Pseudognaphalium Aldunateoides Back in Gnaphalium (Compositae: Gnaphalieae)

Collectanea Botanica 37: e012 enero-diciembre 2018 ISSN-L: 0010-0730 https://doi.org/10.3989/collectbot.2018.v37.012

Pseudognaphalium aldunateoides back in Gnaphalium (Compositae: Gnaphalieae)

A. ACOSTA-MAINDO1 & M. GALBANY-CASALS1,2

1 Unitat de Botànica, Departament de Biologia Animal, Biologia Vegetal i Ecologia, Facultat de Biociències, Universitat Autònoma de Barcelona, ES-08193 Bellaterra, Spain 2 Sistemática y Evolución de Plantas Vasculares (Universitat Autònoma de Barcelona), Unidad Asociada al CSIC

ORCID iD. A. ACOSTA-MAINDO: http://orcid.org/0000-0002-3619-8181, M. GALBANY-CASALS: http://orcid.org/0000-0002-7267-3330

Author for correspondence: M. Galbany-Casals ([email protected])

Editor: A. Susanna

Received 11 May 2017; accepted 10 December 2017; published online 12 July 2018

Abstract PseudognaPhalium aldunateoides back in gnaPhalium (compositae: Gnaphalieae).— Generic classification of some members of the tribe Gnaphalieae (Compositae) and in particular the generic delimitation of the old genus Gnaphali- um has been long problematic. The genus Pseudognaphalium was first split from Gnaphalium based on morphology and later supported by molecular phylogenies. However, the generic identity of some species remains doubtful. Here we provide a morphological study of Pseudognaphalium aldunateoides, which is compared with the type species of Gnaphalium (Gnaphalium uliginosum) and Pseudognaphalium (Gnaphalium oxyphyllum). We also include the three species in a nuclear ribosomal DNA (nrDNA) and a plastid (cpDNA) molecular phylogeny of the tribe Gnaphalieae. Our results show that P. aldunateoides has a dimorphic pappus, character not present in the genus Pseudognaphalium but characteristic of the genus Gnaphalium. It also shows a stereome with intermediate features between the undivided stereome typical of Gnaphalium and the fenestrated stereome typical of Pseudognaphalium. In the nrDNA and the cpDNA trees, P. aldunateoides is placed within the Gnaphalium s. str. clade, not closely related to Pseudognaphalium. With all these evidence, we conclude that P. aldunateoides is correctly placed in Gnaphalium.

Key words: Asteraceae; ETS; ITS; molecular phylogeny; micromorphology; pappus; rpl32-trnL intergenic spacer; stereome.

Resumen PseudognaPhalium aldunateoides de nuevo bajo gnaPhalium (ComPositae: gnaPhalieae).— La clasificación genérica de varios miembros de la tribu Gnaphalieae (Compositae) y en particular la delimitación genérica del antiguo género Gnaphalium ha sido problemática durante mucho tiempo. El género Pseudognaphalium fue escindido de Gnaphalium por su morfología lo que después fue apoyado por filogenias moleculares. Sin embargo, la identidad genérica de algu- nas especies todavía es dudosa. Aquí aportamos un estudio morfológico de Pseudognaphalium aldunateoides, que es comparado con las especies tipo de los géneros Gnaphalium (Gnaphalium uliginosum) y Pseudognaphalium (Gnapha- lium oxyphyllum). También hemos incluido estas tres especies en filogenias moleculares de la tribu Gnaphalieae basa- das en ADN ribosómico nuclear y ADN cloroplástico. Nuestros resultados muestran que P. aldunateoides tiene el vilano dimórfico, carácter que no está presente en el género Pseudognaphalium pero que, sin embargo, es característico del género Gnaphalium. Además, su estereoma muestra características intermedias entre el estereoma no dividido típico del género Gnaphalium y el estereoma fenestrado típico del género Pseudognaphalium. En ambas filogenias, P. aldunateoides se sitúa dentro del clado Gnaphalium s. str., no próximamente emparentado con Pseudognaphalium. Con todas estas evidencias concluimos que P. aldunateoides se clasifica correctamente dentro deGnaphalium .

Palabras clave: Asteraceae; ETS; ITS; espaciador intergénico rpl32-trnL; estereoma; filogenia molecular; micromor- fología; vilano. 2 A. ACOSTA-MAINDO & M. GALBANY-CASALS

Cómo citar este artículo / Citation Acosta-Maindo, A. & Galbany-Casals, M. 2018. Pseudognaphalium aldunateoides back in Gnaphalium (Compositae: Gnaphalie- ae). Collectanea Botanica 37: e012. https://doi.org/10.3989/collectbot.2018.v37.012

INTRODUCTION pappus bristles monomorphic or slightly dimorphic, with the tips scabrid or, when dimorphic, those of the It has long been known that generic classification of hermaphrodite florets barbellate with clavate cells. some genera of the tribe Gnaphalieae (Asteraceae) Pseudognaphalium was one of the segregated is problematic, specially involving the large and het- genera from the old concept of Gnaphalium. This erogeneous Helichrysum Mill. and Gnaphalium L., genus is based on the Mexican species Gnaphalium as well as some other related genera such as Pseudo- oxyphyllum DC. Hilliard & Burtt (1981) proposed gnaphalium Kirp. (Hilliard & Burtt, 1981; Anderberg, that Pseudognaphalium, which at that moment 1991; Galbany-Casals et al., 2014). Hilliard & Burtt only comprised the type species, should addition- (1981) made important advances in establishing new ally include a number of American, Asian and Af- generic delimitations based on a critical and detailed rican species of Gnaphalium, which had a close morphological study of most controversial genera in morphological resemblance to each other, but had Gnaphalieae. In their work, they proved the value of little affinity withGnaphalium s. str. These authors two features, among others: the nature of the stereome considered Pseudognaphalium morphologically of involucral bracts and of the pappus. more similar to Helichrysum, and particularly to The stereome is the thickened lower part of the Helichrysum foetidum (L.) Moench and related involucral bract, often straw-coloured or green, con- species (Hilliard & Burtt, 1981). Although both trasting with the thin lamina and pellucid margins Gnaphalium and Pseudognaphalium have pistillate (Hilliard & Burtt, 1981). The stereome may be un- florets outnumbering the hermaphroditic ones, they divided in some genera: that is, it appears to be uni- differ in the habit and general appearance (Hilliard formly thickened, though sometimes the median vas- & Burtt, 1981), as well as in details of the invo- cular strand is visible and there may occasionally be lucral bracts. Pseudognaphalium species—except minute, thin streaks in the upper part of the stereome Pseudognaphalium oligandrum (DC.) Hilliard & of some bracts. Alternatively, in other genera, the ste- B. L. Burtt—have a fenestrated stereome and a reome may be fenestrated, i.e. some distinct vascular monomorphic the pappus (Hilliard & Burtt, 1981), strands lie in a translucent area of unthickened cells and these characters are shared with the genus He- sharply distinguishable from the thickened cells of the lichrysum. rest of the stereome (Hilliard & Burtt, 1981). Anderberg (1991), based on cladistical analyses The pappus can be monomorphic, that is that bris- of many morphological characters, also conclud- tles of pistillate and hermaphroditic florets have the ed that Pseudognaphalium had little affinity with same morphology, or alternatively, it can be dimor- Gnaphalium, and formally transferred many spe- phic, and in this case bristles of pistillate and her- cies from Gnaphalium to Pseudognaphalium, ris- maphroditic florets have a different morphology. ing the latter to ca. 90 species. Hilliard & Burtt (1981) proposed a new concept for Recent molecular phylogenies have confirmed the old genus Gnaphalium, which they built around that Pseudognaphalium is more closely related to the type species, Gnaphalium uliginosum L. In this Helichrysum than to Gnaphalium (Smissen et al., sense, the genus comprises woolly annual herbs with 2011; Nie et al., 2013, 2016; Galbany-Casals et al., heterogamous capitula arranged in terminal clusters; 2014). Actually, Helichrysum has been recovered capitula with pistillate florets much more numerous as paraphyletic and it includes Pseudognaphalium than hermaphroditic ones; involucral bracts with the and other genera such as Achyrocline (Less.) DC., stereome undivided or with a few thin streaks; Anaphalis DC. and Humeocline Anderb. (Smissen

Collectanea Botanica 37: e012 (enero-diciembre 2018), e012, ISSN-L: 0010-0730, https://doi.org/10.3989/collectbot.2018.v37.012 Pseudognaphalium aldunateoides back in Gnaphalium (Compositae: Gnaphalieae) 3 et al., 2011; Galbany-Casals et al., 2014; Nie et al., and river margins, from sea level to 3700 m (Freire 2016). They all constitute the HAP clade, which is a et al., 2014a). This species was kept in Gnaphali- major component of the tribe Gnaphalieae (Smissen um by Anderberg (1991) as Gnaphalium alduna- et al., 2011; Galbany-Casals et al., 2014). Gnapha- teoides J. Rémy, but it was recently transferred to lium s. str. is not closely related to the HAP clade, Pseudognaphalium by Monti et al. (2013), who ar- since G. uliginosum, together with other species gued that it has a fenestrated stereome, which is the from the genus, are recovered in a clearly indepen- primary morphological character that differentiates dent clade, closely related to the South African ge- both genera. Although Monti et al. (2013) high- nus Vellereophyton Hilliard & B. L. Burtt in plastid lighted that some characters, such as a slightly di- DNA (cpDNA) based trees (Smissen et al., 2011) or morphic pappus, did not match the typical features to Plecostachys Hilliard & B. L. Burtt and Syncar- of Pseudognaphalium, this was not considered suf- pha DC. in nuclear ribosomal DNA (nrDNA) based ficient evidence to keep the species inGnaphalium . trees (Smissen et al., 2011; Nie et al., 2016). This species has never been included in any mo- Although a notable progress has been made in lecular work, so its phylogenetic position, which the last decades regarding the delimitation of these would clarify its generic identity, is unknown. two genera, some particular species remain contro- The aims of this work are to provide an additional versial. One case is Pseudognaphalium aldunate- morphological study of P. aldunateoides and to in- oides (J. Rémy) C. Monti, N. Bayón & Freire (Fig. fer its phylogenetic position in the tribe Gnaphalieae 1), a species native to Central and South Chile and based on nrDNA and plastid DNA markers in order to West Argentina, where it grows in mossy places to clarify its generic identity.

Figure 1. Pseudognaphalium aldunateoides in its natural habitat in Chile, Región IV Coquimbo, Sotaquí, Quebrada Seca (photograph: M. Galbany).

Collectanea Botanica 37: e012 (enero-diciembre 2018), e012, ISSN-L: 0010-0730, https://doi.org/10.3989/collectbot.2018.v37.012 4 A. ACOSTA-MAINDO & M. GALBANY-CASALS

MATERIALS AND METHODS et al. (2010). In total, we included in the analyses 220 rpl32-trnL sequences, of which three are new Taxon sampling for molecular phylogeny (Appendix 1).

Sequences used in this work include newly gener- Purification of PCR products and sequencing ated sequences and previously published sequences. We have included two specimens of Pseudo- PCR products were purified with Exo-SAP-IT gnaphalium aldunateoides, two specimens of (USB Corp., Cleveland, Ohio, USA). Direct se- Pseudognaphalium oxyphyllum—the type species quencing was conducted at the DNA Sequencing of Pseudognaphalium—and 32 additional Pseudo- Core, CGRC/ICBR of the University of Florida, on gnaphalium species. The data set also contained an ABI 3730xl DNA Analyser (Applied Biosyste- eight different Gnaphalium species, including ms) using a Big Dye Terminator v3.1 kit (Applied two specimens of G. uliginosum, the type species Biosystems, Foster City, CA, USA). of Gnaphalium. Finally, we included a selection of 236 species of other Gnaphalieae genera in an Alignments attempt to represent all main clades recovered in previous phylogenetic works for the tribe (Galba- Nucleotide sequences of the two data sets, nrDNA ny-Casals et al., 2010, 2014; Smissen et al., 2011; and cpDNA, were edited using Chromas v2.5.1 Freire et al., 2014b; Nie et al., 2016). Voucher de- (Technelysium, Tewantin, Australia), Bioedit tails for all samples are shown in Appendix 1. v7.2.5 (Hall, 1999) and MEGA v7.0.14 (Kumar et al., 2016), and aligned visually or with the pro- DNA extraction gram Clustal X v2.1 (Thompson et al., 1997) with subsequent visual inspection and manual correc- Total genomic DNA was isolated following the tion. In the cpDNA data set, a fraction of nucleo- CTAB method of Doyle & Dickson (1987) as mod- tides (from 144 to 652 bp, both included) of high ified by Cullings (1992) and Tel-Zur et al. (1999) variability and ambiguously aligned was manual- from silica-gel-dried leaves collected in the field or ly excluded. Additional ambiguously aligned re- from herbarium material (CONC). gions of the two data sets were detected and excluded with the use of Gblocks v0.91 (Castresana, nrDNA amplification strategies 2000; Talavera & Castresana, 2007). It was used with relaxed conditions in order to preserve as The ITS nrDNA region (ITS1, 5.8S ribosomal much information as possible: “Minimum number gene, ITS2) was amplified using the 17SE forward of sequences for a conserved position” and “Min- and the 26SE reverse primers (Sun et al., 1994). The imum number of sequences for a flank position” profile used for amplification was as described in Gal- were half the number of sequences +1, “Maxi- bany-Casals et al. (2004). The conserved 3′ portion of mum number of contiguous nonconserved posi- the ETS nrDNA region was amplified using the for- tions” was 10, “Minimum length of a block” was ward primer Ast-1 and the reverse primer 18S-ETS 5, and “Allowed gap positions” was “With half”. (Markos & Baldwin, 2001). The profile used for am- Final aligned length analysed for each region is plification was as described in Galbany-Casals et al. shown in Table 1, with indication of the percent- (2009). In total, we included in the analyses 277 ITS age of the original data sets that it represents after sequences, of which four are new, and 281 ETS se- the exclusion of ambiguously aligned regions. quences, of which four are new (Appendix 1). Analyses cpDNA amplification strategies The phylogenetic relationships were examined an- The rpl32-trnL intergenic spacer was amplified us- alysing cpDNA and nrDNA separately, given the ing the forward primer rpl32F and the reverse prim- existence of several well-supported incongruent er trnL(UAG) (Shaw et al., 2007). The profile used for patterns in previous phylogenies (Smissen et al., amplification was as described in Galbany-Casals 2011; Galbany-Casals et al., 2014).

For each data set, the best-available model of mo- FigTree 1.4.3 (Rambaut, 2016). Posterior probability lecular evolution required for estimations of phyloge- supports were considered to be significant for nodes ny was selected using the Akaike information criterion with PP ≥ 0.95, and only those are shown. (AIC; Akaike, 1973) as implemented in the software MEGA v7.0.14 (Kumar et al., 2016). The best-fitting Morphological study model was used in each case (see Table 1). In both data sets, Athrixia phylicoides DC., Relha- The sequenced specimens of P. aldunateoides (Ap- nia pungens L’Hérit. and Leysera gnaphalodes (L.) L. pendix 1) were identified using several keys (Re- were coded as outgroup taxa, based on previous stud- iche, 1903; Freire et al., 2014a), and compared to ies (Galbany-Casals et al., 2014; Freire et al., 2014b; the type specimen of the name Gnaphalium al- Nie et al., 2016). Bayesian analyses were conducted dunateoides: “CHILE: Santiago, Feb 1839, Gay using BEAST v1.8.3 (Drummond et al., 2012) with s. n.” (P 00704557 photo!) (Monti et al., 2013; the same parameters for both data sets. Clock model Freire et al., 2014a). Also, for the types of the ge- was coded as an “Uncorrelated relaxed clock” with neric names Pseudognaphalium and Gnaphalium, a “Lognormal” distribution. The data sets were not the two sequenced specimens of P. oxyphyllum and partitioned by locus and the GTR + I + G nucleotide the two of G. uliginosum (Appendix 1) were iden- substitution model (Gu et al., 1995) was used with the tified using several keys (Holub, 1976; McVaugh, following conditions: “Base frequencies” was “Em- 1984; Espinosa-García, 2005; Nesom, 2006; Chen pirical” and “Number of Gamma Categories” was “4”. & Bayer, 2011) and they were compared to the The tree Prior was set as “Speciation: Yule Process” corresponding type specimens: “Herb. Linn. No. (Yule, 1925; Gernhard, 2008) and default prior dis- 989.84” (LINN photo!) for G. uliginosum (Hilliard tributions were chosen for all other parameters. After & Burtt, 1981; Jarvis, 1992) and “Villalpando au operator adjustment, four independent MCMC runs Sud est de Guanajuato, Méndez s. n.” (G-DC pho- were performed, each run of 50 million generations to!) for G. oxyphyllum (McVaugh, 1984). and sampling every 5.000 generations. We verified The stereome and pappus characters were stud- the convergence of chains with Tracer v1.6 (Rambaut ied for the sequenced specimens of P. oxyphyllum et al., 2014) by checking that effective sample size and P. aldunateoides, along with nine addition- values were adequate. The four runs were combined al specimens of P. aldunateoides and a specimen with LogCombiner v1.8.3 (Drummond et al., 2012) of G. uliginosum (Appendix 2). Involucral bracts after discarding the initial 10% generations of each were cleared in a commercial solution of sodium as burn-in. A maximum clade credibility tree was hypochlorite during 3–4 hours (Stace, 1965) to obtained using TreeAnnotator v1.8.3 (Drummond & visualize if the stereome was undivided or fenes- Rambaut, 2007) with the following conditions: “Tar- trated. The stereome and pappus bristles of female get tree type” was “Maximum clade credibility tree”, and hermaphroditic florets were observed using a “Node heights” was “Mean heights” and no burn-in Zeiss Axioscop compound microscope and photo- was performed. The resulting maximum clade cred- graphed with a Jenoptik ProGresC3 digital camera ibility tree was transformed into a cladogram using using the program ProGresCapture7.

