Eriocaulaceae from Serra do Caraça, Minas Gerais, Brazil

Abstract

The Serra do Caraça, located in Minas Gerais state, Brazil, hosts a remarkable biological diversity, comprising rare and endemic plant species. Several naturalists have visited the area since the XIX century and made important botanical collections, many of these established as type specimens to describe different taxa, including Eriocaulaceae (Poales) species and varieties. This work provides a floristic treatment of Eriocaulaceae from Serra do Caraça, presenting species descriptions, distribution map, photographs, identification key, and notes on phenology and distribution. Thirty-three species were recorded in five genera, and Paepalanthus is the most numerous with 21 species, followed by Leiothrix (6 spp.), Syngonanthus (3 spp.), Comanthera (2 spp.), and Eriocaulon (1 sp.). Notable, among those are four species endemic to the Quadrilátero Ferrífero, one micro-endemic to the Serra do Caraça, and 16 endemic to Minas Gerais state. Five new synonyms are proposed. Eighteen taxa have type materials described to Serra do Caraça, corresponding to 12 species, five varieties, and one forma.

Key words:
Brazilian flora; campo rupestre; conservation; Poales; taxonomy

Resumo

A Serra do Caraça, localizada no estado de Minas Gerais, Brasil, possui uma notável diversidade biológica, com espécies raras e endêmicas. Vários naturalistas visitaram a área desde o século XIX e fizeram importantes coleções botânicas, várias destas designadas como espécimes tipo para descrever táxons, incluindo espécies e variedades de Eriocaulaceae. Este trabalho provê um tratamento florístico de Eriocaulaceae para a Serra do Caraça, apresentando a descrição das espécies, mapa de distribuição, fotos, chave de identificação, e informações sobre fenologia e distribuição. Trinta e três espécies foram registradas em cinco gêneros, sendo Paepalanthus o mais numeroso com 21 espécies, seguido por Leiothrix (5 spp.), Syngonanthus (3 spp.), Comanthera (2 spp.) e Eriocaulon (1 sp.). Destacam-se quatro espécies endêmicas para o Quadrilátero Ferrífero, uma micro-endêmica para a Serra do Caraça, e 16 espécies endêmicas para Minas Gerais. Propomos cinco novos sinônimos. Dezoito táxons possuem material tipo descrito para a Serra do Caraça, sendo 12 espécies, cinco variedades, e uma forma.

Palavras-chave:
flora brasileira; campo rupestre; conservação; Poales; taxonomia

Introduction

Eriocaulaceae are distinguished from other Monocots due to its capitulate inflorescences, tiny and unisexual flowers, and leaves usually in rosettes (Giulietti & Hensold 1990; Stützel 1998). Comprising approximately 1,200 species, it is pantropical and most diversified in mountains or plateaus, in open vegetation areas, especially over quartzitic soil (Giulietti & Hensold 1990; Giulietti et al. 2012). The Espinhaço Range located in Minas Gerais and Bahia states (east Brazil) is the center of diversity of the family, specially the Campo Rupestre vegetation (Giulietti & Hensold 1990; Giulietti et al. 1997, 2012; Stützel 1998; Costa et al. 2008). Many species are micro-endemic, restricted to a single locality or even to a single mountain (Giulietti et al. 2005; Costa et al. 2008; Echternacht et al. 2011).

The Quadrilátero Ferrífero (literally “iron quadrangle”) is a region with a quadrangular terrain of unique geological structure (Jacobi & Carmo 2008; Salgado & Carmo 2015), rich in iron ore deposits, being one of the most important reserves in the world (Dorr 1969; Alkmim 1987). This region presents a distinct geological formation from the Espinhaço Range (Uhlein & Noce 2012), but it is located adjacent to it to the south, and biogeographically it might be considered as part of the Espinhaço Range (e.g., UNESCO 2005; Assunção-Silva & Assis 2014). The complex litho-structure of the Quadrilátero Ferrífero promotes habitat heterogeneity, having a rich-associated biota. Thus, as indicated by Drummond et al. (2005), this region has an ​​extreme biological importance, with numerous endemics species (Vincent 2004; Jacobi et al. 2007; Echternacht et al. 2011). As a result of intense mining activity, landscapes are highly disturbed, and many of these species are threatened (Jacobi & Carmo 2008).

The Serra do Caraça is a quartzitic mountain in eastern Quadrilátero Ferrífero, delimited as a conservation unit (Private Reserve of Nature Protection, RPPN), belonging to the Santuário do Caraça (Ordinance 32 30/03/1994). It has a remarkable biological diversity, with more than 1,600 species of phanerogams (Augsten 2015). It is located within the Atlantic Forest domain, with high altitudinal variation, from 750 to 2,072 m elevation, the forest occupying mainly the surrounding valleys and the Campo Rupestre in the highest elevations (Ferreira et al. 1977; Vasconcelos 2000, 2011). Augsten (2015) estimated that 35% of the flora in Serra do Caraça are endemic to the Quadrilátero Ferrífero; as a conservation unit surrounded by mines, it might be considered a refuge for biodiversity.

The Serra do Caraça has an important historical record of biological research and several naturalists have visited it since the XIX century. Several of the collected specimens were used as type materials, including Eriocaulaceae species and varieties (e.g., Koernicke 1863; Ruhland 1903; Silveira 1928; Moldenke 1972, 1973). Since the 1970s, some studies of the flora of Serra do Caraça have been made, some as general floristic surveys (Ferreira et al. 1977; Oliveira 2010; Augsten 2015; Carmo et al. 2018), other on specific groups (Vasconcelos et al. 2002; Morais & Lombardi 2006; Mota 2006; Salino & Viveiros 2012; Castro 2017). They all highlight the area’s importance as irreplaceable for biodiversity conservation. For Eriocaulaceae, the species list generated by the latest floristic survey recorded 36 species, ranking the family among the ten most speciose (Augsten 2015).

Research led by taxonomist specialists has the potential to discover new records, as they have a focused and trained search. For example, Leiothrix gomesii Silveira, previously known from type collection only, was considered presumably extinct (Giulietti 1984; COPAM 085/97; Costa et al. 2008), but during a fieldwork in the highest summit of the Serra do Caraça in 2009, a new population was discovered (voucher: Echternacht, L. 1938. BHCB, UFU). In this scenario, the present work aims to monograph the Eriocaulaceae from Serra do Caraça, as it is one of the richest families in the region (Augsten 2015), and several of its species have type specimens collected there. We provide a taxonomic study, describing the morphology for each species, presenting an identification key, distribution map, photographs, and notes on phenology and distribution.

Material and Methods

The RPPN Santuário do Caraça is enclosed in the municipalities of Catas Altas and Santa Bárbara between the latitudes 20°12’09.7’’S-20º01’08’’S, and longitudes 43º34’53’’W-43°26’04.1’’W, with an area of 10,187.89 ha (Camargo 2001; Mota 2006). The climate is mesothermic (Cwb, rainy temperate) according to the classification of Köppen, classified as subtropical of altitude (Ferreira et al. 1977). The annual temperature average is between 16 °C to 21 °C, and the annual precipitation average is up to 1,500 mm (Brandão et al. 1994). Mountain relief is rugged, formed by a valley with a lower elevation of 750 m surrounded by a mountain ring with high slopes, being the highest summit the Pico do Sol at 2,068 m (Ferreira et al. 1977) (Fig. 1). Other notable peaks are: Pico do Inficcionado (2,068 m), Pico da Carapuça (1,955 m), Pico da Canjerana (1,890 m), and Pico da Conceição (1,800 m) (Ferreira et al. 1977). The vegetations in the RPPN Santuário do Caraça are the Campo Rupestre, Seasonal Semideciduous Forest, Gallery Forests, Riparian Forests, and Nebular Forests (Morais & Lombardi 2006; Vasconcelos 2011) (Fig. 1).

