Beauverdia hirtella subsp. glabrata (Amaryllidaceae), a new subspecies from Rio Grande do Sul State, Brazil Author Deble, Leonardo Paz Universidade Federal do Pampa, Campus Dom Pedrito, Dom Pedrito, Rio Grande do Sul, 96450 - 000, Brazil. & Núcleo Interinstitucional de Pesquisas Botânicas Balduíno Rambo, Santa Maria, Rio Grande do Sul, 97105 - 900, Brazil. text Phytotaxa 2021 2021-11-26 527 1 21 31 journal article 3208 10.11646/phytotaxa.527.1.2 5e205564-bd7d-44be-8995-dd9552353360 1179-3163 5728766 Beauverdia hirtella ( Kunth 1843: 465 ) Herter (1943: 509) subsp. glabrata Deble , subsp. nov. ( Figs. 1 and 2 ). Diagnosis— Beauverdia hirtella subsp. glabrata is distinguished from B. hirtella subsp. hirtella by its glabrous leaves (vs. papillosepubescent), with narrower blade ( 0.2–0.8 mm vs. 0.9–2 mm wide), glabrous scapes or with scarce 1-celled trichomes with uncinate cell (vs. scapes densely covered by 1-celled trichomes with a single straight cell) and nearly spherical or obovate capsules (vs. obovate or obovate-oblong). From B. hirtella subsp. lorentzii differs by its erect-ascendant leaves (vs. patent-recurved), with glabrous and narrowly linear-filiform blade, 0.2–0.8 mm wide (vs. papillose-pubescent, 2–5 mm wide), nearly terete or broadly elliptic in crosssection (vs. flat or slightly convex), scapes glabrous or with few 1-celled trichomes with uncinate cell (vs. scapes densely covered by 1-celled trichomes with straight cell) and nearly spherical or obovate capsules (vs. obovate-oblong). Type : — BRAZIL . Rio Grande do Sul : Dom Pedrito, RS 630, “em afloramento rochoso à esquerda da estrada, após a ponte do rio Tacuarembó , na divisa com São Gabriel e Lavras do Sul ”, 18 May 2019 , L.P. Deble & B.P. Moreira 19451 ( holotype PACA !) Geophyte 4–12 cm high above the soil. Bulb 8–12 × 6–11 mm , nearly spherical, simple, with strong alliaceous smell; outer cataphylls dark-brown, the innermost whitish. Leaves at anthesis 4–10, spirally arranged; leaf sheaths 1.5–6 cm long, subterranean, straw-colored; leaf blades 30–120 × 0.2–0.8 mm , narrowly linear-filiform, nearly terete or broadly elliptic in cross-section, 0.2–0.8 × 0.2–0.5 mm , dark-green, erect-ascendant, glabrous, margin thickened and rigid due to the presence of fibrous vascular bundles. Scapes 1–2, 2– 8 cm long, subcylindrical, erect, then curved at fruit maturity, with alliaceous smell when breaking, glabrous or with scarce 1-celled trichomes with uncinate cell in the distal third, trichomes 60–90 µm long. Bracts 2, ovate-elliptic, 6–10 × 3.5–6 mm , shortly fused at base for 0.5–1 mm . Pedicels 2.5–5 mm long, glabrous. Inflorescence 1-flowered. Flowers 16–24 × 7.5–10 mm , campanulate, goldenyellow, becoming ferruginous at the end of the anthesis, shiny, with a mild mint odor. Tepals 6 (3 + 3), golden-yellow, shortly fused at base for 0.5–2.5 mm , the outer ones oblanceolate or elliptic, blades 11–15 × 4.5–8 mm , base abruptly narrowed into a claw; claws of the outer tepals 1–2 × 0.5–1 mm ; the inner ones oblanceolate, elliptic or broadly elliptic; blades 10–14.5 × 4–7.5 mm ; base abruptly narrowed into a claw; claws of the inner tepals 1–2 × 0.5–1 mm ; tepal middle nerves purplish-brown, conspicuous, thicker towards the base. Staminal filaments 8–9.5 mm long, adnate at their bases for 1.2–1.8 mm , awl-shaped, translucent, yellowish; anthers 1.8–2.2 mm long, curved, pollen goldenyellow. Ovary obovate-oblong, 2.5–3.3 mm long; ovules 12 per locule; style yellow, 4–6.5 mm long, stigmatic portion capitate, greenish-yellow, shiny. Capsule