Diversity of Scleria ( Cyperaceae ) in Santa Catarina , Brazil

Scleria is the fifth largest genus of Cyperaceae, and the third most diverse genus of Cyperaceae in Brazil. In Santa Catarina the floristic account of Cyperaceae is in need of updating, particularly as the Atlantic forest, home to many species of Scleria, is one of the most threatened environments worldwide. Scleria is thus a leading candidate for an updated floristic account, particularly so as to identify the conservation status of its species within the remaining natural vegetation. The survey has confirmed thirteen taxa including a new subspecies, S. georgiana ssp. australis, here described. Scleria sellowiana is lectotypified here. A species identification key, morphological descriptions, geographic distributions, global conservation status according to IUCN Red List criteria, habitats, phenological aspects, taxonomic notes and illustrations are provided for each species.


Introduction
Scleria P.J. Berg. is the fifth largest genus of Cyperaceae (Heywood et al. 2007), comprising 220-250 species (Goetghebeur 1998;Camelbeke et al. 2003) worldwide; it is the third most diverse genus of Cyperaceae in Brazil after Rhynchospora Vahl (147 species) and Cyperus L. (101 species), with 73 species recorded by Alves et al. (2014).Although Cyperaceae occur preferentially in the tropics, in open and wet or flooded environments, species of Scleria mostly prefer forests with diffused light, but also occur in relatively dry environments.
Important taxonomic and floristic studies addressing the family, and treating the genus Scleria in different areas of the Americas, include Core (1936), Adams (1994), Strong (1994Strong ( , 2007)), Kearns et al. (1998), Strong & Acevedo-Rodríguez (2005), Guaglianone et al. (2008) and Alves et al. (2014).In Brazil, however, few recent regional floras focusing on this genus are known (Muniz & Shepherd 1987;Araújo 2009).Barros (1960) reported 15 species from Santa Catarina, based on collections made up until the late 1950s.Since then not only has the vegetation in southern Brazil changed, but species circumscription and nomenclature have also changed.Species new to science have been described (Strong 1994(Strong , 2007;;Araújo & Brummitt 2011), prompting the need for an updated account of Scleria in Santa Catarina, particularly in the light of the effort to produce a complete inventory of the plant species diversity of Brazil.
This study aims to understand the species diversity of Scleria in Santa Catarina, updating the species circumscriptions and nomenclature, enabling their identification and documenting their conservation status worldwide.An identification key, morphological descriptions and general comments on distribution, habitat, ecology and phenology, as well as illustrations, are also presented.

Material and Methods
This investigation was based on primary taxonomic literature and the study of 800 specimens from: BM, CRI, FLOR, FURB, HBR, ICN, JOI, K, MBM, NY (pro part), PACA and RB; collections from MO, NY (pro part) and P were accessed online.Herbarium acronyms follow Thiers (continously updated).The sign "!" after the herbarium abbreviation indicates that the type sheet or image has been seen.Field expeditions were conducted between 2010 and 2011, the top set of vouchers was deposited at FLOR and duplicates sent to: BM, COL, CRI, FURB, GUA, HB, HBR, HUEFS, ICN, K, MBM, MO, NY, P, PACA, R, RB, SI, SP, depending on the availability of each species.A selected specimen is cited bellow the description; the remaining material is listed in the Annex as "index of collections studied".
Species descriptions are based on authors' collections, while the genus description is based on the species found in the area, following Beentje (2010) and Camelbeke et al. (2003).Accepted names follow WCSP (2014), and their respective synonyms are found in Alves et al. (2014).Only synonyms lately used in the herbaria investigated are cited.
Preliminary IUCN categories were established after applying the ArcView GIS 3.3 Conservation Assessment Tool extension (Willis et al. 2003), with values for "extent of occurrence" (EOO) and "area of occupancy" (AOO) calculated in km 2 (IUCN 2012).The results were analysed and final IUCN Red List ratings produced following the guidelines of IUCN (2014).

