Quasiantennaria (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 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 (Luebert et al., 2017). The Gnaphalieae (Cass.) Lecoq & Juill. Species that have been attributed to (Asteraceae) is a small tribe comprising ca. Antennaria Gaertn. in South America, e.g., 185 A. chilensis J. Rémy, A. magellanica Sch. Bip. (= Antennaria chilensis var. magellanica Re- genera and 1240 species (Ward et al., iche), and A. sleumeri Cabrera, have been 2009). In South America, some 20 genera considered as putative disjunctions from and over 100 Northern Hemispheric taxa (Bayer et al., species are recorded with highest di- 1996). Recent next generation sequencing versity in the tropical and subtropical An- using hundreds of loci (Thapa et al., ined.) dean Cordillera shows that Antennaria chilensis is a tradi- tional Antennaria and is closely related (Dillon & Sagástegui-A., 1991a). Many to the Antennaria rosea Greene polyploid have proliferated in upper elevation or agamic complex. It was recovered as sis- alpine habitats in Peru (Dillon, 2005) and ter to Antennaria corymbosa E. Nels., which overwhelmingly they possess reduced, is a putative sexual progenitor of A. rosea small cushions of tightly clustered basal (Bayer, 1990; Thapa et al., ined.). Antennar- densely canescent leaves. Genera can look ia sleumeri is an enigmatic species, known remarkably similar, both in the field and only from scant type material (LP) and un- as collections on herbarium sheets, but ex- til the species can be relocated, collected perience within these groups has shown and studied further its relationships will that upon inspection, microcharacters can likely remain cryptic. The remaining South allow discrimination of genera, especially American species, A. linearifolia Wedd., when combined with DNA sequence data was recently included in analyses by Lu- 10 26 (1): Enero - Abril, 2019 Bayer & Dillon: Quasiantennaria (Asteraceae, Gnaphalieae), a new genus from the central Andes ebert et al. (2017) and Thapa et al. (ined.) apparent in the ITS analysis of Antennaria and notably did not group with other au- (Bayer et al., 1996), because material of A. thentic or traditional Antennaria species, linearifolia was not included in that study. 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 Luebert et al. (2017) found that A. within the Gnaphalieae, viz. Antennaria, linearifolia was resolved in Clade L6 which Loricaria, Mniodes, Cuatrecasasiella H. Rob., included Jalcophila M. O. Dillon & Sagást., Parantennaria Beauverd, Pterygopappus Loricaria Wedd., and Mniodes (A.Gray) Hook. f., Sinoleontopodium Y.-L. Chen Benth. (BPP: 1, MLB: 84). Jalcophila is sister (Anderberg, 1991) and is a good example to a clade with A. linearifolia, Loricaria and of parallel evolution. Antennaria has Mniodes (BPP: 1, MLB: 98). Relationships a Holarctic (mostly North American) within the latter clade remain unresolved. distribution, while Loricaria, Mniodes, Loricaria is well-supported (BPP: 1, MLB: and Cuatrecasasiella are South American, 99) and Mniodes forms a well-supported Parantennaria and Pterygopappus are monophyletic group (BPP: 1, MLB: 90). All Australian, and Sinoleontopodium is Asian other Antennaria were restricted to Clade (China). A recent ITS/ETS phylogeny L5 which contained Diaperia and sister to (Nie et al., 2015) of the Gnaphalieae using Clade L4 which contained Chevreulia Cass. 80% of the genera (835 species) in the tribe and Cuatrecasasiella H. Rob. confirmed the independent origins of the 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 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(-30) corymbose, racemose, or glomerulate, cm tall. Leaves basal, rosulate, sessile; blade terminal, occasionally of a solitary head. oblanceolate-linear, 2-8 cm long, 1.5-3 mm Capitula discoid or rarely disciform; wide, involucres ovoid or campanulate; the lower surface densely tomentose, phyllaries imbricate, scarious, the outer midrib prominent, the upper surface gradually narrowing, the inner prolonged glabrescent, the margins entire. into a petaloid lamina; receptacles convex Capitulescences glomerulate, subtended by to plane, epaleaceous; pistillate florets with foliaceous bracts. Capitula 5-7 mm high, 3-5 corollas filiform, white or lilac, truncate or mm wide, discoid, homogamous, rarely subdentate; staminate florets with corollas disciform and heterogamous; involucres tubular, 5-lobed or 5-dentate, the anther campanulate; phyllaries 4-5-seriate, the bases sagittate, caudate, the terminal outer ovate, 3-4 mm long, ca. 2.5 mm appendages ovate, the styles undivided or wide, apex obtuse, the inner obovate to briefly bifid, the branches truncate.Achenes oblong-lanceolate,