Three new species of Nothoscordum (Amaryllidaceae) from the Campos eco-region of the Grassland Ecosystems of Río de la Plata, Southeast South America

Three new species of Nothoscordum are described: Nothoscordum elongatum Deble & B.P.Moreira, Nothoscordum parvum Deble & B.P.Moreira and Nothoscordum urutauense Deble & H.A.Keller. Nothoscordum elongatum and N. parvum grow in central-eastern Rio Grande do Sul state, while N. urutauense is narrowly endemic in southern Misiones Province, northeast Argentina. The three new taxa are characterized by their strong alliaceous smell, pilose leaves and scapes, and one or two-owered in�orescence. Nothoscordum elongatum is segregated from Nothoscordum marchesii by its densely pilose leaves and scapes (vs. glabrous leaves and scapes), �owers with longer pedicels and ovaries with fewer ovules per locule. Nothoscordum parvum is similar to N. modestum and N. urutauense; however, can be readily separated from both species by its bulbs with bulbils surronding the main bulb, by its narrowly infundibuliform �owers, fused towards the base for 1.8–2.2 mm and by its scapes erect at fruit maturity. Nothoscordum urutauense differs from N. inundatum by its broader leaves, by its �owers with tepals fused at the base for 0.5–1 mm, and by its free �laments at base. These three species of Nothoscordum are described, illustrated, compared with morphologically similar taxa, and featured with their geographic distribution.


Introduction
The Campos eco-region of the Grassland Ecosystems of Río de la Plata (RPG sensu Soriano et al. 1992;Bilenca and Miñarro 2004) comprises in approximate lines the Mesopotamic phytogeographic province and the Uruguayense district of the Pampean phytogeographic province of Cabrera (1976), and thereby are distributed in all Uruguayan territory, part of the Argentine provinces of Entre Ríos, Corrientes and southern Misiones, and the southern half of the state of Rio Grande do Sul in Brazil (in the Brazilian portion it corresponds approximately to the Pampa Biome, sensu IBGE 2004).The mentioned region is one of the main centers of diversity and endemism of Nothoscordum Kunth, having several endemic species, many of which are known only for one or a few places of occurrence (Deble and Moreira 2022).
Botanical collections carried out in 2018 and 2019 revealed three peculiar species of Nothoscordum, which display strong alliaceous smell, dense pilosity on scapes and leaves and one or two-owered in orescences.After the analysis of nomenclatural types, examination of exsiccates, studies of populations in their natural environments, and comparison with closely related taxa it was possible to identify that these three taxa are new, and are herein described.