Table 1. Characteristics of data matrices and substitution models applied in the analyses.

nrDNA (ITS + ETS) cpDNA (rpl32-trnL) Number of taxa 283 220

Number of characters included after removing 973 (87% of the complete aligned 508 (39% of the complete aligned ambiguously aligned regions data set) data set)

Number of MCMC generations 50 million 50 million

Substitution model GTR + I + G (Gu et al., 1995) GTR + I + G (Gu et al., 1995)

Collectanea Botanica 37: e012 (enero-diciembre 2018), e012, ISSN-L: 0010-0730, https://doi.org/10.3989/collectbot.2018.v37.012 6 A. ACOSTA-MAINDO & M. GALBANY-CASALS

RESULTS tles noticeably wider (Fig. 4.1) than those of pistillate florets, which have obtuse cells at the tips Phylogenetic position of Pseudognaphalium al- instead (Fig. 4.2). The stereome is variable. In dunateoides some specimens it fits the description of undivided stereome, i.e. it is uniformly thickened, except for The nrDNA (Fig. 2) and cpDNA (Fig. 3) phylo- some minute, thin translucent streaks in the upper genetic trees placed the genus Pseudognaphali- part of the stereome. In others specimens the trans- um within the HAP clade, being part of the clade lucent area of unthickened cells of the stereome is mainly constituted by the paraphyletic genus He- rather big, and in this cases it resembles a fenestrat- lichrysum, which is in conformity with previous ed stereome (Fig. 5.1). However, in all cases the studies (Smissen et al., 2011; Galbany-Casals et translucent area is limited to the upper part of the al., 2014; Nie et al., 2016). While the nrDNA phy- stereome. logeny showed Pseudognaphalium to be constitut- Gnaphalium uliginosum shows distinctly, though ed by two well-supported and independent clades slightly, dimorphic pappus bristles (Fig. 4.3 and (Fig. 2, clades A.1 and A.2), cpDNA showed it to 4.4) as described by Hilliard & Burtt (1981) and be constituted by three independent lineages (Fig. an undivided stereome with few thin streaks (Fig. 3, clades A.1, A.2 and A.3). In both the nrDNA 5.2). Pseudognaphalium oxyphyllum displays and the cpDNA trees, P. oxyphyllum, the type of a monomorphic pappus (Fig. 4.5 and 4.6) and a Pseudognaphalium, was included in a clade with fenestrated stereome (Fig. 5.3). Hence either spe- other species of Pseudognaphalium. cies agrees in its diagnostic morphological char- Members of Gnaphalium were recovered in a acters with the genus for which it provides the clearly independent and well supported clade: in type of the name. the nrDNA phylogeny, Gnaphalium was sister to some Syncarpha (Fig. 2, clade B), while the cpDNA DISCUSSION phylogeny indicated it to be more closely related to Vellereophyton (Fig. 3, clade B), what in both cases The study of Hilliard & Burtt (1981) demonstrat- agree with previous studies (Smissen et al., 2011; ed the great value of the stereome characters in the Galbany-Casals et al., 2014; Nie et al., 2016). In generic delimitation of Gnaphalieae, as genera are both phylogenies, the Gnaphalium clade included usually uniform—with few exceptions—for ste- the type species G. uliginosum. reome anatomy. Although fenestrated stereomes Both in the nrDNA tree (Fig. 2, clade B) and the present no doubt, Hilliard & Burtt (1981) com- cpDNA tree (Fig. 3, clade B), P. aldunateoides was mented that undivided stereomes, even being most- not placed in the HAP clade with the other Pseudo- ly opaque, can sometimes present median vascular gnaphalium species, but in the Gnaphalium clade. strands visible and there may occasionally be minute, thin streaks in the upper part of the stereome Morphological study of some bracts. This is the case of G. uliginosum (Fig. 5.2) and some specimens of P. aldunateoides. With regards to the habit and general appear- However, in other specimens of P. aldunateoides, ance, the studied specimens of P. aldunateoides the translucent area can be larger and resemble a are dwarf, multistemed, ascending to erect annual fenestrated stereome (Fig. 5.1). Due to the presence plants with concolorous leaves whitish-woolly on of this translucent area, the stereome of P. aldunate- both faces, few capitula arranged in corymbs and oides was interpreted as fenestrated by Monti et al. subtended by linear leaves, and campanulate invo- (2013), and this was the main reason to support the lucre composed of involucral bracts whitish at the combination of this species under Pseudognapha- apex (Fig. 1). They match the descriptions of the lium. However, the translucent area in P. aldunate- species (Reiche, 1903; Freire et al., 2014a) and oides’ bracts is smaller than in a typical fenestrated the type material. stereome (Fig. 5.3) and is limited to the upper part The pappus of P. aldunateoides is slightly to of the stereome. In conclusion, P. aldunateoides’ markedly dimorphic, i.e. the pappus bristles of her- stereomes present intermediate characteristics be- maphroditic florets are subclavate, making the bris- tween undivided and fenestrated stereomes.

Helichrysum hamulosum 1 Pseudognaphalium obtusifolium Pseudognaphalium micradenium Pseudognaphalium macounii Pseudognaphalium oligandrum Pseudognaphalium biolettii Pseudognaphalium leucocephalum 1 Helichrysum lambertianum Pseudognaphalium leucocephalum 2 0.98 Pseudognaphalium californicum Pseudognaphalium ramosissimum Pseudognaphalium semiamplexicaule Pseudognaphalium eggersii Pseudognaphalium oxyphyllum var. oxyphyllum 1 Lucilia acutifolia Pseudognaphalium attenuatum Pseudognaphalium thermale 1 Pseudognaphalium canescens Pseudognaphalium beneolens 1 Pseudognaphalium sylvicola Pseudognaphalium oxyphyllum var. oxyphyllum 2 Lucilia lycopodioides Pseudognaphalium moritzianum 1 Pseudognaphalium hypoleucum Pseudognaphalium flavescens 1 Helichrysum albilanatum 1 Helichrysum aureolum 0.97 Helichrysum setosum Micropsis dasycarpa Helichrysum cooperi 1 Helichrysum elegantissimum Helichrysum aureum var. aureum 0.98 Helichrysum foetidum Helichrysum difficile 0.99 Helichrysum flammeiceps Lucilia nitens Helichrysum transmontanum Pseudognaphalium gaudichaudianum 0.97 Pseudognaphalium melanosphaeroides Pseudognaphalium cheiranthifolium 2 0.99 Pseudognaphalium cheiranthifolium 1 Pseudognaphalium dombeyanum Facelis retusa Pseudognaphalium viravira 1 0.99 1 Pseudognaphalium heterotrichum 0.98 Pseudognaphalium leucopeplum Pseudognaphalium lacteum Pseudognaphalium graveolens Pseudognaphalium badium Facelis plumosa 1 Pseudognaphalium affine 1 1 Pseudognaphalium affine 2 1 Pseudognaphalium sandwicensium Pseudognaphalium luteoalbum 1 0.99 Pseudognaphalium luteoalbum 3 Berroa gnaphalioides Pseudognaphalium montevidense Pseudognaphalium luteoalbum 2 1 Helichrysum confertifolium Helichrysum reflexum 1 Helichrysum mariepscopicum Helichrysum summo-montanum Belloa chilensis 1 Helichrysum formosissimum 1 Helichrysum meyeri-johannis Helichrysum stuhlmannii Helichrysum brownei 1 Helichrysum argyranthum Helichrysum chionoides Chionolaena campestris Helichrysum basalticum 1 Helichrysum nicolai 1 Helichrysum nicolai 2 1 Helichrysum alucense 1 Helichrysum monogynum 1 1 Helichrysum glumaceum 2 Gamochaeta longipedicellata Helichrysum glumaceum 1 1 Helichrysum populifolium Helichrysum trilineatum Helichrysum armenium subsp. armenium 1 Helichrysum thianschanicum Helichrysum mussae Gamochaeta americana 1 Helichrysum rubicundum 1 Helichrysum orientale 1 Helichrysum plicatum 0.98 Helichrysum sibthorpii Helichrysum stoechas 0.96 Helichrysum pendulum Gamochaeta subfalcata 1 Helichrysum crassifolium 0.99 Helichrysum italicum subsp. italicum Helichrysum massanellanum 1 Helichrysum gossypinum 1 Helichrysum melaleucum 0.99 Helichrysum monizii 1 Stuckertiella capitata 1 Helichrysum devium Helichrysum obconicum 1 1 Anaphalis aff. aureopunctata 1 1 Anaphalis aff. busua Anaphalis aff. aureopunctata 2 Anaphalis aff. flavescens Gamochaeta serpyllifolia 1 Anaphalis gracilis 1 1 0.96 Anaphalis aff. virens Anaphalis sp. 1 Anaphalis aff. nepalensis 1 1 Anaphalis aff. nepalensis 2 Anaphalis margaritacea Gamochaetopsis alpina 0.97 Helichrysum plantago 1 Helichrysum nudifolium 1 Helichrysum miconiifolium 0.98 Helichrysum pilosellum Helichrysum globosum Mniodes kunthiana Helichrysum zeyheri 1 Helichrysum psilolepis Helichrysum schimperi Helichrysum pagophilum 1 Helichrysum manopappoides Helichrysum fulvescens 1 Mniodes andina Humeocline madagascariensis Helichrysum forsythii 1 1 Helichrysum neoachyroclinoides Helichrysum cordifolium 1 Helichrysum flagellare 1 Helichrysum triplinerve Mniodes santanica 1 Achyrocline satureioides Achyrocline alata Helichrysum odoratissimum 1 Helichrysum epapposum 1 Achyrocline stenoptera 1 1 Helichrysum forskahlii var. compactum Mniodes longifolia Helichrysum forskahlii var. forskahlii 1 Helichrysum interjacens Helichrysum polycladum 1 Helichrysum simillinum 0.99 Helichrysum melanacme 1 Helichrysum cymosum subsp. calvum Loricaria colombiana Helichrysum anomalum 0.99 Helichrysum griseolanatum Helichrysum mundtii 0.99 Helichrysum auriceps 1 Helichrysum mixtum var. mixtum Helichrysum cephaloideum Jalcophila boliviensis 1 1 Helichrysum albobrunneum 0.99 Helichrysum appendiculatum 1 Helichrysum rosum var. arcuatum 1 Helichrysum rugulosum 1 Helichrysum rosum var. rosum Helichrysum felinum Chevreulia acuminata Helichrysum chrysargyrum 1 1 Helichrysum petiolare 1 Helichrysum patulum 1 Helichrysum patulum 2 1 Helichrysum glomeratum Helichrysum ephelos Cuatrecasasiella argentina Helichrysum truncatum Helichrysum acutatum 1 Helichrysum milfordiae Helichrysum flanaganii Helichrysum umbraculigerum Chionolaena salicifolia 1 Helichrysum argyrophyllum 1 Helichrysum aureonitens Helichrysum platycephalum 1 Helichrysum excisum Helichrysum silvaticum Helichrysum opacum Mexerion sarmentosum Helichrysum herbaceum 1 Helichrysum splendidum Helichrysum montanum Helichrysum marginatum Helichrysum ecklonis Helichrysum paleatum Antennaria dioica 1 Helichrysum confertum Helichrysum sutherlandii Helichrysum natalitium 1 Helichrysum monticola Helichrysum adenocarpum Helichrysum platypterum Filago arizonica 1 Helichrysum mimetes 1 1 Helichrysum lepidissimum 1 Helichrysum obductum Helichrysum homilochrysum Helichrysum lecomtei 0.99 1 Helichrysum selaginifolium 0.99 Filago californica 0.99 Helichrysum benthamii Helichrysum abbayesii 1 Helichrysum cryptomerioides Helichrysum ibityense 1 Helichrysum gymnocephalum Helichrysum xylocladum Stylocline psilocarphoides 1 Helichrysum citrispinum var. citrispinum 1 Helichrysum citrispinum var. hoehnelii 1 Helichrysum gofense 2 1 Helichrysum gofense 1 Helichrysum arwae 1 Helichrysum maranguense Micropus californicus 0.99 Helichrysum chionosphaerum Helichrysum swynnertonii 1 Helichrysum kilimanjari Helichrysum isolepis 1 Helichrysum dasycephalum 0.98 Helichrysum callicomum 1 Logfia minima Helichrysum galpinii 1 Helichrysum zwartbergense 1 Helichrysum litorale 1 Helichrysum spiralepis 1 Helichrysum stellatum Helichrysum candolleanum Logfia gallica 1 Helichrysum argyrosphaerum 1 Helichrysum cerastioides Helichrysum asperum 1 Helichrysum indicum 1 Helichrysum crispum 0.99 Castroviejoa montelinasana Helichrysum retortum Helichrysum hamulosum Helichrysum lambertianum Lucilia acutifolia Lucilia lycopodioides Micropsis dasycarpa Filago carpetana Lucilia nitens 1 1 Facelis retusa Facelis plumosa Berroa gnaphalioides 1 Belloa chilensis Chionolaena campestris Filago congesta 1 1 Gamochaeta longipedicellata 1 1 Gamochaeta americana Gamochaeta subfalcata 1 Stuckertiella capitata 1 Gamochaeta serpyllifolia Gamochaetopsis alpina Filago pygmaea 1 Mniodes kunthiana Mniodes andina 1 Mniodes santanica 1 1 Mniodes longifolia 1 0.99 0.99 Loricaria colombiana Jalcophila boliviensis Filago lutescens subsp. lutescens 1 Chevreulia acuminata Cuatrecasasiella argentina 1 Chionolaena salicifolia Mexerion sarmentosum Antennaria dioica 1 Filago arizonica Bombycilaena discolor 0.99 Filago californica 0.99 Stylocline psilocarphoides 1 Micropus californicus 1 Logfia minima Logfia gallica Castroviejoa montelinasana Micropus supinus Filago carpetana 1 1 Filago congesta Filago pygmaea 1 Filago lutescens subsp. lutescens 0.99 Bombycilaena discolor Micropus supinus Omalotheca supina Omalotheca supina Omalotheca sylvatica Leontopodium alpinum 1 Gnaphalium capense 1 Gnaphalium austroafricanum Omalotheca sylvatica 1 1 Gnaphalium filagopsis Gnaphalium confine 0.99 Gnaphalium declinatum 1 1 Pseudognaphalium aldunateoides 1 0.99 Pseudognaphalium aldunateoides 2 1 Gnaphalium palustre 1 Leontopodium alpinum Gnaphalium uliginosum 1 Gnaphalium uliginosum 2 Syncarpha mucronata 1 1 Syncarpha chlorochrysum Syncarpha paniculata Langebergia canescens Gnaphalium capense Helichrysum dasyanthum 1 1 Petalacte coronata 1 B Edmondia sesamoides 1 Syncarpha canescens Syncarpha staehelina 1 Helichrysum lanceolatum Gnaphalium austroafricanum Raoulia tenuicaulis 1 Ewartia catipes 1 Ozothamnus diosmifolius 1 1 Acanthocladium dockeri Craspedia glauca Vellereophyton dealbatum Gnaphalium filagopsis 1 Lachnospermum fasciculatum 1 1 Metalasia adunca 1 0.98 Dolichothrix ericoides 1 Ifloga spicata 1 Ifloga repens 0.98 1 Amphiglossa corrudaefolia Gnaphalium confine Stoebe aethiopica 1 Lasiopogon glomerulatus Lasiopogon muscoides 0.99 1 Leysera gnaphalodes 1 Relhania pungens Athrixia phylicoides Gnaphalium declinatum 3.0 1 1 Pseudognaphalium aldunateoides 1 0.99 Pseudognaphalium aldunateoides 2 1 Gnaphalium palustre 1 Gnaphalium uliginosum 1 Gnaphalium uliginosum 2 Syncarpha mucronata 1 1 Syncarpha chlorochrysum Syncarpha paniculata Langebergia canescens Helichrysum dasyanthum 1 Petalacte coronata 1 Edmondia sesamoides 1 Syncarpha canescens Syncarpha staehelina 1 Helichrysum lanceolatum Raoulia tenuicaulis Ewartia catipes 1 Ozothamnus diosmifolius 1 1 Acanthocladium dockeri Craspedia glauca Vellereophyton dealbatum 1 Lachnospermum fasciculatum 1 Metalasia adunca 0.98 Dolichothrix ericoides 1 Ifloga spicata 1 Ifloga repens 0.98 1 Amphiglossa corrudaefolia Stoebe aethiopica 1 Lasiopogon glomerulatus Lasiopogon muscoides 1 Leysera gnaphalodes 1 Relhania pungens Athrixia phylicoides 3.0 Figure 2. nrDNA (ITS and 3′ ETS regions) maximum credibility tree obtained from the BEAST analyses of the Gnaphalieae. Only ≥ 0.95 Bayesian posterior probabilities values are shown. Species of the genera Pseudognaphalium and Gnaphalium are highlighted in coloured boxes. Clades discussed in the text are labeled with capital letters and numbers.

Collectanea Botanica 37: e012 (enero-diciembre 2018), e012, ISSN-L: 0010-0730, https://doi.org/10.3989/collectbot.2018.v37.012 8 A. ACOSTA-MAINDO & M. GALBANY-CASALS

