Asteraceae, Gnaphalieae), a New Genus from the Central Andes
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Bayer & Dillon: Quasiantennaria (Asteraceae, Gnaphalieae), a new genus from the central Andes Arnaldoa 26 (1): 9-26, 2019 ISSN: 1815-8242 (edición impresa) http://doi.org/10.22497/arnaldoa.261.26101 ISSN: 2413-3299 (edición online) Quasiantennaria (Asteraceae, Gnaphalieae), a new genus from the central Andes Quasiantennaria (Asteraceae, Gnaphalieae), un nuevo género de los Andes centrales Randall J. Bayer Department of Biological Sciences, University of Memphis, Memphis, TN, USA [email protected] Michael O. Dillon The Field Museum, Integrative Research Center 1400 South Lake Shore Drive, Chicago, IL 60605-2496, USA [email protected] 26 (1): Enero - Abril, 2019 9 Este es un artículo de acceso abierto bajo la licencia CC BY-NC 4.0: https://creativecommons.org/licenses/by-nc/4.0/ Bayer & Dillon: Quasiantennaria (Asteraceae, Gnaphalieae), a new genus from the central Andes Recibido: 2-I-2019; aceptado: 20-II-2019; publicado online: 10-IV-2019; publicado impreso: 30-IV-2019 Abstract Quasiantennaria R. J. Bayer & M. O. Dillon, a new genus from the central Andes distributed from Peru to Bolivia, is described and illustrated. It is characterized among Andean Gnaphalieae by its dioecious or polygamo-dioecious breeding system, cespitose, perennial herbs, basal rosulate leaves, capitulescences scapose, cymose-corymbose. It most closely resembles Antennaria Gaertn., a typically northern hemispheric genus; however, it differs in its achenial trichomes and relationships suggested by DNA sequence data. Keywords: Asteraceae, Gnaphalieae, new genus, Bolivia, Peru, Andean flora. Resumen Quasiantennaria R. J. Bayer & M. O. Dillon, un nuevo género de los Andes centrales con distribución en Perú y Bolivia, se describe e ilustra para Perú. Se caracteriza entre las Gnaphalieae andinas por su sistema de reproducción dioico o polígamo-dioico, cespitosa, hierbas perennes, hojas basales en roseta, capitulescencias escaposas, cimoso-corimbosas. Es muy cercano a Antennaria Gaertn., un género típicamente del hemisferio norte; sin embargo, difiere en los tricomas de los aquenios y las relaciones filogenéticas sugeridas por datos de secuencia de ADN. Palabras clave: Asteraceae, Gnaphalieae, nuevo género, Bolivia, Perú, flora andina. Citation: Bayer, R. & M. Dillon. Quasiantennaria (Asteraceae, Gnaphalieae), a new genus from the central Andes. Arnaldoa 26 (1): 9 - 26. http://doi.org/10.22497/arnaldoa.261.26101 Introduction A. chilensis J. Rémy, A. magellanica Sch. Bip. (= Antennaria chilensis var. magellanica Re- The Gnaphalieae (Cass.) Lecoq & Juill. iche), and A. sleumeri Cabrera, have been (Asteraceae) is a small tribe comprising considered as putative disjunctions from ca. 185 genera and 1240 species (Ward Northern Hemispheric taxa (Bayer et al., et al., 2009). In South America, some 20 1996). Recent next generation sequencing genera and over 100 species are recorded using hundreds of loci (Thapa et al., ined.) with highest diversity in the tropical and shows that Antennaria chilensis is a tradi- subtropical Andean Cordillera (Dillon & tional Antennaria and is closely related Sagástegui-A., 1991a). Many have prolif- to the Antennaria rosea Greene polyploid erated in upper elevation or alpine habitats agamic complex. It was recovered as sis- in Peru (Dillon, 2005) and overwhelming- ter to Antennaria corymbosa E. Nels., which ly they possess reduced, small cushions of is a putative sexual progenitor of A. rosea tightly clustered basal densely canescent (Bayer, 1990; Thapa et al., ined.). Antennar- leaves. Genera can look remarkably simi- ia sleumeri is an enigmatic species, known lar, both in the field and as collections on only from scant type material (LP) and un- herbarium sheets, but experience within til the species can be relocated, collected these groups has shown that upon inspec- and studied further its relationships will tion, microcharacters can allow discrim- likely remain cryptic. The remaining South ination of genera, especially when com- American species, A. linearifolia Wedd., bined with DNA sequence data (Luebert et was recently included in analyses by Lu- al., 2017). ebert et al. (2017) and Thapa et al. (ined.) Species that have been attributed to and notably did not group with other au- Antennaria Gaertn. in South America, e.g., thentic or traditional Antennaria species, 10 26 (1): Enero - Abril, 2019 Bayer & Dillon: Quasiantennaria (Asteraceae, Gnaphalieae), a new genus from the central Andes i.e., A. chilensis, A. dioica (L.) Gaertn., and The distribution of the dioecious A. microphylla Rydb. (see Fig. 1, p. 1188 in breeding syndrome has developed Luebert et al., 2017). independently no fewer than seven times within the Gnaphalieae, viz. Antennaria, Luebert et al. (2017) found that A. Loricaria, Mniodes, Cuatrecasasiella H. Rob., linearifolia was resolved in Clade L6 which Parantennaria Beauverd, Pterygopappus included Jalcophila M. O. Dillon & Sagást., Hook. f., Sinoleontopodium Y.