Results and Discussion
Scleria is represented in Santa Catarina by 13 confirmed taxa, one of which is a new subspecies.They are found on sandbanks, in coastal fields, highlands and in Atlantic forest occupying different environments, especially forest edges, transitional vegetation between forest and grassland, wet grasslands and wetlands.The highest concentration of species and individuals was recorded in the east of the state, especially in the coastal areas, and in the southern and northern plateaux.
Scleria filiculmis is endemic to Brazil (Guaglianone et al. 2008), recorded from the Atlantic forest (Alves et al. 2014) in the highlands of southern Brazil.Although Core (1936) and Ferreira & Eggers (2008) described this species as "lacking a hypogynium", detailed study of the type material revealed a hypogynium with tubercles.Scleria colorata (an accepted species) is not confirmed in Santa Catarina; the collection R. Reitz & R.M. Klein 10067 (HBR) identified as such by Barros in 1963 is actually S. filiculmis.The inflorescence of S. filiculmis resembles S. georgiana Core ssp.australis and S. sellowiana Kunth, differing by the presence and/or shape of the hypogynium, dimension of the leaf blades and the achene surface.Scleria filiculmis has been misidentified as S. balansae Maury, from which differs by the achene and hypogynium morphology.The latter is confirmed from Paraná but not yet known from Santa Catarina.
IUCN Red List category -VULNERABLE (VU B1ab(iii)+B2ab(iii)+D2).The Atlantic Forest is one of the most threatened habitats in the world (Brooks et al. 1999;Myers et al. 2000;Fonseca et al. 2003) and the restricted distribution of S. filiculmis in the Atlantic forest of the southern Brazilian highlands is concerning, particularly when much of this area has been used for pine plantations, cattle ranching and agriculture.The species has so far been found only in the localities of Campo Alegre and Serra Geral in Santa Catarina and Vacaria in Rio Grande do Sul, none of which is protected.The specimen from the vicinity of Vacaria was collected in 2006 by the last author in disturbed Atlantic forest and is the first record for the species outside the state of Santa Catarina.However, the specimen collected in Serra Geral in 1891 (Ule 1939) -the type collection -has no more detailed locality than this.Due to the lack of detail for the type locality it is impossible to be sure of the exact size of the extent of occurrence; however, we can be sure that S. filiculmis does not exceed the threshold of 20,000 km 2 for Vulnerable under Criterion B and also has a restricted Area of Occupancy of less than 20 km 2 if using a cell size of 2x2 km (IUCN 2014).A further locality from Guaratuba in Paraná remains to be confirmed.Threats to this species are from continuing clearance of the native vegetation for livestock and arable agriculture, and for forestry.Recent surveys in known localities have failed to record the species; further surveys in its three known localities and the surrounding areas need to be undertaken to confirm its present status and the number of mature individuals occurring in each location; additional research may reveal that this species should be reassessed as Endangered.Fig. 1i-l Perennial herb, erect, (30)-60-130 cm tall, caespitose-rhizomatous.Culm 1-5 mm wide, not branching, angle scabrous.Sheath 3.5-7.0cm long, winged; wings 1-2 mm wide; ligule absent; contraligule 2-6 × 1-5 mm, triangular (rare obtuse), glabrous, membranous appendage absent; leaf blade 14-25 × 0.5-0.7 cm, linear, apex acute or pseudopremorse, margin scabrous.Inflorescence a paniculodium, elongated, terminal and axillary, 2.5-8.0 × 0.5-2.0cm; bracts 6-11 × 0.3-0.5 cm.Spikelets subandrogynous or staminate, sessile and pedicellate, respectively, staminate glume 3-4 × 1.0-1.5 mm, pistillate glume 2-4 × 1.0-2.5 mm.Achene 1.8-2.3× 1.5-2.0mm, depressed-globose, smooth, usually glabrous, occasionally pilose, white, base muticous, pores absent, basal disk absent; stylobase absent.Hypogynium three-lobed, somewhat corky, lobe-margin entire, smooth.