Figure 1 -
a-f. Vegetations in RPPN Santuário do Caraça, Minas Gerais, Brazil - a. Campo Rupestre formation at the top with forest formations in the surrounding valleys, on the trail to Pico da Carapuça; b. Campo Rupestre formation in Pico do Sol; c. Forest formation on the trail to Gruta de Lourdes; d. Campo Rupestre formation in Pedra da Paciência, on the trail to Bocaina; e. Campo Rupestre formation, view to Cascatinha on the trail to Bocaina; f. Riparian forest, Tabuões. Images: a,c-f. Ambar Soldevila; b. Livia Echternacht

This study was based on fieldwork and analysis of herborized specimens. An initial survey was made consulting databases and images from SpeciesLink (<https://specieslink.net/search/>) and Virtual Reflora Herbarium (<http://reflora.jbrj.gov.br/reflora/herbarioVirtual/>) for all the species of Eriocaulaceae recorded from RPPN Santuário do Caraça. The distribution and phenology of the species were then analyzed from the obtained records. Images available were studied, with particular attention to type specimens. Field expeditions were carried out, covering most of the principal localities of the RPPN Santuário do Caraça, in 2008 (March, May, and June), 2009 (February, March and July), 2012 (June), 2020 (November), and 2021 (July). Information on habitat, habit, phenology, distribution, population size, and conservation were obtained in field and herbaria, together with photographs and geographic coordinates. Specimens collected were deposited in herbaria BHCB, HUFU, OUPR, and SPF (abbreviations according to Thiers, continuously updated).

Identification of the species were made consulting nomenclatural types and key literature, mainly the Flora e Funga do Brasil, specific regional floras, and taxonomic revisions for Eriocaulaceae (Koernicke 1863; Ruhland 1903; Silveira 1928; Giulietti 1984; Sano 1999; Parra 2000; Trovó 2010; Trovó & Sano 2010; Trovó et al. 2013; Echternacht 2012; Watanabe 2015; Echternacht et al. 2015; Chagas 2017; Giulietti 2020b). Classification follows Mabberley (2017) and Christenhusz et al. (2018), considering Paepalanthus as a large monophyletic genus.

Morphological features were analyzed under stereomicroscope; capitula and flowers were rehydrated from dried specimens. Descriptions of vegetative structures follow the terminology of Radford et al. (1974), while for reproductive structures Weberling (1989), Rosa & Scatena (2007), and Scatena et al. (2008). Nomenclatural and taxonomic interpretations and decisions were made on the ground of the extensive research by Dr. Echternacht in American and European herbaria where most of the Eriocaulaceae types are located (B, BM, C, F, G, K, L, LL, LE, M, MO, NY, OXF, P, R, RB, S, SPF, and UPS) (abbreviations according to Thiers, continuously updated).

We assumed the following morphological characteristics as general patterns: plants terrestrial, involucral bracts adaxially glabrous, flowers trimerous and isostemonous, anthers cream, styles and nectariferous portions releasing at the same height, indumentum with simple and patent hairs. Capitula morphology varies during development from obconic to hemispherical. Species that differ in these characters will be specified in their descriptions. Fruits and seeds were often missing and did not provide diagnostic characters, so we opted not to describe them.

All specimens cited as “examined material” were physically analyzed; whenever physical specimens were not available in the visited herbaria, we consulted online databases and imagery and cited them under “local records”. In some label notes, the Pico do Inficcionado is referred to as 2,075 m elevation, but the official altitude is 2,068 m; this difference is probably due to variation in the GPS calibration. The distribution map (Fig. 2) was produced with the support of the QGIS software (QGIS Development Team 2021) using layers available from IBGE (2021). Records without coordinates were georeferenced whenever possible, considering the information on locality in the label notes using Google Earth.

Results and Discussion

A total of 33 species of Eriocaulaceae was recorded in RPPN Santuário do Caraça, belonging to six genera (Tab. 1). Most speciose genus is Paepalanthus, with 21 species; followed by Leiothrix (6 spp.), Syngonanthus (3 spp.), Comanthera (2 spp.), and Eriocaulon (1 sp.). The most widely distributed species in RPPN Santuário do Caraça, with abundant populations, are P. hilarei Körn., Eriocaulon ligulatum (Vell.) L.B.Sm., Leiothrix flavescens (Bong.) Ruhland, L. vivipara (Bong.) Ruhland, and Paepalanthus planifolius (Bong.) Körn. Also, the species P. leucoblepharus and P. mollis Kunth have abundant populations but are restricted to the highest areas, above 1,600 m elevation.

Eriocaulaceae in the RPPN Santuário do Caraça are primarily concentrated at the Campo Rupestre formation, especially in the highest areas and peaks. Localities richer in species are Pico do Inficcionado (13 spp.), and Pico do Sol (7 spp.). Leiothrix gomesii, a rare species with only two known populations, occurs in Pico do Sol, the highest mountain peak in the Quadrilátero Ferrífero. Another species worthy mentioning is Paepalanthus suffruticans, micro-endemic to the RPPN Santuário do Caraça; it is the only species found in riparian forest, in the trail to Capelinha and Gruta de Lourdes, with two abundant populations; it also has records from Campo Rupestre, in the Pico da Carapuça and Pico do Sol. These two species deserve more attention for conservation purposes in RPPN Santuário do Caraça.

Among the 33 species occurring in RPPN Santuário do Caraça, 16 are endemic to Minas Gerais (Tab. 1, Sano et al. 2024), among which four are endemic to the Quadrilátero Ferrífero: Leiothrix gomesii, Paepalanthus camptophyllus, P. leucoblepharus, and P. suffruticans, this last micro-endemic to the RPPN Santuário do Caraça (Rodrigues 2022).

Figure 2 -
Map with the principal localities, mountain peaks, and distribution of Eriocaulaceae records in RPPN Santuário do Caraça, Minas Gerais, Brazil

Table 1
List of species and their respective distributions. State abbreviations: BA = Bahia, ES = Espirito Santo, GO = Goias, PB = Paraíba, PI = Piaui, RJ = Rio de Janeiro, SP = São Paulo, TO = Tocantins. Brazilian regions abbreviations: CW = Central-West, N = North, NE = Northeast, S = South, SE = Southeast.

