swapping with character states unordered and unweighted. In order to locate islands of most parsimonious trees (MPTs) (Maddison 1991), 1000 replications were carried out with random taxon addition, also with TBR branch swapping. The insertion/deletion events (indels) were coded as missing data. Bootstrap (BS, Felsenstein 1985) analyses were performed to obtain branch robustness using 100 replicates in the nrDNA data set and 80 replicates in the cpDNA data set of heuristic search with simple taxon addition and TBR branch swapping. Bayesian analysis was carried out using MrBayes 3.1.2 (Huelsenbeck & Ronquist 2001, Ronquist & Huelsenbeck 2003). The best fit model of nucleotide-substitution for each data set was selected by means of hierarchical likelihood ratio tests (hLRT) and Akaike Information TAXONOMIC STATUS OF THE GENUS ALIELLA Phytotaxa 148 (1) © 2013 Magnolia Press • 3 Criterion (AIC), as implemented in MrModeltest 2.2 (Nylander 2004). The analyses were performed assuming the GTR+Γ substitution model for ETS+ITS data set and F81+ I + Γ for the ycf3-trnS+trnT-trnL data set. Two analyses each with four Markov chains, initiated with random starting trees, were run for 2 × 106 generations and sampled every 100 generations. The first 5000 trees, sampled prior to the stabilization of the log-likelihood value, were ruled out as the burn-in samples. Posterior probabilities (PP) were estimated of the post burn-in trees by constructing a 50% majority-rule consensus tree in PAUP version 4.0b10 (Swofford 2002). Congruence in the phylogenetic signal between the combined nrDNA datasets, nrDNA and cpDNA and combined cpDNA data sets was examined by visual comparison of the tree topologies and by conducting a partition homogeneity test (incongruence length difference, ILD, Farris et al. 1995a, b). The ILD significance values were calculated using 1000 replicates with TNT (Goloboff et al. 2003–2005) using the INCTST script, which was kindly provided by the authors of the program. Morphological and taxonomic study This taxonomic study is based on 90 herbarium specimens of Aliella, including type material, and also 2337 herbarium specimens representing all the species of Phagnalon, in order to evaluate the variability of the diagnostic characters. These collections were from the herbaria ARAN, B, BC, BCN, BM, C, COI, E, FI, HBG, JACA, pers. herb. of J.X. Soler, K, LISU, M, MA, MGC, MPU, RAB, RNG, SALA, SANT, SEV, TFC, TSV, UPS, W and WU. In addition, several photographs of both type and non-type specimens from P and RAB were also examined. Moreover, several specimens of each Aliella taxon were observed and collected in the wild during field surveys (2004 and 2007) and, in some cases, material from private collections (F. Gómiz and L. Sáez) was also studied. Relevant literature was searched for existing typifications and protologues for each taxon studied, in order to designate lectotypes when this was necessary. A Nikon C-PS binocular stereoscopic microscope model was used for gross morphology, whereas for micromorphological features, samples of leaf indumentum, bracts, florets and achenes, pappus, as well as stamens and styles, were soaked in a commercial solution of sodium hypochlorite for 30 to 60 minutes until getting totally transparent. These were subsequently placed on slides, examined under a Zeiss Axioplan microscope and photographed with a digital camera (AxioCam MRc5 Zeiss). Samples of receptacle, styles and achenes were attached to aluminium stubs, coated with a 50 nm layer of gold and examined using a Hitachi 2300 S scanning electron microscope (SEM) at accelerating voltages of 10–15Kv. Terminology for morphological features studied follow Drury & Watson (1966) and Hilliard & Burtt (1981). Diagnostic, reproductive and vegetative characters of recognized Aliella taxa were studied for several herbarium specimens (see additional specimens examined). Comparative analysis of morphological characters was undertaken for a representative sampling of 30 of the 36 Phagnalon taxa (Qaiser & Abid 2003). Samples of selected species of Phagnalon were chosen as well as to cover its whole distribution range: Mediterranean, Macaronesian, Irano-Turanian, Saharo-Arabian or Eritreo-Arabian regions (Appendix 1). Box plots containing means and percentiles and scatter-plots were carried out in order to outline the variability of each quantitative character using STATGRAPHICS 5.0 plus package (Manugistics Inc., Rockville, MD, USA). Results Phylogenetic relationships of Aliella and Phagnalon Results from the ILD test do not found significant incongruities between ITS and ETS and between ycf3trnS and trnT-trnL. However, significant incongruities (P = 0.001) were found between the nrDNA data sets (ITS+ETS) and the cpDNA data sets (ycf3-trnS+trnT-trnL). The aligned length of ETS and ITS data set was 2297 positions, of which 270 were parsimony informative. Maximum parsimony analyses resulted in a strict consensus tree of 541 steps, CI = 0.5970, HI = 0.4030 and RI = 0.8093 (Figure 1). According to results obtained from nuclear phylogeny, the monophyly of Aliella and Phagnalon is not statistically supported. 4 • Phytotaxa 148 (1) © 2013 Magnolia Press MONTES-MORENO ET AL. SA Phagnalon barbeyanum WC-MED Phagnalon sordidum Phagnalon abyssinicum 0.92 1.00 64 Phagnalon phagnaloides 1.00 Ph. stenolepis var. abdulazizianum 0.99 62 100 Phagnalon harazianum Cl1 Phagnalon harazianum Cl2 1.00 100 Phagnalon woodii pop 1 1.00 Phagnalon woodii pop 2 92 0.75 Phagnalon bicolor - Phagnalon viridifolium 1.00 66 Phagnalon purpurascens 1.00 98 0.92 75 Phagnalon umbelliforme Phagnalon melanoleucum 1.00 99 Phagnalon calycinum Phagnalon quartinianum 0.58 0.74 Phagnalon saxatile - - WIDE DISTRIBUTION Phagnalon graecum 1.00 Phagnalon metlesicsii 100 Phagnalon rupestre SAHARO ARABIAN IT Phagnalon schweinfurthii 1.00 100 MEDITERRANEAN - MACARONESIAN 1.00 99 ERITREO - ARABIAN Phagnalon stenolepis 97 1.00 89 Phagnalon sinaicum Phagnalon pygmaeum 1.00 100 - Phagnalon pycnophyllon Phagnalon kotschyi 0.99 60 0.51 Phagnalon niveum Phagnalon persicum - 0.74 66 Phagnalon acuminatum 1.00 100 Phagnalon darvazicum Cl1 1.00 85 Phagnalon darvazicum Cl2 ANTIATLAS: TIZNIT AREA Ph. latifolium pop 1 1.00 100 Ph. latifolium pop 2 Aliella iminouakensis pop 1 1.00 100 1.00 IRANO-TURANIAN 0.86 100 ANTIATLAS: DJEBEL KEST C HIGH ATLAS: IMI N OU AKKA Aliella iminouakensis pop 2 C HIGH ATLAS: M GOUN AREA Aliella ballii subsp. nitida pop 4 TICHKA AREA: W HIGH ATLAS 1.00 99 1.00 1.00 100 OUKAIMEDDEN AREA Aliella ballii subsp. balli pop 3 85 1.00 Aliella embergeri pop 1 100 Aliella embergeri pop 2 1.00 100 C HIGH ATLAS: JAFFAR CIRQUE C HIGH ATLAS: M GOUN AREA Aliella platyphylla pop 2 C HIGH ATLAS: DJEBEL BOU OURIOUL Aliella platyphylla pop 1 OUKAIMEDDEN AREA Pentatrichia avasmontana 1.00 94 RERAYA VALLEY Anisothrix kuntzei Pentatrichia petrosa OUTGROUP 0.88 65 Aliella ballii subsp. balli pop 1 Aliella ballii subsp. balli pop 2 0.01 FIGURE 1. Consensus phylogram of the post-burn-in trees resulting from Bayesian MCMC analysis of the combined ETS and ITS data sets. Numbers above the branches are Bayesian clade credibilities (posterior probability, PP); numbers below the branches indicate parsimony bootstrap percentages (BS). Regional acronyms: C central, MED Mediterranean, SA Saharo-Arabian, W western. Other abbreviations: Cl= clone; pop = population. The aligned length of ycf3-trnS and trnT-trnL combined data set was 1193 positions, of which 31 were parsimony-informative. Maximum parsimony analyses generated a strict consensus tree of 50 steps, CI = 0.66, HI = 0.34 and RI = 0.8547 (Figure 2). Based on results obtained from the combined chloroplast data set, Aliella and Phagnalon are paraphyletic according to Bayesian analyses of the cpDNA dataset, since A. iminouakensis is nested amongst other Phagnalon taxa (PP = 0.95, BS = 63%; Figure 2). TAXONOMIC STATUS OF THE GENUS ALIELLA Phytotaxa 148 (1) © 2013 Magnolia Press • 5 IR Phagnalon darvazicum Phagnalon calycinum NA+HAT Phagnalon graecum CE-MED CV Phagnalon melanoleucum Phagnalon metlesicsii C-MED+CI Phagnalon persicum CI+NA Phagnalon pygmaeum 1.00 93 CR Phagnalon purpurascens CI+NA Phagnalon rupestre MAC+MED+IT Phagnalon saxatile WC-MED Phagnalon umbelliforme CI Phagnalon viridifolium YE Ph. latifolium pop 1 C-HAT + AAT Ph. latifolium pop 2 C-HAT + AAT Phagnalon bicolor HAT+AAT Phagnalon harazianum YE Phagnalon woodii YE SA Phagnalon barbeyanum 0.94 68 SA+ET+WAS Phagnalon schweinfurthii 0.79 YE+ET Phagnalon stenolepis - 0.95 C HIGH ATLAS: IMI N OU AKKA Aliella iminouakensis pop 1 63 0.93 50 0.75 - Phagnalon kotschyi E-MED+WAS Phagnalon pycnophyllon W-AS Phagnalon niveum 1.00 WC-AS Phagnalon sordidum WC-MED 72 Phagnalon sinaicum 0.90 59 ET Phagnalon phagnaloides YE Phagnalon stenolepis var. abdulazizianum Aliella platyphylla pop 1 0.83 74 0.99 - OUKAIMEDDEN AREA Aliella ballii subsp. ballii pop 3 RERAYA VALLEY Aliella platyphylla pop 2 C HIGH ATLAS: DJEBEL BOU OURIOUL 74 Aliella embergeri pop 1 Aliella embergeri pop 2 Aliella ballii subsp. nitida pop 4 71 OUKAIMEDDEN AREA Aliella ballii subsp. ballii pop 1 0.98 0.97 SI+AP Aliella iminouakensis pop 2 C HIGH ATLAS: JAFFAR CIRQUE C HIGH ATLAS: M GOUN AREA TICHKA AREA: W HIGH ATLAS C HIGH ATLAS: M GOUN AREA 0.98 93 Athrixia fontana 1.00 99 Anisothrix kuntzei Pentatrichia petrosa OUTGROUP Athrixia angustissima 0.1 FIGURE 2. Consensus phylogram of the post-burn-in trees resulting from Bayesian MCMC analysis of the combined ycf3-trnS and trnT-trnL data sets. Numbers above the branches are Bayesian clade credibilities (posterior probability, PP); numbers below the branches indicate parsimony bootstrap percentages (BS). Regional acronyms: AS Asian, AAT AntiAtlas, AP Arabian Peninsula, C central, CI Canary Islands, CR Crete, CV Cape Verde, E Eastern, ET Ethiopia, HAT High Atlas, IR Iran, IT Irano-Turanian, MAC Macaronesian, MAT Middle Atlas, MED Mediterranean, NA North African, SA Saharo-Arabian, SI Sinai, W western, YE Yemen. Other abbreviations: pop = population. Morphological features of Aliella and Phagnalon We provide comparative descriptions of Aliella and Phagnalon as well as comments on the variability and usefulness of both diagnostic and other morphological characters. Box plots representing variability of diagnostic characters of Aliella are shown in figures 3–9. 