Helichrysum cymosum subsp. calvum

1 Pseudognaphalium obtusifolium Pseudognaphalium micradenium Pseudognaphalium macounii Pseudognaphalium oligandrum Helichrysum anomalum Pseudognaphalium biolettii Pseudognaphalium leucocephalum 1 Pseudognaphalium leucocephalum 2 0.98 Pseudognaphalium californicum Pseudognaphalium ramosissimum Pseudognaphalium semiamplexicaule Helichrysum griseolanatum Pseudognaphalium eggersii Pseudognaphalium oxyphyllum var. oxyphyllum 1 Pseudognaphalium attenuatum Pseudognaphalium thermale 1 Pseudognaphalium canescens Helichrysum mundtii Pseudognaphalium beneolens 1 Pseudognaphalium sylvicola Pseudognaphalium oxyphyllum var. oxyphyllum 2 Pseudognaphalium moritzianum 1 Pseudognaphalium hypoleucum Pseudognaphalium flavescens Helichrysum auriceps Helichrysum albilanatum 0.99 1 1 Helichrysum aureolum 0.97 Helichrysum setosum Helichrysum cooperi 1 Helichrysum elegantissimum Helichrysum aureum var. aureum 1 Helichrysum mixtum var. mixtum 0.98 Helichrysum foetidum Helichrysum difficile 0.99 Helichrysum flammeiceps Helichrysum transmontanum Pseudognaphalium gaudichaudianum 0.97 Pseudognaphalium melanosphaeroides Helichrysum cephaloideum Pseudognaphalium cheiranthifolium 2 0.99 Pseudognaphalium cheiranthifolium 1 Pseudognaphalium dombeyanum 1 0.99 Pseudognaphalium viravira 1 Pseudognaphalium heterotrichum 0.98 Pseudognaphalium leucopeplum Helichrysum albobrunneum Pseudognaphalium lacteum Pseudognaphalium graveolens Pseudognaphalium badium 1 Pseudognaphalium affine 1 1 Pseudognaphalium affine 2 1 Pseudognaphalium sandwicensium Helichrysum appendiculatum Pseudognaphalium luteoalbum 1 0.99 Pseudognaphalium luteoalbum 3 Pseudognaphalium montevidense 0.99 Pseudognaphalium luteoalbum 2 1 Helichrysum confertifolium Helichrysum reflexum Helichrysum rosum var. arcuatum 1 Helichrysum mariepscopicum 1 Helichrysum summo-montanum 1 Helichrysum formosissimum Helichrysum meyeri-johannis Helichrysum stuhlmannii Helichrysum brownei Helichrysum rugulosum 1 Helichrysum argyranthum 1 Helichrysum chionoides Helichrysum basalticum 1 Helichrysum nicolai 1 Helichrysum nicolai 2 1 Helichrysum alucense Helichrysum rosum var. rosum 1 1 Helichrysum monogynum 1 Helichrysum glumaceum 2 Helichrysum glumaceum 1 Helichrysum populifolium Helichrysum trilineatum Helichrysum felinum Helichrysum armenium subsp. armenium 1 Helichrysum thianschanicum Helichrysum mussae 1 Helichrysum rubicundum 1 Helichrysum orientale 1 Helichrysum plicatum Helichrysum chrysargyrum 0.98 Helichrysum sibthorpii Helichrysum stoechas 0.96 Helichrysum pendulum 1 Helichrysum crassifolium 0.99 Helichrysum italicum subsp. italicum Helichrysum massanellanum Helichrysum petiolare 1 Helichrysum gossypinum 1 1 Helichrysum melaleucum 0.99 Helichrysum monizii 1 Helichrysum devium Helichrysum obconicum Anaphalis aff. aureopunctata 1 1 1 1 Helichrysum patulum 1 1 Anaphalis aff. busua Anaphalis aff. aureopunctata 2 1 Anaphalis aff. flavescens 1 Anaphalis gracilis 0.96 Anaphalis aff. virens Anaphalis sp. Helichrysum patulum 2 1 Anaphalis aff. nepalensis 1 1 Anaphalis aff. nepalensis 2 Anaphalis margaritacea 0.97 Helichrysum plantago 1 Helichrysum nudifolium 1 Helichrysum miconiifolium Helichrysum glomeratum 0.98 Helichrysum pilosellum 1 Helichrysum globosum Helichrysum zeyheri Helichrysum psilolepis Helichrysum schimperi Helichrysum pagophilum Helichrysum ephelos 1 Helichrysum manopappoides Helichrysum fulvescens 1 Humeocline madagascariensis Helichrysum forsythii 1 1 Helichrysum neoachyroclinoides Helichrysum cordifolium Helichrysum truncatum 1 Helichrysum flagellare Helichrysum triplinerve 1 Achyrocline satureioides Achyrocline alata Helichrysum odoratissimum 1 Helichrysum acutatum Helichrysum epapposum Achyrocline stenoptera 1 Helichrysum forskahlii var. compactum Helichrysum forskahlii var. forskahlii 1 Helichrysum interjacens Helichrysum polycladum 1 Helichrysum milfordiae Helichrysum simillinum 1 Helichrysum melanacme 1 Helichrysum cymosum subsp. calvum Helichrysum anomalum Helichrysum griseolanatum Helichrysum mundtii Helichrysum flanaganii 0.99 Helichrysum auriceps 1 Helichrysum mixtum var. mixtum Helichrysum cephaloideum 1 1 Helichrysum albobrunneum 0.99 Helichrysum appendiculatum 1 Helichrysum rosum var. arcuatum Helichrysum umbraculigerum 1 Helichrysum rugulosum 1 Helichrysum rosum var. rosum Helichrysum felinum Helichrysum chrysargyrum 1 Helichrysum petiolare 1 Helichrysum patulum 1 Helichrysum argyrophyllum Helichrysum patulum 2 1 Helichrysum glomeratum Helichrysum ephelos 1 Helichrysum truncatum Helichrysum acutatum 1 Helichrysum milfordiae Helichrysum aureonitens Helichrysum flanaganii Helichrysum umbraculigerum 1 Helichrysum argyrophyllum Helichrysum aureonitens Helichrysum platycephalum 1 Helichrysum excisum Helichrysum platycephalum Helichrysum silvaticum Helichrysum opacum Helichrysum herbaceum 1 Helichrysum splendidum Helichrysum montanum Helichrysum marginatum Helichrysum excisum Helichrysum ecklonis 1 Helichrysum paleatum 1 Helichrysum confertum Helichrysum sutherlandii Helichrysum natalitium Helichrysum monticola Helichrysum silvaticum 1 Helichrysum adenocarpum Helichrysum platypterum 1 Helichrysum mimetes 1 Helichrysum lepidissimum 1 Helichrysum obductum Helichrysum opacum Helichrysum homilochrysum Helichrysum lecomtei 0.99 1 Helichrysum selaginifolium 0.99 Helichrysum benthamii Helichrysum abbayesii 1 Helichrysum cryptomerioides Helichrysum herbaceum Helichrysum ibityense 1 Helichrysum gymnocephalum Helichrysum xylocladum 1 Helichrysum citrispinum var. citrispinum 1 Helichrysum citrispinum var. hoehnelii Helichrysum gofense 2 1 1 Helichrysum splendidum Helichrysum gofense 1 1 Helichrysum arwae Helichrysum maranguense 0.99 Helichrysum chionosphaerum Helichrysum swynnertonii 1 Helichrysum kilimanjari Helichrysum montanum Helichrysum isolepis 1 Helichrysum dasycephalum 0.98 Helichrysum callicomum Helichrysum galpinii 1 Helichrysum zwartbergense 1 Helichrysum litorale Helichrysum marginatum 1 Helichrysum spiralepis 1 Helichrysum stellatum Helichrysum candolleanum 1 Helichrysum argyrosphaerum 1 Helichrysum cerastioides Helichrysum asperum Helichrysum ecklonis 1 Helichrysum indicum 1 Helichrysum crispum 0.99 Helichrysum retortum Helichrysum hamulosum Helichrysum lambertianum Lucilia acutifolia Helichrysum paleatum Lucilia lycopodioides Micropsis dasycarpa Lucilia nitens 1 1 Facelis retusa Facelis plumosa Berroa gnaphalioides Helichrysum confertum Belloa chilensis 1 1 Chionolaena campestris 1 1 Gamochaeta longipedicellata Gamochaeta americana Gamochaeta subfalcata 1 Helichrysum sutherlandii Stuckertiella capitata 1 Gamochaeta serpyllifolia Gamochaetopsis alpina 1 Mniodes kunthiana Mniodes andina 1 Mniodes santanica 1 1 Helichrysum natalitium Mniodes longifolia 0.99 0.99 Loricaria colombiana Jalcophila boliviensis 1 Chevreulia acuminata Cuatrecasasiella argentina 1 Chionolaena salicifolia Helichrysum monticola Mexerion sarmentosum 1 Antennaria dioica 1 Filago arizonica 0.99 Filago californica Stylocline psilocarphoides 1 Micropus californicus Helichrysum adenocarpum 1 Logfia minima Logfia gallica Castroviejoa montelinasana Filago carpetana 1 1 Filago congesta Filago pygmaea Helichrysum platypterum 1 Filago lutescens subsp. lutescens 0.99 Bombycilaena discolor Micropus supinus Omalotheca supina Omalotheca sylvatica Leontopodium alpinum Helichrysum mimetes 1 Gnaphalium capense 1 1 Gnaphalium austroafricanum 1 1 Gnaphalium filagopsis Gnaphalium confine 0.99 Gnaphalium declinatum 1 1 1 Pseudognaphalium aldunateoides 1 Helichrysum lepidissimum 0.99 Pseudognaphalium aldunateoides 2 1 Gnaphalium palustre 1 Gnaphalium uliginosum 1 Gnaphalium uliginosum 2 1 Syncarpha mucronata 1 Syncarpha chlorochrysum Helichrysum obductum 1 Syncarpha paniculata Langebergia canescens Helichrysum dasyanthum 1 Petalacte coronata 1 Edmondia sesamoides Syncarpha canescens Helichrysum homilochrysum 1 Syncarpha staehelina 1 Helichrysum lanceolatum Raoulia tenuicaulis Ewartia catipes 1 Ozothamnus diosmifolius 1 1 Helichrysum lecomtei Acanthocladium dockeri Craspedia glauca Vellereophyton dealbatum 1 Lachnospermum fasciculatum 1 Metalasia adunca 0.98 Dolichothrix ericoides Helichrysum selaginifolium Ifloga spicata 1 1 1 Ifloga repens 0.98 1 Amphiglossa corrudaefolia 0.99 Stoebe aethiopica 1 Lasiopogon glomerulatus Lasiopogon muscoides Helichrysum benthamii 1 Leysera gnaphalodes 1 Relhania pungens Athrixia phylicoides 0.99 3.0 Helichrysum abbayesii 1 Helichrysum cryptomerioides Helichrysum ibityense 1 Helichrysum gymnocephalum Helichrysum xylocladum 1 Helichrysum citrispinum var. citrispinum 1 Helichrysum citrispinum var. hoehnelii 1 Helichrysum gofense 2 1 Helichrysum gofense 1 Helichrysum arwae Helichrysum maranguense 0.99 Helichrysum chionosphaerum Helichrysum swynnertonii 1 Helichrysum kilimanjari Helichrysum isolepis 1 Helichrysum dasycephalum 0.98 Helichrysum callicomum Helichrysum galpinii 1 Helichrysum zwartbergense 1 Helichrysum litorale 1 Helichrysum spiralepis 1 Helichrysum stellatum Helichrysum candolleanum 1 Helichrysum argyrosphaerum 1 Helichrysum cerastioides Helichrysum asperum 1 Helichrysum indicum 1 Helichrysum crispum 0.99 Helichrysum retortum Helichrysum hamulosum Helichrysum lambertianum 3.0 Lucilia acutifolia Figure 2 (cont.). nrDNA (ITS and 3′ ETS regions) maximum credibility tree obtainedLucilia lycopodioidesfrom the BEAST analyses of the Micropsis dasycarpa Gnaphalieae. Only ≥ 0.95 Bayesian posterior probabilities values are shown. SpeciesLucilia of the nitens genera Pseudognaphalium and Gnaphalium are highlighted in coloured boxes. Clades discussed in1 the text1 are labeledFacelis with retusa capital letters and numbers. Facelis plumosa

1 Pseudognaphalium obtusifolium Helichrysum nicolai 2 Pseudognaphalium micradenium Pseudognaphalium macounii Pseudognaphalium oligandrum Pseudognaphalium biolettii Pseudognaphalium leucocephalum 1 1 Pseudognaphalium leucocephalum 2 Helichrysum alucense 0.98 Pseudognaphalium californicum Pseudognaphalium ramosissimum Pseudognaphalium semiamplexicaule Pseudognaphalium eggersii Pseudognaphalium oxyphyllum var. oxyphyllum 1 Pseudognaphalium attenuatum 1 Helichrysum monogynum Pseudognaphalium thermale 1 Pseudognaphalium canescens Pseudognaphalium beneolens 1 Pseudognaphalium sylvicola Pseudognaphalium oxyphyllum var. oxyphyllum 2 Pseudognaphalium moritzianum Helichrysum glumaceum 2 Pseudognaphalium hypoleucum 1 1 Pseudognaphalium flavescens 1 Helichrysum albilanatum 1 Helichrysum aureolum 0.97 Helichrysum setosum Helichrysum glumaceum 1 Helichrysum cooperi 1 Helichrysum elegantissimum Helichrysum aureum var. aureum 0.98 Helichrysum foetidum Helichrysum difficile 0.99 Helichrysum flammeiceps Helichrysum transmontanum Pseudognaphalium gaudichaudianum 0.97 Pseudognaphalium melanosphaeroides Pseudognaphalium cheiranthifolium 2 0.99 Pseudognaphalium cheiranthifolium 1 Pseudognaphalium dombeyanum 0.99 Pseudognaphalium viravira Pseudognaphalium heterotrichum 0.98 Pseudognaphalium leucopeplum Pseudognaphalium lacteum Pseudognaphalium graveolens Pseudognaphalium badium Helichrysum armenium subsp. armenium 1 Pseudognaphalium affine 1 1 Pseudognaphalium affine 2 1 Pseudognaphalium sandwicensium Pseudognaphalium luteoalbum 1 0.99 Pseudognaphalium luteoalbum 3 Pseudognaphalium montevidense Helichrysum thianschanicum Pseudognaphalium luteoalbum 2 1 1 Helichrysum confertifolium Helichrysum reflexum 1 Helichrysum mariepscopicum Helichrysum summo-montanum 1 Helichrysum formosissimum Helichrysum mussae Helichrysum meyeri-johannis Helichrysum stuhlmannii Helichrysum brownei 1 Helichrysum argyranthum Helichrysum chionoides Helichrysum basalticum 1 Helichrysum rubicundum 1 Helichrysum nicolai 1 Helichrysum nicolai 2 1 Helichrysum alucense 1 Helichrysum monogynum 1 Helichrysum glumaceum 2 Helichrysum glumaceum 1 Helichrysum orientale Helichrysum populifolium 1 Helichrysum trilineatum Helichrysum armenium subsp. armenium 1 Helichrysum thianschanicum Helichrysum mussae 1 Helichrysum rubicundum Helichrysum plicatum 1 1 Helichrysum orientale 1 Helichrysum plicatum 0.98 Helichrysum sibthorpii Helichrysum stoechas 0.96 Helichrysum pendulum Helichrysum sibthorpii 1 Helichrysum crassifolium 0.98 0.99 Helichrysum italicum subsp. italicum Helichrysum massanellanum 1 Helichrysum gossypinum 1 Helichrysum melaleucum 0.99 Helichrysum monizii 1 Helichrysum stoechas Helichrysum devium Helichrysum obconicum 1 1 Anaphalis aff. aureopunctata 1 1 Anaphalis aff. busua Anaphalis aff. aureopunctata 2 Anaphalis aff. flavescens 0.96 1 Helichrysum pendulum 1 Anaphalis gracilis 0.96 Anaphalis aff. virens Anaphalis sp. 1 Anaphalis aff. nepalensis 1 1 Anaphalis aff. nepalensis 2 Anaphalis margaritacea 1 Helichrysum crassifolium 0.97 Helichrysum plantago 1 Helichrysum nudifolium 1 Helichrysum miconiifolium 0.98 Helichrysum pilosellum Helichrysum globosum Helichrysum zeyheri Helichrysum italicum subsp. italicum Helichrysum psilolepis 0.99 Helichrysum schimperi Helichrysum pagophilum 1 Helichrysum manopappoides Helichrysum fulvescens 1 Humeocline madagascariensis Helichrysum massanellanum Helichrysum forsythii 1 1 Helichrysum neoachyroclinoides Helichrysum cordifolium 1 Helichrysum flagellare Helichrysum triplinerve 1 1 Achyrocline satureioides Helichrysum gossypinum Achyrocline alata Helichrysum odoratissimum 1 Helichrysum epapposum Achyrocline stenoptera 1 Helichrysum forskahlii var. compactum Helichrysum forskahlii var. forskahlii Helichrysum melaleucum Helichrysum interjacens 1 1 Helichrysum polycladum 1 Helichrysum simillinum Helichrysum melanacme 1 Helichrysum cymosum subsp. calvum 0.99 Helichrysum monizii Helichrysum anomalum Helichrysum griseolanatum Helichrysum mundtii 0.99 Helichrysum auriceps 1 Helichrysum mixtum var. mixtum 1 Helichrysum cephaloideum 1 1 Helichrysum devium Helichrysum albobrunneum 0.99 Helichrysum appendiculatum 1 Helichrysum rosum var. arcuatum 1 Helichrysum rugulosum 1 Helichrysum rosum var. rosum Helichrysum felinum Helichrysum obconicum Helichrysum chrysargyrum 1 Helichrysum petiolare 1 Helichrysum patulum 1 Helichrysum patulum 2 1 Helichrysum glomeratum Helichrysum ephelos Anaphalis aff. aureopunctata 1 Helichrysum truncatum 1 Helichrysum acutatum 1 1 Helichrysum milfordiae Helichrysum flanaganii Helichrysum umbraculigerum Helichrysum argyrophyllum 1 1 Anaphalis aff. busua Helichrysum aureonitens Helichrysum platycephalum 1 Helichrysum excisum Helichrysum silvaticum Helichrysum opacum Helichrysum herbaceum Anaphalis aff. aureopunctata 2 1 Helichrysum splendidum Helichrysum montanum Helichrysum marginatum Helichrysum ecklonis Helichrysum paleatum 1 Helichrysum confertum Anaphalis aff. flavescens Helichrysum sutherlandii Helichrysum natalitium 1 1 Helichrysum monticola Helichrysum adenocarpum Helichrysum platypterum 1 Helichrysum mimetes Anaphalis gracilis 1 Helichrysum lepidissimum 1 1 Helichrysum obductum Helichrysum homilochrysum Helichrysum lecomtei 0.99 1 Helichrysum selaginifolium 0.96 Helichrysum benthamii Anaphalis aff. virens 0.99 Helichrysum abbayesii 1 Helichrysum cryptomerioides Helichrysum ibityense 1 Helichrysum gymnocephalum Helichrysum xylocladum Anaphalis sp. 1 Helichrysum citrispinum var. citrispinum 1 Helichrysum citrispinum var. hoehnelii 1 Helichrysum gofense 2 1 Helichrysum gofense 1 Helichrysum arwae Helichrysum maranguense Anaphalis aff. nepalensis 1 0.99 Helichrysum chionosphaerum 1 Helichrysum swynnertonii 1 Helichrysum kilimanjari Helichrysum isolepis 1 Helichrysum dasycephalum 0.98 Helichrysum callicomum 1 Anaphalis aff. nepalensis 2 Helichrysum galpinii 1 Helichrysum zwartbergense 1 Helichrysum litorale 1 Helichrysum spiralepis 1 Helichrysum stellatum Helichrysum candolleanum Anaphalis margaritacea 1 Helichrysum argyrosphaerum 1 Helichrysum cerastioides Helichrysum asperum 1 Helichrysum indicum 1 Helichrysum crispum 0.99 Helichrysum retortum 0.97 Helichrysum plantago Helichrysum hamulosum Helichrysum lambertianum Lucilia acutifolia Lucilia lycopodioides Micropsis dasycarpa 1 Lucilia nitens Helichrysum nudifolium 1 1 Facelis retusa Facelis plumosa Berroa gnaphalioides 1 Belloa chilensis Chionolaena campestris 1 1 1 Gamochaeta longipedicellata Helichrysum miconiifolium Gamochaeta americana Gamochaeta subfalcata 1 Stuckertiella capitata 1 Gamochaeta serpyllifolia 0.98 Gamochaetopsis alpina Mniodes kunthiana Helichrysum pilosellum 1 Mniodes andina 1 Mniodes santanica 1 1 Mniodes longifolia 0.99 0.99 Loricaria colombiana Jalcophila boliviensis Helichrysum globosum 1 Chevreulia acuminata Cuatrecasasiella argentina 1 Chionolaena salicifolia Mexerion sarmentosum Antennaria dioica 1 Filago arizonica Helichrysum zeyheri 0.99 Filago californica Stylocline psilocarphoides 1 Micropus californicus 1 Logfia minima Logfia gallica Castroviejoa montelinasana Helichrysum psilolepis Filago carpetana 1 1 Filago congesta Filago pygmaea 1 Filago lutescens subsp. lutescens 0.99 Bombycilaena discolor Micropus supinus Helichrysum schimperi Omalotheca supina Omalotheca sylvatica Leontopodium alpinum 1 Gnaphalium capense 1 Gnaphalium austroafricanum 1 1 Gnaphalium filagopsis Helichrysum pagophilum Gnaphalium confine 0.99 Gnaphalium declinatum 1 1 Pseudognaphalium aldunateoides 1 0.99 Pseudognaphalium aldunateoides 2 1 Gnaphalium palustre 1 Gnaphalium uliginosum 1 Helichrysum manopappoides Gnaphalium uliginosum 2 1 Syncarpha mucronata 1 1 Syncarpha chlorochrysum Syncarpha paniculata Langebergia canescens Helichrysum dasyanthum Helichrysum fulvescens 1 Petalacte coronata 1 Edmondia sesamoides 1 Syncarpha canescens Syncarpha staehelina 1 1 Helichrysum lanceolatum Raoulia tenuicaulis Humeocline madagascariensis Ewartia catipes 1 Ozothamnus diosmifolius 1 1 Acanthocladium dockeri Craspedia glauca Vellereophyton dealbatum Lachnospermum fasciculatum Helichrysum forsythii 1 1 Metalasia adunca 0.98 Dolichothrix ericoides 1 Ifloga spicata 1 Ifloga repens 1 0.98 1 Amphiglossa corrudaefolia Helichrysum neoachyroclinoides Stoebe aethiopica 1 1 Lasiopogon glomerulatus Lasiopogon muscoides 1 Leysera gnaphalodes 1 Relhania pungens Athrixia phylicoides Helichrysum cordifolium 3.0 1 Helichrysum flagellare Helichrysum triplinerve 1 Achyrocline satureioides Achyrocline alata Helichrysum odoratissimum 1 Helichrysum epapposum Achyrocline stenoptera 1 Helichrysum forskahlii var. compactum Helichrysum forskahlii var. forskahlii 1 Helichrysum interjacens Helichrysum polycladum 1 Helichrysum simillinum Helichrysum melanacme 1 Helichrysum cymosum subsp. calvum Helichrysum anomalum Helichrysum griseolanatum Helichrysum mundtii

Helichrysum cephaloideum 1 1 Helichrysum albobrunneum 3.0 0.99 Helichrysum appendiculatum 1 Helichrysum rosum var. arcuatum 1 Helichrysum rugulosum 1 Helichrysum rosum var. rosum Helichrysum felinum Helichrysum chrysargyrum 1 Helichrysum petiolare 1 Helichrysum patulum 1 Helichrysum patulum 2 1 Helichrysum glomeratum Helichrysum ephelos Figure 2 (cont.). nrDNA (ITS and 3′ ETS regions) maximum credibility tree obtainedHelichrysum from truncatum the BEAST analyses of the Gnaphalieae. Only ≥ 0.95 Bayesian posterior probabilities values are shown. SpeciesHelichrysum of the genera acutatum Pseudognaphalium and 1 Helichrysum milfordiae Gnaphalium are highlighted in coloured boxes. Clades discussed in the text are labeledHelichrysum with capital flanaganii letters and numbers.