	Species	Author	Distribution
1	Comanthera centauroides	(Bong.) L.R.Parra & Giul.	BA, MG, RJ
2	Comanthera nivea	(Bong.) L.R.Parra & Giul.	ES, MG, RJ, SP
3	Eriocaulon ligulatum	(Vell.) L.B.Sm.	ES, MG, SP
4	Leiothrix curvifolia	(Bong.) Ruhland	MG
5	Leiothrix flavescens	(Bong.) Ruhland	South America
6	Leiothrix gomesii	Silveira	Southeastern QF
7	Leiothrix longipes	Silveira	MG
8	Leiothrix mucronata	(Bong.) Ruhland	MG
9	Leiothrix vivipara	(Bong.) Ruhland	MG
10	Paepalanthus calvus	Körn	MG, SP
11	Paepalanthus camptophyllus	Ruhland	QF
12	Paepalanthus ciliolatus	Ruhland	MG
13	Paepalanthus conduplicatus	Körn	MG
14	Paepalanthus dianthoides	Mart.	MG
15	Paepalanthus elongatus	(Bong.) Körn.	BA, GO, MG, SP, TO
16	Paepalanthus erectifolius	Silveira	BA, MG
17	Paepalanthus flaccidus	(Bong.) Kunth	GO, MG, SP
18	Paepalanthus freyreissii	(Thunb.) Körn.	Endemic to MG
19	Paepalanthus hilairei	Körn.	CW, N, NE, SE
20	Paepalanthus leucoblepharus	Körn.	Caraça and vicinities
21	Paepalanthus manicatus	Poulsen ex Malme	BA, GO, MG, PI, SP
22	Paepalanthus melaleucus	(Bong.) Kunth	BA, MG
23	Paepalanthus mollis	Kunth.	MG
24	Paepalanthus planifolius	(Bong.) Körn.	CW, SE, S
25	Paepalanthus polyanthus	(Bong.) Kunth	CW, NE, SE, S
26	Paepalanthus pubescens	Körn.	MG, SP
27	Paepalanthus scleranthus	Ruhland	BA, MG
28	Paepalanthus suffruticans	Ruhland	Caraça
29	Paepalanthus vaginatus	Körn.	MG
30	Paepalanthus vellozioides	Körn.	MG
31	Syngonanthus anthemiflorus	(Bong.) Ruhland	MG
32	Syngonanthus caulescens	(Poir.) Ruhland	South America
33	Syngonathus gracilis	(Bong.) Ruhland	BA, ES, MG, PB, RJ

The floristic survey by Augsten (2015) recorded 36 species of Eriocaulaceae for RPPN Santuário do Caraça. The review of this list with updated determinations led to the exclusion of eight species (Tab. 2). On the other hand, five (5) species were added to the list, with new records: Leiothrix mucronata (Bong.) Ruhland, Paepalanthus calvus Körn., P. erectifolius Silveira, P. pubescens Körn., and P. vaginatus Körn.

In our research, we found a total of 18 taxa with type specimens described to RPPN Santuário do Caraça, corresponding to 12 species, five varieties, and one forma (Tab. 3). The morphological descriptions of these taxa provided here are especially relevant for taxonomy, as they represent the morphology of populations in the type locality.

We point out some taxa that we have doubts about their presence in RPPN Santuário do Caraça. Eriocaulon macrocephalum Bong. [basionym of Paepalanthus macrocephalus (Bong.) Körn.] was described by Bongard (1831), who cited in the protologue the type material from Serra da Lapa and Serra do Caraça (“Habitat pratis humidis Serra da Lapa et Serra da Carassa”). However, the material from Serra do Caraça was never found; Andrino et al. (2021) revised and recircumscribed P. macrocephalus [= Actinocephalus macrocephalus (Bong.) F.N. Costa & Sano], and the Serra do Caraça was not considered as part of the species distribution. Eriocaulon exiguum Bong. [basionym of Paepalanthus exiguus (Bong.) Körn.] was also described by Bongard (1831); Koernicke (1863), in the Flora brasiliensis cited a specimen from Serra do Caraça (Riedel s.n.). However, this material was not found, and no other records were found to confirm it. We consider that it is probable that P. exiguus occurs in Serra do Caraça because this species is abundant in the Quadrilátero Ferrífero and there are several records from Catas Altas and Santa Bárbara, that are localities very close to RPPN Santuário do Caraça. Sano (2004) designated the material collected by Martius (s.n., M) as the lectotype of Paepalanthus ithyphyllus Mart. [= Actinocephalus ithyphyllus (Mart.) Sano], from an uncertain location, cited as “In ferruginosos ad Capão, Pires, et Serra do Caraça rel., prov. Minarum”; as different locations are mentioned, we do not know the origin of the type specimen, and no other material of this species was ever collected from RPPN Santuário do Caraça.

Quadrilátero ferrífero has no other survey published for the Eriocaulaceae flora to compare with our results; however, comparing it with other floras from Minas Gerais state, the number of species is expressive. It has a similar species richness to Serra de Grão Mogol (32 species) (Sano et al. 2010), it is richer than the core Serra da Mantiqueira (24 species) (Trovó et al. 2015), Serra do Ibitipoca (22 species) (Ferreira et al. 2011), and Serra da Canastra (26 species) (Freitas 2018). Numbers from Serra do Caraça are lower compared only with areas from the core Serra do Espinhaço, as the Biribiri State Park in the Diamantina Plateau (33 species only of Paepalanthus; Andrino et al. 2015), and the Serra do Cipó [83 species of Eriocaulaceae (Giulietti et al. 1987), including ten species of Paepalanthus sub. Xeractis (Hensold 1998), 20 species of Syngonanthus (Parra 1998), and nine species of Paepalanthus sect. Actinocephalus (Körn.) Ruhland (Sano 1998)].

Table 2
Species previously reported for Caraça (Augsten 2015) that were excluded from our updated list.

Species previously determined for Caraça	Reason for exclusion
Leiothrix propinqua	Updated determination (Oliveira 411) as Leiothrix longipes
Paepalanthus exiguus	Lost specimen (Riedel s.n. not found) and updated determination (Badini s.n.) as Paepalanthus manicatus
Paepalanthus globulifer	Updated determination (Oliveira 296) by L.J.Sautier as Paepalanthus conduplicatus
Paepalanthus ithyphyllus	Doubtful location of type, never recollected in Caraça, and updated determination (Badini s.n.) as Paepalanthus hilairei
Paepalanthus langsdorffii	Described variety (P. langsdorffii var. caracensis) under synonymy of P. suffructicans, and updated determination (Lisboa s.n.) as Paepalanthus dianthoides
Comanthera squarrosa	The name is synonymized under Comanthera centauroides
Paepalanthus chlorocephalus	The name is synonymized under Paepalanthus suffruticans
Paepalanthus bryoides	Typing error of the exsicata (Braga 215), wrong location.

Table 3
Taxa with types from Caraça and their respective accepted names. HeS = Heterotypic synonym.