nearly spherical or obovate 5–7.5 × 4.5–6 mm . Seeds 1.2–2 mm , black, shiny, tetrahedrally angled. Etymology:— From Latin glabratus (= glabrous), a reference to the leaves and floral pedicels that are glabrous, a distinctive feature in relation to the other subspecies of B. hirtella . Material Examined:— BRAZIL . Rio Grande do Sul , Dom Pedrito , localidade da Serrinha , entre rochas, 16 June 2018 , L.P. Deble & B.P. Moreira 19245 ( PACA !) . Lavras do Sul , Rincão do Inferno , na divisa com Dom Pedrito , nos paredões rochosos do rio Tacuarembó , 17 May 2017 , L.P. Deble & R.C. Pontes 19181 ( PACA ! SMDB !) . São Gabriel, RS 630, à direita da Estrada , após o cerro do Graxaim , 2 June 2018 , L.P. Deble et al. 19244 ( PACA !) . Phenology:— Flowering time May–June, fruiting time June–July. The flowers open around midday and close in the late afternoon, withering after two or three days. Distribution and habitat: — Beauverdia hirtella subsp. glabrata was found in Dom Pedrito, Lavras do Sul and São Gabriel municipalities, in south-central Rio Grande do Sul state , Brazil . The plants grow on rocky grasslands, mainly volcanoclastic rocks of the formation Acampamento Velho and granitic rocks of the formation Santa Maria ( Sommer et al. 1999 , 2011 ). This region is the most diverse in number of taxa of Beauverdia in Brazil , having in addition to this new subspecies, the occurrence of B. dyalistemon , B. hirtella subsp. hirtella , B. felipponei and B. vittata . B. hirtella subsp. glabrata and B. hirtella subsp. hirtella can be found sympatrically near the urban area of Dom Pedrito municipality, in a location called “Serrinha”, which corresponds to the northern limit of geographic distribution of the subspecies hirtella . In this location the subspecies hirtella occurs in deeper soils and associated with cespitose grasses, among the grasslands, while the subspecies glabrata stays restricted to concavity of rocks, which occur in a mosaic within these grasslands. During our field study, intermediate forms between both subspecies were not observed. Beauverdia hirtella subsp. glabrata displays its bulbs developed in the concavity or directly above of the rocks, associated with mosses, others herbaceous angiosperm species, and the lycophyta Sellaginella selloi Hieronymus (1900: 306) . In the same environments of occurrence of B. hirtella subsp. glabrata many others bulbous were registered, including the Iridaceae Calydorea riograndensis Deble (2011: 36) , Cypella fucata Ravenna (1981a: 18) , Cypella pusilla ( Link & Otto 1828: 125 ) Jackson (1893: 689) , Herbertia darwinii Roitman & Castillo (2008: 311) , H. lahue ( Molina 1810: 110 ) Goldblatt (1978: 379) , H. pulchella Sweet (1829 : pl. 222), Herbertia sp. , Kelissa brasiliensis ( Baker 1877: 134 ) Ravenna (1981b: 106) , and Onira unguiculata ( Baker 1892: 72 ) Ravenna (1983: 204) , some Oxalidaceae , as Oxalis perdicaria ( Molina 1782: 146 ) Bertero (1829: 739) and O. sellowiana Zuccarini (1831: 198) , and several others Amaryllidaceae , mainly of the genera Nothoscordum and Zephyranthes , among them Nothoscordum collinum Ravenna (1989: 41) , N. gaudichaudianum Kunth (1843: 458) , N. minarum Beauverd (1908: 1001) , N. muscorum Ravenna (1991: 41) , N. ostenii Beauverd (1908: 996) , Zephyranthes bifida ( Herbert 1825 : pl. 2597) García & Meerow in García et al. (2019: 492) , Z. caerulea ( Grisebach 1879: 320 ) Baker (1888: 37) , Z. gracilis Herbert (1837: 172) , Z. flavissima Ravenna (1967b: 28) , Z. minima Herbert (1837: 172) , and