Collectanea Botanica 37: e012 (enero-diciembre 2018), e012, ISSN-L: 0010-0730, https://doi.org/10.3989/collectbot.2018.v37.012 10 A. ACOSTA-MAINDO & M. GALBANY-CASALS

1 Pseudognaphalium obtusifolium Pseudognaphalium micradenium Pseudognaphalium macounii Pseudognaphalium oligandrum Pseudognaphalium biolettii Pseudognaphalium leucocephalum 1 Pseudognaphalium leucocephalum 2 0.98 Pseudognaphalium californicum Pseudognaphalium obtusifolium A.1 Pseudognaphalium ramosissimum 1 Pseudognaphalium semiamplexicaule Pseudognaphalium eggersii Pseudognaphalium oxyphyllum var. oxyphyllum 1 Pseudognaphalium attenuatum Pseudognaphalium thermale Pseudognaphalium micradenium 1 Pseudognaphalium canescens Pseudognaphalium beneolens 1 Pseudognaphalium sylvicola Pseudognaphalium oxyphyllum var. oxyphyllum 2 Pseudognaphalium moritzianum Pseudognaphalium hypoleucum Pseudognaphalium macounii 1 Pseudognaphalium flavescens 1 Helichrysum albilanatum 1 Helichrysum aureolum 0.97 Helichrysum setosum Helichrysum cooperi Pseudognaphalium oligandrum 1 Helichrysum elegantissimum Helichrysum aureum var. aureum 0.98 Helichrysum foetidum Helichrysum difficile 0.99 Helichrysum flammeiceps Helichrysum transmontanum Pseudognaphalium biolettii Pseudognaphalium gaudichaudianum 0.97 Pseudognaphalium melanosphaeroides Pseudognaphalium cheiranthifolium 2 0.99 Pseudognaphalium cheiranthifolium 1 Pseudognaphalium dombeyanum 0.99 Pseudognaphalium viravira Pseudognaphalium leucocephalum 1 Pseudognaphalium heterotrichum 0.98 Pseudognaphalium leucopeplum Pseudognaphalium lacteum Pseudognaphalium graveolens Pseudognaphalium badium 1 Pseudognaphalium affine 1 Pseudognaphalium leucocephalum 2 1 Pseudognaphalium affine 2 1 Pseudognaphalium sandwicensium Pseudognaphalium luteoalbum 1 0.99 Pseudognaphalium luteoalbum 3 Pseudognaphalium montevidense Pseudognaphalium luteoalbum 2 0.98 Pseudognaphalium californicum 1 Helichrysum confertifolium Helichrysum reflexum 1 Helichrysum mariepscopicum Helichrysum summo-montanum 1 Helichrysum formosissimum Helichrysum meyeri-johannis Pseudognaphalium ramosissimum Helichrysum stuhlmannii Helichrysum brownei 1 Helichrysum argyranthum Helichrysum chionoides Helichrysum basalticum 1 Helichrysum nicolai 1 Pseudognaphalium semiamplexicaule Helichrysum nicolai 2 1 Helichrysum alucense 1 Helichrysum monogynum 1 Helichrysum glumaceum 2 Helichrysum glumaceum 1 Helichrysum populifolium Pseudognaphalium eggersii Helichrysum trilineatum Helichrysum armenium subsp. armenium 1 Helichrysum thianschanicum Helichrysum mussae 1 Helichrysum rubicundum Helichrysum orientale Pseudognaphalium oxyphyllum var. oxyphyllum 1 1 1 Helichrysum plicatum 0.98 Helichrysum sibthorpii Helichrysum stoechas 0.96 Helichrysum pendulum 1 Helichrysum crassifolium Pseudognaphalium attenuatum 0.99 Helichrysum italicum subsp. italicum Helichrysum massanellanum 1 Helichrysum gossypinum 1 Helichrysum melaleucum 0.99 Helichrysum monizii 1 Helichrysum devium Pseudognaphalium thermale Helichrysum obconicum 1 1 Anaphalis aff. aureopunctata 1 1 Anaphalis aff. busua Anaphalis aff. aureopunctata 2 1 1 Anaphalis aff. flavescens 1 Anaphalis gracilis Pseudognaphalium canescens 0.96 Anaphalis aff. virens Anaphalis sp. 1 Anaphalis aff. nepalensis 1 1 Anaphalis aff. nepalensis 2 Anaphalis margaritacea 0.97 Helichrysum plantago Pseudognaphalium beneolens 1 Helichrysum nudifolium 1 Helichrysum miconiifolium 0.98 Helichrysum pilosellum Helichrysum globosum Helichrysum zeyheri Helichrysum psilolepis Pseudognaphalium sylvicola Helichrysum schimperi Helichrysum pagophilum Helichrysum manopappoides 1 1 Helichrysum fulvescens 1 Humeocline madagascariensis Helichrysum forsythii Pseudognaphalium oxyphyllum var. oxyphyllum 2 1 1 Helichrysum neoachyroclinoides Helichrysum cordifolium 1 Helichrysum flagellare Helichrysum triplinerve 1 Achyrocline satureioides Achyrocline alata Pseudognaphalium moritzianum Helichrysum odoratissimum 1 Helichrysum epapposum Achyrocline stenoptera 1 Helichrysum forskahlii var. compactum Helichrysum forskahlii var. forskahlii Helichrysum interjacens Pseudognaphalium hypoleucum 1 Helichrysum polycladum 1 1 Helichrysum simillinum Helichrysum melanacme 1 Helichrysum cymosum subsp. calvum Helichrysum anomalum Pseudognaphalium flavescens Helichrysum griseolanatum Helichrysum mundtii 0.99 Helichrysum auriceps 1 Helichrysum mixtum var. mixtum Helichrysum cephaloideum 1 1 Helichrysum albobrunneum Helichrysum albilanatum 0.99 Helichrysum appendiculatum 1 1 Helichrysum rosum var. arcuatum 1 Helichrysum rugulosum 1 Helichrysum rosum var. rosum Helichrysum felinum Helichrysum chrysargyrum 1 Helichrysum aureolum 1 Helichrysum petiolare 1 Helichrysum patulum 1 Helichrysum patulum 2 1 Helichrysum glomeratum Helichrysum ephelos Helichrysum truncatum Helichrysum setosum Helichrysum acutatum 1 Helichrysum milfordiae Helichrysum flanaganii 0.97 Helichrysum umbraculigerum 1 Helichrysum argyrophyllum Helichrysum aureonitens Helichrysum cooperi Helichrysum platycephalum 1 Helichrysum excisum Helichrysum silvaticum Helichrysum opacum Helichrysum herbaceum 1 Helichrysum splendidum Helichrysum elegantissimum Helichrysum montanum Helichrysum marginatum 1 Helichrysum ecklonis Helichrysum paleatum 1 Helichrysum confertum Helichrysum sutherlandii Helichrysum aureum var. aureum Helichrysum natalitium 1 Helichrysum monticola Helichrysum adenocarpum Helichrysum platypterum 1 Helichrysum mimetes 1 Helichrysum lepidissimum Helichrysum foetidum 1 Helichrysum obductum 0.98 Helichrysum homilochrysum Helichrysum lecomtei 0.99 1 Helichrysum selaginifolium 0.99 Helichrysum benthamii Helichrysum difficile Helichrysum abbayesii 1 Helichrysum cryptomerioides Helichrysum ibityense 1 Helichrysum gymnocephalum Helichrysum xylocladum 1 Helichrysum citrispinum var. citrispinum Helichrysum flammeiceps 1 Helichrysum citrispinum var. hoehnelii 0.99 1 Helichrysum gofense 2 1 Helichrysum gofense 1 Helichrysum arwae Helichrysum maranguense 0.99 Helichrysum chionosphaerum Helichrysum transmontanum Helichrysum swynnertonii 1 Helichrysum kilimanjari Helichrysum isolepis 1 Helichrysum dasycephalum 0.98 Helichrysum callicomum Helichrysum galpinii Pseudognaphalium gaudichaudianum 1 Helichrysum zwartbergense 1 Helichrysum litorale 1 Helichrysum spiralepis A.2 1 Helichrysum stellatum Helichrysum candolleanum Helichrysum argyrosphaerum 0.97 1 Pseudognaphalium melanosphaeroides 1 Helichrysum cerastioides Helichrysum asperum 1 Helichrysum indicum 1 Helichrysum crispum 0.99 Helichrysum retortum Helichrysum hamulosum Pseudognaphalium cheiranthifolium 2 Helichrysum lambertianum Lucilia acutifolia Lucilia lycopodioides Micropsis dasycarpa 0.99 Lucilia nitens 1 Facelis retusa Pseudognaphalium cheiranthifolium 1 1 Facelis plumosa Berroa gnaphalioides 1 Belloa chilensis Chionolaena campestris 1 1 Gamochaeta longipedicellata Gamochaeta americana Pseudognaphalium dombeyanum Gamochaeta subfalcata 1 Stuckertiella capitata 1 Gamochaeta serpyllifolia Gamochaetopsis alpina 1 Mniodes kunthiana Mniodes andina Pseudognaphalium viravira 1 Mniodes santanica 0.99 1 1 Mniodes longifolia 0.99 0.99 Loricaria colombiana Jalcophila boliviensis 1 Chevreulia acuminata Pseudognaphalium heterotrichum Cuatrecasasiella argentina 1 Chionolaena salicifolia Mexerion sarmentosum Antennaria dioica 1 Filago arizonica 0.99 Filago californica Pseudognaphalium leucopeplum Stylocline psilocarphoides 1 Micropus californicus 0.98 1 Logfia minima Logfia gallica Castroviejoa montelinasana Filago carpetana Pseudognaphalium lacteum 1 1 Filago congesta Filago pygmaea 1 Filago lutescens subsp. lutescens 0.99 Bombycilaena discolor Micropus supinus Omalotheca supina Pseudognaphalium graveolens Omalotheca sylvatica Leontopodium alpinum 1 Gnaphalium capense 1 Gnaphalium austroafricanum 1 1 Gnaphalium filagopsis Gnaphalium confine Pseudognaphalium badium 0.99 Gnaphalium declinatum 1 1 Pseudognaphalium aldunateoides 1 0.99 Pseudognaphalium aldunateoides 2 1 Gnaphalium palustre 1 Gnaphalium uliginosum 1 Gnaphalium uliginosum 2 Pseudognaphalium affine 1 Syncarpha mucronata 1 1 1 Syncarpha chlorochrysum Syncarpha paniculata Langebergia canescens Helichrysum dasyanthum 1 Petalacte coronata Pseudognaphalium affine 2 1 Edmondia sesamoides 1 1 Syncarpha canescens Syncarpha staehelina 1 Helichrysum lanceolatum Raoulia tenuicaulis Ewartia catipes Pseudognaphalium sandwicensium 1 Ozothamnus diosmifolius 1 1 1 Acanthocladium dockeri Craspedia glauca Vellereophyton dealbatum 1 Lachnospermum fasciculatum 1 Pseudognaphalium luteoalbum 1 Metalasia adunca 0.98 Dolichothrix ericoides 1 Ifloga spicata 1 Ifloga repens 0.98 1 Amphiglossa corrudaefolia Stoebe aethiopica Pseudognaphalium luteoalbum 3 1 Lasiopogon glomerulatus 0.99 Lasiopogon muscoides 1 Leysera gnaphalodes 1 Relhania pungens Athrixia phylicoides 3.0 Pseudognaphalium montevidense Pseudognaphalium luteoalbum 2 1 Helichrysum confertifolium Helichrysum reflexum 1 Helichrysum mariepscopicum Helichrysum summo-montanum 1 Helichrysum formosissimum Helichrysum meyeri-johannis Helichrysum stuhlmannii Helichrysum brownei 1 Helichrysum argyranthum Helichrysum chionoides Helichrysum basalticum 1 Helichrysum nicolai 1 Helichrysum nicolai 2 1 Helichrysum alucense 1 Helichrysum monogynum 1 Helichrysum glumaceum 2 Helichrysum glumaceum 1 Helichrysum populifolium Helichrysum trilineatum

3.0 Helichrysum mussae 1 Helichrysum rubicundum 1 Helichrysum orientale 1 Helichrysum plicatum 0.98 Helichrysum sibthorpii Helichrysum stoechas 0.96 Helichrysum pendulum 1 Helichrysum crassifolium 3.0 0.99 Helichrysum italicum subsp. italicum Helichrysum massanellanum Figure 2 (cont.). nrDNA (ITS and 3′ ETS regions) maximum1 credibilityHelichrysum tree obtained gossypinum from the BEAST analyses of the Gnaphalieae. Only ≥ 0.95 Bayesian posterior probabilities values are1 shown.Helichrysum Species melaleucumof the genera Pseudognaphalium and Gnaphalium are highlighted in coloured boxes. Clades discussed0.99 in the textHelichrysum are labeled monizii with capital letters and numbers. 1 Helichrysum devium

Helichrysum truncatum

Helichrysum interjacens Helichrysum rugulosum Helichrysum rosum var. rosum Anaphalis aff. virens Helichrysum maranguense Helichrysum rosum var. arcuatum Helichrysum schimperi Helichrysum pilosellum Helichrysum forskahlii var. compactum Anaphalis aff. flavescens Achyrocline satureioides Helichrysum forskahlii var. forskahlii Helichrysum griseolanatum Helichrysum anomalum Achyrocline alata Anaphalis gracilis Achyrocline stenoptera Helichrysum polycladum Helichrysum chrysargyrum Helichrysum patulum 0.98 Anaphalis aff. aureopunctata 1 Helichrysum populifolium Helichrysum homilochrysum Helichrysum nudifolium Helichrysum patulum Helichrysum miconiifolium Anaphalis sp. Helichrysum simillinum Helichrysum petiolare Helichrysum platycephalum Helichrysum odoratissimum Helichrysum epapposum Anaphalis aff. busua Helichrysum aureonitens Helichrysum melanacme Helichrysum triplinerve Helichrysum neoachyroclinoides Helichrysum flagellare Anaphalis aff. aureopunctata 2 Helichrysum fulvescens Helichrysum manopappoides 0.97 Helichrysum cordifolium Helichrysum forsythii 1 Humeocline madagascariensis 1 Helichrysum gofense 2 Helichrysum herbaceum Helichrysum mimetes Helichrysum confertifolium Helichrysum lepidissimum 1 Helichrysum appendiculatum Helichrysum gofense 1 Helichrysum cephaloideum Helichrysum auriceps Helichrysum silvaticum Helichrysum obductum Helichrysum monogynum Helichrysum citrispinum var. citrispinum Helichrysum glumaceum 1 Helichrysum alucense 1 Helichrysum glumaceum 2 Helichrysum mundtii Helichrysum glomeratum Helichrysum citrispinum var. hoehnelii 1 Helichrysum ephelos 0.97 Helichrysum dasycephalum Helichrysum callicomum Helichrysum albilanatum Anaphalis aff. nepalensis Helichrysum arwae Helichrysum excisum Helichrysum transmontanum Helichrysum setosum Anaphalis margaritacea Helichrysum trilineatum Helichrysum hamulosum Helichrysum aureum var. aureum Helichrysum cooperi Helichrysum foetidum Helichrysum splendidum Helichrysum elegantissimum Helichrysum retortum Helichrysum aureolum Helichrysum argyranthum 0.99 Helichrysum chionoides 1 Helichrysum brownei Helichrysum montanum Helichrysum thianschanicum Helichrysum marginatum Pseudognaphalium oligandrum Anaphalis aff. nepalensis Helichrysum ecklonis Pseudognaphalium oxyphyllum var. oxyphyllum 1 Helichrysum sibthorpii Pseudognaphalium oxyphyllum var. oxyphyllum 2 Pseudognaphalium ramosissimum Pseudognaphalium californicum Pseudognaphalium leucocephalum 1 Pseudognaphalium biolettii Helichrysum armenium Pseudognaphalium beneolens Pseudognaphalium macounii 0.96 Pseudognaphalium badium Pseudognaphalium cheiranthifolium 1 Pseudognaphalium obtusifolium Helichrysum orientale Pseudognaphalium thermale Pseudognaphalium heterotrichum 0.99 Pseudognaphalium lacteum 1 Helichrysum psilolepis 0.99 Helichrysum nicolai 2 Helichrysum plicatum Helichrysum nicolai 1 Helichrysum basalticum Pseudognaphalium luteoalbum 1 Helichrysum milfordiae Helichrysum rubicundum Helichrysum umbraculigerum Helichrysum albobrunneum Helichrysum argyrophyllum Helichrysum flanaganii Helichrysum confertum Helichrysum pendulum Helichrysum adenocarpum Helichrysum summo-montanum Helichrysum lambertianum Helichrysum reflexum 1 Helichrysum isolepis Helichrysum crassifolium Helichrysum kilimanjari Helichrysum gymnocephalum Helichrysum felinum Helichrysum pagophilum Helichrysum globosum Helichrysum stoechas 0.96 Helichrysum crispum Helichrysum indicum 1 Helichrysum litorale Helichrysum zwartbergense Helichrysum stellatum Helichrysum devium Helichrysum argyrosphaerum Helichrysum asperum 1 Helichrysum candolleanum Helichrysum cerastioides Helichrysum abbayesii Helichrysum monizii Helichrysum lecomtei Helichrysum benthamii Helichrysum cryptomerioides Helichrysum selaginifolium Helichrysum natalitium Helichrysum obconicum Helichrysum platypterum Helichrysum sutherlandii Helichrysum ibityense Helichrysum truncatum Anaphalis aff. virens H. italicum subsp. italicum Anaphalis aff. flavescens Anaphalis gracilis 1 0.98 Anaphalis aff. aureopunctata 1 Anaphalis sp. Anaphalis aff. busua Helichrysum gossypinum Anaphalis aff. aureopunctata 2 1 Helichrysum gofense 2 Helichrysum gofense 1 H. citrispinum var. citrispinum H. citrispinum var. hoehnelii Helichrysum melaleucum Helichrysum arwae Helichrysum hamulosum Helichrysum retortum Helichrysum thianschanicum Helichrysum sibthorpii Helichrysum massanellanum Helichrysum armenium Helichrysum orientale 0.99 Helichrysum plicatum Helichrysum rubicundum Helichrysum pendulum 1 Helichrysum formosissimum 1 Helichrysum crassifolium Helichrysum stoechas 1 Helichrysum devium Helichrysum monizii Helichrysum obconicum Helichrysum meyeri-johannis H. italicum subsp. italicum Helichrysum gossypinum Helichrysum melaleucum Helichrysum massanellanum 1 Helichrysum formosissimum Helichrysum stuhlmannii Helichrysum meyeri-johannis Helichrysum stuhlmannii Helichrysum galpinii 0.98 Petalacte coronata Helichrysum dasyanthum Helichrysum galpinii Helichrysum lanceolatum Facelis retusa Facelis plumosa Micropsis dasycarpa Gamochaeta longipedicellata 0.98 Petalacte coronata Mniodes longifolia Lucilia linearifolia Gamochaeta serpyllifolia Chionolaena campestris Helichrysum dasyanthum Jalcophila boliviensis Omalotheca supina Cuatrecasasiella argentina Gamochaetopsis alpina Luciliocline subspicata Helichrysum lanceolatum Berroa gnaphalioides Castroviejoa montelinasana Gamochaeta subfalcata Stuckertiella capitata Belloa chilensis Facelis retusa Antennaria dioica Mniodes santanica Lucilia acutifolia Gamochaeta americana Mniodes kunthiana Facelis plumosa Lucilia lycopodioides Lucilia nitens Chevreulia acuminata 1 Filago lutescens subsp. lutescens Filago carpetana Micropsis dasycarpa 1 Filago pygmaea Bombycilaena discolor Filago congesta Leontopodium alpinum 11 Filago arizonica Gamochaeta longipedicellata 0.98 Filago californica Stylocline psilocarphoides 11 Micropus californicus Logfia gallica 1 Logfia minima Mniodes longifolia Micropus supinus Loricaria colombiana Syncarpha mucronata 1 Gnaphalium uliginosum 1 1 1 1 Pseudognaphalium aldunateoides 1 Lucilia linearifolia Gnaphalium austroafricanum Vellereophyton dealbatum 1 Ifloga spicata Ifloga repens 0.99 Lachnospermum fasciculatum Gamochaeta serpyllifolia Dolichothrix ericoides Lasiopogon muscoides Relhania pungens 11 Athrixia phylicoides Leysera gnaphalodes Chionolaena campestris 3.0 Jalcophila boliviensis Omalotheca supina Cuatrecasasiella argentina Gamochaetopsis alpina Luciliocline subspicata Berroa gnaphalioides Castroviejoa montelinasana Gamochaeta subfalcata Stuckertiella capitata Belloa chilensis Antennaria dioica Mniodes santanica Lucilia acutifolia Gamochaeta americana Mniodes kunthiana Lucilia lycopodioides Lucilia nitens Chevreulia acuminata 1 Filago lutescens subsp. lutescens Filago carpetana 1 Filago pygmaea Bombycilaena discolor Filago congesta Leontopodium alpinum 1 Filago arizonica 0.98 Filago californica Stylocline psilocarphoides 1 Micropus californicus Logfia gallica 1 Logfia minima Micropus supinus Loricaria colombiana Syncarpha mucronata 1 1 Gnaphalium uliginosum 1 1 Pseudognaphalium aldunateoides 1 B 1 Gnaphalium austroafricanum Vellereophyton dealbatum 1 Ifloga spicata Ifloga repens 0.99 Lachnospermum fasciculatum Dolichothrix ericoides Lasiopogon muscoides Relhania pungens 1 Athrixia phylicoides Leysera gnaphalodes 3.0

Figure 3. cpDNA (rpl32-trnL intergenic spacer) maximum credibility tree obtained from the BEAST analyses of the Gnaphalieae. Only ≥ 0.95 Bayesian posterior probabilities values are shown. Species of the genera Pseudognaphalium and Gnaphalium are highlighted in coloured boxes. Clades discussed in the text are labeled with capital letters and numbers.