6 • Phytotaxa 148 (1) © 2013 Magnolia Press MONTES-MORENO ET AL. General habit:—All Aliella species have robust woody stems of (2.3)5–11(33) cm × (4.5)6.5–8.5(28) mm. The stem is branched into several procumbent, vegetative stems, usually marked with the scars of older leaves. Capitula and leaves are produced from numerous robust brachyblasts arising from stems in A. ballii, A. embergeri and A. iminouakensis. In A. platyphylla, capitula and leaves are produced directly from vegetative stems. All Phagnalon species are also woody perennials, but they are usually branched with numerous, ascending or procumbent (in some taxa), vegetative stems measuring (0.8)7–26(51) cm × (1)1.5–3(10) mm, and these are usually narrower than those of taxa included in Aliella. Capitula and leaves are produced directly from the stems, not from brachyblasts, in all Phagnalon species except for Ph. latifolium and Ph. pygmaeum (Sieber 1823: 322) Greuter (1975: 23), the leaves and capitula of which are produced from robust brachyblasts arising from stems. Leaves:—All Aliella species have imbricate leaves, concolourous in A. ballii and A. embergeri and discolourous in A. iminouakensis and A. platyphylla. Leaves may be petiolate (A. embergeri, A. iminouakensis and A. platyphylla) to decurrent (A. ballii). Leaf shape, size and margin are rather fixed for some species. For example, they are lyrate with lobulate and revolute margins in A. iminouakensis. However, leaf shape, size and margin are variable in A. ballii, A. embergeri and A. platyphylla. Phagnalon species also have imbricate leaves that are generally discolourous and sessile to decurrent. Leaves are linear to lanceolate or oblanceolate to spathulate, and the margins are generally revolute, ocasionally undulate and entire to dentate. Leaf shape and size are rather fixed in some species (e.g. Ph. stenolepis Chiovenda (1911: 71)—lanceolate and dentate; Ph. purpurascens Schultz Bipontinus in Webb & Berthelot (1844: 212) and Ph. sordidum (Linnaeus 1753: 857) Reichenbach (1831–1832: 224)—linear and entire), but variable in the others (e.g. Ph. rupestre (Linnaeus 1767: 113) Candolle (1836: 396)and Ph. saxatile (Linnaeus 1753: 857) Cassini 1819:174). Indumentum:—Hairs found on leaves, stems and bracts correspond to three types. The first one is a long, cylindrical, multicellular, biseriate glandular hair, typically consisting of 4–7 cells per series in the Compositae (Freire & Katinas 1995), and is present on the outside of corolla lobes, throughout the corolla tube and leaves, bracts, peduncles and receptacles. In all Aliella species these hairs consist of 4–12 cells per series and are (53.32)80–120(226.39) µm long. In Phagnalon, they consist of 5–9 cells per series and are 60– 481.08 µm long (Figure 10; A–D). The second one is an eglandular, filiform twin hair, composed of two parallel cells and a third basal cell (Hess 1938, Anderberg 1991, Freire & Katinas 1995). These hairs are found on the surface of the achenes of both Aliella and Phagnalon (Figure 10; E–F). Finally, the third type is an eglandular hair composed of a single short, basal cell and a very long terminal cell. This hair corresponds to hair-type “B” of Drury & Watson (1966) (Figure 10; G–H) and has been found in both Aliella and Phagnalon on the adaxial and abaxial surfaces of leaves, on stems and peduncles and on the adaxial surface of bracts. Bracts on the peduncle:—The peduncle bears one capitulum in all species of Aliella, and this is subtended by (0)3–9(25) bracts measuring (0.9)3–5(12) mm × (0.4)1–2(3.4) mm. The peduncles may be densely to scarcely bracteate throughout their length in A. platyphylla and A. ballii, these having (3)6–12(25) and (0)3–8(19) bracts, respectively. In both species, there is a wide range of variability, and the shape and size of the bracts are similar to those of involucral bracts. Conversely, the peduncles of A. embergeri and A. iminouakensis are provided with few bracts: 1–3 or (0)1–2(5) bracts, respectively. When present, the bracts of the peduncle are always smaller than the involucral ones. In Phagnalon species, the peduncle may be branched, bearing one to several capitula. The bracts on the peduncle are usually few, always smaller than the involucral bracts, and cartilaginous (Figures 3–4). The peduncle bears (0)1–3(12) bracts measuring (0.2)1.2– 1.8(9.5) × (0.2)0.4–0.6(3) mm. However, specimens of Ph. darvazicum Krascheninnikov (1936: 343) and Ph. kotschyi Schultz Bipontinus ex Boissier (1875: 221) with densely bracteate peduncles bearing 7–12 bracts have been found. TAXONOMIC STATUS OF THE GENUS ALIELLA Phytotaxa 148 (1) © 2013 Magnolia Press • 7 FIGURES 3–9. Box plots representing the variability of diagnostic characters of Aliella. Fig. 3. Number of bracts on the peduncle. Fig. 4. Ratio of length of the bracts on the peduncle/Length of the middle bracts. Fig. 5. Female corolla width. Fig. 6. Length of sterile appendages of anthers. Fig. 7. Ratio length of cilia/bristle width of the base of the bristle. Fig. 8. Ratio length of cilia/bristle width of the middle of the bristle. Fig. 9. Ratio length of cilia/bristle width of the apex of the bristle. 8 • Phytotaxa 148 (1) © 2013 Magnolia Press MONTES-MORENO ET AL. FIGURE 10. Indumentum. (A–B). Multicellular, biseriate glandular hairs on corolla. A. Aliella iminouakensis (Sáez 6182, Sáez pers. herb.), scale bar = 50 µm. B. Aliella embergeri (Maire 602, MPU), scale bar = 50 µm. (C–D). Multicellular, biseriate glandular hairs on the receptacle. C. Aliella iminouakensis (Sáez 6182, Sáez pers. herb.), scale bar = 500 µm. D. Aliella iminouakensis (Sáez 6182, Sáez pers. herb.), scale bar = 20 µm. (E–F). Achene and eglandular, filiform twin hairs. E. Aliella iminouakensis (Sáez 6182, Sáez pers. herb.), scale bar = 500 µm. F. Achene surface and eglandular, filiform twin hairs. Aliella platyphylla (Fernández Casas 3329, MA), scale bar = 100 µm. (G–H). Eglandular hair, hair type “B” of Drury & Watson (1966). G. Phagnalon schweinfurthii (Hedge et al. W7790, E), scale bar = 100 µm. H. Phagnalon acuminatum (Bowes Lyon 631, E). TAXONOMIC STATUS OF THE GENUS ALIELLA Phytotaxa 148 (1) © 2013 Magnolia Press • 9 Peduncles:—In Aliella species the peduncles are (0.1)1–2.5(5) cm long. In Phagnalon, the peduncles are slightly longer, being (0.4)5–8(20) cm long. As the measures are overlapping, this morphological character is not useful to distinguish Aliella and Phagnalon. Capitula:—In Aliella species the capitula are heterogamous, solitary, terminal, disciform, (4.2)7.8– 10(15) × (3.1)4–8(13) mm and campanulate, generally with female florets outnumbering hermaphrodite florets, with the exception of A. iminouakensis, where hermaphrodite florets can outnumber the female florets. In all Phagnalon species the capitula are also heterogamous, disciform, solitary or arranged in synflorescences, (4)7–8.2(14) × 3.8–6.7(11) mm, cylindrical to campanulate, with female florets outnumbering the hermaphrodite florets. Involucral bracts:—All Aliella species have imbricate involucral bracts with flat margins, and these are arranged in several rows. The outermost are tomentose, the middle bracts are tomentose to glabrous, whereas the innermost are usually glabrous to subglabrous in both Aliella and Phagnalon. Bracts, in both genera, are composed of a proximal stereome and a distal lamina, and may vary in texture from entirely papery in A. platyphylla to scarious in A. ballii and herbaceous in A. embergeri and A. iminouakensis. In Phagnalon, the texture of the lamina of the involucral bracts is coriaceous, herbaceous or scarious, and the margins are flat or undulate. FIGURE 11. Female florets. A. Aliella ballii (Sáez 6191, Sáez pers. herb.) B. Phagnalon latifolium (Charpin et al. MAR 206, W). C. Phagnalon stenolepis (Collenette 9135, E), (A–C) scale bar = 1 mm. Waxy cushions on the corolla lobes. D. Phagnalon carolipaui (Sauvage 12374, MA), scale bar = 20 µm. Florets:—The corolla is tubular or more or less campanulate at the apex in both female and hermaphrodite florets of Aliella (Figure 11; A). In Phagnalon, the female florets are narrowly tubular, and the hermaphrodite florets are tubular to more or less campanulate at the apex (Figure 11; B–C). Female florets are 10 • Phytotaxa 148 (1) © 2013 Magnolia Press MONTES-MORENO ET AL. located in the outermost rows, and hermaphrodite florets are distributed at the centre of capitula of both Aliella and Phagnalon. In Aliella, (64)115–160(194) female florets and (36)60–90(137) hermaphrodite florets occur in each capitulum. Female florets measure (3.2)4.4–5(5.9) × (0.2)0.3–0.5(0.8) mm, and hermaphrodite florets measure (2.8)5–5.8(6.5) × (0.2)0.5–0.8(1.5) mm. However, the corolla of the female florets is slightly narrower in Phagnalon than in Aliella (Figure 5). In Phagnalon, female florets measure (2.4)4.4–6(8.6) × (0.075)0.1–0.2(0.57) mm, and hermaphrodite florets measure (3)4.8–6.8(9) × (0.2)0.4–0.55(0.95) mm. In Phagnalon, female florets occur in larger numbers per capitulum than in Aliella, and (15)75–250(627) female florets and (4)20–60(226) hermaphrodite florets per capitulum have been recorded for the former genus. However, differences between Aliella and Phagnalon regarding the number of female and hermaphrodite florets and the width of female florets are not statistically significant. Corolla lobes and waxy cushions:—Corolla lobes are strongly reflexed in both Aliella and Phagnalon. Waxy cushions have been observed on the abaxial surface of all species of Aliella, as well as the following species of Phagnalon: Ph. bicolor Ball (1873: 364), Ph. carolipaui Font Quer (1927:13), Ph. garamantum Maire (1929: 185), Ph. kotschyi, Ph. latifolium, Ph. persicum Boissier (1846: 74), Ph. phagnaloides (Schultz Bipontinus ex Richard 1847: 393) Cufodontis (1966: 1105), Ph. pygmaeum, and Ph. stenolepis (Figure 11; D and Table 2). Stamens:—Ecalcarate stamens are frequently found in Gnaphalieae (Ward et al. 2009). This means that the filament is inserted at the base of the thecae, and the polliniferous part of the anther does not protrude below the insertion point (Galbany-Casals et al. 2004). Stamens are ecalcarate in all Aliella species (Figure 12; A–B, Table 2). The anthers are caudate and distinctly tailed in Aliella, that is, they have distinct sterile appendages below the insertion point (anthers are 2.5–3.2 times longer than the sterile appendages; being (313.04)430–690(807.64) µm long). Stamens are also ecalcarate in all Phagnalon species studied, the sterile appendages ranges (54.36)140–230(490.93) µm long. However, some species have caudate and distinctly tailed anthers, as occurs in Aliella species: Ph. harazianum Deflers (1889: 150), Ph. viridifolium Decaisne in Boissier (1875: 220), Ph. woodii Qaiser & Lack (1985: 6)and Ph. yerrimense Qaiser & Lack (1985: 17) (anthers 2.2–3 times longer than the sterile appendages; being (147.65)260–370(490.93) µm long) (Figure 12; C–D, Table 2). The remaining species also have caudate anthers, but are provided with medium-length tails (anthers 6–10 times longer than the sterile appendages; being (54.36)130–200(309.18) µm long, Figure 12; E– H, Table 2). Therefore, the range of length of the sterile appendages is greater in Aliella than in Phagnalon (Figure 6). Styles:—The style is bifid with two separate stigmatic lines, but there is an apically confluent area in all species of both Aliella and Phagnalon (Figure 13; A–D). The sweeping hairs are distributed apically and extend dorsally in both genera. The style branches are rounded with obtuse, sweeping hairs in A. ballii, A. embergeri and A. platyphylla (Figure 13; A–C), and truncate with acute, sweeping hairs in A. iminouakensis (Figure 13; D). Those of Phagnalon show variation; some species have acute sweeping hairs (for example Ph. persicum, whereas others have obtuse sweeping hairs (Table 2). Achenes:—Achenes of both Aliella and Phagnalon are ovoid to cylindrical, with a scalariform epidermis and a pubescent indumentum, densely covered with equally distributed, acute, filiform, non-myxogenic, twin hairs (Figure 10; E–F). In both genera, 4–5 vascular bundles were observed. In Aliella species achenes measure (0.7)1.1–1.5(1.8) × (0.2)0.3–0.35(0.5) mm. Similarly, in Phagnalon species achenes measure (0.7)1– 1.5(1.7) × (0.125)0.2–0.4(0.5) mm. Pappus:—The pappus of both Aliella and Phagnalon is monomorphic and composed of a single row of free bristles that cohere slightly at the base and have acute apical cells. In Aliella, it consists of a single row of 4–9 free bristles, barbellate from the base to the apex. In Phagnalon, the pappus is formed of (4)5–8(17) bristles, and these are barbellate from the base to the apex in most Phagnalon species. However, basally scabrid bristles were found in some Phagnalon species (Table 2, Figures 7–9). TAXONOMIC STATUS OF THE GENUS ALIELLA Phytotaxa 148 (1) © 2013 Magnolia Press • 11 FIGURE 12. Stamens and sterile basal appendages. (A–B) Aliella platyphylla (Ait Lafkin et al., MA). A. Stamens, scale bar = 200 µm. B. Sterile basal appendages, scale bar = 100 µm. (C–D) Phagnalon yerrimense (Brunt 2422, BM). C. Stamens, scale bar = 500 µm. D. Sterile basal appendages, scale bar = 200 µm. (E–F) Phagnalon sinaicum (Fayed 1384, E). E. Stamens, scale bar = 500 µm. F. Sterile basal appendages, scale bar = 200 µm. (G–H). Phagnalon barbeyanum (Harris? 