Z. tubispatha (L’Heritier: 1788: 6) Herbert (1821: 36) . Conservation: — Beauverdia hirtella subsp. glabrata is known only from four localities, and the populations spread in an extension of occurrence lesser than 5,000 km ² and area of occupancy smaller than 500 km ². The subspecies occurs in specific habitat, as it only grows on rocky grasslands, mainly volcanoclastic rocks and granitic rocks, and the bulbs develop in rocky concavities. These environments are being modified by economic activities, as mining and agriculture. The urban expansion and the invasion of exotic species in the region of occurrence of the subspecies, notably “capim-anonni-2” ( Eragrostis plana Nees von Esenbeck 1841: 390 ), an exotic invasive species native to Africa, result in a direct threat to this taxon. In accordance with the criteria of IUCN (2019) , Beauverdia hirtella subsp. glabrata can be considerate Endangered (EN), based on direct threat, geographic distribution, occupation area and population size (EN B1+B2ab (i, ii, iii) + c (ii, iv) + C1+C2a (i), b). FIGURE 1. Beauverdia hirtella subsp. glabrata . A–B. Habit. C . Flower, upper view. D . Flower, lateral view, evidencing bracts and scape. E. Flower, showing the pistil and stamens. F . Capsule. G. seed. H. Pistil. I. Ovary, cross-section. J. Leaf, proximal part, evidencing the apex of the leaf-sheaths. K. Leaf, distal part. L. Leaf, cross-section of the proximal part. M. Leaf, cross-section of the distal part. N. Ovules, removed of the locule (A, C–E, H–N, from L. P. Deble & B.P. Moreira 19451; B, F–G, from L. P. Deble & B.P. Moreira 19245 ). FIGURE 2 . Beauverdia hirtella subsp. glabrata . A . Habitat of B. hirtella subsp. glabrata , growing with Beauverdia vittata , Oxalis perdicaria and O. sellowiana and several others delicate Poaceae and Cyperaceae . B . Habit. C . Flower, upper view. D . Flower, inclined view. E . Flower, lateral view. F . Capsule (A–E from L. P. Deble & B.P. Moreira 19451 ; F, from L. P. Deble & B.P. Moreira 19245 ). FIGURE 3. Beauverdia hirtella subsp. hirtella (A–C) and B. hirtella subsp. lorentzii (D–E). A. Habit. B. Flower, upper view. C. Flower, lateral view. D. Flower, upper view. E. Flower, inclined view (A–C from L. P. Deble et al. 18851 , D–E from L.P. Deble & F.S. Alves14739 ). Discussion: — Beauverdia hirtella subsp. glabrata is easily recognized by its bulbs and scapes with strong alliaceous smell, narrowly linear-filiform and erect-ascendant leaves, with glabrous blades, scapes glabrous or with scarce 1-celled trichomes with uncinate cell in the distal third, and flowers with perigone composed by golden-yellow tepals. Beauverdia hirtella subsp. glabrata is morphologically close to both currently recognized subspecies of B. hirtella , since all these taxa have alliaceous smell and flowers with similar shape and size, and perigone composed by golden-yellow tepals ( Fig. 3 ). However, B. hirtella subsp. glabrata is distinguished from B. hirtella subsp. hirtella by its glabrous leaves (vs. papillose-pubescent), with narrower blade ( 0.2–0.8 mm vs. 0.9–2 mm wide), scapes glabrous or with scarce 1-celled trichomes with uncinate cell and 60–90 µm long (vs. scapes densely covered by 1-celled trichomes with straight cell and 110–180 µm long), and nearly spherical or obovate capsules (vs. obovate-oblong or oblong). The new subspecies is easily differentiated from B. hirtella subsp. lorentzii , due its erect-ascendant leaves (vs. patent-recurved), linear-filiform blade, 0.2–0.8 mm wide (linear blade, 2–5 mm wide), nearly