Collectanea Botanica 37: e012 (enero-diciembre 2018), e012, ISSN-L: 0010-0730, https://doi.org/10.3989/collectbot.2018.v37.012 12 A. ACOSTA-MAINDO & M. GALBANY-CASALS

Helichrysum mundtii Helichrysum glomeratum Helichrysum interjacens Helichrysum rugulosum Helichrysum rosum var. rosum Helichrysum maranguense 1 Helichrysum ephelos Helichrysum rosum var. arcuatum Helichrysum schimperi Helichrysum pilosellum 0.97 Helichrysum forskahlii var. compactum Achyrocline satureioides Helichrysum dasycephalum Helichrysum forskahlii var. forskahlii Helichrysum griseolanatum Helichrysum anomalum Achyrocline alata Achyrocline stenoptera Helichrysum callicomum Helichrysum polycladum Helichrysum chrysargyrum Helichrysum patulum Helichrysum populifolium Helichrysum homilochrysum Helichrysum albilanatum Helichrysum nudifolium Helichrysum patulum Helichrysum miconiifolium Helichrysum simillinum Helichrysum petiolare Anaphalis aff. nepalensis Helichrysum platycephalum Helichrysum odoratissimum Helichrysum epapposum Helichrysum aureonitens Helichrysum melanacme Helichrysum excisum Helichrysum triplinerve Helichrysum neoachyroclinoides Helichrysum flagellare Helichrysum fulvescens Helichrysum transmontanum Helichrysum manopappoides 0.97 Helichrysum cordifolium Helichrysum forsythii Humeocline madagascariensis 1 Helichrysum herbaceum Helichrysum setosum Helichrysum mimetes Helichrysum confertifolium Helichrysum lepidissimum 1 Helichrysum appendiculatum Helichrysum cephaloideum Anaphalis margaritacea Helichrysum auriceps Helichrysum silvaticum Helichrysum obductum Helichrysum monogynum Helichrysum glumaceum 1 Helichrysum trilineatum Helichrysum alucense Helichrysum glumaceum 2 Helichrysum mundtii Helichrysum glomeratum 1 Helichrysum ephelos 0.97 Helichrysum aureum var. aureum Helichrysum dasycephalum Helichrysum callicomum Helichrysum albilanatum Anaphalis aff. nepalensis Helichrysum excisum Helichrysum cooperi Helichrysum transmontanum Helichrysum setosum Anaphalis margaritacea Helichrysum trilineatum Helichrysum aureum var. aureum Helichrysum foetidum Helichrysum cooperi Helichrysum foetidum Helichrysum splendidum Helichrysum elegantissimum Helichrysum aureolum Helichrysum splendidum Helichrysum argyranthum 0.99 Helichrysum chionoides 1 Helichrysum brownei Helichrysum montanum Helichrysum marginatum Helichrysum elegantissimum Pseudognaphalium oligandrum Anaphalis aff. nepalensis Helichrysum ecklonis Pseudognaphalium oxyphyllum var. oxyphyllum 1 Pseudognaphalium oxyphyllum var. oxyphyllum 2 Helichrysum aureolum Pseudognaphalium ramosissimum Pseudognaphalium californicum Pseudognaphalium leucocephalum 1 Pseudognaphalium biolettii Pseudognaphalium beneolens Helichrysum argyranthum Pseudognaphalium macounii 0.96 Pseudognaphalium badium Pseudognaphalium cheiranthifolium 1 0.99 Pseudognaphalium obtusifolium Pseudognaphalium thermale Helichrysum chionoides Pseudognaphalium heterotrichum Pseudognaphalium lacteum 1 Helichrysum psilolepis 0.99 Helichrysum nicolai 2 1 Helichrysum nicolai 1 Helichrysum brownei Helichrysum basalticum Pseudognaphalium luteoalbum 1 Helichrysum milfordiae Helichrysum umbraculigerum Helichrysum albobrunneum Helichrysum montanum Helichrysum argyrophyllum Helichrysum flanaganii Helichrysum confertum Helichrysum adenocarpum Helichrysum summo-montanum Helichrysum marginatum Helichrysum lambertianum Helichrysum reflexum Helichrysum isolepis Helichrysum kilimanjari Helichrysum gymnocephalum Pseudognaphalium oligandrum A.1 Helichrysum felinum Helichrysum pagophilum Helichrysum globosum 0.96 Helichrysum crispum Anaphalis aff. nepalensis Helichrysum indicum 1 Helichrysum litorale Helichrysum zwartbergense Helichrysum stellatum Helichrysum argyrosphaerum Helichrysum ecklonis Helichrysum asperum Helichrysum candolleanum Helichrysum cerastioides Helichrysum abbayesii Helichrysum lecomtei Pseudognaphalium oxyphyllum var. oxyphyllum 1 Helichrysum benthamii Helichrysum cryptomerioides Helichrysum selaginifolium Helichrysum natalitium Helichrysum platypterum Pseudognaphalium oxyphyllum var. oxyphyllum 2 Helichrysum sutherlandii Helichrysum ibityense Helichrysum truncatum Anaphalis aff. virens Anaphalis aff. flavescens Pseudognaphalium ramosissimum Anaphalis gracilis 1 0.98 Anaphalis aff. aureopunctata 1 Anaphalis sp. Anaphalis aff. busua Anaphalis aff. aureopunctata 2 Pseudognaphalium californicum 1 Helichrysum gofense 2 Helichrysum gofense 1 H. citrispinum var. citrispinum H. citrispinum var. hoehnelii Helichrysum arwae Pseudognaphalium leucocephalum 1 Helichrysum hamulosum Helichrysum retortum Helichrysum thianschanicum Helichrysum sibthorpii Helichrysum armenium Pseudognaphalium biolettii Helichrysum orientale 0.99 Helichrysum plicatum Helichrysum rubicundum Helichrysum pendulum 1 Helichrysum crassifolium Pseudognaphalium beneolens Helichrysum stoechas 1 Helichrysum devium Helichrysum monizii A.2 Helichrysum obconicum H. italicum subsp. italicum Pseudognaphalium macounii Helichrysum gossypinum Helichrysum melaleucum Helichrysum massanellanum 1 Helichrysum formosissimum Helichrysum meyeri-johannis Pseudognaphalium badium Helichrysum stuhlmannii 0.96 Helichrysum galpinii 0.98 Petalacte coronata Helichrysum dasyanthum Helichrysum lanceolatum Pseudognaphalium cheiranthifolium 1 Facelis retusa Facelis plumosa Micropsis dasycarpa Gamochaeta longipedicellata Mniodes longifolia Pseudognaphalium obtusifolium Lucilia linearifolia Gamochaeta serpyllifolia Chionolaena campestris Jalcophila boliviensis Omalotheca supina Pseudognaphalium thermale Cuatrecasasiella argentina Gamochaetopsis alpina Luciliocline subspicata Berroa gnaphalioides Castroviejoa montelinasana Pseudognaphalium heterotrichum Gamochaeta subfalcata Stuckertiella capitata Belloa chilensis Antennaria dioica Mniodes santanica Pseudognaphalium lacteum Lucilia acutifolia Gamochaeta americana Mniodes kunthiana 1 Lucilia lycopodioides Lucilia nitens Helichrysum psilolepis Chevreulia acuminata 1 Filago lutescens subsp. lutescens Filago carpetana 1 Filago pygmaea 0.99 Helichrysum nicolai 2 Bombycilaena discolor Filago congesta Leontopodium alpinum 11 Filago arizonica 0.98 Filago californica Helichrysum nicolai 1 Stylocline psilocarphoides 11 Micropus californicus Logfia gallica 1 Logfia minima Micropus supinus Helichrysum basalticum Loricaria colombiana Syncarpha mucronata 1 1 Gnaphalium uliginosum 1 1 Pseudognaphalium aldunateoides 1 1 Gnaphalium austroafricanum Pseudognaphalium luteoalbum 1 Vellereophyton dealbatum A.3 1 Ifloga spicata Ifloga repens 0.99 Lachnospermum fasciculatum Dolichothrix ericoides Helichrysum milfordiae Lasiopogon muscoides Relhania pungens 11 Athrixia phylicoides Leysera gnaphalodes 3.0 Helichrysum umbraculigerum Helichrysum albobrunneum Helichrysum argyrophyllum Helichrysum flanaganii Helichrysum confertum Helichrysum adenocarpum Helichrysum summo-montanum Helichrysum lambertianum Helichrysum reflexum Helichrysum isolepis Helichrysum kilimanjari Helichrysum gymnocephalum Helichrysum felinum Helichrysum pagophilum Helichrysum globosum 0.96 Helichrysum crispum Helichrysum indicum 1 Helichrysum litorale Helichrysum zwartbergense Helichrysum stellatum Helichrysum argyrosphaerum Helichrysum asperum Helichrysum candolleanum Helichrysum cerastioides Helichrysum abbayesii Helichrysum lecomtei Helichrysum benthamii Helichrysum cryptomerioides Helichrysum selaginifolium Helichrysum natalitium Helichrysum platypterum Helichrysum sutherlandii Helichrysum ibityense Helichrysum truncatum Anaphalis aff. virens Anaphalis aff. flavescens Anaphalis gracilis 0.98 Anaphalis aff. aureopunctata 1 Anaphalis sp. Anaphalis aff. busua Anaphalis aff. aureopunctata 2 1 Helichrysum gofense 2 Helichrysum gofense 1 1 Helichrysum citrispinum var. citrispinum Helichrysum citrispinum var. hoehnelii Helichrysum arwae 3.0 Helichrysum hamulosum Helichrysum retortum Helichrysum thianschanicum Helichrysum sibthorpii Figure 3 (cont.). cpDNA (rpl32-trnL intergenic spacer) maximum credibilityHelichrysum tree obtained armenium from the BEAST analyses of the Gnaphalieae. Only ≥ 0.95 Bayesian posterior probabilities values are shown. Species of the genera Pseudognaphalium and Gnaphalium are highlighted in coloured boxes. Clades discussed in the text are labeled with capital letters and numbers.

Helichrysum interjacens Helichrysum rugulosum Helichrysum rosum var. rosum Helichrysum maranguense Helichrysum rosum var. arcuatum Helichrysum schimperi Helichrysum pilosellum Helichrysum forskahlii var. compactum Achyrocline satureioides Helichrysum forskahlii var. forskahlii Helichrysum griseolanatum Helichrysum interjacens Helichrysum anomalum Achyrocline alata Achyrocline stenoptera Helichrysum polycladum Helichrysum chrysargyrum Helichrysum rugulosum Helichrysum patulum Helichrysum populifolium Helichrysum homilochrysum Helichrysum nudifolium Helichrysum rosum var. rosum Helichrysum patulum Helichrysum miconiifolium Helichrysum simillinum Helichrysum petiolare Helichrysum platycephalum Helichrysum maranguense Helichrysum odoratissimum Helichrysum epapposum Helichrysum aureonitens Helichrysum melanacme Helichrysum triplinerve Helichrysum rosum var. arcuatum Helichrysum neoachyroclinoides Helichrysum flagellare Helichrysum fulvescens Helichrysum manopappoides 0.97 Helichrysum cordifolium Helichrysum schimperi Helichrysum forsythii Humeocline madagascariensis 1 Helichrysum herbaceum Helichrysum mimetes Helichrysum confertifolium Helichrysum pilosellum Helichrysum lepidissimum 1 Helichrysum appendiculatum Helichrysum cephaloideum Helichrysum auriceps Helichrysum silvaticum Helichrysum forskahlii var. compactum Helichrysum obductum Helichrysum monogynum Helichrysum glumaceum 1 Helichrysum alucense Helichrysum glumaceum 2 Achyrocline satureioides Helichrysum mundtii Helichrysum glomeratum 1 Helichrysum ephelos 0.97 Helichrysum dasycephalum Helichrysum callicomum Helichrysum forskahlii var. forskahlii Helichrysum albilanatum Anaphalis aff. nepalensis Helichrysum excisum Helichrysum transmontanum Helichrysum setosum Helichrysum griseolanatum Anaphalis margaritacea Helichrysum trilineatum Helichrysum aureum var. aureum Helichrysum cooperi Helichrysum foetidum Helichrysum anomalum Helichrysum splendidum Helichrysum elegantissimum Helichrysum aureolum Helichrysum argyranthum 0.99 Helichrysum chionoides Achyrocline alata 1 Helichrysum brownei Helichrysum montanum Helichrysum marginatum Pseudognaphalium oligandrum Anaphalis aff. nepalensis Achyrocline stenoptera Helichrysum ecklonis Pseudognaphalium oxyphyllum var. oxyphyllum 1 Pseudognaphalium oxyphyllum var. oxyphyllum 2 Pseudognaphalium ramosissimum Pseudognaphalium californicum Helichrysum polycladum Pseudognaphalium leucocephalum 1 Pseudognaphalium biolettii Pseudognaphalium beneolens Pseudognaphalium macounii Pseudognaphalium badium Helichrysum chrysargyrum 0.96 Pseudognaphalium cheiranthifolium 1 Pseudognaphalium obtusifolium Pseudognaphalium thermale Pseudognaphalium heterotrichum Pseudognaphalium lacteum Helichrysum patulum 1 Helichrysum psilolepis 0.99 Helichrysum nicolai 2 Helichrysum nicolai 1 Helichrysum basalticum Pseudognaphalium luteoalbum 1 Helichrysum populifolium Helichrysum milfordiae Helichrysum umbraculigerum Helichrysum albobrunneum Helichrysum argyrophyllum Helichrysum homilochrysum Helichrysum flanaganii Helichrysum confertum Helichrysum adenocarpum Helichrysum summo-montanum Helichrysum lambertianum Helichrysum nudifolium Helichrysum reflexum Helichrysum isolepis Helichrysum kilimanjari Helichrysum gymnocephalum Helichrysum felinum Helichrysum patulum Helichrysum pagophilum Helichrysum globosum 0.96 Helichrysum crispum Helichrysum indicum 1 Helichrysum litorale Helichrysum miconiifolium Helichrysum zwartbergense Helichrysum stellatum Helichrysum argyrosphaerum Helichrysum asperum Helichrysum candolleanum Helichrysum simillinum Helichrysum cerastioides Helichrysum abbayesii Helichrysum lecomtei Helichrysum benthamii Helichrysum cryptomerioides Helichrysum petiolare Helichrysum selaginifolium Helichrysum natalitium Helichrysum platypterum Helichrysum sutherlandii Helichrysum ibityense Helichrysum platycephalum Helichrysum truncatum Anaphalis aff. virens Anaphalis aff. flavescens Anaphalis gracilis 1 0.98 Anaphalis aff. aureopunctata 1 Helichrysum odoratissimum Anaphalis sp. Anaphalis aff. busua Anaphalis aff. aureopunctata 2 1 Helichrysum gofense 2 Helichrysum gofense 1 Helichrysum epapposum H. citrispinum var. citrispinum H. citrispinum var. hoehnelii Helichrysum arwae Helichrysum hamulosum Helichrysum retortum Helichrysum aureonitens Helichrysum thianschanicum Helichrysum sibthorpii Helichrysum armenium Helichrysum orientale 0.99 Helichrysum plicatum Helichrysum melanacme Helichrysum rubicundum Helichrysum pendulum 1 Helichrysum crassifolium Helichrysum stoechas Helichrysum devium Helichrysum triplinerve 1 Helichrysum monizii Helichrysum obconicum H. italicum subsp. italicum Helichrysum gossypinum Helichrysum melaleucum Helichrysum neoachyroclinoides Helichrysum massanellanum 1 Helichrysum formosissimum Helichrysum meyeri-johannis Helichrysum stuhlmannii Helichrysum galpinii Helichrysum flagellare 0.98 Petalacte coronata Helichrysum dasyanthum Helichrysum lanceolatum Facelis retusa Facelis plumosa Helichrysum fulvescens Micropsis dasycarpa Gamochaeta longipedicellata Mniodes longifolia Lucilia linearifolia Gamochaeta serpyllifolia Helichrysum manopappoides Chionolaena campestris Jalcophila boliviensis Omalotheca supina 0.97 Cuatrecasasiella argentina Helichrysum cordifolium Gamochaetopsis alpina Luciliocline subspicata Berroa gnaphalioides Castroviejoa montelinasana Gamochaeta subfalcata Helichrysum forsythii Stuckertiella capitata Belloa chilensis Antennaria dioica Mniodes santanica Lucilia acutifolia Humeocline madagascariensis Gamochaeta americana Mniodes kunthiana Lucilia lycopodioides 1 Lucilia nitens Chevreulia acuminata Helichrysum herbaceum 1 Filago lutescens subsp. lutescens Filago carpetana 1 Filago pygmaea Bombycilaena discolor Filago congesta Helichrysum mimetes Leontopodium alpinum 11 Filago arizonica 0.98 Filago californica Stylocline psilocarphoides 11 Micropus californicus Helichrysum confertifolium Logfia gallica 1 Logfia minima Micropus supinus Loricaria colombiana Syncarpha mucronata Helichrysum lepidissimum 1 1 Gnaphalium uliginosum 1 1 Pseudognaphalium aldunateoides 1 1 Gnaphalium austroafricanum Vellereophyton dealbatum 1 Ifloga spicata 1 Helichrysum appendiculatum Ifloga repens 0.99 Lachnospermum fasciculatum Dolichothrix ericoides Lasiopogon muscoides Relhania pungens Helichrysum cephaloideum 11 Athrixia phylicoides Leysera gnaphalodes 3.0 Helichrysum auriceps Helichrysum silvaticum Helichrysum obductum Helichrysum monogynum Helichrysum glumaceum 1 Helichrysum alucense Helichrysum glumaceum 2 Helichrysum mundtii Helichrysum glomeratum 1 Helichrysum ephelos 0.97 Helichrysum dasycephalum Helichrysum callicomum 3.0 Anaphalis aff. nepalensis Helichrysum excisum Helichrysum transmontanum Helichrysum setosum Anaphalis margaritacea Helichrysum trilineatum Helichrysum aureum var. aureum Helichrysum cooperi Helichrysum foetidum Helichrysum splendidum Helichrysum elegantissimum Helichrysum aureolum Helichrysum argyranthum 0.99 Helichrysum chionoides 1 Helichrysum brownei Helichrysum montanum Helichrysum marginatum Pseudognaphalium oligandrum Anaphalis aff. nepalensis Helichrysum ecklonis Pseudognaphalium oxyphyllum var. oxyphyllum 1 Pseudognaphalium oxyphyllum var. oxyphyllum 2 Pseudognaphalium ramosissimum Pseudognaphalium californicum Pseudognaphalium leucocephalum 1 Pseudognaphalium biolettii Pseudognaphalium beneolens Pseudognaphalium macounii 0.96 Pseudognaphalium badium Pseudognaphalium cheiranthifolium 1 Pseudognaphalium obtusifolium Figure 3 (cont.). cpDNA (rpl32-trnL intergenic spacer) maximum credibilityPseudognaphalium tree obtained from thermale the BEAST analyses of the Pseudognaphalium heterotrichum Gnaphalieae. Only ≥ 0.95 Bayesian posterior probabilities values are shown.Pseudognaphalium Species of the genera lacteum Pseudognaphalium and Gnaphalium are highlighted in coloured boxes. Clades1 discussed in the textHelichrysum are labeled withpsilolepis capital letters and numbers. 0.99 Helichrysum nicolai 2 Helichrysum nicolai 1

Collectanea Botanica 37: e012 (enero-diciembre 2018), e012, ISSN-L: 0010-0730, https://doi.org/10.3989/collectbot.2018.v37.012 14 A. ACOSTA-MAINDO & M. GALBANY-CASALS

1 2

3 4

5 6

Figure 4. Pappus bristles showing tip cells. Left column: pappus bristle of hermaphroditic florets. Right column: pappus bristle of pistillate florets. (1) and (2) Pseudognaphalium aldunateoides (Galbany 2525 & Arrabal, BC); (3) and (4) Gnaphalium uliginosum (Galbany 2090 & Arrabal, BC); (5) and (6) Pseudognaphalium oxyphyllum (Galbany 2480 & Arrabal, BC). Scale bar 1–6 = 40 µm.