1688, E). G. Stamens, scale bar = 500 µm. H. Sterile basal appendages, scale bar = 200 µm. 12 • Phytotaxa 148 (1) © 2013 Magnolia Press MONTES-MORENO ET AL. FIGURE 13. Styles. A. Aliella ballii (Sáez 6186, Sáez pers. herb.), scale bar = 200 µm. B. Aliella embergeri (Gómiz 5650, Gómiz pers. herb.), scale bar = 500 µm. C. Aliella platyphylla (Fernández Casas 3329, MA), scale bar = 200 µm. D. Aliella iminouakensis (Sáez 6182, Sáez pers. herb.), scale bar = 100 µm. Discussion Phylogeny Nuclear and chloroplast phylogenies indicate that only two species of Aliella form a monophyletic group (A. ballii and A. embergeri; Figures 1 and 2). Aliella iminouakensis is not nested within this clade and form monophyletic group with other Phagnalon species, according to cpDNA phylogeny. This confirms the paraphyly of Aliella and Phagnalon, as previously suggested by Anderberg (1991). However, Aliella and Phagnalon altogether form a strongly supported monophyletic group (Figures 1 and 2), as previous results have already shown (Montes-Moreno et al. 2010). Aliella ballii subsp. nitida forms a highly supported, monophyletic group with A. ballii subsp. ballii, according to nrDNA phylogeny (PP = 1.00, BS = 99%; Figure 1). This group is also supported by a number of morphological characters: obovate and scarious bracts (silvery white in A. ballii subsp. nitida) and sessile and oblong to linear leaves (densely tomentose in A. ballii subsp. nitida). Morphologically, A. embergeri is close to A. ballii, since both species share lobulate leaves and lanceolate bracts and, in fact, the morphological characters that separate both species are variable. For example, the lobules of the leaves of many herbarium specimens of A. ballii studied are strongly reflexed, so that they appear entire. There is also morphological variability in the shape of the involucral bracts which, in some specimens of A. ballii studied, are lanceolate and acute. Furthermore, on the basis of nrDNA data set phylogeny, A. ballii and A. embergeri are sister taxa (Figure 1). The type of the genus, A. platyphylla, shares morphological characters with A. ballii (densely bracteate peduncles) and A. embergeri (petiolate leaves with lobulate margins). However, it forms only a weakly supported clade with these two species. TAXONOMIC STATUS OF THE GENUS ALIELLA Phytotaxa 148 (1) © 2013 Magnolia Press • 13 TABLE 2. Comparison of micromorphological characters in Phagnalon and Aliella. Base of the bristle TAXON Distribution Middle of the bristle Apex of the bristle Length Width of Ratio Length Width of Ratio Width of Sweeping Length of Waxy Length Width of Ratio the Length the Length of cilia female hairs of sterile cushions of cilia the Length of cilia (Micres) bristle of cilia/ of florets style appendages (Micres) bristle of cilia/ (Micres) bristle (Micres) Width (Micres) cilia/ corolla of stamens (Micres) Width of the Width (mm) (Micres) of the bristle of the bristle bristle Phagnalon acuminatum Irano-Turanian (Afghanistan, Pakistan,Iran) 0.1–0.4 Obtuse 139.26– 197.71 Absent 43.17 16.77 2.57 59.42 19.51 3.05 123.21 20.49 6.01 Phagnalon abyssinicum Eritreo Arabian (Ethiopia) 0.075– 0.175 Obtuse 93.02– 95.23 Absent 44.21 21.98 2.01 38.56 16.05 2.40 98.62 26.53 3.72 Aliella ballii subsp. ballii Morocco: High Atlas 0.2–0.8 Obtuse 390.33– 807.64 Present 35.09 24.25 1.45 54.77 20.48 2.67 114.46 23.17 4.94 Aliella ballii subsp. nitida Morocco: High Unknown Atlas-W Obtuse Unknown Present 60.89 22.98 2.65 91.38 24.93 3.67 82.45 29.93 2.75 Phagnalon barbeyanum Saharo-Arabian 0.1–0.25 (Saudi Arabia, Egypt, Jordan, Iraq) Obtuse 107.11– 265.42 Absent 50.12 32.34 1.55 62.95 27.54 2.29 72.61 31.65 2.29 Phagnalon bicolor North Africa (Morocco) 0.1–0.4 Obtuse 158.06– 270.95 Present 47.26 38.33 1.23 70.18 33.68 2.08 114.71 22.95 5.00 Phagnalon calycinum North Africa (Morocco) 0.1–0.22 Obtuse 113.636– 133.33 Present 45.71 29.96 1.53 56.44 22.41 2.52 135.6 22.58 6.01 Phagnalon carolipaui Morocco: Rif 0.1–0.2 Obtuse 156.3– 250.37 Present 52.69 19.08 2.76 64.83 20.37 3.18 111.73 22.74 4.91 Phagnalon darvazicum Irano-Turanian (Afghanistan, Pakistan,Iran) 0.1–0.2 Obtuse 95.83– 222.33 Absent 28.92 20.9 1.38 48.25 17.58 2.74 119.12 26.21 4.54 Aliella embergeri Morocco: High Atlas 0.45–0.5 Obtuse 477.52– 695.14 Present 47.84 35.45 1.35 32.39 21.88 1.48 103.77 29.48 3.52 Phagnalon graecum Central Eastern Mediterranean 0.1–0.2 Obtuse 150– 192.57 Absent 57.8 42.39 1.36 42.53 34.22 1.24 146.27 48.04 3.05 Phagnalon harazianum Eritreo Arabian (Yemen) 0.1–0.2 Obtuse 147.65– 350.59 Absent 38.29 29.93 1.28 61.32 15.57 3.94 88.39 22.65 3.90 Aliella iminouakensis Morocco: High Central Atlas 0.3–0.5 Acute 482.76– 760.12 Present 33.73 19.13 1.76 35.78 16.31 2.19 98.98 19.85 4.99 Phagnalon kotschyi Eastern Mediterranean + IranoTuranian (Iran, Iraq, Lebanon, Turkey) 0.15–0.4 Obtuse 168.88– 255.81 Present 47.05 42.5 1.11 58.5 17.79 3.29 104.93 12.98 8.08 Phagnalon latifolium North Africa (Morocco) 0.2–0.4 Obtuse 159.09– 173.91 Present 51.51 45.98 1.12 46.05 32.11 1.43 95.8 20.79 4.61 Phagnalon lavranosii Saharo Arabian: Djibouti 0.1–0.2 Obtuse 165.77– 185.85 Absent 34.4 31.25 1.10 50.92 21.95 2.32 115.25 28.84 4.00 Cape Verde (Macaronesia) 0.1–0.25 Obtuse 76.66 Absent 31.69 17.13 1.85 64.66 16.01 4.04 62.84 23.36 2.69 Phagnalon nitidum Saharo Arabian: Egypt, Ethiopia 0.1–0.15 Obtuse 110.25– 199.05 Absent 33.43 29.63 1.13 53.51 21.51 2.49 64.12 21.02 3.05 Phagnalon niveum Irano-Turanian (Afghanistan, China, India, Nepal, Pakistan) 0.1–0.2 Obtuse 150– 205.27 Absent 37.9 18.49 2.05 56.62 11.52 4.91 68.3 19.78 3.45 Phagnalon persicum Irano-Turanian (Iran) 0.1–0.3 Acute 150– 225.41 Present 48.52 23.59 2.06 56.46 20.21 2.79 119.86 23.48 5.10 Phagnalon phagnaloides Eritreo-Arabian 0.1–0.175 (Ethiopia) Obtuse 156.05– 296.15 Present 54.48 19.14 2.85 66.49 17.96 3.70 105.48 16.14 6.54 Aliella platyphylla Morocco: High Atlas-C 0.2–0.8 Obtuse 316.94– 492.77 Present 59.15 26.4 2.24 67.73 29.45 2.30 167.02 27.38 6.10 Phagnalon purpurascens Canary Islands+North Africa (Mediterranean -Macaronesian) 0.1–0.2 Obtuse 116.6– 134.91 Absent 51 26.18 1.95 67.19 22.54 2.98 102.71 19.97 5.14 Phagnalon melanoleucum ...... continued on the next page 14 • Phytotaxa 148 (1) © 2013 Magnolia Press MONTES-MORENO ET AL. TABLE 2 (continued) Base of the bristle TAXON Distribution Middle of the bristle Apex of the bristle Width of Sweeping Length of Waxy Length Width of Ratio Length Width of Ratio Length Width of Ratio the Length female sterile the hairs of cushions of cilia Length of cilia the Length of cilia (Micres) bristle of cilia/ florets appendages style (Micres) bristle of cilia/ (Micres) bristle of (Micres) Width corolla of stamens (Micres) Width (Micres) cilia/ of the (mm) (Micres) of the Width bristle bristle of the bristle Phagnalon pycnophyllon Irano-Turanian (Pakistan, Afghanistan) 0.15–0.3 Acute 66.6–83.33 Absent 35.5 24.54 1.44 56.95 19.14 2.98 101.09 21.04 4.80 Phagnalon pygmaeum Mediterranean (Crete) 0.2–0.57 Obtuse 150– 166.66 Present 38.15 32.86 1.16 76.48 31.93 2.40 90.53 26.72 3.39 Phagnalon quartinianum Eritreo-Arabian (Ethiopia, Eritrea) 0.075– 0.15 Obtuse 178.88– 296.99 Absent 35.61 22.72 1.57 56.55 21.47 2.63 88.14 19.57 4.50 Phagnalon Macaronesian+ 0.1–0.25 rupestre subsp. Mediterranean+ rupestre Irano-Turanian Obtuse 100– 159.59 Absent 47.11 26.2 1.80 58.87 19.82 2.97 128.94 25.62 5.03 Phagnalon rupestre subsp. illyricum Central Mediterranean 0.1–0.2 Obtuse 110.4–200 Absent 47.65 25.31 1.88 55.35 18.32 3.02 84.1 20.92 4.02 Phagnalon saxatile Western Central Mediterranean 0.1–0.2 Obtuse 150–240 Absent 53.58 28.87 1.86 70.12 28.64 2.45 111.61 29.6 3.77 Phagnalon schweinfurthii var. androssovii Irano-Turanian 0.1–0.2 Obtuse 131.42– 163.81 Absent 52.62 26.87 1.96 63.55 25.88 2.46 75.77 19.43 3.90 Phagnalon schweinfurthii var. schweinfurthii SaharoArabian: Sudan 0.1–0.2 Obtuse 123.81– 150 Absent 34.27 34.27 1.00 57.26 17.42 3.29 103.16 18.1 5.70 Phagnalon sinaicum Saharo-Arabian (Egypt, Arabian Peninsula) + Hoggar 0.1–0.2 Obtuse 131.56– 201.22 Present 46.79 42.38 1.10 59.24 33.14 1.79 107.26 28.43 3.77 Phagnalon sordidum Western Central Mediterranean 0.1–0.2 Obtuse 60–100 Absent 53.2 26.4 2.02 48.87 28.14 1.74 74.51 26.58 2.80 Phagnalon stenolepis Eritreo-Arabian (Ethiopia, Sudan, Saudi Arabia,Yemen) 0.1–0.2 Obtuse 212.38– 296.57 Present 31.8 16.81 1.89 51.86 15.5 3.35 85.45 10.37 8.24 Macaronesian (Canary Islands) 0.1–0.2 Obtuse 95.29– 168.68 Absent 45.66 19.09 2.39 42.46 15.49 2.74 68.12 17.05 4.00 Phagnalon viridifolium Eritreo-Arabian (Saudi Arabia, Oman) 0.1–0.3 Obtuse 163– 468.39 Absent 49.25 42.86 1.15 49.78 30.01 1.66 84.41 18.56 4.55 Phagnalon woodii Eritreo Arabian (Yemen) 0.15– 0.225 Obtuse 268.99– 490.93 Absent 45.19 20.83 2.17 41.54 18.05 2.30 49.52 30.28 1.64 Phagnalon yerrimense Eritreo Arabian (Yemen) 0.1–0.2 Obtuse 227.96– 471.76 Absent 59.7 56.69 1.01 29.77 24.46 1.22 72.18 34.89 2.07 Phagnalon umbelliforme There are incongruities between the molecular and morphological data obtained for A. iminouakensis. These incongruities are indicated by the ILD test and the tree topologies, as this taxon does not group with the other Aliella species, according to our phylogenetic results. However, A. iminouakensis displays all the diagnostic characters found in other species of Aliella, as Emberger (in Quézel 1951) and Dobignard (1997) have previously pointed out. Moreover, according to cpDNA data set phylogeny, two haplotypes are present in the populations of A. iminouakensis. The first is similar to Phagnalon haplotypes (population 1, corresponding to type locality, Imi-n-Ouaka), whereas the other is similar to Aliella haplotypes (population 2, corresponding to High Central Atlas, Tizi-n-Tirguist locality). The haplotype of population 1 forms a monophyletic group with other Phagnalon haplotypes of the Irano-Turanian and Saharo-Arabian area, according to cpDNA phylogeny. However, the other haplotype of A. iminouakensis (population 2) forms a monophyletic group with A. ballii subsp. nitida. In the same line, the unresolved position of this taxon, the TAXONOMIC