terete or broadly elliptic in cross-section (vs. flat or slightly convex), flowers composed by six tepals (vs. eight tepals), and nearly spherical or obovate capsules (vs. obovate-oblong or oblong), and the same pubescence features mentioned to distinguish from B. hirtella subsp. hirtella . Additional features to distinguish the subspecies of Beauverdia hirtella can be assessed in Table 1 . From others species in the genus, B. hirtella subsp. glabrata clearly differs in several aspects. From B. dyalistemon the new subspecies contrasts by its strong alliaceous smell (vs. absent), erect-ascendant leaves (vs patent or patentrecurved), narrower leaf blade ( 0.2–0.8 mm vs 1–5 mm wide), sulcate or nearly terete in cross-section (vs. flat or slightly convex), golden-yellow tepals (vs. light yellow), and stamens with adnate filaments at their bases (vs. free). B. hirtella subsp. glabrata distinguishes from B. felipponei by its strong alliaceous smell (vs. absent), erect-ascendant leaves (vs patent or ascendant-recurved), with 0.2–0.8 mm wide (vs. 1–5 mm wide), and tepals oblanceolate or broadly elliptic in shape (vs. lanceolate or elliptic-lanceolate). Beauverdia hirtella subsp. glabrata is readily distinguished from B. vittata by its erect-ascendant leaves (vs. ascendant or ascendant-recurved), blades with 0.2–0.8 mm (vs. 1–4 mm wide), flowers with golden-yellow tepals (vs. white), oblanceolate or broadly elliptic in shape (vs. lanceolate or elliptic-lanceolate), and stamens with adnate filaments (vs. free). TABLE 1. A comparison of selected characters differing among the subspecies of Beauverdia hirtella .

Beauverdia hirtella
Character
subsp. glabrata	subsp. hirtella	subsp. lorentzii
Plant height (cm)	4–12	6–20	6–25
Arrangement and shape of leaves	spirally arranged, erect- ascendant, blades narrowly linear-filiform	spirally arranged, erect-ascendant or ascendant-reflexes, blades narrowly linear-filiform	spirally arranged, patentrecurved, blades linear
Leaf-blades size (mm)	40–120 × 0.2–0.8	40–180 × 0.9–2	80–210 × 2–5
Shape and size of leaf-blade in cross section (mm)	nearly terete or broadly elliptic, 0.2–0.8 × 0.2–0.5	elliptic or broadly elliptic, 0.9–2 × 0.2–0.6	flat or slightly convex, 2–5 × 0.2–0.5
Indument	Absent or rarely present, scarce in the distal third of scapes	densely papillose-pubescent, covering the leaves, scapes and pedicels	densely papillose-pubescent, covering the leaves, scapes and pedicels
Type of trichomes and size (µm)	1-celled with terminal cell uncinate, 60–90	1-celled and terminal cell straight, 110–180	idem subsp. glabrata
Scapes length (cm)	2–8	3.5–13	4–12
Number of tepals	6 (3 + 3)	6 (3 + 3)	8 (4 + 4)
Outer tepal shape and size (mm)	oblanceolate or elliptic; 11–15 × 4.5–8	oblanceolate or elliptic; 12–18 × 6–10	oblanceolate or elliptic; 15–23 × 8–10
Inner tepal shape and size (mm)	oblanceolate, elliptic or broadly elliptic; 10–14.5 × 4–7.5	elliptic; 11–17 × 3.5–8	elliptic; 10–20 × 5–8
Stamens number; and filaments length (mm)	6; 8–9.5	6; 8–11	8; 7.5–10
Ovary shape and size (mm)	obovate-oblong, 2.5–3.3	obovate-oblong or oblong; 2.7–5	oblong; 3.5–5
Number of ovules per locule	12	9–12	13–16
Capsule shape and size (mm)	nearly spherical or obovate, 5–7.5 × 4.5–6	obovate-oblong; 7–10 × 4–6	obovate-oblong or oblong; 7.5–11 × 4–6
Geographic distribution and habitat	On rocky grasslands in south- central Rio Grande do Sul state, Brazil	On rocky grasslands and grasslands in Uruguay and southern Rio Grande do Sul state, Brazil	On grasslands in southeastern Corrientes and eastern Entre Ríos provinces, Argentina