Figure 5. Involucral bracts showing the stereome: (1) Pseudognaphalium aldunateoides (Galbany 2525 & Arrabal, BC); (2) Gnaphalium uliginosum (Galbany 2090 & Arrabal, BC); (3) Pseudognaphalium oxyphyllum (Galbany 2480 & Arrabal, BC). Scale bar 1–3 = 100 µm.

Collectanea Botanica 37: e012 (enero-diciembre 2018), e012, ISSN-L: 0010-0730, https://doi.org/10.3989/collectbot.2018.v37.012 16 A. ACOSTA-MAINDO & M. GALBANY-CASALS

Regarding pappus morphology, our morpholog- it is multistemmed, branched from the base and 6 ical study shows that the pappus of P. aldunateoi- to 10 cm high. These two species were presented des is always dimorphic. While this character is separately in the morphological study of Monti et variable in Gnaphalium (Hilliard & Burtt, 1981), al. (2013), in which they were both transferred to the pappus is always monomorphic in the genus Pseudognaphalium. In that work, the studied spec- Pseudognaphalium (Anderberg, 1991). Monti et imens of G. perpusillum were reported to have a al. (2013) already noticed the dimporphic pappus dimporphic pappus and a fenestrated stereome, but of P. aldunateoides, but they considered it to be an the figure showed it has intermediate characteris- exception in Pseudognaphalium. tics, being similar to that of G. aldunateoides. Later Considering the phylogenetic position of P. al- on, Freire et al. (2014a) stated that the diagnostic dunateoides within the Gnaphalium clade, both in characters of G. perpusillum are variable features the nrDNA (Fig. 2, clade B) and the cpDNA (Fig. 3, throughout the range of G. aldunateoides and syn- clade B) phylogenetic trees, it is clear that this species onymised both taxa under the latter. With the evi- should be classified as a member of Gnaphalium. In dence of morphological characters, P. perpusillum the nrDNA tree (Fig. 2), Gnaphalium is composed of should be also placed under Gnaphalium as G. per- two main clades, one constituted by all the African pusillum, if considered as a separate species. species included—from Gnaphalium capense Hil- liard to Gnaphalium declinatum L. f.—and the other ACKNOWLEDGMENTS constituted by European and American species. With- in the latter, G. aldunateoides is sister to Gnaphalium M. G.-C. greatly appreciates S. Arrabal’s help, enthusiastic support, dedication and joy in all the shared field expeditions. palustre Nutt., a North American species. Although O. Hinojosa, D. Juárez and S. Teillier provided very valuable there seems to be a geographical pattern in the phy- help and advice for field trips and plant collections and they are logenetic relationships obtained, the sampling of the gratefully acknowledged. We thank S. Magallón for organising genus is too scarce to venture any conclusions. the stay of M. G.-C. at the “Universidad Nacional Autónoma de México”. We thank the staff of the herbaria BC (N. Nua- Our results confirm the predictive value of mor- lart), CONC (A. Marticorena), MEXU (D. Gernandt and L. phological characters in the generic delimitation Villaseñor) and SGO (G. Rojas) for kindly receiving M. G.-C., of Gnaphalieae, as Hilliard & Burtt (1981) already providing specimens for this study and/or institutional support for plant collections. L. Guàrdia provided the microscope highlighted with insight, taking into account that and camera and technical support necessary for using them. J. they did not counted with molecular phylogenies Requena provided technical support in the lab. P. Carnicero at that time. Moreover, our results also agree with provided helpful advice for phylogenetic analyses. G. Acos- Anderberg’s (1991) proposal based on morpholog- ta and M. Balaguer provided informatics support for running phylogenetic analyses. We thank S. Freire and C. Monti for ical characters, in which more than 70 species were critically reading the manuscript and providing interesting dis- transferred from Gnaphalium to Pseudognaphalium cussion. Two anonymous reviewers contributed to improve the but P. aldunateoides was left as G. aldunateoides. paper. M. G.-C. benefitted from a “Systematics Research fund” from the Systematics Association and the Linnean Society of However, we show that the stereome can sometimes London and a “Beca Iberoamérica Jóvenes Profesores Inves- be variable within a species, and present interme- tigadores” from “Banco Santander”, which partly financed the diate characteristics that can make it difficult to be stays and field work in Chile and Mexico, respectively. The assigned to one of the two described states. In con- Catalan government (“Ajuts a grups consolidats” 2014-SGR 514) partly financed this work. clusion, this character should always be examined in several specimens, and should no be used alone for REFERENCES generic delimitation, but in combination with other morphological characters and with molecular data. Akaike, H. 1973. Information theory as an extension of the maxi- Gnaphalium perpusillum Phil. was described on mum likelihood principle. In: Petrov, B. N. & Csaki, F. 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Appendix 1. Species included in molecular anal- HG797723, HG797987, HG798144. Antennaria yses with voucher information and INSDC (Inter- dioica (L.) Gaertn.: Spain, Huesca, Santos-Vicente national Nucleotide Sequence Database Collabo- et al. MS 428 (SALA); FN645833, FN645610, ration) accession numbers (ITS, ETS, rpl32-trnL FN649336. Athrixia phylicoides DC.: South Africa, intergenic spacer). Newly generated sequences Eastern Cape Province, Romo 14395 et al. (BC); FN645816, FN645634, FN649330. Belloa chilen- are marked with an asterisk (*), hyphens (-) in- sis (Hook. & Arn.) J. Rémy: Argentina, Rio Negro, dicate missing sequences. Acronyms of the her- Bariloche, PN Nahuel Huapi, Co. Tronador, C. baria follow Index Herbariorum (Thiers, 2017). Ezcurra 2262 (BCRU); KM091388, KM091349, KM091343. Berroa gnaphalioides (Less.) Beau- Acanthocladium dockeri F. Muell.: Australia, Bayer verd: Uruguay, San José de Mayo, Kiyú, 17-12- 99011 (SA); JQ95655, JQ956648, -. Achyrocline 2008, Urtubey et al. 391 (SI); KM091386, alata (Kunth) DC.: Brasil, Rio Grande du Sul, São KM091355, KM091325. Bombycilaena discolor Francisco de Paula, Grazziotin 8941 et al. (W 1995- (Pers.) M. Laínz: Spain, Zamora, Cañizal, 4937); HG797714, HG797978, HG798135. Achy- Martínez-Ortega 1819 & Andrés-Sánchez (SALA rocline satureioides (Lam.) DC.: Argentina, Entre 134225), FN645843, FN645560, FN649364. Cas- Ríos, Departamento Colón, D. G. Gutiérrez 658 troviejoa montelinasana (Schmid) Galbany, L. (LP); HG797713, HG797976, HG798133. Achyro- Sáez & Benedí: Italy, Sardinia, Monte Línas, Gal- cline stenoptera (DC.) Hilliard & B. L. Burtt: Tanza- bany & Sáez s. n. (BCN 4644); AY445229, nia, Olmoti, Galbany & Arrabal s. n. (BC 867819); FN645559, FN649341. Chevreulia acuminata FJ211449 and FJ211507, HG797977, HG798134. Less.: Argentina, Jujuy, Palpalá, Villa de Las Ser- Amphiglossa corrudaefolia DC.: South Africa, ranías de Zapla, Urtubey & Freire 425 (SI); Prince Albert, Koekemoer 1291 (BOL); JF911388, KM091392, KM091361, KM091322. Chionolaena FM173124, - . Anaphalis aff. aureopunctata Lin- campestris Deble: Brasil, Minas Gerais, Parque Es- gelsh. & Borza (1): China, Sichuan Province, Bouf- tadual de Ibitipoca, R. Mello-Silva et al. 3257 (SPF); ford 35751 et al. (HUH); HG797715, HG797979, KM091407, KM091377, KM091338. Chionolaena HG798136. Anaphalis aff. aureopunctata Lingelsh. salicifolia (Bertol.) G. L. Nesom: Mexico, Oaxaca, & Borza (2): China, Sichuan Province, Boufford Ernst 2764 (US); KT865467, KT865263, - . Cras- 32622 et al. (HUH); HG797716, HG797980, pedia glauca Spreng.: Australia, Tasmania, Eagle- HG798137. Anaphalis aff. busua DC.: China, Sich- hawk Neck, Ford & Purves 21/03 (CHR 565520); uan Province, Boufford 37045 et al. (HUH); EF187655, EF187629, -. Cuatrecasasiella argenti- HG797717, HG797981, HG798138. Anaphalis aff. na (Cabrera) H. Rob.: Bolivia, La Paz, Tamayo, Ul- flavescens Hand.-Mazz.: China, Xizang (Tibet) la-Ulla, reserva de Apolobamba, Co. Puntani, Ur- Province, Boufford 31246 et al. (HUH); HG797718, tubey et al. 497 (SI); KM091401, KM091366, HG797982, HG798139. Anaphalis aff. nepalensis KM091337. Dolichothrix ericoides (Lam.) Hilliard (Spreng.) Hand.-Mazz. (1): China, Sichuan Prov- & B. L. Burtt: South Africa, Western Cape Prov- ince, Boufford 36783 et al. (HUH); HG797720, ince, Romo 14514 et al. (BC); FN645828, HG797984, HG798141. Anaphalis aff. nepalensis FN645622, FN649332. Edmondia sesamoides (L.) (Spreng.) Hand.-Mazz. (2): China, Xizang (Tibet) Hilliard: South Africa, Jonaskop, Bergh 1130 Province, Boufford 31830 et al. (HUH); HG797721, (NBG); JF893882, FM173129, - . Ewartia catipes HG797985, HG798142. Anaphalis aff. virens C. C. Beauverd: Australia, Tasmania, Ben Lomond, Ward Chang: China, Xizang (Tibet) Province, Boufford 94098/9 (CANU 37226); U95290, FJ404694, - . 31568 et al. (HUH); HG797722, HG797986, Facelis plumosa (Wedd.) Sch. Bip.: Bolivia, La HG798143. Anaphalis gracilis Hand.-Mazz.: Chi- Paz, Murillo, Camino antiguo a Unduavi, subiendo na, Sichuan Province, Boufford 34675 et al. (HUH); el valle de Chuquiaguillo, Urtubey et al. 470 (SI); HG797719, HG797983, HG798140. Anaphalis KM091394, KM091372, KM091345. Facelis retu- margaritacea (L.) Benth. & Hook. f.: Canada, J. M. sa (Lam.) Sch. Bip.: Argentina, Córdoba, Cala- Blanco & E. Blanco s. n. (BC); FN645827, muchita, Urtubey & Baztarrica 202 (SI); KM091385, FN645632, FN649352. Anaphalis sp.: China, Sich- KM091352, KM091321. Filago arizonica A. Gray: uan Province, Boufford 32750 et al. (HUH); Mexico, Baja California, Valle Las Palmas, Cerro

Collectanea Botanica 37: e012 (enero-diciembre 2018), e012, ISSN-L: 0010-0730, https://doi.org/10.3989/collectbot.2018.v37.012 20 A. ACOSTA-MAINDO & M. GALBANY-CASALS

Bola, Boyd 10377 et al. (RSA 657500); FN645839, Zwettl, Wallnofer 13757 (US); KT865482, FN645615, FN649343. Filago californica Nutt.: KT865288, - . Helichrysum abbayesii Humbert: United States of America, Esmeralda Co., Tule Can- Madagascar, Fianarantsoa Province, Maharumona yon, Tiehm 14663 (RSA 713363); FN645840, road, Bayer MAD-04026 et al. (CANB 660363); FN645616, FN649344. Filago carpetana (Lange) HG797724, HG797988, HG798145. Helichrysum Chrtek & Holub; Spain, Cáceres, Logrosán, Las acutatum DC.: South Africa, Mpumalanga Prov- Chamizas, Santos-Vicente 566 et al. (SALA ince, Romo 14596 et al. (BC 867799); FJ211457 134319); FN645858, FN645568, FN649372. Fila- and FJ211515, HG797989, - . Helichrysum adeno- go congesta DC.: Spain, Granada, Baza, An- carpum DC.: South Africa, Mpumalanga Province, drés-Sánchez 52 & Martínez-Ortega (SALA 134203), Romo 14611 et al. (BC 867814); HG797725, FN645848, FN645577, FN649382. Filago lute- HG797990, HG798146. Helichrysum albilanatum scens Jord. subsp. lutescens: Spain, Ávila, Nava- Hilliard: South Africa, Mpumalanga Province, cepeda de Tormes, Martínez-Ortega 1829 (SALA Romo 14603 et al. (BC 867806); HG797726, 134165); FN645882, FN645596, FN649396. Fila- HG797991, HG798147. Helichrysum albobrunne- go pygmaea L.: Spain, Badajoz, Monesterio, San- um S. Moore: Lesotho, ex Roy. Bot. Gard. Kew tos-Vicente 563 et al. (SALA 134315); FN645868, (BCN 6097); AY445215, HG797992, HG798148. FN645574, FN649379. Gamochaeta americana Helichrysum alucense García-Cas., S. Scholz & E. (Mill.) Wedd.: Argentina, Neuquén, Aluminé, cami- Hernández: Spain, Canary Islands, La Gomera, no de Moquehue a Norquinco, Urtubey 410 et al. García Casanova s. n. (TFC 36682); AY445223, (SI); Argentina, Buenos Aires, La Plata, Urtubey HG797993, HG798149. Helichrysum anomalum 187 (LP); KM091411, KM091382, KM091348. Less.: South Africa, Eastern Cape Province, Romo Gamochaeta longipedicellata Cabrera: Bolivia, La 14463 et al. (BC 867691); HG797727, HG797994, Paz, Murillo, Urtubey 473 et al. (SI); KM091410, HG798150. Helichrysum appendiculatum (L. f.) KM091381, KM091347. Gamochaeta serpyllifolia Less.: South Africa, Kwazulu-Natal Province, Romo Wedd.: Chile, XIV Región de Los Ríos, Valdivia, 14362 et al. (BC 867635); HG797728, HG797995, comuna de Panguipulli, faldeos Volcán Chosuenco, HG798151. Helichrysum argyranthum O. Hoffm.: M. Baeza 4357 (CONC); KM091409, KM091380, Tanzania, Olmoti, Galbany & Arrabal s. n. (BC KM091346. Gamochaeta subfalcata (Cabrera) 867822); FJ211465 and FJ211523, HG797996, Cabrera: Spain, Girona, between Mollet de Perelada HG798152. Helichrysum argyrophyllum DC.: and St. Climent, Galbany et al. s. n. (BCN); FN645834, South Africa, Eastern Cape Province, Romo 14472 FN645557, FN649338. Gamochaetopsis alpina et al. (BC 867699); HG797729, HG797997, (Poepp.) Anderb. & Freire: Argentina, Río Negro, HG798153. Helichrysum argyrosphaerum DC.: Bariloche, Co. Tronador, N. Bayón 1339 (LPAG); South Africa, Free State Province, Koekemoer 3532 KM091390, KM091356, KM091326. Gnaphalium (BC); HG797730, HG797998, HG798154. Heli- austroafricanum Hilliard: South Africa, Kwazu- chrysum armenium DC. subsp. armenium: Tur- lu-Natal Province, Romo 14365 et al. (BC); key, Adiyaman, Susanna 2346 et al. (BCN 6127); FN645830, FN645630, FN649353. Gnaphalium AY445208, FJ211577, HG798155. Helichrysum capense Hilliard: South Africa, Western Cape, arwae J. R. I. Wood: Yemen, ex Roy. Bot. Gard. Koekemoer 3117 (US); KT865479, KT865281, - . Kew (BCN 6103); AY445219, HG797999, Gnaphalium confine Harv.: Sudwest Afrika, Volk HG798156. Helichrysum asperum (Thunb.) Hil- 667 (US); KT865480, KT865282, - . Gnaphalium liard & B. L. Burtt var. albidulum (DC.) Hilliard: declinatum L. f.: South Africa, Western Cape, Elim, South Africa, Western Cape Province, Romo 14526 F, N. G. Bergh 1073; - , FR821617, - . Gnaphalium et al. (BC 867744); FJ211470 and FJ211528, filagopsis Hilliard & B. L. Burtt: Namibia, Seydel HG798000, HG798157. Helichrysum aureolum 2674 (US); - , KT865283, - . Gnaphalium palustre Hilliard: South Africa, Mpumalanga Province, Nutt.: United States of America, Nevada, Arnold Koekemoer 3494 (BC); HG797731, HG798001, 11941 (US); KT865481, KT865285, - . Gnaphali- HG798158. Helichrysum aureonitens Sch. Bip.: um uliginosum L. (1): Armenia, Shirak Province, South Africa, Mpumalanga Province, Romo 14579 Vitek et al. s. n. (BCN 39933); FN645823, FN645624, et al. (BC 867786); HG797732, HG798002, FN649359. Gnaphalium uliginosum L. (2): Austria, HG798159. Helichrysum aureum (Houtt.) Merrill