STATUS OF THE GENUS ALIELLA Phytotaxa 148 (1) © 2013 Magnolia Press • 15 disagreement between morphology and molecular data, and the incongruities between nrDNA and cpDNA data sets indicate that A. iminouakensis has a complex diversification history. An ancient hybridization event could be involved in the diversification of A. iminouakensis, as previously proposed by Montes-Moreno et al. (2010). This hybridization event would involve an ancestor of A. iminouakensis as a paternal (nrDNA) donor and an ancestor of Irano-Turanian and Saharo-Arabian Phagnalon (Figure 2) as the maternal (cpDNA) donor. The position of Ph. latifolium based on combined nrDNA and cpDNA phylogeny remains unresolved as in previous results of Montes-Moreno et al. (2010), since it displays intermediate morphology between that of Aliella and Phagnalon (thick stems, dense caespitose habit and presence of robust brachyblasts arising from stems similar to species included in Aliella). However, Ph. latifolium does not share other diagnostic characters with species included in Aliella, such as the presence of long-tailed anthers, or the wide, tubular to campanulate female florets. Moreover, its relationship to Aliella is not supported by molecular data. Morphology According to the taxonomic evaluation, the diagnostic characters used to describe Aliella by Qaiser & Lack (1986) are variable, and they are shared with Phagnalon species (Figures 3–9). Firstly, not all diagnostic characters which are ascribed to Aliella are shared by all Aliella taxa, as the peduncles are not densely bracteate in A. embergeri and A. iminouakensis. Moreover, the bracts of A. iminouakensis and A. embergeri, like in many species of Phagnalon, are smaller and cartilaginous. Secondly, waxy cushions have also been observed in some Phagnalon species (see results for waxy cushions). Thirdly, some Phagnalon species have distinctly caudate anthers (Ph. harazianum, Ph. viridifolium, Ph. woodii and Ph. yerrimense), while the others display a combination of both caudate anthers and waxy cushions: Ph. bicolor, Ph. carolipaui, Ph. garamantum, Ph. kotschyi, Ph. latifolium, Ph. persicum, Ph. phagnaloides, Ph. pygmaeum and Ph. stenolepis. Finally, the pappus barbellate from base to apex is identical in both Aliella and Phagnalon. Conversely, there are other morphological characters that could contribute to the segregation of Aliella. Female florets are campanulate and wider in Aliella than in Phagnalon, as Qaiser & Lack (1986) have already pointed out (Figure 5), and the range of length of the sterile appendages is greater in Aliella than in Phagnalon (Figure 6). However, these differences are not statistically significant. Moreover, there is a considerable morphological overlap between the remaining diagnostic characters of Aliella and Phagnalon (Figures 3–4; Figures 7–9). The width of stems and robustness of habit also differ between Aliella and Phagnalon, but these characters could be a morphological adaptation in response to herbivore pressure, which is particularly intense throughout the Atlas Mountains. In conclusion, even though there is some evidence for considering Aliella a different group, morphological differences found regarding the length of sterile appendages and the width of female florets are not significant. The remaining diagnostic characters show variability and are shared by both Aliella and Phagnalon. In addition, current molecular evidence indicates paraphyly of both genera, although, together, these form a strongly supported, monophyletic group. Therefore, the species of Aliella do not constitute a natural group considering the mophology and phylogeny, and therefore we propose merging Aliella with Phagnalon without formal taxonomic recognition at any rank. Taxonomic Treatment 1. Phagnalon ballii (Klatt) N. Montes, N. Garcia, Benedí & L. Sáez, comb. nov. Basionym: Helichrysum ballii Klatt (1896: 836). Gnaphalium helichrysoides Ball (1873: 364), nom. illeg., non Gnaphalium helichrysoides Weddell (1856: 146). Phagnalon helichrysoides Cosson ex Maire (1928: 51), nom. illeg. (Art. 52.1). Aliella helichrysoides Qaiser & Lack (1986: 492), nom. illeg. (Art. 52.1). Aliella bracteata Anderberg (1991: 54), nom. inval. (Art. 41.5). Aliella ballii (Klatt) Greuter (2003: 241). Ind. loc.: “Hab. in regione superiori Atlantis Majoris. In jugo Tagherot! circa 3000 m”. Type:—MOROCCO. In regione superiori Atlantis Majoris in jugo Tagherot, 2530–3630 m, 15 May 1871, Ball (lectotype FI!, designated by Qaiser & Lack 1986: 492; isolectotypes K-000274293!, P-00093473 photo!). 16 • Phytotaxa 148 (1) © 2013 Magnolia Press MONTES-MORENO ET AL. Caespitose plants forming dense cushions, tomentose. Stems branched to form several robust, vegetative, procumbent stems up to (3.9)5.5–12(27.5) cm × (4.6)6.5–8.5(28) mm, scarcely to densely leafy. Leaves (1.1)5–10(31.5) × (0.2)1–2(10) mm, arising from numerous robust brachyblasts, decurrent, sessile to petiolate, oblong-linear to oblanceolate or spathulate; margins entire, crenate to lobulate, not mucronate, revolute; concolourous, adaxial surface pubescent, tomentose to densely tomentose (rarely glabrescent) and abaxial surface densely tomentose; eglandular or with scattered, multicellar, biseriate, glandular hairs. Peduncles (0.1)1.95–2.6(4.7) cm long, scarcely or densely bracteate, provided with (0)3–8(19) bracts; bracts (1.5)2.9–3.6(4.5) × (0.4)1–2(2.2) mm, bract lamina ovate, obovate to linear-lanceolate or spathulate, apex obtuse or acute, scarious or coriaceous, tomentose, eglandular. Capitulum arising from numerous robust brachyblasts, (4.2)7.5–9.5(15) × (3.2)3.8–6.5(13) mm, campanulate to poculiform, with (58)84–107(127) bracts, 102–190 female florets and 36–102 hermaphrodite florets. Involucral bracts imbricate in 7–8 rows, lamina scarious to coriaceous, brown or silvery white. Outermost bracts (2)3.35–4.2(5.7) × (0.55)1.4–0.8(2.5) mm, ovate, obovate to lanceolate, obtuse or acute, tomentose on the stereome, eglandular; middle bracts (3.4)5.4–6.7(8.2) × (0.6)1.4–0.8(2.8) mm, oblong, obovate-spathulate to linear-lanceolate, obtuse and acuminate to acute, subglabrous to tomentose on the stereome, eglandular; innermost bracts (4.1)6.2–7(7.8) × (0.25)0.5–0.5(1) mm, oblong, linear to lanceolate, acute, subglabrous to glabrous. Corolla of female florets (3.5)4.8–5(5.5) × (0.2)0.3–0.8 mm; biseriate hairs (53.32)81–109(133.19) µm, of 4–10 cells per series. Stamens with basal sterile appendages (390.33)410–747(807.64) µm. Style branches 266.5–818 µm long, rounded, with obtuse sweeping hairs. Achenes (0.7)1.3–1.5(1.8) × (0.25)0.3–0.4 mm, duplex hairs 253.16– 430.37 µm long, shorter or sligthly longer than the achene width. Pappus consisting of (5)5–7(9) free bristles, (3.6)4.55–5.55(6.5) mm. Distribution:—Western (Tichka tableland, Djebel Ousdim), Central (Ourika and Reraya valleys, Djebel Oukaimeden, Djebel Toubkal, Djebel Bou Ourioul and surroundings) and Eastern areas of the High Atlas (Ari Ayachi and surroundings, Djebel Mesrough, Djebel M’Goun, Zahouia Ahansal) and Middle Atlas (Bou Naceur massif, Djebel Guelb-er-Rahal). Figure 14. Habitat:—Calcareous or siliceous, granitic or sandstone crevices, from 1800–3630 m. Phenology:—Flowers from May to July. Variability:—Phagnalon ballii is a variable species, especially in terms of morphology and size of leaves, bracts and indumentum, as would be expected, since it is the most widely distributed taxon of the group of study in the Atlas mountains. Specimens with shorter leaves have been recognized at varietal level [Phagnalon helichrysoides var. microphyllum (Maire) Emb.], and specimens with lobulate margins and lanceolate bracts have been recognized as separate species [Phagnalon embergeri Humbert & Maire]. However, there are many specimens with intermediate characters to those of Ph. ballii and Ph. embergeri. Specimens from High Central Atlas (Aït Bouguemmez area, see additional specimens examined) have sessile leaves and strongly reflexed, lobulate margins, giving the appearance of entire leaves. In addition, some other specimens identified as Ph. ballii have lobulate margins and lanceolate bracts (Reraya valley, see additional specimens examined). Owing to this variability in morphology, as well as molecular phylogenetic data which indicate that Ph. ballii and Ph. embergeri are sister taxa, we propose that Ph. embergeri be recognized as a subspecies of Ph. ballii. The indumentum on the adaxial surface of the leaves is also variable, and may be glabrescent to pubescent. Specimens possessing a pubescent indumentum on the adaxial surface of the leaves have also been recognized at varietal level [Gnaphalium helichrysoides subvar. lanatum Litard. & Maire]. Based on herbarium and field-collected material, it is proposed that variation merely reflects phenotypic plasticity and does not merit taxonomic recognition. Taxonomic affinities:—see taxonomic affinities under the remaining accepted species. TAXONOMIC STATUS OF THE GENUS ALIELLA Phytotaxa 148 (1) © 2013 Magnolia Press • 17 1a. Phagnalon ballii subsp. ballii Gnaphalium helichrysoides var. microphyllum Maire (1924: 86). Phagnalon helichrysoides var. microphyllum (Maire) Emberger (1935: 224). Ind. loc.: “Habitat in monte Afougueur Atlantis Majoris, ubi leg. Ibrahim”. Type:— MOROCCO. M. Grand Atlas Djebel Afougueur, 1876, Ibrahim (holotype P, isotypes MPU-001561!, K-00274294!). Gnaphalium helichrysoides subvar. lanatum Litardière & Maire (1924: 12). Ind. loc.: “Hab. in rupibus porphyricis alpinis atlantis majoris: sub jugo Tagherat ditionis Reraya, ad alt. 3.200 m, ubi julio floret”. Type:—MOROCCO. Reraya: rochers porphyriques sur le Tizi-n-Tagherat, 3200 m, 23 July 1922, Maire (lectotype MPU-000290!, designated by Qaiser & Lack 1986: 492 ; isolectotype RAB-41639 photo!). Stems (3.9)5–12.5(27.5) cm × (4.7)5–8(28) mm, scarcely to densely leafy. Cauline leaves (1.1)2–10(31.5) × (0.2)0.8–1.4(4) mm, oblong to linear, decurrent; margins entire to crenate; adaxial surface of leaves generally pubescent (rarely glabrescent), abaxial surface densely tomentose. Peduncles (0.5)1.2–3.2(4.2) cm, densely bracteate, with (5)6–8(19) bracts, bract lamina obovate to spathulate. Capitulum (6.9)7.8–9.5(10.5) × (3.5)3.6–4(13) mm, consisting of (77)93–109(111) bracts, 116–190 female florets and 50–80 hermaphroditic florets. Involucral bracts imbricate in 7–8 rows, lamina scarious, brown. Outermost bracts (2.8)3.5–4.5(5.7) × (1.1)1.35–1.8(2.2) mm, obovate, tomentose, eglandular, aplicate, obtuse; middle bracts (3.4)5.5–7(8.2) × (0.7)1.4–1.9(2.8) mm, obovate to spathulate, obtuse to acuminate; innermost bracts (4.8)6–7(7.8) × (0.25)0.5– 0.7(1) mm, linear to lanceolate, acute. Corolla of female florets (4.1)4.8–5(5.5) × (0.2)0.3–0.8 mm, corolla with biseriate hairs (53.32)81–109(133.19) µm long, of 4–10 cells per series. Illustrations:—Ball (1878: pl. XXIII), as Gnaphalium helichrysoides; Qaiser & Lack (1986: 494), as Aliella helichrysoides. Distribution:—Central area of the High Atlas (Ourika and Reraya valleys, Djebel Oukaimeden, Djebel Toubkal, Djebel Bou Ourioul and surrounding areas). Figure 14. Habitat:—Calcareous, siliceous, granitic or sandstone crevices, from 2300–3630 m. Phenology:—Flowers from May to July. FIGURE 14. Distribution of Phagnalon ballii subsp. ballii, Phagnalon ballii subsp. embergeri, Phagnalon ballii subsp. nitidum, Phagnalon platyphyllum and Phagnalon iminouakense. 