Material and Methods
The research was part of the revision of the genus Nothoscordum in the Campos eco-region of the Grassland Ecosystems of Río de la Plata, Southeast South America, which is still in progress, and was carried out by eld surveys (central, north and northeastern Argentina, southern Brazil, Paraguay and Uruguay), and analysis of herbarium specimens of the herbaria CTES, FCQ, HDCF, ICN, MVHM, MVM, MVFA, MVJB, P, PACA, PY, SI and SMDB and digital imagens of the Herbaria B, G, K, MBM, NY, P and US (acronyms according to Thiers 2023).Collected specimens were deposited into CTES and PACA herbaria.The descriptions of the three new taxa are based on morphological characteristics of plants observed in natural habitat and on dry material.Figures 1, 2 and 3 was elaborated using Indian ink, and the illustrations are based on both live specimens and dry material.Figures 4 and 5 were made from photos of specimens in nature or in cultivation and the boards were edited with the Adobe photoshop software, version 24.5.For the elaboration of Figs. 6 and 7, it was utilized the software ArcMap version 10.7, and the features were created from the map base titled Topographic.Diagnosis Nothoscordum elongatum is morphologically similar to N. marchesii; however, can be promptly distinguished by its densely pilose leaves and scapes (vs.glabrous), owers with 15-32 mm long pedicels, (vs.3-8 mm long), and 5-6 ovules per locule (vs.13-16).
Etymology The adjective elongatus in Latin means elongated, a reference to the delicate and slender habit of the new species.
Phenology Flowering time April-May, fruiting time May-June.The owers open around midday and close in the late afternoon, withering after one or two days.During the owering period, populations exude a strong alliaceous odor, which becomes the predominant smell in the places and around areas of occurrence of specimens of Nothoscordum elongatum.
Conservation Based on the analysis of exsiccates and collections in the possible places of occurrence, only two small populations were identi ed, which indicates the rarity of the species.The extent of occurrence (EOO) is less than 100 km² and the Area of Occupancy (AOO) is less than 10 km².This species occurs in speci c habitat, in environments with poor drainage capacity in places of shallow soils and the bulbs developed in crevices of sandstone and conglomerate rocks.These environments are being modi ed by economic activities, as mining and agriculture.In accordance with the criteria of IUCN (2019), Nothoscordum elongatum can be considered Critically Endangered (CR), based in direct threat, geographic distribution, occupation area and population size Diagnosis Nothoscordum parvum is morphologically related to N. modestum and N. urutauense; however, it can be readily separated from N. modestum by its bulbs surrounded by bulbils (vs.simple), with strong alliaceous smell (vs.mild alliaceous smell when crushing the cataphylls and foliage), leaves and scapes densely pilose (vs.glabrous), owers infundibuliform, fused towards the base for 1.8-2.2mm (vs.broadly campanulate, fused towards the base for ca.0.5 mm long), and scapes erect at fruit maturity (vs.humifuse).Nothoscordum parvum differs from Nothoscordum urutauense by its bulbs surrounded by bulbils (vs.simple), by its narrowly liform leaves (0.4-0.5 mm vs. 0.8-1.6 mm), by its erect scapes at fruit maturity (vs.humifuse) and by its nearly obovate and slightly 3-humped fruits (vs.spherical and strongly 3-humped).

Discussion
The study of new species and eld analysis of other previously described taxa, which were known only from herbarium material, have helped to reveal the limits of the genus Nothoscordum (Deble 2022; Deble and Moreira 2022).Important morphological characteristics, such as the position of the scapes at fruit maturation (humifuse vs. erect), the number of owers in the in orescence (solitary vs. numerous) and the union or not of the laments is more diversi ed within the genus than previously recognized.and N. urutauense.On the other hand, these species have long pedicellate owers, a morphological characteristic commonly found in Nothoscordum.Nothoscordum parvum, in turn, although morphologically related to the other species described here, has erect scapes during fruit maturation, as in most Nothoscordum species.

From a morphological point
of view, the description of these three new species corroborates the inclusion of Beauverdia in Nothoscordum, as previously suggested by other authors (for example, Crosa 2006; Pellicier et al. 2017; Sassone and Giussani 2018; Deble and Moreira 2022).Distinctive morphological characteristics for the recognition of Beauverdia, such as solitary owers and humifuse scapes during fruit maturation are found in Nothoscordum elongatum

Figures
Figures

Figure 1 NothoscordumFigure 2 Figure 3 Figure 4
Figure 1 Nothoscordum elongatum a Flowering specimens bFruiting specimen.c Flower, lateral view, showing pedicels, bracts and apex of scape d Flower, upper view e Flower open, showing pistil and stamens f Proximal part of the leaf blade and leaf sheaths gDistal part of leaf blade h Capsule i Seeds j-k Cross section of leaf blade l Trichome of the scape (a, c-g, j-l from Deble and Moreira 20153; b, h-i from Deble and Moreira 20294)