Collectanea Botanica 37: e012 (enero-diciembre 2018), e012, ISSN-L: 0010-0730, https://doi.org/10.3989/collectbot.2018.v37.012 Pseudognaphalium aldunateoides back in Gnaphalium (Compositae: Gnaphalieae) 21 var. aureum: South Africa, Eastern Cape Province, DC.: Madagascar, Antananarivo Province, Bayer Romo 14414 et al. (BC 867667); HG797733, MAD-04003 et al. (CANB 660340); HG797744, HG798003, HG798160. Helichrysum auriceps Hil- HG798020, HG798176. Helichrysum crassifolium liard: South Africa, Kwazulu-Natal Province, Romo (L.) D. Don: Spain, Balearic Islands, Majorca, Gal- 14371 et al. (BC 867644); HG797734, HG798004, bany & Sáez s. n. (BCN 6117); AY445190, HG798161. Helichrysum basalticum Hilliard: Le- FJ211540, HG798177. Helichrysum crispum (L.) sotho, ex Roy. Bot. Gard. Kew (BCN 6095); D. Don: South Africa, Western Cape Province, AY445211, HG798005, HG798162. Helichrysum Romo 14532 et al. (BC 867748); HG797746, benthamii R. Vig. & Humbert: Madagascar, Fian- HG798022, HG798179. Helichrysum cryptomer- arantsoa Province, Massif de Itremo, Bayer MAD- ioides Baker: Madagascar, Fianarantsoa Province, 04031 et al. (CANB 660368); HG797735, Massif de Itremo, Bayer MAD-04045 et al. (CANB HG798006, HG798163. Helichrysum brownei S. 660382); HG797745, HG798021, HG798178. He- Moore: Kenya, Mount Kenya, Galbany & Arrabal lichrysum cymosum (L.) D. Don subsp. calvum Hil- s. n. (BC 867843); FJ211464 and FJ211522, liard: South Africa, Eastern Cape Province, Romo HG798007, HG798164. Helichrysum callicomum 14410 et al. (BC 867663); HG797748, HG798024, Harv.: South Africa, Free State Province, Romo - . Helichrysum dasyanthum (Willd.) Sweet: Ex J. 14342 et al. (BC 867623); FJ211473 and FJ211531, Bot. Mar i Murtra, Blanes (BCN 6107); AY445226, HG798008, HG798165. Helichrysum candollea- HM450869, HG798181. Helichrysum dasycepha- num Buek: Mozambique, Gaza Province, Burrows lum O. Hoffm.: South Africa, Eastern Cape Prov- 8560 (Buffelskloof herb.); HG797736, HG798009, ince, Romo 14411 et al. (BC 867664); HG797749, HG798166. Helichrysum cephaloideum DC.: HG798025, HG798182. Helichrysum devium J. Y. South Africa, Mpumalanga Province, Romo 14578 Johnson: Portugal, Madeira Island, Jardim s. n. et al. (BC 867785); HG797737, HG798010, (MADJ); FJ211441 and FJ211499, FJ211579, HG798167. Helichrysum cerastioides DC.: South HG798183. Helichrysum difficile Hilliard: South Africa, Mpumalanga Province, Burrows 8508 (Buf- Africa, Mpumalanga Province, Romo 14588B et al. felskloof herb.); HG797738, HG798011, HG798168. (BC 867793); HG797750, HG798026, - . Helichry- Helichrysum chionoides Philipson: Kenya, Mount sum ecklonis Sond.: South Africa, Eastern Cape Kenya, Galbany & Arrabal s. n. (BC 867840); Province, Romo 14485 et al. (BC 867712); FJ211463 and FJ211521, HG798012, HG798169. HG797751, HG798027, HG798184. Helichrysum Helichrysum chionosphaerum DC.: South Africa, elegantissimum DC.: South Africa, Eastern Cape Eastern Cape Province, Romo 14450 et al. (BC Province, Romo 14435 et al. (BC 867676); 867682); FJ211461 and FJ211519, HG798013, - . HG797752, HG798028, HG798185. Helichrysum Helichrysum chrysargyrum Moeser: South Africa, epapposum H. Bol.: South Africa, Mpumalanga Mpumalanga Province, McMurtry 8871 (Buffelsk- Province, Romo 14581 et al. (BC 867788); FJ211459 loof herb.); HG797739, HG798014, HG798170. and FJ211517, HG798029, HG798186. Helichry- Helichrysum citrispinum Del. var. citrispinum: sum ephelos Hilliard: South Africa, Mpumalanga Ethiopia, Mount Choke, Aldasoro 9952 & Alarcón Province, Romo 14601 et al. (BC 867804); (BC); HG797740, HG798015, HG798171. Heli- HG797753, HG798030, HG798187. Helichrysum chrysum citrispinum Del. var. hoehnelii (Sch- excisum (Thunb.) Less.: South Africa, Western weinf.) Hedberg: Ethiopia, Bale Mountains plateau, Cape Province, Koekemoer 3433 (BC); HG797754, Aldasoro 10378 & Alarcón (BC); HG797741, HG798031, HG798188. Helichrysum felinum Less.: HG798016, HG798172. Helichrysum confertifoli- South Africa, Eastern Cape Province, Romo 14469 et um Klatt: South Africa, Mpumalanga Province, al. (BC 867696); HG797755, HG798032, HG798189. Romo 14599 et al. (BC 867802); HG797742, Helichrysum flagellare Bak: Madagascar, Anta- HG798017, HG798173. Helichrysum confertum nanarivo Province, Bayer MAD04004 et al. (CANB N. E. Br.: Ex Roy. Bot. Gard. Kew (BCN 6096); 660341); HG797756, HG798033, HG798190. He- AY445214, HG798018, HG798174. Helichrysum lichrysum flammeiceps Brenan: Malawi, Nyika cooperi Harv.: South Africa, Free State Province, National Park, Koekemoer 1854 (BC); HG797757, Romo 14345 et al. (BC 867626); HG797743, HG798034, - . Helichrysum flanaganii H. Bol.: HG798019, HG798175. Helichrysum cordifolium South Africa, Eastern Cape Province, Romo 14424

Collectanea Botanica 37: e012 (enero-diciembre 2018), e012, ISSN-L: 0010-0730, https://doi.org/10.3989/collectbot.2018.v37.012 22 A. ACOSTA-MAINDO & M. GALBANY-CASALS et al. (BC 867673); HG797758, HG798035, HG797766, HG798050, HG798207. Helichrysum HG798191. Helichrysum foetidum (L.) Moench: hamulosum DC.: South Africa, Western Cape Prov- Ex Dresden Bot. Gard. (BCN 8219); AY445221, ince, Romo 14540 et al. (BC 867751); HG797767, HG798036, HG798192. Helichrysum formosissi- HG798051, HG798208. Helichrysum herbaceum mum (Sch. Bip.) A. Rich.: Kenya, Mount Kenya, (Andr.) Sweet: South Africa, Kwazulu-Natal Prov- Galbany & Arrabal s. n. (BC 867832); FJ211446 ince, Romo 14369 et al. (BC 867642); HG797768, and FJ211504, HG798037, HG798193. Helichry- HG798052, HG798209. Helichrysum homilochrysum forskahlii (J. F. Gmel.) Hilliard & B. L. Burtt sum S. Moore: South Africa, Mpumalanga Prov- var. compactum (Vatke) Mesfin: Kenya, Mount ince, Burrows 7748 (Buffelskloof herb.); HG797769, Kenya, Galbany & Arrabal s. n. (BC 867830); HG798053, HG798210. Helichrysum ibityense HG797759, HG798038, HG798194. Helichrysum Humbert: Madagascar, Fianarantsoa Province, Am- forskahlii (J. F. Gmel.) Hilliard & B. L. Burtt var. batosira, Bayer MAD-04052 et al. (CANB 660389); forskahlii: Tanzania, Empaakai, Galbany & Arrabal HG797770, HG798054, HG798211. Helichrysum s. n. (BC 867823); FJ211447 and FJ211505, indicum (L.) Grierson: South Africa, Western Cape HG798039, HG798195. Helichrysum forsythii Province, Romo 14547 et al. (BC 867758); Humbert: Madagascar, Fianarantsoa Province, 30 HG797771, HG798055, HG798212. Helichrysum km from Ivato, Bayer MAD-04021 et al. (CANB interjacens Hilliard: South Africa, Mpumalanga 660358); HG797760, HG798040, HG798196. Heli- Province, Romo 14570 et al. (BC 867777); chrysum fulvescens DC.: Madagascar, Antananari- HG797772, HG798056, HG798213. Helichrysum vo Province, Bayer MAD04001 et al. (CANB isolepis H. Bol.: South Africa, Eastern Cape Prov- 660338); HG797761, HG798041, HG798197. Heli- ince, Romo 14477 et al. (BC 867704); HG797773, chrysum galpinii N. E. Br.: South Africa, Mpuma- HG798057, HG798214. Helichrysum italicum langa Province, Romo 14569 et al. (BC 867776); (Roth) G. Don subsp. italicum: Bosnia-Herzegovi- HG797762, HG798042, HG798198. Helichrysum na, Herzegovina, Redžić et al. s. n. (BCN 20756); globosum A. Rich.: Kenya, Mount Kenya, Galbany FJ211422 and FJ211480, FJ211548, HG798215. & Arrabal s. n. (BC 867838); FJ211454 and Helichrysum kilimanjari Oliv.: Kenya, Mount FJ211512, HG798043, HG798199. Helichrysum Kenya, Galbany & Arrabal s. n. (BC 867836); glomeratum Klatt: South Africa, Kwazulu-Natal FJ211453 and FJ211511, HG798058, HG798217. Province, Romo 14390 et al. (BC 867657); FJ211474 Helichrysum lambertianum DC.: South Africa, and FJ211532, HG798044, HG798200. Helichry- Western Cape Province, Romo 14556 et al. (BC sum glumaceum DC. (1): Oman, Musadam, Mc- 867767); FJ211472 and FJ211530, HG798059, Leish 3727 (E 121502); FJ211469 and FJ211527, HG798218. Helichrysum lanceolatum (Buchanan) HG798045, HG798201. Helichrysum glumaceum Kirk: New Zealand, Nelson, Pelorus Valley, Glenny DC. (2): Ethiopia, SW of Negele, Aldasoro 10262 & 8969 (CHR 574730); HM244720, HM450855, Alarcón (BC); HG797763, HG798046, HG798202. HG798219. Helichrysum lecomtei R. Vig. & Hum- Helichrysum gofense Cufod. (1): Ethiopia, Bale bert: Madagascar, Fianarantsoa Province, Anala- Mountains plateau, Aldasoro 10336 & Alarcón may, Bayer MAD-04016 et al. (CANB 660353); (BC); HG797764, HG798047, HG798203. Heli- HG797774, HG798060, HG798220. Helichrysum chrysum gofense Cufod. (2): Ethiopia, Bale Moun- lepidissimum S. Moore: South Africa, Mpumalanga tains plateau, Aldasoro 10376 & Alarcón (BC); Province, Burrows s. n. (Buffelskloof herb.); HG797765, HG798048, HG798204. Helichrysum HG797775, HG798061, HG798221. Helichrysum gossypinum Sch. Bip.: Spain, Canary Islands, Lan- litorale H. Bol.: South Africa, Eastern Cape Prov- zarote, Galbany & Arrabal s. n. (BCN 25226); ince, Romo 14500 et al. (BC 867722); HM244706, FJ211440 and FJ211498, FJ211578, HG798205. HG798062, HG798222. Helichrysum manopap- Helichrysum griseolanatum Hilliard: South Africa, poides Humbert: Madagascar, Bayer MAD-04023 et Eastern Cape Province, Romo 14413 et al. (BC al. (CANB 660360); HG797776, HG798063, 867666); FJ211476 and FJ211534, HG798049, HG798223. Helichrysum maranguense O. Hoffm.: HG798206. Helichrysum gymnocephalum (DC.) Tanzania, Empaakai, Galbany & Arrabal s. n. (BC Humbert: Madagascar, Antananarivo Province, Mt. 867825); FJ211452 and FJ211510, HG798064, Ibity, Bayer MAD-04053 et al. (CANB 660390); HG798224. Helichrysum marginatum DC.: South

Africa, Eastern Cape Province, Romo 14434 et al. Mt. Ibity, Bayer MAD-04067 et al. (CANB (BC 867675); FJ211460 and FJ211518, HG798065, 660404); HG797785, HG798077, HG798237. He- HG798225. Helichrysum mariepscopicum Hil- lichrysum nicolai N. Kilian, Galbany & Oberpr. liard: South Africa, Mpumalanga Province, Romo (1): Cape Verde, São Nicolau, Alto das Cabaças, 14592 et al. (BC 867795); HG797777, HG798066, Galbany 2111-3 & Molero (BC); FN691030, - . Helichrysum massanellanum Herrando, J. M. HG798078, HG798238. Helichrysum nicolai N. Blanco, L. Sáez & Galbany: Spain, Balearic Is- Kilian, Galbany & Oberpr. (2): Cape Verde, São lands, Majorca, Galbany & Sáez s. n. (BCN 6115); Nicolau, Alto das Cabaças, Galbany 2111-17 & AY445195, FJ211546, HG798216. Helichrysum Molero (BC); FN691031, HG798079, HG798239. melaleucum Rchb. ex Holl: Portugal, Madeira Is- Helichrysum nudifolium (L.) Less.: Kenya, Mount land, Jardim s. n. (MADJ); FJ211443 and FJ211501, Kenya, Galbany & Arrabal s. n. (BC 867834); FJ211581, HG798226. Helichrysum melanacme FJ211456 and FJ211514, HG798080, HG798240. DC.: South Africa, Free State Province, Romo Helichrysum obconicum DC.: Portugal, Madeira 14334 et al. (BC 867616); HG797778, HG798067, Island, Jardim s. n. (MADJ); FJ211442 and HG798227. Helichrysum meyeri-johannis Engl.: FJ211500, FJ211580, HG798241. Helichrysum Kenya, ex Roy. Bot. Gard. Kew (BCN 6104); obductum H. Bol.: South Africa, Mpumalanga AY445216, HG798068, HG798228. Helichrysum Province, Romo 14573 et al. (BC 867780); miconiifolium DC.: South Africa, Free State Prov- HG797786, HG798081, HG798242. Helichrysum ince, Romo 14348 et al. (BC 867629); HG797779, odoratissimum (L.) Sweet: Tanzania, Olmoti, Gal- HG798069, HG798229. Helichrysum milfordiae bany & Arrabal s. n. (BC 867820); FJ211448 and Killick: Lesotho, ex Roy. Bot. Gard. Kew (BCN FJ211506; HG798082, HG798243. Helichrysum 6101); AY445212, FJ211537, HG798230. Heli- opacum Klatt: South Africa, Mpumalanga Prov- chrysum mimetes S. Moore: South Africa, Mpum- ince, Romo 14593 et al. (BC 867796); HG797787, alanga Province, Romo 14610 et al. (BC 867813); HG798083, - . Helichrysum orientale (L.) Gaertn.: FJ211462 and FJ211520, HG798070, HG798231. Greece, Crete, ex Roy. Bot. Gard. Kew (BCN Helichrysum mixtum (O. Kuntze) Moeser var. 6098); AY445205, FJ211567, HG798244. Heli- mixtum: South Africa, Kwazulu-Natal Province, chrysum pagophilum M. D. Hend.: Lesotho, ex Romo 14374 et al. (BC 867647); HG797780, Roy. Bot. Gard. Kew (BCN 6100); AY445217, HG798071, - . Helichrysum monizii Lowe: Portu- HG798084, HG798245. Helichrysum paleatum gal, Madeira Island, Jardim s. n. (MADJ); FJ211444 Hilliard: South Africa, Kwazulu-Natal Province, and FJ211502, FJ211582, HG798232. Helichry- Romo 14385 et al. (BC 867652); HG797788, sum monogynum B. L. Burtt & Sunding: Spain, HG798085, - . Helichrysum patulum (L.) D. Don Canary Islands, Lanzarote, Galbany & Arrabal (1): South Africa, Western Cape Province, Romo s. n. (BCN 25227); FJ211468 and FJ211526, 14507 et al. (BC 867729); HG797789, HG798086, HG798072, HG798233. Helichrysum montanum HG798246. Helichrysum patulum (L.) D. Don (2): DC.: South Africa, Kwazulu-Natal Province, Romo South Africa, Western Cape Province, Romo 14527 14392 et al. (BC 867659); HG797781, HG798073, et al. (BC 867745); HG797790, HG798087, HG798234. Helichrysum monticola Hilliard: HG798247. Helichrysum pendulum (C. Presl) C. South Africa, Mpumalanga Province, Romo 14591 Presl: Spain, Balearic Islands, Ibiza, Sáez s. n. et al. (BC 867794); HG797782, HG798074, - . He- (BCN 6118); AY445189, FJ211539, HG798248. lichrysum mundtii Harv.: South Africa, Kwazu- Helichrysum petiolare Hilliard & B. L. Burtt: Ex lu-Natal Province, Romo 14368 et al. (BC 867641); J. Bot. Mar i Murtra, Blanes (BCN 6110); HG797783, HG798075, HG798235. Helichrysum AY445213, HG798088, HG798249. Helichrysum mussae Nevski: Tadzhikistan, Zeravshchan Mts., pilosellum (L. f.) Less.: South Africa, Mpumalan- Filatov et al. 81 (LE); FJ211426 and FJ211484, ga Province, Romo 14597 et al. (BC 867800); FJ211555, - . Helichrysum natalitium DC.: South HG797791, HG798089, HG798250. Helichrysum Africa, Kwazulu-Natal Province, Burrows 8431 plantago DC.: Madagascar, Antananarivo Prov- (Buffelskloof herb.); HG797784, HG798076, ince, Mt. Ibity, Bayer MAD-04062 et al. (CANB HG798236. Helichrysum neoachyroclinoides 660399); HG797792, HG798090, - . Helichrysum Humbert: Madagascar, Antananarivo Province, platycephalum Baker: Madagascar, Fianarantsoa

Collectanea Botanica 37: e012 (enero-diciembre 2018), e012, ISSN-L: 0010-0730, https://doi.org/10.3989/collectbot.2018.v37.012 24 A. ACOSTA-MAINDO & M. GALBANY-CASALS

Province, Bayer MAD-04022 et al. (CANB 660359); HG798268. Helichrysum spiralepis Hilliard & B. HG797793, HG798091, HG798251. Helichrysum L. Burtt: South Africa, Kwazulu-Natal Province, platypterum DC.: South Africa, Kwazulu-Natal Romo 14372 et al. (BC 867645); FJ211477 and Province, Romo 14360 et al. (BC 867634); FJ211458 FJ211535, HG798105, - . Helichrysum splen- and FJ211516, HG798092, HG798252. Helichry- didum (Thunb.) Less.: Ex Roy. Bot. Gard. Kew sum plicatum DC.: Turkey, Sivas, Susanna 2419 et (BCN 6102); AY445218, HG798106, HG798269. al. (BCN 6129); AY445201, FJ211556, HG798253. Helichrysum stellatum (L.) Less.: South Africa, Helichrysum polycladum Klatt: South Africa, Northern Cape Province, Koekemoer 3513 (BC); Mpumalanga Province, Romo 14598 et al. (BC HG797801, HG798107, HG798270. Helichrysum 867801); HG797794, HG798093, HG798254. Heli- stoechas (L.) Moench: Spain, Lleida, Galbany s. n. chrysum populifolium DC.: South Africa, ex Silver- (BCN 6114); AY445193, FJ211543, FN649351. hill Seeds (BCN 8218); AY445210, FJ211538, Helichrysum stuhlmannii O. Hoffm.: Uganda, HG798255. Helichrysum psilolepis Harv.: South Rwenzori Mts., Roquet s. n. (BC 867841); Africa, Eastern Cape Province, Romo 14461 et al. FJ211466 and FJ211524, HG798108, HG798271. (BC 867689); HG797795, HG798094, HG798256. Helichrysum summo-montanum Verdoorn: South Helichrysum reflexum N. E. Br.: South Africa, Africa, Mpumalanga Province, Burrows 7370 Mpumalanga Province, Romo 14571 et al. (BC (Buffelskloof herb.); HG797802, HG798109, 867778); FJ211445 and FJ211503, HG798095, HG798272. Helichrysum sutherlandii Harv.: HG798257. Helichrysum retortum (L.) Willd.: South Africa, Kwazulu-Natal Province, Romo South Africa, ex Silverhill Seeds (BCN 6112); 14370 et al. (BC 867643); HG797803, HG798110, AY445222, HG798096, HG798258. Helichrysum HG798273. Helichrysum swynnertonii S. Moore: rosum (Berg.) Less. var. arcuatum Hilliard: South South Africa, Mpumalanga Province, Camacho Africa, Eastern Cape Province, Romo 14494 et al. s. n. (Buffelskloof herb.); HG797804, HG798111, (BC 867718); HG797797, HG798098, HG798260. - . Helichrysum thianschanicum Regel: Ex Hortus Helichrysum rosum (Berg.) Less. var. rosum: South Botanicus Táhor (BCN 10337); AY445200, Africa, Eastern Cape Province, Romo 14462 et al. FJ211554, HG798274. Helichrysum transmonta- (BC 867690); HG797796, HG798097, HG798259. num Hilliard: South Africa, Mpumalanga Prov- Helichrysum rubicundum (K. Koch) Bornm.: Iran, ince, Burrows 7543 (Buffelskloof herb.); Azarbaidjan, Termeh et al. s. n. (IRAN 35924,4); HG797805, HG798112, HG798275. Helichrysum FJ211437 and FJ211495, FJ211573, HG798261. trilineatum DC.: South Africa, Eastern Cape Prov- Helichrysum rugulosum Less.: South Africa, Free ince, Romo 14416 et al. (BC 867669); HG797806, State Province, Romo 14331 et al. (BC 867613); HG798113, HG798276. Helichrysum triplinerve FJ211471 and FJ211529, HG798099, HG798262. DC.: Madagascar, Fianarantsoa Province, Bayer Helichrysum schimperi (A. Rich.) Moeser: Tanza- MAD-04057 et al. (CANB 660394); HG797807, nia, Olmoti, Galbany & Arrabal s. n. (BC 867821); HG798114, HG798277. Helichrysum truncatum FJ211451 and FJ211509, HG798100, HG798263. Burtt Davy: South Africa, Mpumalanga Province, Helichrysum selaginifolium R. Vig. & Humbert: Romo 14574 et al. (BC 867781); HG797808, Madagascar, Antananarivo Province, Mt. Ibity, Bay- HG798115, HG798278. Helichrysum umbracu- er MAD-04074 et al. (CANB 660411); HG797798, ligerum Less.: South Africa, Kwazulu-Natal Prov- HG798101, HG798264. Helichrysum setosum ince, Romo 14366 et al. (BC 867639); FJ211450 Harv.: Tanzania, Empaakai, Galbany & Arrabal s. n. and FJ211508, HG798116, HG798279. Helichry- (BC 867824); FJ211467 and FJ211525, HG798102, sum xylocladum Humbert: Madagascar, Antanan- HG798265. Helichrysum sibthorpii Rouy: Greece, arivo Province, Mt. Ibity, Bayer MAD-04073 et al. ex Roy. Bot. Gard. Kew (BCN 6099); AY445203, (CANB 660410); HG797809, HG798117, - . Heli- FJ211561, HG798266. Helichrysum silvaticum chrysum zeyheri Less.: South Africa, Western Hilliard: Mozambique, Licuati Sand Forest, Mc- Cape Province, Romo 14542 et al. (BC 867754); Murtry 11424 (Buffelskloof herb.); HG797799, FJ211478 and FJ211536, HG798118, - . Helichry- HG798103, HG798267. Helichrysum simillinum sum zwartbergense H. Bol.: South Africa, Western DC.: South Africa, Kwazulu-Natal Province, Romo Cape Province, Romo 14520 et al. (BC 867739); 14364 et al. (BC 867637); HG797800, HG798104, HM244707, HG798119, HG798280. Humeocline