Taxa	Author	Accepted name	Voucher
Syngonanthus caracensis (HeS)	Silveira	Comanthera centauroides	Silveira 430
Syngonanthus squarrosus (HeS)	Ruhland	Comanthera centauroides	Glaziou 15541
Paepalanthus camptophyllys	Ruhland	Paepalanthus camptophyllys	Glaziou 15520
Paepalanthus chloroblepharus (HeS)	Ruhland	Paepalanthus suffruticans	Ule s.n.
Paepalanthus ciliolatus	Ruhland	Paepalanthus ciliolatus	Glaziou 15527
Paepalanthus conduplicatus	Körn.	Paepalanthus conduplicatus	Sellow s.n.
Paepalanthus corymboides (HeS)	Ruhland	Paepalanthus melaleucus	Ule 2725
Paepalanthus globosus (HeS)	Ruhland	Paepalanthus leucoblepharus	Glaziou 15537
Paepalanthus scleranthus	Ruhland	Paepalanthus scleranthus	Ule 2717
Paepalanthus suffruticans	Ruhland	Paepalanthus suffruticans	Glaziou 15529
Paepalanthus vaginatus	Körn.	Paepalanthus vaginatus	Riedel s.n.
Paepalanthus vellozioides	Körn.	Paepalanthus vellozioides	Sellow 1368, Claussen 96
Leiothrix curvifolia var. setacea (HeS)	Ruhland	Leiothrix mucronata	Glaziou 15542, 15543
Leiothrix flavescens var. parvifolia (HeS)	Moldenke	L. flavescens var. flavescens	Irwin 28946
Leiothrix vivipara var. angusta (HeS)	Ruhland	Leiothrix vivipara	Mendonça 323, Claussen 1163, Glaziou 15515
Paepalanthus langsdorffii var. caracensis (HeS)	Moldenke	Paepalanthus suffruticans	Irwin 29110
Paepalanthus suffruticans var. angustifolia (HeS)	Silveira	Paepalanthus suffruticans	Silveira 331
Paepalanthus manicatus f. robusta (HeS)	Ruhland	Paepalanthus manicatus	Ule 2715

In this work, we propose five new synonyms for Eriocaulaceae: Paepalanthus chloroblepharus Ruhland, P. langsdorffii var. caracensis Moldenke, and P. suffruticans var. angustifolius Silveira ex Moldenke as synonyms of P. suffruticans Ruhland; P. globosus Ruhland under P. leucoblepharus Körn; and P. manicatus f. robusta under P. manicatus. Justifications are under each species taxonomic treatment.

Key to the species of Eriocaulaceae in Serra do Caraça

1. Plants with erect fertile central axis and lateral branches (paraclades) 2

1’. Plants without an erect fertile central axis or, if present, without lateral branches (paraclades) and with scapes in umbels 3

2. Paraclades verticillate along the central axis; scapes pilose when young 19. Paepalanthus hilairei

2’. Paraclades alternate spiraled along the central axis; scapes villous when young 25. Paepalanthus polyanthus

3. Plants without spathes and scapes 27. Paepalanthus scleranthus

3’. Plants with spathes and scapes 4

4. Scapes bearing numerous capitula 5

4’. Scapes with a single capitulum 8

5. Scapes splitted at apex 6

5’. Scapes not splitted at apex 7

6. Aerial stem elongated; involucral bracts brown, apex acute, not ciliate; leaves green at the adaxial base, sparsely ciliate with long hairs 22. Paepalanthus melaleucus

6’. Aerial stem short, restricted to the leaf rosette, involucral bracts golden to light castaneous, apex long acuminate, ciliate; leaves brown at the adaxial base, ciliate at the apex only, with tiny hairs 30. Paepalanthus vellozioides

7. Rosette leaves distichous, conduplicate; involucral bracts abaxially glabrous, ciliate; anther cream 13. Paepalanthus conduplicatus

7’. Rosette leaves spiraled, flat; involucral bracts abaxially densely pilose, not ciliate; anther castaneous 24. Paepalanthus planifolius

8. Involucral bracts series discolor and dimorphic, internal series spathulate 9

8’. Involucral bracts series with similar or gradual color and shape, internal series with various shapes, not spathulate 10

9. Scapes emerging from the stem at the leaf axil; involucral bracts straw-colored in the external series; petals of pistillate flowers with rounded apex 2. Comanthera nivea

9’. Scapes emerging from a reproductive axis; involucral bracts castaneous in the external series; petals of pistillate flowers with acuminate apex 31. Syngonanthus anthemiflorus

10. Plants with leaves in basal rosette or caespitose, stem subterranean, reptant or, if aerial, then restricted to the leaf rosette 11

10’. Plants with leaves spiraled along an elongated aerial stem or with an elongated fertile axis 25

11. Spathe opening truncate 12

11’. Spathe opening oblique 13

12. Plants amphibian; fenestrate leaves; diplostemonous flowers; petals of staminate and pistillate flowers with black glands; black anther 3. Eriocaulon ligulatum

12’. Plants terrestrial; striate leaves; isostemonous flowers; petals of staminate and pistillate flowers without glands; cream anther 5. Leiothrix flavescens

13. Involucral bracts cream, straw-colored or golden 14

13’. Involucral bracts castaneous, brown or blackish 20

14. Capitula presenting pseudoviviparity, forming clonal clumps 9. Leiothrix vivipara

14’. Capitula without pseudoviviparity 15

15. Leaves caespitose, apex obtuse to round; capitula cyathiform 1. Comanthera centauroides

15’. Leaves rosulate, apex acute, acuminate or mucronate; capitula hemispheric 16

16. Internal series of involucral bracts surpassing the flower disc, adaxially pilose in the distal half 6. Leiothrix gomesii

16’. Internal series of involucral bracts not surpassing the flower disc, adaxially glabrous 17

17. Plants with long and wide leaves, 3-5 cm × 0.5-0.7 mm 7. Leiothrix longipes

17’. Plants with short and thin leaves, 0.5-2 cm × 0.2-0.5 mm 18

18. Involucral bracts with apex obtuse to round, abaxially glabrous, not ciliate, straw-colored to golden, elliptic to oblong, 5-6 series 33. Syngonanthus gracilis

18’. Involucral bracts with apex acute, abaxially pilose to glabrescent, ciliate, cream to straw-colored, lanceolate, 3-4 series 19

19. Leaves with acute apex; spathes shorter than the leaves, green; plants 13-17 cm tall; scapes 12-17 cm long 4. Leiothrix curvifolia

19’. Leaves with mucronate apex; spathes longer than the leaves, castaneous; plants 8-13.5 cm tall; scapes 6.5-13.5 cm long 8. Leiothrix mucronata

20. Flowers dimerous 21

20’. Flowers trimerous 22

21. Leaves caespitose; involucral bracts recurvate, dark brown at center with light brown margin, densely ciliate, slightly surpassing the flower disc, abaxially glabrous 15. Paepalanthus elongatus

21’. Leaves rosulate; involucral bracts appressed, homogeneously brown and sparsely ciliate, not surpassing the flower disc, abaxially densely pilose 29. Paepalanthus vaginatus

22. Leaves and scapes pilose to glabrescent; involucral bracts castaneous, abaxially densely pilose at the apex; anther cream 26. Paepalanthus pubescens

22’. Leaves and scapes glabrous; involucral bracts brown or blackish, abaxially glabrous; anther pigmented 23

23. Plants with leaves entirely and densely ciliate; spathes with striate indumentum, densely pilose abaxially, adaxially tufted at apex 20. Paepalanthus leucoblepharus

23’. Plants with leaves not ciliate or ciliate at the base only; spathes abaxially glabrous or sparsely pilose; adaxially glabrous or pilose 24

24. Plants with delicate rosette, leaves small and thin, 3.5-10 × 3-4 mm, striate, ciliate at the base; spathes adaxially pilose, small capitula 4-6 mm wide 12. Paepalanthus ciliolatus