-L. Chen Loricaria Wedd., and Mniodes (A.Gray) (Anderberg, 1991) and is a good example Benth. (BPP: 1, MLB: 84). Jalcophila is sister of parallel evolution. Antennaria has to a clade with A. linearifolia, Loricaria and a Holarctic (mostly North American) Mniodes (BPP: 1, MLB: 98). Relationships distribution, while Loricaria, Mniodes, within the latter clade remain unresolved. and Cuatrecasasiella are South American, Loricaria is well-supported (BPP: 1, MLB: Parantennaria and Pterygopappus are 99) and Mniodes forms a well-supported Australian, and Sinoleontopodium is Asian monophyletic group (BPP: 1, MLB: 90). All (China). A recent ITS/ETS phylogeny other Antennaria were restricted to Clade (Nie et al., 2015) of the Gnaphalieae using L5 which contained Diaperia and sister to 80% of the genera (835 species) in the tribe Clade L4 which contained Chevreulia Cass. confirmed the independent origins of the and Cuatrecasasiella H. Rob. dioecious Gnaphalieae. Sinoleontopodium Thapa et al. (ined.) conducted an anal- was described because its dioecious nature ysis using hundreds of loci to reconstruct made it an “atypical” Leontopodium (Chen, the phylogeny of 34 species of Antennaria 1985), however, it is clearly shown to be including, A. linearifolia. Antennaria lineari- imbedded within traditional Leontopodium folia fell clearly outside a strongly support- (Nie et al., 2015) by the molecular data. , have ed monophyletic traditional Antennaria, in been upheld as distinct from Antennaria an outgroup that contained four outgroup The evolution of the dioecious trait in taxa, Mniodes schultzii (as Belloa), M. subspi- the Gnaphalieae seems to be associated cata (as Luciliocline). Gamochaetopsis alpina with the high elevation, usually cushion (Poepp. & Endl.) Anderb. & S. E. Freire, plant habit, and is therefore not always and Facelis lasiocarpa (Griseb.) Cabrera. a trustworthy indicator of phylogenetic The relationships of A. linearifolia to any of relatedness. Much taxonomic confusion in the outgroup taxa was unresolved (Thapa the Gnaphalieae has been caused because et al., ined.), however it is manifestly not an dioecy has been used as a strong taxonomic Antennaria. character, assumed to be homologous. Therefore, molecular data has Therefore, as A. linearifolia cannot suggested that Antennaria linearifolia, remain in traditional Antennaria and its regardless of its overall morphology, was relationships to other genera remain recovered in a clade together with Loricaria unclear, we propose that the taxon be and Mniodes, while the remainder of moved to a new genus, Quasiantennaria. Antennaria species form a separate clade (Luebert et al., 2017). This result was not apparent in the ITS analysis of Antennaria (Bayer et al., 1996), because material of A. linearifolia was not included in that study. 26 (1): Enero - Abril, 2019 11 Bayer & Dillon: Quasiantennaria (Asteraceae, Gnaphalieae), a new genus from the central Andes Results Leontopodium linearifolium (Wedd.) Benth. & Hook., Gen. Pl. 2: 303. 1873. Quasiantennaria R. J. Bayer & M. O. Dillon, gen. nov. Leontopodium linearifolium (Wedd.) Britton, Bull. Torrey Bot. Club 19(5): 148. TYPE: Quasiantennaria linearifolia 1892. (Wedd.) R. J. Bayer & M. O. Dillon Gnaphalium linearifolium (Wedd.) Dioecious or more rarely polygamo- Franchet, Bull. Soc. Bot. France 39: 135. dioecious, perennial herbs; stems simple, 1892. ascending or erect, tomentose or lanate. Basal leaves rosulate, oblanceolate to Description: Dioecious or rarely spatulate, entire; cauline leaves alternate, polygamo-dioecious, perennial herbs, smaller. Capitulescences cymose- rhizomatous; stems unbranched, 5-15(- corymbose, racemose, or glomerulate, 30) cm tall. Leaves basal, rosulate, sessile; terminal, occasionally of a solitary head. blade oblanceolate-linear, 2-8 cm long, Capitula discoid or rarely disciform; 1.5-3 mm wide, the lower surface densely involucres ovoid or campanulate; tomentose, midrib prominent, the upper phyllaries imbricate, scarious, the outer surface glabrescent, the margins entire. gradually narrowing, the inner prolonged Capitulescences glomerulate, subtended by into a petaloid lamina; receptacles convex foliaceous bracts. Capitula 5-7 mm high, 3-5 to plane, epaleaceous; pistillate florets with mm wide, discoid, homogamous, rarely corollas filiform, white or lilac, truncate or disciform and heterogamous; involucres subdentate; staminate florets with corollas campanulate; phyllaries 4-5-seriate, the tubular, 5-lobed or 5-dentate, the anther outer ovate, 3-4 mm long, ca. 2.5 mm bases sagittate, caudate, the terminal wide, apex obtuse,