Collectanea Botanica 37: e012 (enero-diciembre 2018), e012, ISSN-L: 0010-0730, https://doi.org/10.3989/collectbot.2018.v37.012 Pseudognaphalium aldunateoides back in Gnaphalium (Compositae: Gnaphalieae) 25 madagascariensis (Humb.) Anderb.: Madagascar, KM091353, KM091331. Luciliocline subspicata Antananarivo Province, Mt. Ibity, Bayer MAD- (Wedd.) Anderb. & Freire: Bolivia, Oruro, Sajama, 04061 et al. (CANB 660398); HG797810, colina Pacollo, S. Beck 30693 (LPB); - , - , HG798120, HG798281. Ifloga repens (L.) Hil- KM091344. Metalasia adunca Less.: South Afri- liard: Republic of South Africa, Northern Cape, ca, Karis & Arnerup 996 (S); JF893847, JF893752, Namakwa N. P., Koekemoer 3277 (BC); FN645826, - . Mexerion sarmentosum (Klatt) G. L. Nesom: FN645628, FN649357. Ifloga spicata (Forssk.) Mexico, Mexico state, Beaman 1932 (US); Sch. Bip.: Spain, Almería, Cuevas de Almanzora, KT865561, KT865380, - . Micropsis dasycarpa Santos-Vicente 534 et al. (SALA 134240); (Griseb.) Beauverd: Argentina, Buenos Aires, FN645825, FN645627, FN649356. Jalcophila bo- Tandil, La Cascada, Urtubey & Freire 543 (SI); liviensis Anderb. & Freire: Bolivia, La Paz, Ta- KM091408, KM091378, KM091339. Micropus mayo, Ulla-Ulla, reserva de Apolobamba, Co. Pun- californicus Fischer & C. A. Meyer: United States, tani, Urtubey 488 et al. (SI); KM091402, Butte Co., N side of Bardees Bar Road, Ahart KM091370, KM091342. Lachnospermum fascic- 12624 (RSA 712992); FN645821, FN645621, ulatum (Thunb.) Baill.: Republic of South Africa, FN649350. Micropus supinus L.: Spain, Sala- Western Cape Province, between Op-die-berg and manca, Martín de Yeltes, Rico 7883 (SALA); Citrusdal, Romo 14559 et al. (BC), FN645829, FN645818, FN645609, FN649335. Mniodes andi- FN645623, FN649333. Langebergia canescens na (A. Gray) Cuatrec.: Bolivia, Oruro, X. Men- (DC.) Anderb.: South Africa, Western Cape Prov- hofer 1514 (LPB); - , KM091367, - . Mniodes ince, Bayer & Puttock s. n. (CANB 499988); kunthiana (DC.) Freire, Chemisquy, Anderb. & HG797811, HG798121, - . Lasiopogon glomerula- Urtubey: Bolivia, La Paz, Murillo, Urtubey 475 et tus (Harv.) Hilliard: South Africa, SAF-96180 (F); al. (SI); KM091393, KM091365, KM091329. KT865529, KT865345, - . Lasiopogon muscoides Mniodes longifolia (Cuatrec. & Aristeg.) Freire, (Desf.) DC.: Spain, Almería, Cuevas de Almanzo- Chemisquy, Anderb. & Urtubey: Bolivia, La Paz, ra, Santos-Vicente 499 et al. (SALA), FN645831, Nor-Yungas, camino a Coroico, Urtubey 507 et al. FN645629, FN649334. Leontopodium alpinum (SI); KM091400, KM091364, KM091335. Mniodes Cass.: Spain, Huesca, Posets, Roquet s. n. (BC); santanica (Cabrera) Freire, Chemisquy, Anderb. FN645824, FN645625, FN649348. Leysera & Urtubey: Argentina, Catamarca, Abato, Co. gnaphalodes (L.) L.: South Africa, Western Cape Manchado, N. Bayón & J. Vera Bahima 1519 Province, Romo 14546 et al. (BC 867757); (LPAG); KM091397, KM091359, KM091336. Om- FN645815, FN645636, FN649329. Logfia gallica alotheca supina (L.) DC.: Andorra, Port Creussans, (L.) Coss. & Germ.: Spain, Almería, Sorbas, Galbany & Lluent s. n. (BCN 6121); AY445230, Martínez-Ortega 1796 et al. (SALA 134224), FN645558, FN649354. Omalotheca sylvatica (L.) FN645838, FN645556, FN649339. Logfia minima Sch. Bip. & F. W. Schultz: Switzerland, Charpin & (Sm.) Dumort.: Spain, Salamanca, San Miguel de Hainard s. n. (US); KT865568, KT865387, - . Valero, Martínez-Ortega 1805 (SALA 134219); Ozothamnus diosmifolius (Vent.) DC.: Australia, FN645817, FN645613, FN649347. Loricaria co- New South Wales, Bayer NSW 94-006 (NSW, lombiana Cuatrec.: Colombia, Tolima, J. C. Ospi- ALTA); HM244716, HM450851, - . Petalacte cor- na 6400 (SI). Colombia, Caldas, D. Sanin E 89 onata (L.) D. Don: South Africa, Western Cape (SI); KM091404, KM091371, KM091330. Lucilia Province, Bayer & Chandler SAF-01095 (CANB acutifolia (Poir.) Cass.: Argentina, Buenos Aires, 634589); HG797812, HG798122, HG798282. Tandil, Urtubey 512 (SI); KM091396, KM091374, Pseudognaphalium affine (D. Don) Anderb. (1): KM091332. Lucilia linearifolia Baker: Argentina, Korea, Mt. Baika-san, Cho YH et al. 090506-135 Jujuy, Villa de la Sierras de Zapla, Co. de La Cruz, (KUN); KT865569, KT865390, - . Pseudognapha- Urtubey & Freire 427 (SI); - , - , KM091324. Lu- lium affine (D. Don) Anderb. (2): China, Hubei, cilia lycopodioides (Less.) Freire: Argentina, Jujuy, Nie 714 (KUN); JQ895515, JQ895378, - . Pseudo- Villa de la Sierras de Zapla, Co. de La Cruz, Ur- gnaphalium aldunateoides (J. Rémy) C. Monti, tubey & Freire 426 (SI); KM091384, KM091357, N. Bayón & Freire [Gnaphalium aldunateoides J. KM091323. Lucilia nitens Less.: Uruguay, Monte- Rémy] (1): Chile, Región IV Coquimbo, llegan- video, Carrasco, Urtubey 394 (SI); KM091399, do a Sotaquí desde Monte Patria o Combarbalá,

Collectanea Botanica 37: e012 (enero-diciembre 2018), e012, ISSN-L: 0010-0730, https://doi.org/10.3989/collectbot.2018.v37.012 26 A. ACOSTA-MAINDO & M. GALBANY-CASALS

Quebrada Seca, 30º 39′ 29,1′′ S, 71º 05′ 15,8′′ W, Hilliard & B. L. Burtt: China, Sichuan, Nie 1191 98 m, Galbany 2525 & Arrabal (BC); *MG649409, (KUN); JQ895519, JQ895382, - . Pseudognapha- *MG649413, *MG649416. Pseudognaphalium lium lacteum (Meyen & Walp.) Anderb.: Argenti- aldunateoides (J. Rémy) C. Monti, N. Bayón & na, Jujuy, Tumbaya, RP52, entre Purmamarca y Freire [Gnaphalium aldunateoides J. Rémy] (2): Susques, pasando Ayo. Potrerillos, Urtubey & Chile, Región Metropolitana, Prov. Chacabuco, Al- Freire 436 (SI); KM091387, KM091362, tos de Chicauma, sector tranque, 33º 10′ S 70º 58′ KM091340. Pseudognaphalium leucocephalum W, 2050 m, N. García & L. Faúndez 177476 (A. Gray) Anderb. (1): United States of America, (CONC); *MG649408, *MG649412, - . Pseudo- California, Orange Co., Roberts 6088 (RSA gnaphalium attenuatum (DC.) Anderb.: Mexico, 703730); HG797817, HG798128, HG798288. Veracruz, Robert Merrill King 4131 (US); Pseudognaphalium leucocephalum (A. Gray) An- KT865570, KT865391, - . Pseudognaphalium ba- derb. (2): Mexico, Sinaloa, Kimnqch et al. 1992 dium (Wedd.) Anderb.: Bolivia, La Paz, Murillo, (US); KT865578, KT865398, - . Pseudognaphali- Urtubey 485 et al. (SI); KM091405, KM091375, um leucopeplum (Cabrera) Anderb.: Bolivia, KM091333. Pseudognaphalium beneolens (Da- Franz, Tamayo, Beck 32545 (US); KT865579, vidson) Anderb. : United States of America, Cali- KT865399, - . Pseudognaphalium luteoalbum fornia, San Diego Co., Rebman 10825 (RSA (L.) Hilliard & B. L. Burtt (1): Portugal, Marinha 705579); HG797813, HG798123, HG798283. Grande, Susanna 2435 & Garcia-Jacas (BCN Pseudognaphalium biolettii Anderb.: United 6125); AY445227, FN645633, FN649358. Pseudo- States of America, California, Los Angeles Co., gnaphalium luteoalbum (L.) Hilliard & B. L. Burtt Sanders 25759 (RSA 712717); HG797814, (2): Australia, Victoria, Robert Merrill King 9742 HG798124, HG798284. Pseudognaphalium cali- (US); KT865581, KT865401, - . Pseudognaphali- fornicum (DC.) Anderb.: United States of Ameri- um luteoalbum (L.) Hilliard & B. L. Burtt (3): ca, California, Riverside Co., Boyd 10845 (RSA Kenya, Robert Merrill King 9800 (US); KT865580, 701560); HM244709; HG798125, HG798285. KT865400, - . Pseudognaphalium macounii Pseudognaphalium canescens (DC.) Anderb.: (Greene) Kartesz: United States of America, Utah, Mexico, Sonora, Spooner 2423 (US); JQ895522, Daggett Co., Goodrich 22823 (RSA 490532); JQ895385, - . Pseudognaphalium cheiranthifoli- HG797821, HG798132, HG798293. Pseudo- um (Lam.) Hilliard & B. L. Burtt (1): Chile, region gnaphalium melanosphaeroides (Sch. Bip. ex VI, Pichilemu, Carnicero-Campmany s. n. (SI); Wedd.) Anderb.: Bolivia, Cochabamba, Gonzalo HG797816, HG798127, HG798287. Pseudo- 58 (US); KT865582, KT865402, - . Pseudo- gnaphalium cheiranthifolium (Lam.) Hilliard & gnaphalium micradenium (Weath.) G. L. Nesom: B. L. Burtt (2): Brazil, Curitiba, Cordeiro and Bar- United States of America, Tennessee, Roane Co., bosa 2173 (US); KT865571, KT865392, - . Pseudo- Patrick 3813; JX524594, - , - . Pseudognaphalium gnaphalium dombeyanum (DC.) Anderb.: Peru, montevidense (Spreng.) Anderb.: Argentina, Rio Pampa, Carlos 16094 (US); KT865572, - , - . Negro, Simon Pablo 860 (US); - , KT865403, - . Pseudognaphalium eggersii (Urb.) Anderb.: Dom- Pseudognaphalium moritzianum (Klatt) V. M. Ba- inica, Baoruco, Acevedo-Rdgz 13172 (US); dillo: Venezuela, Merida, Robert Merrill King KT865573, KT865393, - . Pseudognaphalium fla- 10417 (US); - , KT865404, - . Pseudognaphalium vescens (Kitam.) Anderb.: China, Xizang, Xia 2730 obtusifolium (L.) Hilliard & B. L. Burtt: United (KUN); KT865574, KT865394, - . Pseudognapha- States of America, Iowa, Lucas Co., Neese 11074 lium gaudichaudianum (DC.) Anderb.: Bolivia, (RSA 299987); HG797818, HG798129, HG798290. Murillo, Solomon 16478 (US); KT865576, Pseudognaphalium oligandrum (DC.) Hilliard & KT865396, - . Pseudognaphalium graveolens B. L. Burtt: South Africa, Free State Province, (Kunth) Anderb.: Peru, Cajamarca, Becher & Ter- Romo 14336 et al. (BC 867618); HM244708, rones 2097 (US); KT865575, KT865395, - . HM450843, HG798289. Pseudognaphalium oxy- Pseudognaphalium heterotrichum (Phil.) An- phyllum (DC.) Kirp. var. oxyphyllum (1): Mexico, derb.: Chile, region III, Coquimbo, Carnice- Estado de México, Alrededores de Xalatlaco, 19º ro-Campmany s. n. (SI); HG797815, HG798126, 11′ 31,5′′ N 99º 24′ 38,2′′ W, 2831 m, Galbany 2431 HG798286. Pseudognaphalium hypoleucum (DC.) & Arrabal (BC); *MG649410, *MG649414,

*MG649417. *Pseudognaphalium oxyphyllum South Africa, Western Cape, Koekemoer 3481 (DC.) Kirp. var. oxyphyllum (2): Mexico, Distrito (US); KT865600, KT865428, - . Vellereophyton Federal, Ciudad de México, Ciudad Universitaria, dealbatum (Thunb.) Hilliard & B. L. Burtt: South cerca de la estación de metrobus “Ciudad Universi- Africa, Western Cape Province, Romo 14549 et al. taria”, entre la estación y la parada del Pumabus, (BC); FN645832, FN645631, FN649355. 19º 19′ 25,6′′ N 99º 11′ 19,8′′ W, 2304 m, Galbany 2480 & Arrabal (BC); *MG649411, *MG649415, Appendix 2. Material examined for morpho- *MG649418. Pseudognaphalium ramosissimum logical studies. (Nutt.) Anderb.: United States of America, Califor- nia, San Diego Co., Sanders 31665 (RSA 713933); Gnaphalium uliginosum L.: France: Gruey-les- HG797819, HG798130, HG798291. Pseudo- Surance, vores de camps i camins, Arrabal & Gal- gnaphalium sandwicensium (Gaudich.) Anderb.: bany 2090 (BC). Pseudognaphalium aldunateoi- United States of America, Hawaii, Funk 12765 des (J. Rémy) C. Monti, N. Bayón & Freire: Chile: (US); KT865585, KT865409, - . Pseudognaphali- Prov. de Bío-Bío, Negrete, a orillas del Biobío, um semiamplexicaule (DC.) Anderb.: Guatemala, en la ribera Norte del río, A. Pfister s. n. (CONC Alta Verapaz, Fother 11090 (US); KT865586, 10502); Prov. Aconcagua, Putaendo, Laguna del KT865410, - . Pseudognaphalium sylvicola Copin, 2620 m, crece en las planicias húmedas de (McVaugh) G. L. Nesom: Mexico, Chapingo, Koch las cordilleras de Aconcagua, Parra 37 (CONC 82168 (US); KT865587, KT865411, - . Pseudo- 116325); Prov. Cautin, Temuco, Río Cautin, 120 gnaphalium thermale (E. E. Nelson) G. L. Nesom: m, Montero 5728 (CONC 75310); Prov. Valpara- United States of America, California, T26N, R11E, iso, Dpto. Valparaiso, Lago Peñuelas, Ricardi Ahart 12405 (RSA 716686); HG797820, 5445 (CONC 89745); VII Región, Prov. Linares, HG798131, HG798292. Pseudognaphalium vira- Quella Sur, 36° 05′ S 72° 08′ W, 120 m, en culti- vira (Molina) Anderb.: Chile, Mahu 10213 (US); vo de arroz, a orillas del pretil, escasa, Matthei & KT865589, KT865413, - . Raoulia tenuicaulis Quezada 1420 (CONC 110827); Cauquenes, ap- Hook. f.: New Zealand, Gisborne, mouth of Motu proximately 2 km south of Chanco, on west side River, Smissen s. n. (CHR 607934); HM244728, of road, 35° 45′ S 72° 32′ W, 50 m, Bliss 2272 HM450865, - . Relhania pungens L’Hérit.: South (CONC 145334); Reg. Meropolitana, San José Africa, Western Cape Province, Koekemoer 3427 de Maipo, Cajón del Morales, 33° 46′ S 70° 04′ (BC); FN645814; FN645635; FN649331. Stoebe W, 2450 m, entre la laguna y el glaciar, Teillier & aethiopica L.: South Africa, Cederberg, Karis & Márquez 5316 (CONC 156533); Reg. Metropoli- Swenson 919 (S); JF893878, FM173147, - . Stuck- tana, Prov. Chacabuco, Altos de Chicauma, sec- ertiella capitata (Wedd.) Beauverd: Bolivia, La tor tranque, 33° 10′ S 70° 58′ W, 2050 m, García Paz, Murillo, Camino antiguo a Unduavi, subiendo & Faúndez 3613 (CONC 177476);VIII Región, el valle de Chuquiaguillo, Urtubey 469 et al. (SI); Prov. Ñuble, Ninhue, Río Ninhue, km 32 desde KM091398, KM091369, KM091341. Stylocline San Gregorio a Pocillas, 36° 08′ S 72° 05′ W, 190 psilocarphoides M. Peck: United States, Lyon co., m, Mihoc, Marticorena, Gómez & Negritto 6227 Pine Nut Mts., Tiehm 14828 (RSA 712497); (CONC 161086); Región IV Coquimbo, llegando FN645819, FN645619, FN649346. Syncarpha ca- a Sotaquí desde Monte Patria o Combarbalá, Que- nescens (L.) B. Nord.: South Africa, Cape Town, brada Seca, 30º 39′ 29,1′′ S, 71º 05′ 15,8′′ W, 98 m, Bergh 1222 (NBG); JF893884, FM173153, - . Syn- Galbany 2525 & Arrabal (BC). Pseudognapha- carpha chlorochrysum (DC.) B. Nord.: South Af- lium oxyphyllum (DC.) Kirp. var. oxyphyllum: rica, Western Cape, Koekemoer 3424 (US); Mexico: Estado de México, Alrededores de Xa- KT865598, KT865426, - . Syncarpha mucronata latlaco, 19º 11′ 31,5′′ N 99º 24′ 38,2′′ W, 2831 m, (P. J. Bergius) B. Nord.: South Africa, Western Galbany 2431 & Arrabal (BC); Distrito Federal, Cape Province, Romo 14511 et al. (BC 867732); Ciudad de México, Ciudad Universitaria, cerca de FJ211421 and FJ211479, FN645626, FN649360. la estación de metrobus “Ciudad Universitaria”, Syncarpha paniculata (L.) B. Nord.: South Africa, entre la estación y la parada del Pumabus, 19º 19′ Western Cape, Koekemoer 3428 (US); KT865599, 25,6′′ N 99º 11′ 19,8′′ W, 2304 m, Galbany 2480 KT865427, - . Syncarpha staehelina (L.) B . Nord.: & Arrabal (BC).

Collectanea Botanica 37: e012 (enero-diciembre 2018), e012, ISSN-L: 0010-0730, https://doi.org/10.3989/collectbot.2018.v37.012