24’. Plants with robust tank rosette, leaves long and wide, 16-23 cm × 12-32 mm, not striate, not ciliate; spathes adaxially glabrous, large capitula 14-16 mm wide 10. Paepalanthus calvus

25. Involucral bracts cream to gold 26

25’. Involucral bracts light castaneous, brown or blackish 27

26. Flowers trimerous; involucral bracts surpassing the flower disc 0.5-1 mm long, apex acute; petals of staminate flowers fleshy; petals of pistillate flowers fused in the middle 32. Syngonanthus caulescens

26’. Flowers dimerous; involucral bracts not surpassing the flower disc, apex obtuse to round; petals of staminate flowers membranous; petals of pistillate flowers free 17. Paepalanthus flaccidus

27. Involucral bracts adaxially pilose; corolla of the staminate flower adaxially pilose 28

27’. Involucral bracts adaxially glabrous; corolla of the staminate flower adaxially glabrous 29

28. Involucral bracts with apex acuminate to long acuminate in the external series, internal series pilose adaxially at the apex only, sometimes surpassing the flower disc 0.5-1 mm long; floral bracts apiculate; petals of pistillate flowers cream, anther cream 14. Paepalanthus dianthoides

28’. Involucral bracts with apex obtuse to acute in the external series, internal series adaxially densely pilose from middle to the top, always surpassing the flower disc 1.5-3 mm long; floral bracts lanciform; petals of pistillate flowers blackish at apex, anther pigmented 23. Paepalanthus mollis Kunth var. mollis

29. Plants with an elongated fertile axis; flowers dimerous; scapes in spherical umbels 16. Paepalanthus erectifolius

29’. Plants without elongated fertile axis; flowers trimerous; scape not in umbels, or with umbels non spherical 30

30. Aerial stem branched 31

30’. Aerial stem non-branched 32

31. Leaves pilose to glabrescent on both surfaces, ciliate with long hairs; spathes abaxially sparsely pilose to glabrescent; scapes densely pilose; involucral bracts abaxially pilose; anther cream 11. Paepalanthus camptophyllus

31’. Leaves glabrous on both surfaces, not ciliate; spathes abaxially glabrous; scapes glabrous; involucral bracts abaxially glabrous; anther black 18. Paepalanthus freyreissii

32. Plants small and delicate, 6-8.5 cm tall; involucral bracts abaxially glabrous, not ciliate; tiny leaves 0.2-1.5 cm × 0.5-1 mm 21. Paepalanthus manicatus

32’. Plants tall and robust, 30-100 cm tall; involucral bracts abaxially pilose with adpressed hairs, ciliate; long leaves 6.5-23 cm × 1.8-32 mm 28. Paepalanthus suffruticans

1. Comanthera centauroides (Bong.) L.R. Parra & Giul., Taxon 59: 1143. 2010.Fig. 3a-b

Plants 9-21 cm tall. Rhizome present. Subterranean stem 1.5-4 cm long, oblique, densely branched. Rosette absent, leaves caespitose 2-3.5 × 0.1-0.3 cm, linear, obtuse to rounded apex, pilose to glabrescent on both surfaces, ciliate at the base. Inflorescences with scapes emerging from the rhizome at the leaf axil. Spathes 1.5-3 cm long, abaxially pilose to glabrescent, oblique opening. Scapes 6.5-19 cm long, free, 1-3 per stem branch, densely pilose to glabrescent. Capitula cyathiform, 3-7 mm wide. Involucral bracts straw-colored, 2-3 mm long, not surpassing the flower disc, 7-8 series, elliptic, obovate to narrow-elliptic, rounded apex, abaxially glabrous, not ciliate, cream and spathulate internal series. Floral bracts absent. Staminate flowers 3-5 mm long; cream sepals, 2.5-3 mm long, fused in the lower third, narrow-elliptic, concave, apex obtuse, glabrous; petals cream, 2.5-3 mm long, fused in the lower third, narrow-elliptic, concave, obtuse apex, glabrous; stamens 3, anther dorsifixed; 3 nectariferous carpellodes. Pistillate flowers 3-4 mm long; cream sepals cream, 2.5-3 mm long, free, ovate to narrow-ovate, concave, obtuse apex, glabrous; cream petals, 3-3.5 mm long, fused at the middle, free at the base and top, narrow-spathulate, flat, rounded apex, hairy on both surfaces at the middle; gynoecium 2-3 mm long, 3 nectariferous branches, stigmatic branches simple.

Examined material: Campos de Fora, 20°07’03”S, 43°31’08”W, 1,504 m, 17.VII.2021, L. Echternacht & P. Mendes 2891 (OUPR); 19.VIII.2000, J. Ordones et al. 257 (BHZB, OUPR); 20°04’13”S, 43°18’39”W, 1,497 m, 24.XI.2020, A. Soldevila & P. Vanucci 328 (OUPR); Mirante 1 da Piscina, 20°03’37”S, 43°18’00”W, 1,339 m, 23.XI.2020, A. Soldevila & P. Vanucci 327 (OUPR); Pico do Inficcionado, 7.VIII.2002, J. Ordones 929 (BHZB, OUPR).

Comanthera centauroides is distributed through the Espinhaço Range from Bahia to Minas Gerais and in the restingas in Rio de Janeiro (Echternacht et al. 2015; Echternacht & Parra 2024). In RPPN Santuário do Caraça, it forms abundant populations in the following localities: Pico da Canjerana, trail to Padre Caio cave, Pico do Sol, Pico do Inficcionado, Campos de Fora, and Mirante 1 da Piscina. It grows in quartzitic Campo Rupestre, in open areas over sandy to organic, constantly or seasonally flooded soils, among herbaceous vegetation. Individuals are perennial, flowering between April and October.

Comanthera centauroides is mainly recognized by its caespitose habit with a densely branched rhizome forming clumps, leaves linear and tortuous, floral bracts absent, and the capitula cyathiform (i.e., urceolate capitula with involucral bracts lower that the flower disc, forming a fringe at top). Syngonanthus caracensis Silveira and S. squarrosus Ruhland have type materials from RPPN Santuário do Caraça (Silveira 430, R; Glaziou 15541, B, NY, US; respectively), and both are currently under synonymy of Comanthera centauroides.

Figure 3 -
a-o. Species of Eriocaulaceae - a-b. Comanthera centauroides - a. habit; b. cyathiform capitula; c-d. C. nivea - c. habit; d. capitulum with cream involucral bracts, surpassing the flower disc; e. Eriocaulon ligulatum - habit; f-g. Leiothrix curvifolia - f. habit, forming clumps; g. capitulum; h-i. L. flavescens - h. habit; i. capitulum; j-m. L. gomesii - j. habitat and habit; k. rosette with wooly indumentum at the leaf base; l-m. capitulum - l. involucral bracts adaxially pilose and flower disc; m. straw-colored to golden involucral bracts; n. L. mucronata - habit; o. L. vivipara - sprouting scapes. Images: a-g,j-n. Livia Echternacht; h,i,o. Ambar Soldevila.