Figure 5 Nothoscordum
Figure 5 Nothoscordum urutauense a Flowering specimen b Flower, lateral view, showing bracts and pedicels c Open ower, evidencing Stamens and pistil d Flower upper view e Capsule, partially opened and showing the seeds 1.8-2.2mm, the outers ones oblanceolate, 10-11 × 2.1-2.3 mm, apex slight acute, base attenuate; the inners ones elliptic or oblanceolate, 9-10 × 2-2.2 mm; apex slight acute, base attenuate; tepal middle nerves greenish-white, little pronounced, thicker towards the base.Staminal laments 4.5-5.5 mm long, awl-shaped, translucent, yellowish-white, free at their bases and attached directly to the tepal; anthers 1-1.3 mm long, curved, pollen golden-yellow.Cyperaceae and Poaceae species.Nothoscorum collinum Ravenna is another endemic in the same area, but this species occurs in more dry places and often the bulbs developed in crevices of rocks.Conservation Based on the analysis of exsiccates and collections in the possible places of occurrence, only three small populations were identi ed, which indicates the rarity of the species.The extent of occurrence (EOO) is less than 100 km² and the Area of Occupancy (AOO) is less than 10 km².This species occurs in speci c habitat, grows in environments with poor drainage capacity in places of wet shallow soils, with abundant organic matter and rich in clay minerals, developed over concavity of granitic rocks.These environments are being modi ed by economic activities, as mining and agriculture.In accordance with the criteria of IUCN (2019), Nothoscordum parvum can be considered Critically Endangered (CR), based in direct threat, geographic distribution, occupation area and population size [CR B1 + B2b (i, ii, iii, iv, v) + c (iii, iv)].
Phenology Flowering and fruiting time September.The owers open in the middle afternoon during one or two days.During the owering period, populations exude a strong alliaceous odor, which becomes the predominant smell in the places and around areas of occurrence of specimens of Nothoscordum parvum.Additional material examined BRAZIL.Rio Grande do Sul: São Gabriel, Três Cerros, 410m, 18 Sep 2021, LP Deble and BP Moreira 20233 (PACA!); trajeto a vila de Palma, 280m, 18 Sep 2021, L.P. Deble and BP Moreira 20234 (PACA!).Distribution and Habitat Nothoscordum parvum occurs in central-southern Rio Grande do Sul state, Brazil in the border between São Gabriel and Lavras do Sul municipalities.The specimens grow between 280-410m, in environments with poor drainage capacity, in places of wet shallow soils, with abundant organic matter and rich in clay minerals, developed over concavities of granitic rocks from the domain of undeformed granitic rocks (CPRM 2009).The individuals grow associated with mosses, Drosera brevifolia Pursh (Droseraceae), Nothoscordum dyalistemon (Guagl.)Crosa (Amaryllidaceae) and small and delicate Geophyte 8-14 cm high above the soil.Bulb 6-8 × 6-8 mm, nearly spherical or ovoid, simple, with strong alliaceous smell; outer cataphylls dark-brown, the innermost whitish.Leaves at anthesis 3-10, spirally arranged; leaf sheaths 2-6 cm long, subterranean, straw-colored; leaf blades 60-130 × 0.8-1.6 mm, canaliculate in cross-section, dark-green, erect-ascendant or ascendant re exes, margin thickened and rigid, apex slight acute or obtuse, abundant pilosity composed of 1-celled trichomes with straight or slightly curved cell, trichomes 150-200 µm long.Scapes 1-5, thicker than the leaves, 4-8 cm long, subcylindrical, green or purplish-green, erect, then humifuse at fruit maturity, with alliaceous smell when breaking, densely covered by 1-celled trichomes with straight terminal cell, trichomes 200-240 µm long.Bracts 2, ovate-elliptic, 6-10 × 3-4.5 mm, shortly fused at base for ca. 1 mm.Pedicels 15-25 mm long, light-green, densely covered by 1-celled trichomes shorter than the trichomes of scapes.In orescence 2-owered.Flowers 8-10 × 7-8 mm, campanulate, Conservation The species occurs in the Urutaú Reserve site closest to a human population (next to the suburbs of the city of Candelaria and a short distance from an improvised garbage disposal site).The only known population has about 50 individuals distributed in about 200 m 2 .Other similar environments present in the Urutaú Reserve, in the Candelaria Department and in several localities in the south of Misiones Province and north of Corrientes Province have been exhaustively explored, but no other populations have been found.All this suggests that according to criteria B1ab(iii) + D (IUCN 2019), the species is critically endangered (CR).

Table 1
Brief history of the generic and infrageneric interpretation of 1-2-owered species of Nothoscordum and some morphologically related taxa