By its general aspect, B. hirtella subsp. glabrata is very similar to Nothoscordum setaceum ( Baker 1871: 381 ) Ravenna (1968: 58) , considering the leaves with glabrous and narrowly linear-filiform blade and size and shape of scapes, and fruits. However, B. hirtella subsp. glabrata can be immediately recognized by the larger size of the flowers, the shape of the tepals (oblanceolate or broadly elliptic vs. lanceolate or elliptic-lanceolate) and the stamens with filaments adnate at their bases (vs. free). Nothoscordum setaceum is an enigmatic species, given that the location of the type collection is doubtful and because it is known only from a single herbarium voucher. This species was recognized under Ipheion by Traub & Moldenke (1955) , and as Nothoscordum by Ravenna (1968) . More recently, it was considered a doubtful species of Beauverdia , or even a possible valid name for B. vittata ( Sassone et al. 2014a ) . Beauverdia subsessilis is another taxon which must be compared with B. hirtella subsp. glabrata , since this species is glabrous and was mistakenly included in the synonymy of B. hirtella (Deble unp. data); however, in B. subsessilis the alliaceous smell is absent (vs. present), the bulbs emit lateral rhizomes (vs. bulbs without rhizomes) and its leaves display 0.9–2 mm wide, and are flat in cross-section (vs. leaves 0.2–0.8 mm wide, nearly terete or broadly elliptic in cross-section). Nothoscordum marchesii Crosa (2005: 254) is another taxon similar with B. hirtella subsp. glabrata , due the strong alliaceous smell and narrowly linear-filiform and erect-ascendent or erect leaves, with up to 0.8 mm wide. However, B. hirtella subsp. glabrata can be easily distinguished by its bigger flowers (16–24 × 7.5–10 mm vs. 8–15 × 4.5–8 mm ), golden-yellow colored (vs. white-cream), and larger outer tepals (11–15 × 4.5–7 mm vs. 10–12 × 4–4.5 mm ) and inner tepals (10–14.5 × 4–6.5 mm vs. 8–10 × 4–4.5 mm ). Other materials examined:— Beauverdia hirtella subsp. hirtella . BRAZIL . Rio Grande do Sul : Aceguá, BR 473, no campo, 14 May 2017 , L.P. Deble et al. 16988 (PACA!); Dom Pedrito, Serrinha, 18 May 2018 , L.P. Deble et al. 18851 (PACA!); Santana do Livramento, estrada aeroporto, no campo, 16 June 2014 , L.P. Deble et al. 15317 (PACA!). URUGUAY . Maldonado : Piriápolis, Cerro Inglés, inter saxa, floreis aureis, C. Osten 5499 (MVM); Sierra de las Ánimas, 5 June 1968 , P. Izaguirre et al. s. n. (MVFA 2736!); Cerro Pan de Azúcar, 18 May 1973 , B. Rosengurtt 1845 (MVFA!); Cerro Arequita, May 1964 , Del Puerto et al . 3479 (MVM!); Minas: Cerro Verdún, 18 May 1960 , P. Izaguirre 1340 (MVFA); Cerro Verdún, 24 May 1959 , O. Crosa s. n . (MVFA 2154); Montevideo : sur la Bonite, April 1836 , M . Gaudichaud ( holotype of Triteleia hirtella B100247568 photo!, isotype P00852554 photo!); Rivera : Cerro junto al Miriñaque, 3 May 1960 , B. Rosengurtt B-8180 (MVFA!); Rocha : F. Felippone 3491 (SI photo!); Without precise locality, F. Sellow 3664 (K00524638, left specimen photo!). Beauverdia hirtella subsp. lorentzii . ARGENTINA . Corrientes : Curuzú Cuatiá, 3 May 1986 , T.M. Pedersen 14553 (SI!); Monte Caseros, 19 May 1952 , E.G. Nicora 6247 (SI, photo!); Monte Caseros, cercanías de, E.G. Nicora 5360 (SI!); Trés Bocas, no campo, 22 May 2014 , L.P. Deble & F.S. Alves 14739 (PACA!); Entre Ríos : Concepción del Uruguay, en suelo arenoso, 24 May 1877 , P.G. Lorentz 1024 (CORD); Campichuelo, orilla del río Uruguay, 10 April 1963 , A. Burkart et al. 24138 (SI).