2. Comanthera nivea (Bong.) L.R. Parra & Giul., Taxon 59: 1141. 2010.Fig. 3c-d

Plants 12-25 cm tall. Rhizome present. Subterranean stem 0.5-4 cm long, erect, branched or not. Rosette leaves 1.5-4 cm × 0.3-0.6 mm, linear, acute apex, pilose to glabrescent with adpressed and patent hairs on both surfaces, ciliate. Inflorescences with scapes emerging from the stem at the leaf axil. Spathes 1.2-3.5 cm long, abaxially pilose with adpressed or patent hairs, oblique opening. Scapes 6-25 cm long, free, 4-30 per rosette, pilose with adpressed hairs. Capitula 3-4 mm wide. Involucral bracts straw-colored to cream, 4-6 series, rounded apex, abaxially glabrous, not ciliate, straw-colored bracts of the external series, 1.5-2 mm long, obovate, internal series straw-colored at base and cream from middle to top, 3-4 mm long, spathulate, internal series surpassing the flower disc 2-3 mm long. Floral bracts absent. Staminate flowers 2-2.2 mm long; cream sepals, 0.8-1 mm long, fused at base, elliptic, concave, obtuse to rounded apex, glabrous; cream petals, 0.8-1 mm long, fused at base, elliptic, concave, rounded apex, glabrous; stamens 3, anthers dorsifixed; 3 nectariferous carpellodes . Pistillate flowers 1.8-2.2 mm long, pedicel 0.2-0.3 mm long, increasing in size during fruit ripening, reaching about 1 mm long; cream sepals, 1-1.2 mm long, free, elliptic, concave, rounded apex, glabrous; cream petals, 2-2.2 mm long, fused at middle, free at base and top, spathulate, rounded apex, abaxially hairy at middle, adaxially glabrous; gynoecium 1.5-1.8 mm long, 3 nectariferous branches, stigmatic branches simple.

Local records: caminho para Padre Caio, 27.V.2002, J. Ordones 854 (BHZB); Pico do Inficcionado, 20°08’18”S, 43°27’19”W, 2,001 m, 8.III.2008, L. Echternacht et al. 1615 (BHCB); sandstone summit of Serra da Caraça, 25.I.1971, H.S. Irwin et al. 29129 (NYBG, US); Serra do Caraça, 21.VII.1972, L.E. Mello Filho 3554 (NYBG); 11.IX.1990, J.R. Stehmann & D.C. Zappi (BHCB 28423); Serrote de Cangerana, 8.III.1982, N. Hensold 540 (NYBG); trilha para Capelinha e Gruta de Lourdes, 9.I.1982, N. Hensold (RB 317741); trilha para o Pico do Sol, 3.III.2010, L.N. Perillo 465 (BHCB).

Examined material: Ouro Preto, Cachoeira das Andorinhas, 20°17’15”S, 43°30’29”W, 7.VI.2018, L.G. Pedrosa 405 (OUPR); Serra das Camarinhas, 20°22’05”S, 43°30’21”W, 1,420 m, 22.VII.2016, L. Echternacht & T.V. Bastos 2697 (OUPR); 20°17’15”S, 43°30’29”W, 1.XII.2002, M.C.T.B. Messias 807 (OUPR); serra de Ouro Preto, 20°21’50”S, 43°31’14”W, 8.II.2012, R.C. Cardoso & T.C. Silva 67 (OUPR); 3.V.2012, R.C. Cardoso & T.C. Silva 101 (OUPR).

Although there are some recent records of Comanthera nivea from RPPN Santuário do Caraça hosted in BHCB reported in SpeciesLink (<https://specieslink.net/search/>), we did not have access to these specimens - they were not retrieved during this research. The description, therefore, is based on the work of Parra (2000) and materials from the municipality of Ouro Preto, the closest sites to Serra do Caraça available.

Comanthera nivea is distributed in all four states of southeastern Brazil, in open mountainous areas, especially in Serra do Espinhaço, Quadrilátero Ferrífero, Serra da Canastra, Serra da Mantiqueira, and in the Restinga (Parra 2000; Echternacht & Parra 2024). The species was found in the following localities in RPPN Santuário do Caraça: Capelinha and Gruta de Lourdes trail, Pico do Sol, Padre Caio cave trail, Pico do Inficcionado, and Pico da Canjerana. It grows in quartzitic Campo Rupestre, as rupicolous or terricolous or over sandy soils (Parra 2000). Specimens with flowers were recorded in January, March, May, July, and September.

Comanthera nivea is mainly recognized by its involucral bracts cream, spathulate, and delicate, surpassing the flower disc, floral bracts absent, and rosettes with linear leaves.

3. Eriocaulon ligulatum (Vell.) L.B. Sm., Contrib. Gray Herb.Harvard Univ. 124: 5. 1939.Fig. 3e

Plants 50-120 cm tall. Rhizome absent. Stem restricted to the leaf rosette, non-branched. Rosette leaves 7-35 × 0.5-4 cm, lanceolate, obtuse to rounded apex, glabrous on both surfaces, not ciliate. Inflorescences with scapes emerging from the stem at the leaf axil. Spathes 12-30 cm long, abaxially glabrous, truncate opening. Scapes 27-80 cm long, free, 4-18 per rosette, glabrous, rarely pilose. Capitula 7-15 mm wide. Involucral bracts straw-colored, 4-8 mm long, not surpassing the flower disc, ovate to elliptic, obtuse to rounded apex, abaxially glabrous, not ciliate. Floral bracts cream, 3.8-4.2 mm long, elliptic to oblong, concave, acute apex, tufted. Flowers diplostemonous. Staminate flowers 4-4.2 mm long; gray to blackish sepals, 3.8-4 mm long, free, elliptic to obovate, concave, obtuse apex, tufted; cream petals, 3-3.2 mm long, fused at base, with black glands, elliptic, concave, obtuse apex, tufted; stamens 6; black anthers, dorsifixed, 3 black nectariferous carpellodes. Pistillate flowers 3.2-4.2 mm long; gray to blackish sepals, 3-4 mm long, free, obovate, concave, obtuse apex, tufted; cream petals, 2-2.2 mm long, free with black glands, spathulate, obtuse apex, abaxially hairy in the distal half; gynoecium 2-2.2 mm long, nectariferous branches absent; stigmatic branches simple.

Examined material: trilha para Banho do Belchior, 20°03’45”S, 43°17’33”W, 1,254 m, 25.XI.2020, A. Soldevila & P. Vanucci 338 (OUPR); trilha para Cachoeira Bocaina, 20°03’48”S, 43°16’59”W, 1,255 m, 4.XI.2020, A. Soldevila & P. Vanucci 313 (OUPR); trilha para Cascatinha, 20°06’10”S, 43°28’57”W, 1,246 m, 16.VII.2021, L. Echternacht & P. Mendes 2888 (OUPR); Serra do Caraça, VI.1907, L.B. Damazio (OUPR 13104).

Eriocaulon ligulatum is distributed in in Espírito Santo, Minas Gerais, São Paulo, and Rio de Janeiro (Chagas et al. 2024). It was found in the following localities in RPPN Santuário do Caraça: trail to Cascatinha, Tanque Grande, the trail to Bocaina, and the trail to Banho do Belchior. It grows in temporarily flooded locations and river banks, and we observed populations with more than 30 individuals, which are perennial. Specimens with flowers were recorded in January, June, July, August, September, and November. In July, individuals with young flowers were recorded, and in November, those with old flowers, suggesting a flowering period synchronous with the dry season, which is common in aquatic Eriocaulaceae.