18 • Phytotaxa 148 (1) © 2013 Magnolia Press MONTES-MORENO ET AL. Variability:—Phagnalon ballii subsp. ballii is a morphologically variable taxon, especially with regards to indumentum and size of leaves. The specimens with shorter leaves have been recognized at varietal level [Phagnalon helichrysoides var. microphyllum (Maire) Emb.], and plants with an indumentum containing a high density of eglandular hairs have also been recognized as varieties [Gnaphalium helichrysoides subvar. lanatum Litard. & Maire]. However, a detailed study of herbarium specimens and personal observations made by individuals at different field localities did not reveal any constant morphological discontinuities. Consequently, this variability does not merit systematic recognition. Additional specimens examined:—MOROCCO. High Atlas, Tagherot, 2530–3630 m, 15 May 1871, Ball (K 000274293; P 00093473 photo!); Reraya, porphyric rocks in jugo Tagherat, 3200 m, no date, Litardière (MPU); High Atlas, Djebel Afougueur, 1876, Ibrahim (MPU 001561, K 00274294); Reraya: SidiChamarouch near Arroum, rocks, 2300 m, 19 June 1921, Jahandiez 778 & Maire (MPU); Reraya: porphyric rocks, in Tizi-n-Tagherat, 2500 m, 21 July 1922, Maire (MPU); High Central Atlas, Reraya, Tizi-n-Tagherot, porphyric rocks, 3200 m, 23 July 1922, Maire (MPU 000290, RAB 41639 photo!); High Atlas, Mesfioua region, Aouljdid mountain, granitic rocks, 2700 m, 12 July 1924, Maire (MPU); High Atlas, Erdouz mountain above Amismiz, calcareous rocks, 2600 m, 1 July 1925, Maire (MPU); High Atlas, Aouljdid mountain, 2600– 2800 m, 23 July 1926, Maire (MPU); High Atlas, Anremer mountain, calcareous rocks, 3000 m, 27 July 1926, Maire (MPU); High Atlas, Seksaoua, Mount Chaif Jarmelt, granitic rocks, 2700 m, 9 August 1933, anonymous (MPU); Grand Atlas (Seksaoua) granitic rocks, Tizi-n-Ousdim, 2800 m, 10 August 1933, Emberger (MPU); High Atlas, Tizi-Sekoumt, Jashdient, rock, basalt, 2880 m, 23 June 1936, Balls (BM 000889352); High Atlas, Djebel Bou Ourioul above Tizi-n-Tichka, 2500-3000 m, 10 May 1937, ChaworthMusters 236 (BM 000889353); High Central Atlas, cliffs near Isougane-n-Guagouns, 2800 m, 1 August 1938, L. Faurel (MPU); High Atlas, Sidi Chamarouch, 2400 m, 25 August 1951, Polunin 2187 (BM 000889359); High Central Atlas, Tizi-Oukaïmeden, vertical sandstone rocks, 2680 m, 20 July 1973, Davis 5515 (BM 000889357); High Atlas, Jebel Toubkal between Neltner hut & Sidi Chamarouch, 24 July 1973, Davis 55527B (BM 000889358); High Atlas, Oukaïmeden, Jebel Angour, 3200 m, 16 June 1974, Reading Univ. B.M. Expedition 765 (BM 000889356); High Atlas, Oukaïmeden, Jebel Oukaïmeden, 3100 m, no date, Reading Univ. B.M. Expedition 771 (BM 000889355); High Atlas, Jebel Angour, south face, 3300 m, 21 July 1976, Humphries & Humphries 85 (BM 000889354); High Atlas, Oukaïmedene, 2600–2700 m, 21 May 1985, Blanché & al. (MA 340297, SALA 44888); Alto Atlas: Jebel Angour, pr. Oukaimeden, granitic rocks, 31º10’58’’N, 7º50’37’’W, 3000 m, 6 July 1997, Güemes & al. 1672 (MA 594269); High Central Atlas, Tizi-nOu-Addi, 2900 m, 22 July 2004, Sáez 6186 (Sáez pers. herb.); High Central Atlas, Jbel Angour, 3450 m, 26 July 2004, Sáez 6191 (Sáez pers. herb.); High Central Atlas, Oukaïmeden, between Tizi-n-Ou-Addi and Jebel Angour, porphyric southern rock crevices towards Bou Iguenouane, 3000 m, 5 September 2007, MontesMoreno & Rodríguez-Gómez (BC); High Central Atlas, Reraya valley, between Sidi Chamarouch and Netlner Hut, rock crevices towards south west, 2400 m, 6 September 2007, Montes-Moreno & Rodríguez-Gómez (BC). 1b. Phagnalon ballii subsp. embergeri (Humbert & Maire) N. Montes, N. Garcia, Benedí & L. Sáez, comb. nov. Basionym: Phagnalon embergeri Humbert & Maire (1928: 52). Aliella embergeri (Humbert & Maire) Qaiser & Lack (1986: 493). Ind. loc.: “Hab. in rupibus calcareis excelsis Atlantis Majoris nec non Atlantis Medii, maio et junio florens: in monte Mesrouh prope Gourrama (Humbert, 1926), ad alt. 2700 m; in monte Guelb-er-Rahal supra Reggou, ad. alt. 2300 m (Emberger & Maire 1927)”. Type:—MOROCCO. In Atlantis Medii montibus Guelb-erRahal, 2300 m, 25 June 1927, Maire (lectotype MPU-001885!, designated here; isolectotypes MPU-010699!, P0093472 photo!). Montis Aït Mesrough, rochers calcaires 2500-2700 m, June 1926, H. Humbert & Maire (syntype MPU-001886 photo!, P-0093471 photo!). Phagnalon lepineyi Emberger (1935: 224) nom. inval. (Art. 39.1). Ph. embergeri subsp. lepineyi Maire (1941: 1142), nom. inval. (Art. 39.1). Ph. embergeri var. genuinum Maire (1941: 1142), nom. inval. (Art. 24.3). TAXONOMIC STATUS OF THE GENUS ALIELLA Phytotaxa 148 (1) © 2013 Magnolia Press • 19 Stems (5.6)5.85–9.1(9) cm × (5.5)7–10.5(12) mm, scarcely to densely leafy. Leaves (2.5)4–14(19) × (0.5)2– 3.5(10) mm, generally oblanceolate to spathulate, sometimes oblong, petiolate; margins lobulate, revolute, sometimes appearing entire; adaxial surface tomentose, abaxial surface densely tomentose. Peduncles (1.2)1.9–3.2(4.7) cm, scarcely bracteate with 1–3 bracts, bract lamina linear to lanceolate, apex acute, coriaceous. Capitulum 10–15 × 3.2–4.1 mm with 127 bracts, 102 female florets and 36–50 hermaphrodite florets. Involucral bracts imbricate in 8 rows, lamina coriaceous, brown. Outermost bracts (2.5)3.25–4.2(4.6) × (0.55)0.7–0.8(1) mm, lanceolate, acute; middle bracts (4.9)5.4–6.7(7.0) × (0.6)0.7–0.8(0.9) mm, linear to lanceolate, acute; innermost bracts (5.9)6.2–7.5(7.7) × (0.3)0.35–0.5(0.55) mm, linear, acute. Corolla of female florets 3.5–4 × 0.45–0.5 mm, corolla with hairs biseriate (70.48)81–109(110.55) µm, with (6)7–8(9) cells per series. Distribution:—Eastern areas of the High Atlas (Ari Ayachi and surroundings, Djebel Mesrough, Djebel M’Goun, Zahouia Ahansal) and Middle Atlas (Bou Naceur massif, Djebel Guelb-er-Rahal). Figure 14. Habitat:—Calcareous rock crevices 1850–3300 m. Phenology:—Flowers from May to July. Variability:—Phagnalon ballii subsp. embergeri is a variable taxon, particularly in its leaf shape, size and margin. Specimens with tomentose, longer and wider leaves have been found in eastern areas of the High Atlas: Zahouia Ahansal and Ari Ayachi. Specimens found at higher altitudes are smaller in habit with reduced leaves; the leaf margins are strongly revolute and appear to be entire, a detail which may be an adaptation to the extreme habitat of these plants. This morphological adaptation to high altitude habitats has also been documented for other species (Dobignard 1997). Additional specimens examined:—MOROCCO. Montis Aït Mesrough, calcareous rock crevices, 2500– 2700 m, June 1926, Humbert & Maire (MPU 001886 photo!, P 0093471 photo!); Middle Atlas, Guelb-erRahal, 2300 m, 25 June 1927, Maire (MPU 001885, MPU 010699, P 0093472 photo!) ; Eastern High Atlas, Ghat, Tizi-n-Mallal, calcareous rocks, 3200 m, 29 June 1931, anonymous (MPU); Zaouia Ahansal: cliffs of Akka-n-Tazzert, 1850 m, 16 July 1934, Emberger (MPU 006148); High Atlas, near Imilchil, 2300 m, 26 June 1939, Maire 541 (MPU Maire); M’Goun, 2800 m, no date, anonymous (MPU Maire); High Atlas, Tizi-nTighoughizin, calcareous rocks, 2600 m, 26 June 1939, Maire 602 (MPU s.n.); Bou Naceur, 3000 m, July 1953, Guinet & al. (MPU Maire); Ari Ayachi, vanaf Tagouilelt, 2200 m, 23 June 1954, Van Steenis 19478 (BM 000889361, RNG 122005); Bou Naceur, 2700 m, July 1954, Emberger (MPU Maire); High Atlas, Ayachi facing cliffs S of Cirque du Jaffar, 20 km from Midelt, 2211 m, 22 June 1974, B.M. Expedition 963 (BM 000889360, RNG s.n.); High Central Atlas, Gorge at the entrance of the Cirque of Jaffar (Midelt), 9 May 2000, Gómiz 5650 (Gómiz pers. herb.); High Atlas, M’Goun, Oumskik pass to Plateau of Tarkeddit, calcareous rock crevices, 3300 m, 14 July 2004, Galbany & al. (BCN); High Central Atlas: near from Agoutí, between Agori pass towards Plateau of Tarkeddit, calcareous rock crevices, 3300 m, 9 September 2007, Montes-Moreno & Rodríguez-Gómez (BC). 1c. Phagnalon ballii subsp. nitidum (Emb.) N. Montes-Moreno, N. Garcia, Benedí & L. Sáez, comb. nov. Basionym: Phagnalon helichrysoides var. nitidum Emberger (1935: 224). Aliella helichrysoides subsp. nitida (Emb.) Qaiser & Lack (1986: 493). Phagnalon helichrysoides subsp. nitidum (Emb.) Ibn Tattou in Fennane & Ibn Tattou (1998: 46). Aliella ballii subsp. nitida (Emb.) Greuter (2003: 241). Ind. loc.: “Hab. in rupibus graniticis in jugo Ousdim (Seksaoua)”. Type:—MOROCCO. Grand Atlas (Seksaoua) rochers graniticis du Tizi-n-Ousdim, 2800 m, 10 August 1933, Emberger (lectotype MPU-006147!, designated by Qaiser & Lack 1986: 493; isolectotypes BC84882!, MPU010683 photo!, RAB-044241 photo!). Stems 3.5–7 cm × 4.6–7 mm, densely leafy. Leaves (3.5)4.2–4.5(4.7) × (1.2)1.3–1.5 mm, oblong to linear, sessile; margins entire and revolute, adaxial and abaxial surface of leaves densely tomentose. Peduncles 0.1– 0.2(0.6) cm, scarcely bracteate with 0–1 bracts, bract lamina ovate to obovate, obtuse or acute, scarious. Capitulum 4.2–6.8 × 3.5–5 mm, with 58–65 bracts. Involucral bracts imbricate in 7 rows, bract lamina scarious, silvery white. Outermost bracts (2)2.4–2.5(2.7) × (1)1.8–2.1(2.5) mm, ovate to obovate, obtuse or acute; middle bracts (4)4.2–4.6(4.8) × 1.5–2(2.3) mm, oblong to obovate, obtuse; innermost bracts (4.1) 4.2– 20 • Phytotaxa 148 (1) © 2013 Magnolia Press MONTES-MORENO ET AL. 5.9 × (0.5)0.6–0.7(0.8) mm, oblong to lanceolate, acute. Corolla with biseriate hairs 56.05–83.34 µm long, with 5–6 cells per series. Distribution:—Western area of the High Atlas (Tichka tableland, Djebel Ousdim, Djebel Moulay Ali). Figure 14. Habitat:—Granitic rock crevices at 2600–2800 m. Phenology:—Flowers from August to September. Variability:—Phagnalon ballii subsp. nitidum is not a very variable taxon. It has a consistently short and dense habit, short peduncles and leaves, the involucral bracts scarious, always silvery white. Populations with this constant combination of morphological characters are geographically circumscribed on the Western area of the High Atlas, and therefore, it is concluded that these morphological discontinuities deserve systematic recognition at subspecific level. Additional specimens examined:—MOROCCO. High Atlas, Seksaoua, granitic rocks of plateau of Tichka, 2400 m, 8 August 1933, Emberger (BC 84882); Seksaoua, Ousdim mountain, granitic rocks, 2800 m, 10 August 1933, Emberger (BC 84882, MPU 006147, MPU010683 photo!, RAB 044241 photo!); High Occidental Atlas: Assif-n-Aït Tament, near Ida Ou Gailal, Moulay Ali, ascending from Tizi-Igourramene, between granitic rock crevices, 2600 m, 3 September 2007, Montes-Moreno, Rodríguez-Gómez & Muhammad (BC). 