Scleria gaertneri
Scleria gaertneri is a pantropical species (Camelbecke et al. 2003, recorded as S. melaleuca Rchb.ex Schltdl.& Cham.); in Brazil it has been recorded in all biomes (Alves et al. 2014).In this State it occurs from the northern coast to as far as the Island of Santa Catarina, including inland areas of the Vale do Itajaí on sunnier edges of humid forests, or less often occupying wet grassland, the edges of sandbank vegetation and disturbed forest remnants.In herbaria it is usually identifyed as S. pterota C. Presl (nomen nudum) or S. melaleuca (synonym, Longhi-Wagner et al. 2010).It differs from the other species by the elongated paniculodium, by its smooth and globose achenes and by the corky hypogynium, trilobed and with an entire margin.It flowers and fruits equally all year long.
IUCN Red List category -LEAST CONCERN (LC).A common, widely distributed species, also found in protected areas, with an EOO of more than 15,000,000 km 2 and an AOO of more than 17,000,000 km 2 .Fig. 1m-p Diagnosis: Rhizome slender, culms isolated, contraligule truncate, achenes slightly sulcate vertically towards the base; it differs from the North American taxon which shows a nodulose rhizome, culms in tufts, the contraligule is absent, and the achene ribbed with three ridges (Ball et al. 2014).
Scleria georgiana ssp.australis has been found only in Santa Catarina state, although it has been suggested to also occur in Paraguay (Barros 1960); however, this has never been confirmed and no specimens are cited with which this can be checked.The species is recorded from scarce populations near the coastline, in the Parque Estadual do Maciambú (Palhoça), and formerly in Jurerê beach (Florianópolis) and in Sombrio, the latter being the southern known limit for the species; it is possibly restricted to the Parque Estadual do Maciambú.This taxon inhabits moist sandbanks and sandy marshes.Until very recently both taxa were recognized as one, performing a rather odd disjunction between the neotropical North and Central America and the subtropical region of Southern South America.A careful morphological and ecological study proved to be essential in recognizing this new taxon.Scleria georgiana ssp.australis differs from others described here based on the achene shape, surface and ornamentation, which has pores present at the base.It was collected with flowers and fruits in January, September and October.
Scleria latifolia is a neotropical species (Core 1936;Camelbeke et al. 2003), present in Amazon and Atlantic forests, Caatinga and Cerrado of all Brazilian regions (Alves et al. 2014).In Santa Catarina it is distributed throughout, occupying the edges of moist forests, shaded and humid environments of shrubby to tall sandbank vegetation, well adapted to anthropogenic environments such as edges of trails, roadsides and forest remnants near watercourses.Scleria latifolia has pioneer potential in degraded environments.Scleria latifolia resembles S. panicoides Kunth, differing by the shape, colour and ornamentation of the achene.The achene's white colour when dry turns dark violet when re-hydratated.Locally known as monkey razor (= "navalha-de-macaco", due to sharp leaf blades), bird nuts (= "baga-decurió", staple food for birds) or wild wheat (= "trigo-guarani", eaten by Guarani people).
IUCN Red List category -LEAST CONCERN (LC).A common, widely distributed species found in protected areas, with an EOO of more than 17,000,000 km 2 and an AOO of more than 16,000,000 km 2 .

Scleria leptostachya
Scleria leptostachya occurs in South America (Camelbeke et al. 2003); in Brazil it spreads from the northeast to the south (Alves et al. 2014).Santa Catarina is the southern limit of its occurrence where fewer records have been found, two from the coast, and three to the middle-west, those latter ones collected between 1959 and 1962.This species differs from all other species by its panicle of fascicles, lax here whereas more congested in other species, and also by its achene ornamentation, unique to this species.
A morphological comparison between the type collections of Scleria catharinensis and S. leptostachya, supported by the protologue of these two species, confirmed that they are the same taxon, as stated by Guaglianone et al. (2008).The species flowers and fruits in January and February.
IUCN Red List category -LEAST CONCERN (LC).A common, widely distributed species, also found in protected areas, with an EOO of more than 4,000,000 km 2 and an AOO of nearly 3,000,000 km 2 .

Scleria microcarpa Nees ex
Scleria microcarpa is mostly neotropical (Camelbeke et al. 2003); in Brazil it is found in all biomes (Alves et al. 2014).Recorded in Santa Catarina once in 1955, at the forest's edge in Itajaí; it may no longer be found in this State.Scleria microcarpa and S. uleana Boeck.share similarities of the achene and hypogynium, but the latter is a climbing species while S. microcarpa is not, besides which they differ in inflorescence pattern, in the dimensions of the plant overall and of the achene, and hypogynium margins.It flowers and fruits in April.
IUCN Red List category -LEAST CONCERN (LC).A common, widely distributed species, found in protected areas, with an EOO of almost 17,000,000 km 2 and an AOO of more than 18,000,000 km 2 .

Scleria panicoides
Scleria panicoides is a neotropical species extending to southern South America (Camelbeke et al. 2003); in Brazil it is found in the Cerrado, Atlantic forest and Pampa (Alves et al. 2014).In Santa Catarina it occurs along coastal areas and inland in highlands, forests and humid environments, especially within and on the edges of healthy forest, hillside forest and from restored mined areas.Recorded recently in coastal forest (Florianópolis) and mountain forest (Joinville), in healthy, well-developed secondary forest, indicating perhaps some tolerance to degraded environments.Besides S. latifolia, mentioned above, S. panicoides also resembles S. plusiphylla Steud.differing in the number and density of spikelets, and achene ornamentation.It flowers and fruits throughout the year.
IUCN Red List category -LEAST CONCERN (LC).A widely distributed species occurring within protected areas, with an EOO of more than 4,000,000 km 2 and an AOO of more than 2,700,000 km 2 .