Eriocaulon ligulatum is recognized by its amphibian habit, with robust individuals, fenestrate leaves, truncate spathe opening, straw-colored involucral bracts not surpassing the flower disc, diplostemonous flowers, gray to blackish sepals of staminate and pistillate flowers, petals with a black gland, and black anthers.

4. Leiothrix curvifolia (Bong.) Ruhland, Pflanzenr. 4(30): 233. 1903.Fig. 3f-g

Plants 13-17 cm tall, forming clumps. Rhizome present. Subterranean stem branched. Rosette leaves 10-20 × 0.2-0.3 mm, linear, acute apex, pilose to glabrescent on both surfaces, ciliate or not. Inflorescences with scapes emerging from the stem at the leaf axil. Spathes 1-2.2 cm long, lax, abaxially pilose to glabrescent, oblique opening. Scapes 12-17 cm long, free, 1-2 per rosette, pilose to glabrescent. Capitula 5-6 mm wide. Involucral bracts cream to straw-colored, 1.5-2 mm long, not surpassing the flower disc, lanceolate, 3-4 series, acute apex, abaxially pilose to glabrescent, ciliate. Floral bracts cream, 3-3.2 mm long, oblanceolate, falcate, concave, acute apex, abaxially densely pilose, tufted. Staminate flowers 3-3.2 mm long; cream sepals, 2-2.2 mm long, fused at base, oblanceolate, falcate, concave, acute apex, abaxially pilose, tufted; cream corolla, 2 mm long, tubular, acute lobes, glabrous; stamens 3, anthers basifixed; 3 nectariferous carpellodes. Pistillate flowers 2.7-3 mm long; cream sepals, 2.2-2.5 mm long, free, oblong, falcate, concave, acute apex, abaxially hairy from middle to top, tufted; cream petals, 2-2.2 mm long, free, oblong, falcate, flat, acute apex, tufted; gynoecium 2.2 mm long, 3 nectariferous branches, releasing from the column below the simple stigmatic branches.

Examined material: trilha para Pico da Conceição, 20°06’29”S, 43°31’21”W, 1,503 m, 17.VII.2021, L. Echternacht & P. Mendes 2896 (OUPR).

Leiothrix curvifolia is endemic to the Espinhaço Range in Minas Gerais, distributed from Barão de Cocais to Diamantina (Giulietti 2020, 2024). In RPPN Santuário do Caraça, it was found in the Pico da Conceição only. It grows in quartzitic Campo Rupestre, over dry to humid sandy soils, among open vegetation. Plants are perennial and individuals with flowers were recorded in March and in July. As noted by Giulietti (1984), the peak of flowering occurs from January to March.

Leiothrix curvifolia is remarkable by the presence of rhizome forming clumps with linear rosette leaves. This species is very similar to Leiothrix mucronata, as noted by Giulietti (1984, 2020b); flowers are very similar, but L. curvifolia can be distinguished by leaves with acute apex (vs. mucronate), green spathes shorter than the leaves (vs. castaneous and longer), with taller plants and longer scapes.

5. Leiothrix flavescens (Bong.) Ruhland, Pflanzenr. 4(30): 231. 1903.Fig. 3h-i

Plants 30-82 cm tall. Rhizome present. Stem short, 0.5-2 cm long, restricted to the leaf rosette, sometimes branched. Rosette leaves 3.5-11 × 0.5-1 cm, lanceolate, obtuse apex, pilose to glabrescent on both surfaces, ciliate. Inflorescences with scapes emerging from the stem at the leaf axil. Spathes 4.5-13.5 cm long, abaxially densely pilose to glabrescent, truncate opening. Scapes 17-80 cm long, free, 3-30 per rosette, pilose to glabrescent. Capitula 5-10 mm wide. Involucral bracts castaneous to golden, 2-4 mm long, not surpassing the flower disc, 4-5 series, ovate to oblong, obtuse apex, abaxially pilose, ciliate. Floral bracts cream, 3.5-4 mm long, oblanceolate, concave, acute apex, abaxially pilose, tufted. Staminate flowers 3.2-4 mm long; cream sepals, 2 mm long, free, obovate, concave, truncate to rounded apex, abaxially hairy in the distal half, tufted; cream corolla, 2 mm long, tubular, obtuse lobes, glabrous; stamens 3, anthers basifixed; 3 nectariferous carpellodes. Pistillate flowers 4.5-5 mm long; cream sepals, 2.5-2.8 mm long, free, elliptic, concave, acute apex, abaxially hairy in the distal half, tufted; cream petals, 2.5-3 mm long, free, narrow-lanceolate, flat, acute apex, glabrous; gynoecium 3-3.2 mm long, 3 nectariferous branches, releasing from the column below the simple stigmatic branches.

Examined material: 20.XI.1985, M.M.N. Braga et al. (BHCB 10376); 1.XI.1997, P.H.C. Corgosinho & M.F. Vasconcelos (BHCB 40047); 10.IX.1990, J.R. Stehmann (BHCB 28435); subida para o Pico do Inficcionado, no primeiro platô após a subida mais íngreme, 20°08’05”S, 43°27’49”W, 1,829 m, 8.III.2008, L. Echternacht et al. 1622 (BHCB); trilha para Cachoeira da Bocaina, próximo a cachoeira, 20°04’21”S, 43°16’33”W, 1,277 m, 4.XI.2020, A. Soldevila & P. Vanucci 317 (OUPR); trilha para Cachoeira Cascatinha, 20°06’24”S, 43°28’36”W, 1,250 m, 16.VII.2021, L. Echternacht & P. Mendes 2887 (OUPR); 20°03’44”S, 43°17’00”W, 1,251 m, 4.XI.2020, A. Soldevila & P. Vanucci 312 (OUPR); trilha para piscina, 9.I.1982, N. Hensold et al. (BHCB 36782).

Leiothrix flavescens is the most widespread species in the genus, widely distributed in Brazil, also occurring in Peru, Venezuela, and Guianas (Giulietti 2020, 2024). It is widely distributed in RPPN Santuário do Caraça, being one the most abundant species of the family. It was found in the following localities: trail to Cascatinha, trail to Bocaina, trail to Piscina, Tanque Grande, ascent to Pico do Inficcionado, ascent to Pico da Canjerana and Cascudo. It grows in temporarily flooded locations, river banks, over moisty sandy to organic soils, among open herbaceous vegetation. Individuals are perennial; specimens with flowers were recorded in January, March, July, August, September, November, and December. The peak of flowering occurs from November to March, in the rainy season.

Leiothrix flavescens inhabits flooded environments such as river banks or swamps. It is recognized by its leaves with obtuse apex, truncate spathe opening, and castaneous to gold involucral bracts.

Leiothrix flavescens var. parvifolia Moldenke was described based on type material from Serra do Caraça, “on the lower slopes of the Serra do Caraça about 10 km west of Barão de Cocais”, collected by Irwin (28946, NY, UB) (Moldenke 1972). Giulietti and Hensold (1991) synonymized L. flavescens var. parvifolia as L. flavescens, and in the nomenclatural treatment of Leiothrix (Giulietti 2020b), L. flavescens var. parvifolia is considered synonymous with L. flavescens var. flavescens.