2. Phagnalon platyphyllum (Maire) Maire (1928: 52). Gnaphalium helichrysoides var. platyphyllum Maire (1924: 85). Aliella platyphylla (Maire) Qaiser & Lack (1986: 490). Ind. loc.: “Ourika, rochers granitiques audessus de Iabassen, 2400–2600 m, Maire; 1921; Reraya, cirque d’Around, rochers porphyriques du versant W. de l’Imserdine, 2500 m (R. de Litardière, 1923)”. Type:—MOROCCO. Reraya, cirque d’Arround, vers W du Djebel Imserdine, rochers porphyriques, 9 August 1923, Litardière (lectotype MPU-s.n.!, designated here; isolectotypes P-00093475 photo!, RAB-s.n. photo!). Grand Atlas, Ourika: rochers granitiques entre Abessem et le Tizi-n-Tachdirt, 11 July 1921, Maire (syntype MPU-001559!, P-00084148 photo!, P-00093474 photo!, RABs.n. photo!). Illustration: Qaiser & Lack (1986: 491), sub Aliella platyphylla. Decumbent, chasmophytic, tomentose perennials branching into several vegetative stems (2.3)6–14(33) cm × 5–12 mm. Leaves (2.1)11–26(40) × (2.8)5–14(19) mm, subopposite to alternate, petiolate, margins dentate to rarely lobulate, mucronate and flat, usually obovate (rarely spathulate), discolourous, adaxial surface green and abaxial surface white to greyish, glabrescent to tomentose on adaxial surface and densely tomentose on the abaxial surface, eglandular. Peduncles (0.4)1.1–1.7(5) cm, densely bracteate up to the capitula, provided with (3)6–12(25) bracts; bracts (2.3)4–5(7.5) × (0.7)1.4–2.5(3.4) mm; bract lamina obovate to lanceolate, papery, tomentose, eglandular, acuminate. Capitulum arising from the vegetative stems, (6.7)8.2–9.9(12.5) × (3.5)4.5–8.5(11) mm, campanulate to poculiform, with (59)66–109(127) bracts, (94)120–150(194) female florets and (59)73–90(91) hermaphrodite florets. Involucral bracts imbricate in 6–8 rows, lamina papery. Outermost bracts (2.4)4.6–5.8(7) × (1.1)2.1–2.7(3.2) mm, obovate, acuminate, acute to obtuse, tomentose, eglandular, appressed; middle bracts (4.4)5.4–6.5(8.5) × (0.9)1.6–2.3(3.7) mm, obovate-spathulate to lanceolate, acute to acuminate, glabrescent, eglandular; innermost bracts (3.9)5–6(7.5) × (0.3)0.7–0.9(1.3) mm, linear, glabrous. Corolla of female florets (3.3)4.5–5.1(5.9) × (0.2)0.3–0.5(0.8) mm, with biseriate hairs (62.74)100–165(226.39) µm, of 7–11(12) cells per series. Stamens with sterile basal appendages 316.94– 492.77 µm long. Style branches 327.2–474.4 µm long, truncate to rounded, with obtuse sweeping hairs. Achenes (0.7)1.2–1.5(1.7) × (0.2)0.3–0.4(0.5) mm, duplex hairs 203.39–390.6 µm long, usually shorter than the achene width. Pappus consisting of 5–6 bristles, bristles (3.1)3.9–4.8(5.7) mm. Illustration:—Qaiser & Lack (1986: 491), as Aliella platyphylla. Distribution:—Central area of the High Atlas (Ourika and Reraya valleys, Djebel Oukaimeden, Djebel Toubkal, Djebel Bou Ourioul and surroundings). Figure 14. Habitat:—Chasmophytic on porphyric, granitic and sandstone crevices on vertical cliffs at 2000–3300 m. TAXONOMIC STATUS OF THE GENUS ALIELLA Phytotaxa 148 (1) © 2013 Magnolia Press • 21 Phenology:—Flowers from May to July. Variability:—Phagnalon platyphyllum is a morphologically variable species with regards to leaves, particularly their shape, size and margin. Specimens studied have obovate to spathulate leaves, in some cases, on the same specimen. Leaf size also shows variability. Specimens with shorter and longer leaves have also been studied and, in some cases, short and long leaves are present on the same specimen. Finally, leaves are usually dentate, but the shortest and youngest leaves may be entire. Taxonomic affinities:—Phagnalon platyphyllum is very different from Ph. ballii and Ph. iminouakense, mainly due the presence of dentate and mucronate leaves and the papery bracts. However, it is morphologically close to Ph. ballii subsp. ballii due the presence of densely bracteate peduncles. This affinity is not supported by results obtained from nrDNA and cpDNA phylogenies, as A. platyphylla does not form a strongly supported monophyletic group with Ph. ballii. Additional specimens examined:—MOROCCO. Ourika, granitic rocks above Iebassen, 2400–2600 m, 1921, Maire (MPU); Reraya, cirque d’Around, porphyric rocks, Imserdine, 2500 m, 1923, Litardière (MPU); High Central Atlas, Djebel Bou Ourioul, July 2007, Molero & al. (BCN); Reraya Valley: Cirque d’Arround, towards west of Djebel Imserdine, porphyric rocks, 2500 m, 9 August 1923, Litardière (MPU s.n., MPU 001559, P 00093475 photo!, P 00093474 photo!, RAB photo!); High Atlas, Mesfioua region, granitic rocks towards south, Aouljdid mountain, 2700 m, 12 July 1924, Maire (MPU); Sidi-Chamarouch, porphyric rocks towards South, 2400–2500 m, 21 July 1924, Maire (MPU); Reraya valley, porphyric rocks, Immouzer, 2800– 2900 m, 22 July 1924, Maire (MPU); Grand Atlas: Reraya, porphyric rocks, 2000 m, 29 June 1936, Maire & Balls 2976 (MPU); High Atlas, Tizi-n-Tagalat, Arround, rock faces-sun, 2460 m, 29 June 1936, Balls 2976 (BM 000889362); High Atlas, Djebel Bou Ourioul above Tizi-n-Tichka, 2500-3000 m, 2 May 1937, Chaworth-Musters 210 (BM 000889363); High Atlas, Oukaïmeden, Jebel Angour, 2700 m, 15 June 1974, Reading Univ. B.M. Expedition (BM 000889365); High Atlas, Jebel Angour, rock crevices, 3300 m, 21 July 1976, Humphries & Humphries (BM 000889364); Marrakech: Tazerala cliffs above Oukaïmeden, 2700-2740 m, 5 June 1980, Charpin & al. (MA 244401, SALA 48502); High Atlas, Oukaïmeden, Tazenah, vertical rocks, 2750 m, 2 July 1980, Fernández Casas 3329 (MA257764, GDA-23501; SALA 48502); High Atlas, Oukaïmedene, 2600–2700 m, 27 May 1985, Blanché & al. (MA 299321); High Atlas, 2 km below Oukaïmeden on road to Ourika Valley, 18 July 1989, Ait Lafkin & al. (MA 499433); Marruecos, Alto Atlas, Jbel Angour, near Oukaimeden, 2800 m, granitic rocks, 6 July 1997, Güemes 1642 (MA 594288); High Central Atlas, Tizi-n-Ou-Addi, 3000 m, 22 July 2004, Sáez 6188 (Sáez pers. herb.); High Central Atlas, Adrar Oulaouine, porphyric rocks, 2700 m, 23 July 2004, Sáez 6189 (BCN); High Central Atlas, Oukaïmeden, Tizin-Ou-Addi, silicic rocks towards North, 2900 m, 4 September 2007, Montes-Moreno & Rodríguez-Gómez (BC); High Central Atlas, Reraya Valley, between Sidi Chamarouch and Netler Hut, rock crevices towards North East, 2400 m, 6 September 2007, Montes-Moreno & Rodríguez-Gómez (BC). 3. Phagnalon iminouakense Emberger in Maire (1932: 189). Aliella iminouakensis (Emb.) Dobignard & Jeanmonod (1997: 143). Ind. loc.: “Falaises calcaires du Grand Atlas à Imi-n-Ouaka, 1800 m”. Type:— MOROCCO. Grand Atlas de Demnat: Falaises calcaires d’Imi-n-Ouaka près Tirsal, 1800 m, 29 June 1931, Emberger (lectotype MPU-001098!, designated here). Caespitose, tomentose plants forming dense cushions and branching to form several procumbent vegetative stems (2.5)4.75–6.5(10.7) cm × 4.5–6.5 mm, scarcely to densely leafy. Leaves (3.5)9–20(27.5) × (2)3–5(7) mm, arising from numerous robust brachyblasts, alternate, petiolate; margins revolute, lobulate to lyrate, discolourous, adaxial surface green and abaxial surface white to greyish, glabrescent to tomentose on the abaxial surface and densely tomentose on the adaxial surface, eglandular. Peduncles (0.4)1–2.6(3.5) cm, scarcely bracteate with (0)1–2(5) bracts; bracts (0.9)1.25–9(12) × (0.4)0.5–0.55(0.7) mm; bract lamina linear to lanceolate, coriaceous, tomentose, eglandular, acute. Capitulum arising from robust brachyblasts, (4.7)5.2– 7.7(8) × (3.1)3.3–5(6.4) mm, campanulate, composed of (59)79–117(127) bracts, 64–136 female florets and 85–137 hermaphrodite florets. Involucral bracts imbricate in 7–8 rows, lamina coriaceous. Outermost bracts 22 • Phytotaxa 148 (1) © 2013 Magnolia Press MONTES-MORENO ET AL. (1.5)1.6–2.1(2.6) × (0.5)0.6–0.8(1) mm, lanceolate, acute, tomentose, eglandular; middle bracts (2.4)3.75– 5(6.7) × (0.6)0.7–0.8(1.0) mm, linear to lanceolate, acute, tomentose, eglandular; innermost bracts (4.5)4.9– 5.6(6.3) × (0.2)0.3–0.5(0.55) mm, linear, acute, glabrous. Corolla of female florets (3.2)3.4–4.8(5) × 0.3– 0.45(0.5) mm, with biseriate hairs (91.85)101–131(140.2) μm, with 11 cells per series. Stamens with sterile basal appendages 482.76–760.12 μm long. Style branches 178.33–96.12 μm long, truncate, with acute sweeping hairs. Achenes 0.8–1.5 × (0.2)0.25–0.3(0.4) mm, with 3 vascular bundles, duplex hairs 208.86– 323.53 μm long, shorter or as long as the achene width. Pappus consisting of one row of 5–7 bristles; bristles (3)3.2–4.8(5.3) mm long. Distribution:—Eastern area of the High Atlas (Imi-n-Ouaka near to Djebel Ghat and Djebel MGoun). We found this species growing also in the Tizi-n-Tirguist (High Central Atlas), which is a new chorological record (see additional specimens examined). Figure 14. Habitat:—Grows in calcareous rock crevices at 1800–2900 m (Dobignard 1997). Phenology:—Flowers from June to September. Variability:—Although Phagnalon iminouakense is not a very variable species, it shows variation in the leaf size. Specimens found in the type locality, Gorges of Imi-n-Ouaka, 1800–1950 m, had wider and longer, lyrate to lobulate leaves (3.5)10–20(27.5) × (2)3–5(7) mm. Conversely, specimens found in Tizi-n-Tirguist (M’Goun surroundings, 2642 m) had smaller leaves, which were narrower and shorter (8.5)9.5–11(12) × (2.1)2.2–3(3.6) mm. This morphological adaptation to high altitude habitats has been pointed out by Dobignard (1997) and has been also found in Ph. ballii subsp. embergeri. This morphological polymorphism merely reflects a phenotypic plasticity which should not receive taxonomic recognition. Taxonomic affinities:—Phagnalon iminouakense resembles Ph. ballii subsp. embergeri in the shape of the involucral bracts and leaves, as both taxa have leaves with lobulate margins and acute, scarious, linear to lanceolate involucral bracts. Phagnalon iminouakense shares the rest of diagnostic characters with Ph. ballii and Ph. embergeri. However, it differs from these in having acute sweeping hairs on the stigmatic surface. Additional specimens examined:—MOROCCO. High Central Atlas, Demnate, calcareous cliffs of Imin-Ouaka, near Tirsal, 1800 m, 29 June 1931, Emberger (MPU 001098); High Atlas, Demnate, gorges of Imin-Ouaka, 1800 m, 30 June 1931, Emberger (MPU 006906 photo!); Imi-n-Ouaka, July 1935, Quézel (MPU Maire); High Atlas, Djebel Ghat, cliffs of gorges of Imi n’Ouaka, 1800 m, 3 August 1935, Gattefossé (MPU Maire; BC 86415); Gorge d’Imi-n-Ouaka, 1950 m, 10 July 1936, Balls 3041 (MPU Maire, E 00191203); Imin-Ouaka, no date, Quézel (MPU Maire); M’Goun, Gorges above M’Goun, July 1954, anonymous (MPU Maire); High Central Atlas, between Imi-n-Ouaka and Tirsal, calcareous rocks, 1800 m, 21 July 2004, Sáez 6182 (Sáez pers. herb.); High Central Atlas, Ait Bouguemmez, near Agouti, Tizi-n-Tirguist, calcareous rock crevices, 2642 m, 8 September 2007, Montes-Moreno & Rodríguez-Gómez (BC); High Central Atlas, near Demnate, Gorge of Imi-n-Ouaka, entrance of the gorge in rock crevices, 1800 m, 7 September 2007, MontesMoreno & Rodríguez-Gómez (BC). Acknowledgements The authors thank the curators of all herbaria visited and/or consulted (B, BC, BCN, BM, E, FI, JACA, K, LE, MA, MPU, RAB, RNG, SEV, TFC, W and WU.). We also acknowledge M. Aixart, M. Galbany-Casals, F. Gómiz, O. Hidalgo, J. Molero, J.M. Montserrat, R. Rodríguez-Gómez, A. Romo and M. Salvà for providing material and/or help in field collections. In addition, we also thank M. Galbany-Casals and A. Susanna for helpful comments and again J. Molero for providing the RAB herbarium photographs. This work has been partly financed by the Spanish government (REN2002-04634-C05-01, CGL2007-60781/BOS, CGL200505471-C04-04) and the Catalan government (“Ajuts a grups de recerca consolidats”2009/SGR/00439). N. Montes-Moreno benefited from a FPI (“Formación de Personal Investigador”) predoctoral grant from the “Ministerio de Educación y Ciencia”. TAXONOMIC STATUS OF THE GENUS ALIELLA Phytotaxa 148 (1) © 2013 Magnolia Press • 23 References Anderberg, A.A. (1988) The genus Anisothrix O. 