Scleria plusiophylla
Scleria plusiophylla is widely distributed in South America (Camelbeke et al. 2003); present in Atlantic rain forest and the Brazilian central savanna (Alves et al. 2014).In Santa Catarina it has been recorded in the Vale of Itajaí and the northern and southern plateaux, found in shaded environments inside moist forest.It flowers and fruits throughout the year.
IUCN Red List category -LEAST CONCERN (LC).A common, widely distributed species found within protected areas, with EOO of more than 2,800,000 km 2 and an AOO of more than 2,000,000 km 2 .
Scleria secans is a widespread neotropical species (Camelbeke et al. 2003), found in all biomes in Brazil (Alves et al. 2014).In Santa Catarina it has been recorded from the Vale do Itajaí and the northern plateau, within moist forests and at their edge.Local populations may form a thicket reaching ca.7m tall, covering trees of relatively open, disturbed forests, promoting the establishment of other shade-loving species.It is known locally as dog's razor ("capa-cão") because its leaves and culms are strongly scabrous and very sharp.It differs from all other species cited here by its branched culm and presence of a ligule.It flowers and fruits all year round.
IUCN Red List category -LEAST CONCERN (LC).A common, widely distributed species, also found within protected areas, with an EOO of more than 16,700,000 km 2 and an AOO of more than 14,400,000 km 2 .
Scleria sellowiana is a subtropical species known from southern South America (Barros 1960) and within Brazil only recorded from the Atlantic Forest of southern Brazil (Alves et al. 2014).In Santa Catarina it is distributed along the coastal area, in the northern and southern plateaux, on wet grasslands, along the edges of streams and on sunny slopes.Sellow and Kunth, respectively the collector of the type of S. sellowiana and the author of the species, were based in the Berlin herbarium (B).Although Kunth (1837) did not specify where the type collection was placed, it is reasonable to believe that the holotype was placed in the Berlin herbarium (B).Stafleu & Cowan (1981) state that Sellow's Cyperaceae type collection remains extant at B; however, a recent search was conducted for S. sellowiana type material but proved fruitless.It is therefore likely to have been destroyed.Since the K sheet is so far the only Sellow collection found bearing Kunth's handwriting, and it also provides a perfect match of the species diagnosis, it is therefore chosen as the lectotype.Scleria sellowiana resembles S. balansae Maury ex Micheli, S. ciliata Michx., S. filiculmis and S. georgiana ssp.australis, all of which it is often misidentified as in herbarium collections.However, diagnostic taxonomic characters are found in the achene: smooth in S. balansae and S. georgiana ssp.australis as opposed to reticulate or rugose in S. sellowiana.Scleria filiculmis has a disc compressing the tubercles of the hypogynium, which is lacking in S. sellowiana, while S. ciliata has sharp projections and cavities at the base of the achene, which are absent from the achene of S. sellowiana.The species has been collected in flower and fruit from October to January.
IUCN Red List category -LEAST CONCERN (LC).A widespread species across southern South America, including within protected areas, with an EOO of more than 4,200,000 km 2 and an AOO of more than 1,500,000 km 2 .Camelbeke et al. 2000).
Scleria uleana is endemic to Brazil, occurring in the Atlantic Forest domain of the South and Southeast regions (Alves et al. 2014).Records from Santa Catarina report populations only in coastal areas, occurring at the edges of streams in preserved or disturbed sandbank vegetation.Flowers and fruits from April to December in Santa Catarina.
Scleria variegata is endemic to southern Brazil (Core 1936;Muniz & Shepherd 1987), occurring in the Atlantic Forest domain (Alves et al. 2014).In Santa Catarina, a single population has been found in Florianópolis, recorded in 2007 and again in 2010, at the edges of coastal forest.The species is distinguished by its climbing habit, IUCN Red List category -LEAST CONCERN (LC).This species is widely distributed although uncommon within its range, yet it has been recorded from several localities, including within protected areas.It has an EOO of almost 900,000 km 2 and an AOO of more than 430,000 km 2 .
Barros (1960)crocarpa Nees ex Kunth and S. secans (L.) Urban are well distributed across the whole American continent, while S. distans Poir., S. gaertneri Raddi and S. latifolia Sw. are pantropical, extending into subtropical areas.However, four taxa are confirmed as endemic to Brazil: S. filiculmis Boeck., S. georgiana Core ssp.australisA.C.Araujo, S. uleana and S. variegata (Nees) Steud., the first being microthermic and the latter being macrothermic with its austral distribution limit in Santa Catarina; similarly, S. leptostachya Kunth also has its southernmost distribution in this state.Scleria filiculmis and S. georgiana subsp.australishavebeen assessed as Vulnerable under IUCN criteria, both being endemic to southern Brazil; the remaning taxa are assessed as Least Concern.Scleria balansae Maury, S. hirtella Sw., S. myricocarpa Kunth and S. pauciflora Muhl.exWilld.citedbyBarros(1960)were all misidentifications, and have not been confirmed in Santa Catarina.This paper presents as a taxonomic novelty a new subspecies endemic to Santa Catarina, and proposes a lectotype for S. sellowiana.
. It flowers and fruits all year.