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Academic Press, San Diego, pp. 315–322. 26 • Phytotaxa 148 (1) © 2013 Magnolia Press MONTES-MORENO ET AL. Appendix 1. Material examined from Phagnalon s. str. included in the taxonomic study, voucher information. Phagnalon abyssinicum Sch. Bip. ex A. Rich., ETHIOPIA. Teum Tsch’enna, Scholoda mountain, 5 September 1837, Schimper 251 (W 1889-65825, W 1889-51974, W 1889-278833, W 0006781); Mountains near Adoam, 1852, Schimper & Hohemack? 854 (MPU); Abyssinia, ubr Lotha, 2400 m, 10 May 1854, Schimper (W 0011729); Abyssinia, Choa, no date, Petit (W 0006779, W 0006730); Africa boreal, no date, Petit (W 1889-131939); Auf Bergen 1800-2700 m Meer. Ambea Harres 2250 m, inter meer, 22-?-1862, anonymous (BM 000929703); Near Village of Bachianà (Tucur Dinghiá-Godar), 24 January 1937, anonymous (W 1961-19161). Phagnalon acuminatum Boiss., AFGHANISTAN. Bagram Tal, 1900 m, 4 August 1964, Neubauer 4607 (W 1965-15830); East of Afghanistan: In declibibus australibus jugi Salang, 2300 m, 25 June 1965, Rechinger 31358 (W 1966-2408); South East Afghanistan, Tirin: in pass between lake “Arghndab Reservoir” and Tirin, 1400-1700 m, 23 May 1967, Rechinger 35017 (W 1980-792); East of Afghanistan, Ghazni, Inter Ghazni, and Sariab, near Dasht-I Nawar, substr. granit, 2500 m, 17 July 1967, Rechinger 37133 (W 1980-785); Kabul: Obere Tang-e-Gharu, 1 km unterhalb des Schluchteingangs, 1740 m, 19 May 1970, Anders 3639 (W 1971-6423); Bamian Province, 15 miles South of Doab, road to Bamian, rocky slopes, 1800 m, 26 May 1971, Grey Wilson & Hewer 921 (W 1973-12778, E 00191176); Kapisa: Untereres Panjsher Tal gegenueber Korawa, 1700 m, 24 May 1973, Anders 9784 (W 1980-01745); Kabul: Ostseite des Paghman-Gebirges, Umgebung von Khushak, 2200-2400 m, 8 June 1973, Anders 10146 (W 1980-1744). IRAN. Chitral, dry southern cliff face, 1440 m, 12 May 1958, Bowes Lyon 631 (W 1969-2612, E 00191224); Chitral, Drosh, crevices in dry cliffs, 3 May 1958, Stainton 2363 (E00191223). Phagnalon barbeyanum Asch. & Schweinf., SAUDI ARABIA. 180 km South Weast of Turayf Camp, in basalt, among rocks, 690 m, 18 March 1982, Collenette 3439 (E 00191192); Harrat al Harrah, South of Turayf, Wadi Tarbah, steep rocky bands of wadi, 660 m, 12 March 1988, Collenette 6550 (E 00191212); Harrat al Harrah, South of Turayf, rocky runnel near summit, 900 m, 13 March 1988, Collenette 6561 (E 00191191); EGYPT. In oberen Wadi Dugla (Dégeli) swischen Cair und Sués, 3 May 1879, Schweinfurth 277 (W1879-467); Goreiah Wadi Birched?, 25 August 1900, anonymous (BM 000929708); Hilcohstal una Mayer-Tymartal bei Kairo; 10 April 1904, Keller 22 (WU-04-129/4, BM 000929707); Ain Musa? Bei Kairo, Keller 146 (BM 000929706); Wadi Terrranir ? (Stelwan), 1 April 1945, Davis 10516 (E 00191170); IRAQ. Al Salman, Cliffs in rock of steep valley from plateau to aoasis, 24 March 1960, Harris? 1688 (E 00191172). Phagnalon bicolor Ball, MOROCCO. Agadir: Tizi-n-Tagounit pass, acid vertical rocks, 1800 m, 27 May 1980, Fernández Casas 3231 (MA 282437); Agadir: near Tafraoute, Tizi-Mlil, silicic rocks, 1600 m, 26 May 1985, Fernández Casas 9370 (MA 299344); Agadir: Souk Tleta de Tasserirt (Anti-Atlas), silicic rocks, 1800 m, no date, Charpin & al. (MA 244383). Phagnalon calycinum (Cav.) DC., MOROCCO. Kenitra, Dunes of Mehdiya-Plage, sand, 2 May 1987, Lippert 23126 (W1999-05044); Kenitra, Mehdiya-Plage, sand, 2 May 1987, Podlech 43509 (W2003-13718); Kenitra, Medhdiya-Plage, 2 May 1987, Lippert 23125 (W 1999-5044); Agadir, Atlantic coast, 15 km S Tamanar, 1 May 2002, Staudinger 6797 (W 2004-03365). Phagnalon carolipaui Font Quer, MOROCCO. Above Targuist, Zarkat, 1200 m, not above Bu-Meziat, 1700 m, between 29 May 1927 and 17 June 1927, Font Quer 654 (BC 31481 ; MA 124734); Near Asib de Ketama, 1450 m, 1 July 1929, Font Quer 434 (BC 810468, MA 124733, GDA-5106); Djebel Khesena (= Jesana), 1550 m, 23 June 1930, Font Quer (BC 810479); Tazekka: forest of Bab-Azhar, rocks in O. Rhanef Valley, 15 June 1954, Sauvage 12374 (MA 244791). Phagnalon graecum Boiss. & Heldr., CYPRUS. Limassol, Montes Troodos: in mountain Khionistra between Pano Platres and Prodhromos, 100-1950 m, 7 May 1986, Rechinger 62015 (MA 418873); Auf Felsen amb Buffavento Gebirge, 5-1889, Pichler (WU-04-123/163); Petra tou Romiou, 16 April 1992, Lambinon 92/ 377 (FI 4652/35, MA 563657). Phagnalon harazianum Deflers, YEMEN. By Sana’a Hodeida road side “Monaco” village 91 km from Sana’a, rocky hillside, 2 December 1971, Brunt 2488 (BM 000929696); Bait al Alama, J. Al Mahdad, 20 km W. of Amran, on terrace walls, 3000 m, 26 September 1978, Miller 238 (E 00191190); Wadi Bana, Qal Haqul, basalt rock faces in gorge, 14º11’N, 44º21’E, 2500 m, 19 June 1979, Wood Y1277 (E 00191189); North of Yemen, 27 km SW Sana’a an der Strasse nach Menacha, 2720 m, 5 October 1981, Podlech 36371 (W 2003-04530). Phagnalon kotschyi Sch. Bip. ex Boiss., IRAQ. Mosul (Kurdistan) TAXONOMIC STATUS OF THE GENUS ALIELLA Phytotaxa 148 (1) © 2013 Magnolia Press • 27 near Turkey, prov. Hakari, Sharanish in calcareous mountain A Zakho, 4 July 1957, Rechinger 11524 (WUs.n., MA 418592, E 00191164, W 1960-3914). TURKEY: Hakkari, Cilo Dag, in Diz deresi, 6 August 1954, Davis 24004 (K s.n.). Phagnalon latifolium Maire, MOROCCO. High Atlas, region Mesfioua, Aouljdid mountain, granitic rocks, 2700 m, 12 July 1924, Maire (MPU 001948); Agadir, Souk Tleta de Tasserirt (AntiAtlas), silicic rocks, 1800-1820 m, 27 May 1980, Charpin & al. MAR 206 (W1982-00318); W de Tizi-nTagounit (Tiznit), 4 May 2000, Gómiz (Gómiz pers. herb.); Djebel el Kest, June 2007, Montserrat & al. (BC). Phagnalon lavranosii M. Qaiser & H.W. Lack, DJIBOUTI. Goda Mountains, small steep peaks immediately below and N of Peak Deloncle, 1500 m, 9 April 1974, Lavranos 11141 (E, K 000274284). Phagnalon melanoleucum Webb, CAPE VERDE. Sao Nicolau, Mte. Gordo, East-South East facing slope, 1000-1100 m, 13 January 1994, Kilian 3156 & Leyens (B-100172671); Sao Nicolau, Upper Rib. da Prata, path from Fregatta over the Weast ridge to Praia Blanca, North-North East facing, rocky slopes into Rib. da Prata, 550-650 m, 18 January 1994, Kilian 3248 & Leyens (B-100172674); S. Nicolau, small wet ribera at the foot of Mte. Gordo, 1060 m, 24 January 1995, Leyens CV-95-201 (B-100172678). Phagnalon nitidum Fresen., EGYPT. Inter rupes Sinai, 3 May 1835, Schimper (W1889-102773, W1889-9215, W1889-65824, W1889-278846, W1889186744); Wadi Ashar, mittelegyptische Wüste, arabische Seite, 8 April 1877, Schweinfurth 224 (K 1742/84-9, K 2001/02144-25). UNLOCATED: no locality, no date, Kaiser 273 (W1931-11623). Phagnalon niveum Edgew., PAKISTAN. Dera Ismail Khan: Montes Sulaiman, 80 km East Fort Sandeman, calcareous rocks, Dhana Sar, 21 May 1965, Rechinger 30136 (MA 416843); Kohat Cherat, South West Peshawar, sandy soil, 700-950 m, 27 May 1965, Rechinger 30256 (W 1966-2414). AFGHANISTAN: Nangarhar: between Chigha Serai and Kandai, 16 July 1965, Rechinger 32431a (W 1966-2416). Phagnalon persicum Boiss., IRAN. Lorestan, Tolpayegan, Korssan, 2500-2800 m, 30 June 1969, Iransharhr (W 1971-935); Fars: Karkan: Yassnj (forest), 10 September 1971, Zargani (W 1972-09183); Fars: Bamu Protected Region, Tang-e Chah Mahaki, in crevices, 1800-2000 m, 1 June 1975, Wendelbo & Foroughi 17724 (E 00191198); Goischen Felspatten auf dem Gipfel des Kuh Sacbs Buschom, 30 June 1985, Stapf 1312 (WU). Phagnalon phagnaloides (Sch. Bip. ex A. Rich.) Cufod., ETHIOPIA. In montibus elatis sterilibus ad montm Silke, and in Silke mountain, 2700 m above sea, 9 February 1840, Schimper 685 (BM 000888073); In Abyssinia, Schimper (UPS); Abyssinien Berg Silke, Schimper (B-100097164). Phagnalon pygmaeum (Sieber) Greuter, GREECE. Crete, auf den Sudlichen Abfallen gegen Mavrous Lakous, 21/22 July 1882, Spreitzenhofer (WU); Lasithi, Felsen der alpinen Region des Aphendi Khrislos, 25 July 1904, Dörfler 1059 (WU); Lefka Ori, near Amoutsera hut and water cistern in Amoutsera valley N of Anooli, 1850 m, 30 June 1994, Turland 795 (BM 00098221). Phagnalon purpurascens Sch. Bip., SPAIN. Tenerife, El Bailadero-San Andrés-Santa Cruz, 150-300 m, 18 April 1968, Ehrendorfer & al. (W 1974-13391, W 1974-13768); Tenerife, Montañas de Teno, 100 m, 16 March 1969, Bramwell (E 00191204); Fuerteventura, Mña. Cardones, 31 March 1975, Pérez & Acebes (TFC 4770); Lanzarote, Haría, Malpaís de la Corona, 17 March 1997, Reyes-Betancort (TFC 40435). Phagnalon pycnophyllon Rech. f., PAKISTAN. Loralai, betwen 18-19 miles from Sanyari and Harnai, 14 May 1965, Kazmi 1698 (M 0098770); Baluchistan, Quetta, Loralai to Harnai, Torkan Pass, Top Section of Pass above Dil Kuna village, 1400 m, 14 May 1965, Lamond 1246 (C 4, E 00191217); Baluchistan: Landa, porus in crevices limestone rocks, 2100 m, 23 May 1988, Lace (E 00191214); Quetta, Landra 2100 m, no date, Lace? (E 00191215). Phagnalon quartinianum A. Rich., ETHIOPIA. Teum Tsch’enna, ad declivia regionis australis montis Scholoda, 5 December 1837, Schultz 251 (BM 000929699). ERITREA. Asmara to Arbaroba, 12 October 1952, Scott 200 (BM 000929687); Cusai, Torrente Mai Aini, 1800 m, 25 September 1902, Pappi 1867 (BM 000929701); Lungo il Fiume Mareb presso Debarroa, 1900 m, 8 October 1902, Pappi 440 (BM 0009297000). Phagnalon rupestre (L.) DC., CROATIA. An trocknen Mauern ud Ackerdämmen am Wege nach s. Girolamo, und in verschiedenen andern derlei Orten des südlichen Abhanges des Mariàn-Berges (W 0011679, W 1896-7791, W 0011671). SPAIN. Near Raules, 30SWH5239, rock crevices, 23 June 1976, Castroviejo & Valdés-Bermejo (MA 444604); Gata de Gorgos, path to Font de la Mata, 16 May 2004, Montes-Moreno s. n. (BCN). Phagnalon saxatile (L.) Cass., SPAIN. Cómpeta, Sierra de Almijara, towards the pass, 740-920 m, no date, Cubas, López & Moreno (MA444970); Alelo-Peña larga, 560 m, 5 July 1980, Palasí (MA329978); Montserrat, towards Capella de la Sta. Cova, calcareous rocks, 850 m, 10 July 2004, 28 • Phytotaxa 148 (1) © 2013 Magnolia Press MONTES-MORENO ET AL. Montes-Moreno & Rodríguez-Gómez (BCN). Phagnalon schweinfurthii Sch. Bip. ex Schweinf., PAKISTAN. Peshawar, Tor Khama, 700 m, 29 May 1965, Rechinger 30360 (W 1966-2419). AFGHANISTAN. Herat, between Obeh and Khodja Chisht near latter Pardu Khan, rock crevices, 1600 m, 10 May 1969, Hedge & al. W7790 (E 00191196). IRAN. Tehran, mountains near Amninabad, 15 km a Tehran, 1200-1400 m, 17 May 1974, Rechinger (MA 420246, W 1980-01166); Turkmenistán: Badjyz, 6 May 1977, Botchantzev s. n. (LE). Phagnalon sinaicum Bornm. & Kneuck., EGYPT. Sinai, s.d., Kaiser 124 (W1931-11616). SAUDI ARABIA. Medina, Al Figra area, 55-95 km SW of Medina, 550-1650 m, 3 March 1988, Fayed 1384 (E 00191201); Hema Figra, 60 km West of Madinah near Hutlag’s House, rocky hillside Juniper zone, 1740 m, 22 April 1989, Collenette 7138 (E 00191200). Phagnalon sordidum (L.) Rchb., SPAIN. La Población, above ground, calcareous rocks, 1050-1100 m, 18 July 1973, Montserrat & Villar (JACA 481973); Villaseca, El Barranco, calcareous rocks, 960 m, 2 July 1983, Romero (MA 568542, SALA 39829); Torres del Obispo, near from Jusen, Graus, 730-780 m, 29 May 1987, Montserrat (JACA 0450487). Phagnalon stenolepis Chiov., YEMEN. On a steep rocky bank, Jebel Badaan, Ibb, 2700 m, 10 April 1978, Wood 2308 (E 00191207); Amran to Huth road, 5 km. N of Khamir, 21 May 1983, Miller 3155 (E 00191208); Wadi Adoof, near Turbah. 1700 m, 17 March 1984, Miller & King 5234 (E 00191209). SAUDI ARABIA.View point between Mindala and Jebal Ibrahim, 2100 m, 10 May 1994, Collenette 9135 (E 0092209). Phagnalon umbelliforme DC., SPAIN. La Palma, Fuencaliente, common amongst rocks around Volcan de San Antonio, 350 m, 8 June 1969, Bramwell (E 00191206); Tenerife, Puertito de Güimar, 2 March 1975, Wildpret & Rivas Goday (TFC 4509); La Palma, Colada de Martín, Fuencaliente, 750 m, 6 April 1990, Pérez de Paz & al. (TFC 30025); Costas de Jedey, Barranco de Las Palmas,10 April 1990, Pérez de Paz (TFC 30031). Phagnalon viridifolium Decne. ex Boiss., SAUDI ARABIA. Jebel al Ahmar, a granite massif, 143 km SW of Braydeh, in crevices on summit, 1260 m, 13 February 1980, Collenette 1801 (E 00191199). OMAN. Vicinity of Masjid Mu’alla, steep rocky slope, 1680 m, 10 April 1975, Mandaville Jr. 6426 (BM 000929691); Jabal al-Akhdar: Jabal al-Akhdar: Vicinity of Birkat Sharaf, 2070 m, 13 April 1975, Mandaville Jr. 6545 (BM 000929692); Jabal al-Akhdar, Vicinity of Birkat Sharaf, cliff face, 1373 m, 18 April 1975, Mandaville Jr. 6618 (BM 000929690); Northern Oman, Jabal Akhdar, 12 April 1993, Mc Leish 1724 (E). Phagnalon woodii M. Qaiser & H.W. Lack, YEMEN. IBB, Jabel Tarqa, Jibla, 3000 m, on rocks on the middle summit, 13 October 1972, Wood 72/164 (BM 000929714); Jabal Sabir, near Taizz, amphibole granite, 2500–3000 m, 23 September 1977, Lavranos & Newton 15959 (E 00191202); 45 km from Taizz along road to Ibb, rocky slope, 13º50’N, 44º11’E, 2150 m, 15 November 1995, Thulin & al. 9376 (E 0034581). Phagnalon yerrimense M. Qaiser & H.W. Lack, YEMEN. gorge at South East corner of Yerrim plain, cleft in cliff side of gorge, 20 November 1971, Brunt 2422 (BM 000888069). TAXONOMIC STATUS OF THE GENUS ALIELLA Phytotaxa 148 (1) © 2013 Magnolia Press • 29 APPENDIX 2. Species included in the molecular analysis, voucher information and Genbank accession numbers (ETS, ITS, ycf3-TrnS, TrnT-TrnL). An asterisk indicates sequences previously published. An short dash indicates DNA region not sequenced. Aliella ballii (Klatt) Greuter, Morocco: High Central Atlas, Tizi-n-Ou-Addi, 2900 m, 22 July 2004, Sáez 6186 (Sáez pers. herb.), Population 1, HM245966*; HM246006*; HM246102*; HM246068*. Aliella ballii (Klatt) Greuter, Morocco: High Central Atlas, Jbel Angour, 3450 m, 26 July 2004, Sáez 6191 (Sáez pers. herb.), Population 2, HM245967*; HM246007*;-;-. Aliella ballii (Klatt) Greuter, Morocco: High Central Atlas: Reraya Valley, betwen Sidi Chamarouch and Netler Hut, 2400 m, rock cervices towards South-West, 6 September 2007, Montes-Moreno & Rodríguez-Gómez (BC), Population 3, KC899260; KC899254; KC96131; KC961305. Aliella ballii (Klatt) Greuter subsp. nitida (Emb.) Greuter, Morocco: High Occidental Atlas: Assifn-Aït Tament, near Ida Ou Gailal, Moulay Ali, 2600 m, ascending from Tizi Igourramene, between granitic rock crevices, 3 September 2007, Montes-Moreno & Rodríguez-Gómez (BC), Population 4, KC899261; KC899255; KC961312; KC961306. Aliella embergeri (Humbert & Maire) Qaiser & Lack, Morocco: High Central Atlas, Gorge at the entrance of the Cirque of Jaffar (Midelt), 9 May 2000, Gómiz 5650 (Gómiz pers. herb.), Population 1, HM245968*; HM246008*; HM246103*; HM246069*. Aliella embergeri (Humbert & Maire) Qaiser & Lack, Morocco: High Central Atlas: near from Agoutí, between Agori towards Plateau of Tarkeddit, 3300 m, calcareous rock crevices, 9 September 2007, Montes-Moreno & Rodríguez-Gómez (BC), Population 2, KC899262; KC899256; KC961313; KC961307. Aliella iminouakensis (Emb.) Dobignard & Jeanm., Morocco: between Imi-n-Ouaka and Tirsal, 21 July 2004, Sáez 6182 (Sáez pers. herb.), Population 1, HM245969*; HM246009*; HM246104*; HM246070*. Aliella iminouakensis (Emb.) Dobignard & Jeanm., Morocco: High Central Atlas: Ait Bouguemmez, near from Agoutí, Tizi-n-Tirguist, 2642 m, calcareous rock crevices, 8 September 2007, Montes-Moreno & Rodríguez-Gómez (BC), Population 2, KC899263; KC899257; KC961314; KC961308. Aliella platyphylla (Maire) Qaiser & Lack, Morocco: Tizi-n-Ou-Addi, 22 July 2004, Sáez 6188 (Sáez pers. herb.), Population 1, HM245970*; HM246010*; HM246105*; HM246071*. Aliella platyphylla (Maire) Qaiser & Lack, Morocco: Djebel Bou Ourioul, VII-2007, J. Molero & al. (BCN), Population 2, KC899264; KC899258; KC961315; KC961309. Anisothrix kuntzei O. Hoffm., South Africa: Ladismith District longe berg Range Kogmons Kloof Pass cliffs on Side of Road near Old English fort, 22 September 1996, Bayer & Puttock s. n. (CSIRO, SAF), HM245971*; HM246011*; HM246106*; HM246072*. Athrixia angustissima DC., South Africa: E Cape Province: between Rhodes and Naudesnek, 2 February 2007, Galbany-Casals, Koekemoer & Romo 14419 (BC),-;-; HM246107*; HM246073*. Athrixia fontana MacOwan, South Africa, Mpumalanga Province: Lydenburg district, Mokobulaan Plantations, 16 February 2007, Arrabal, Burrows, Galbany-Casals, Koekemoer, Romo 14587 & Unarine (BC),-;-; HM246108*; HM246074*. Pentatrichia avasmontana Merxm., Namibia: Windhoek Bergland, foothills of Ausberge on reefs and rocks, Halbstrauch, Klein, 1600 m, 21 September 1964, Seydel s.n. (B), HM245972*; HM246012*; –; –;. Pentatrichia petrosa Klatt., South Africa: Klein-Windhoek, mica schist slope, Foothills of the Erosberge, 23 May 1962, Giess 3949 (W), HM245973*; HM246013*; HM246109*; HM246075*. Phagnalon abyssinicum Sch. Bip. ex A. Rich, Ethiopia, Prov. Begemden: Larger Sabra into Tällak Valley of Rocks, 3140 m, 16 March 1966, Sebald 1277 (WU), HM245974*; HM246014*;-;-. Phagnalon acuminatum Boiss., Afghanistan: East Afghanistan, Kabul, 13 May 1967, Rechinger 34484 (MA), HM245975*; HM246015*; –; –;. Phagnalon barbeyanum Asch. & Schweinf., Saudi Arabia: Harrat al Harrah S of Turayf, Jabal Liss, 13 March 1988, Collenette 6561 (E), HM245976*; HM246016*; HM246110*; HM246076*. Phagnalon bicolor Ball, Morocco: About 10 km south of Skhour- Rehamna (Marrakech), 03 April 1999, Gómiz s.n. (Gómiz pers. herb.), HM245977*; HM246017*; HM246111*; HM246077*. Phagnalon calycinum (Cav.) DC., Morocco: About 3 km south of Jorf Lasfar (El Jadida), 30 April 1995, Gómiz s.n. (Gómiz pers. herb.), HM245978*; HM246018*; HM246112*; HM246078*. Phagnalon darvazicum Krasch., Tajikistan: Daphaz., 31 May 1986, Kamelin, Kasparov &. Xanvinov s.n. (LE), HM245979*-HM245980*; HM246020*; HM246113*; HM246079*. Phagnalon graecum Boiss. & Heldr., Italy: Puglia, Foggia, Gargano, Monte San Angelo, 07 July 2002, Aldasoro 3308 (MA), HM245981*; HM246021*; HM246114*; HM246080*. Phagnalon harazianum Deflers, Yemen: Wadi Bana, Qal haqul, 19 June 1979, Wood Y1277 (E), HM245982*-HM245983*; HM246022*; HM246115*; HM246081*. Phagnalon kotschyi Sch. Bip. ex Boiss., Iraq: distr. Mosul (Kurdistan) near Turkey, prov. Hakari, Sharanish in calcareous mountain A Zakho, 4 July 1957, Rechinger 11524 (MA), HM245984*; HM246024*; HM246116*; HM246082*. Phagnalon latifolium Maire, Morocco: W de Tizi -NTagounit (Tiznit), 4 May 2000, Gómiz s. n. (Gómiz pers. herb.), Population 1, HM245985*; HM246025*; 30 • Phytotaxa 148 (1) © 2013 Magnolia Press MONTES-MORENO ET AL. HM246117*; HM246083*. Phagnalon latifolium Maire, Morocco: Djebel el Kest, June 2007, Montserrat & al. (BC), Population 2, KC899265; KC899259; KC961316; KC961310. Phagnalon melanoleucum Webb, Cape Verde Islands: São Nicolau Upper Rib de Prata path from Fregatta over W ridge to Praia Branca, 550560 m, 18 January 1994, Kilian 3248 & Leyens (B), HM245986*; HM246026*; HM246118*; HM246084*. Phagnalon metlesicsii Pignatti, Spain, Lanzarote: San Bartolomé Tomaren, 24 February 1995, Reyes Betancort & León-Arencibia s.n. (TFC), HM245987*; HM246027*; HM246119*; HM246085*. Phagnalon niveum Edgew., Pakistan: Western Pakistan, Dera Ismail Khan, 21 May 1965, Rechinger 30136 (MA), HM245988*; HM246028*; HM246120*; HM246086*. Phagnalon persicum Boiss., Iran, Prov Kerman: Kuh-e Kabr., 08 June 1977, Assadi, Edmonson & Miller 1763 (E), HM245989*; HM246029*; HM246121*; HM246087*. Phagnalon phagnaloides (Sch. Bip. ex A. Rich.) Cufod., Ethiopia: In Abyssinia, Schimper (UPS), HM245990*; HM246030*; HM246122*; HM246088*. Phagnalon purpurascens Sch. Bip., Spain, Gran Canaria: San Nicolás de Tolentino, cliff of Pino Gordo, 31 March 1998, A. Marrero & González Martín s.n. (MA), HM245991*; HM246031*; HM246123*; HM246089*. Phagnalon pycnophyllon Rech. f., Pakistan: Baluchistan: Quetta: Loralai to Harnai, Torkhan Pass, top section of pass above Dil Kuna Village, 1965, Lamond s.n. (E), HM245992*; HM246032*; HM246124*; HM246090*. Phagnalon pygmaeum (Sieber) Greuter, Greece: Creta, Kakovoli, September 2000, ex Mediterranean Agronomic Institute of Chania, HM245993*; HM246033*; HM246125*; HM246091*. Phagnalon quartinianum A. Rich., Ethiopia: Asmara to Arbaroba, 12 October 1952, Scott 200 (BM), HM245994*; HM246034*; –; –;. Phagnalon rupestre (L.) DC., Spain: Gata de Gorgos, path to Font de la Mata, 16 May 2004, Montes-Moreno s. n. (BCN), HM245995*; HM246035*; HM246126*; HM246092*. Phagnalon saxatile (L.) Cass., Spain: Collserola, Carretera de les Aigües to St. Pere Màrtir, 02 May 2004, Montes-Moreno & R. Rodríguez-Gómez (BCN), HM245996*; HM246036*; HM246127*; HM246093*. Phagnalon schweinfurthii Sch. Bip. ex Schweinf., Turkmenistan: Badjyz, 6 May 1977, Botchantzev s. n. (LE), HM245997*; HM246037*; HM246128*; HM246094*. Phagnalon sinaicum Bornm. & Kneuck., Saudi Arabia: Hema Figra, 60 Km W of Madinah near Hublag’s house, 22 April 1989, Collenette 7138 (E), HM245998*; HM246038*; HM246129*; HM246095*. Phagnalon sordidum (L.) Rchb., Spain: Montserrat, path from station to Sta. Cova church, 10 July 2004, Montes-Moreno & R. Rodríguez-Gómez (BCN), HM245999*; HM246039*; HM246130*; HM246096*. Phagnalon stenolepis Chiov., Yemen: Amran to Huth road, 5 km. North of Khamir, 21 May 1983, Miller 3155 (E), HM246001*; HM246041*; HM246132*; HM246098*. Phagnalon stenolepis Chiov. var. abdulazizianum Chaudhary, Saudi Arabia: Mindala & Jabal Ibrahim, 10 May 1994, Collenette 9135 (E), HM246000*; HM246040*; HM246131*; HM246097*. Phagnalon umbelliforme DC., Spain, El Hierro: El Pinar a Restinga, ex Botanical Garden Viera & Clavijo, HM246002*; HM246042*; HM246133*; HM24609*. Phagnalon viridifolium Decne. ex Boiss., Oman: Northern Oman, Jabal Akhdar, 12 April 1993, Mc Leish 1724 (E), HM246003*; HM246043*; HM246134*; HM246100*. Phagnalon woodii Qaiser & Lack, Yemen: Jabal Sabir, near Taizz, amphibole granite, 23 September 1977, Lavranos & Newton 15959 (E), Population 1, HM246004*; HM246044*; –; –;. Phagnalon woodii Qaiser & Lack, Yemen: Ibb, 45 km from Taizz along road to Ibb, 1995, Thulin, Gjebrehiwet & Gifri s.n. (E), Population 2, HM246005*; HM246045*; HM246135*; HM246101*. TAXONOMIC STATUS OF THE GENUS ALIELLA Phytotaxa 148 (1) © 2013 Magnolia Press • 31

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Evaluation of the taxonomic status of the genus Aliella (Compositae, Gnaphalieae): a recircumscription of the genus Phagnalon

Evaluation of the taxonomic status of the genus Aliella (Compositae, Gnaphalieae): a recircumscription of the genus Phagnalon

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