6. Leiothrix gomesii Silveira, Floral. Mont. 1: 289. 1928.Fig. 3j-m

Plants 30-70 cm tall, forming clumps. Rhizome present. Subterranean stem branched, with a wooly indumentum. Rosette leaves 0.8-5.5 × 0.15-0.4 cm, linear, acute apex, pilose to glabrescent on both surfaces, ciliate. Inflorescences with scapes emerging from the stem, at the leaf axil. Spathes 5-12 cm long, abaxially sparsely pilose to glabrescent, oblique opening. Scapes 28-68 cm long, free, 1-3 per rosette, densely pilose to glabrescent. Capitula 10-15 mm wide. Involucral bracts straw-colored to golden, 4-8 mm long, surpassing the flower disc 1-2 mm long, 4-5 series, lanceolate, acuminate apex, abaxially pilose to glabrescent, adaxially pilose in the distal half, ciliate. Floral bracts cream, 5-6 mm long, linear to oblanceolate, concave, acute apex, tufted. Staminate flowers 5-5.5 mm long; cream sepals, 3-4 mm long, fused at base, oblanceolate, concave, acute apex, tufted; petals white, 3-3.5 mm long, free, linear, apex acuminate, glabrous; stamens 3, anther basifixed; 3 nectariferous carpellodes. Pistillate flowers 5-5.5 mm long; sepals white, 4-4.2 mm long, free, elliptic, concave, acute apex, tufted; cream petals, 4-4.2 mm long, free, linear, flat, acuminate apex, glabrous; gynoecium 3.8-4 mm long, with long hairs at base, 3 nectariferous branches, releasing from the column below the simple stigmatic branches.

Examined material: Pico do Sol, 1.III.2009, L. Echternacht et al. 1938 (BHCB, HUFU); 20°06’22”S, 43°27’12”W, 1,937 m, 29.VI.2009, C.T. Oliveira et al. 487 (BHCB).

Leiothrix gomesii is endemic to the southeastern Quadrilátero Ferrífero. It was known from type collection only, from Serra de Capanema, and has been considered presumably extinct (Giulietti 1984; COPAM 085/97; Biodiversitas 2007; Costa et al. 2008). One population was discovered in the RPPN Santuário do Caraça in Pico do Sol in 2009 (voucher: Echternacht, L. 1938. BHCB, UFU; Oliveira, C. 487. BHCB); then, another population was discovered in 2018 in Ouro Preto, with no precise locality (voucher: Pedrosa, L. 363. OUPR) (Giulietti 2020). This species in the RPPN Santuário do Caraça is restricted to the Pico do Sol, in quartzitic Campo Rupestre, at the peak plateau above 2,000 m elevation, over sandy soils. Considering that the region is highly disturbed by mining, agricultural and urban activities, these populations might be a relict from former more abundant and spread populations. Specimens were recorded in March with young and mature flowers and in June with old flowers, suggesting a flowering peak probably in the first semester of the year.

Leiothrix gomesii is recognized by its branched rhizome, stem with wooly indumentum, straw-colored to golden involucral bracts, surpassing the flower disc by 1-2 mm long, and adaxially pilose.

7. Leiothrix longipes Silveira, Floral. Mont. 1: 303. 1928.

Plants 5-20 cm tall. Rhizome present. Stem short, restricted to the leaf rosette, non-branched. Rosette leaves 3-5 cm × 0.5-0.7 mm, linear, acute apex, pilose to glabrescent on both surfaces, ciliate or not. Inflorescences with scapes emerging from the stem at the leaf axil. Spathes 2.5-6 mm long, abaxially pilose to glabrescent, oblique opening. Scapes 3-25 cm long, free, 30-40 per rosette, densely pilose to glabrescent. Capitula 4-6 mm wide. Involucral bracts straw-colored to golden, 2 mm long, not surpassing the flower disc, ovate, 3-4 series, acute apex, pilose abaxially, ciliate. Floral bracts linear-lanceolate, acute apex, abaxially pilose. Staminate flowers 2.4 mm long; sepals fused at base, concave, obtuse to acute apex, abaxially pilose; tubular corolla, acute lobes, glabrous; stamens 3, anthers basifixed; 3 nectariferous carpellodes. Pistillate flowers 2.8 mm long; free sepals, concave, pilose abaxially; free petals, linear, flat, acute apex, ciliate; gynoecium with 3 nectariferous branches, releasing from the column below the simple stigmatic branches.

Local records: Pico da Conceição, 6.III.1982, N. Hensold 538 (RB).

Examined material: Pico do Sol, 20°06’43”S, 43°26’55”W, 1,918 m, 25.IV.2009, C.T. Oliveira et al. 411 (BHCB).

Leiothrix longipes is endemic to Minas Gerais, distributed in the Espinhaço Range from Diamantina to Caeté (Giulietti 2020). In RPPN Santuário do Caraça, it was found in Pico da Conceição and Pico do Sol. It grows in quartzitic Campo Rupestre, over sandy organic soil, between 1,400 to 1,900 m elevation, among grasses, sedges, small trees, and shrubs. Specimens were recorded in March with mature and young flowers and in April with immature inflorescences.

Leiothrix longipes is mainly recognized by linear leaves ca. 3-5 cm long, usually longer than L. vivipara and L. curvifolia (both ca. 0.7-2 cm long). As noted by Giulietti (1984, 2020b), L. longipes has large variations in the leaf length and width. It can be also recognized by its pilose scapes and involucral bracts with acute apex, pilose abaxially.

Two records were found in RPPN Santuário do Caraça, but only one was available at the herbarium (BHCB) for analysis, and it has immature flowers (C.T. Oliveira et al. 411). Therefore, the description of fertile parts was based on Giulietti (1984), who determined the specimen N. Hensold 538 (RB); a photo of this specimen was obtained from Virtual Reflora Herbarium (<http://reflora.jbrj.gov.br/reflora/herbarioVirtual/>).

8. Leiothrix mucronata (Bong.) Ruhland, Pflanzenr. 4(30): 232. 1903.Fig. 3n

Plants 8-13.5 cm tall, forming clumps. Rhizome present. Subterranean stem branched. Rosette leaves 9-15 × 0.3-0.5 mm, linear, mucronate apex, pilose to glabrescent on both surfaces, ciliate or not. Inflorescences with scapes emerging from the rhizome at the leaf axil. Spathes 1-2.5 cm long, lax, abaxially pilose to glabrescent, oblique opening. Scapes 6.5-13.5 cm long, free, 2-4 per rosette, pilose to glabrescent. Capitula 4-6 mm wide. Involucral bracts straw-colored, 1.5-2.5 mm long, not surpassing the flower disc, lanceolate, 3-4 series, acute apex, abaxially pilose to glabrescent, ciliate. Floral bracts cream, 3-3.2 mm long, oblanceolate, concave, acute apex, abaxially densely pilose, tufted. Staminate flowers 3-3.2 mm long; cream sepals, 2-2.2 mm long, fused at base, obovate, concave, acute apex, tufted; cream corolla, 2-2.2 mm long, tubular, acute lobes, glabrous; stamens 3, anthers basifixed; 3 nectariferous carpellodes. Pistillate flowers 2.2-2.5 mm long; cream sepals, 2-2.2 mm long, free, obovate, concave, acute apex, abaxially hairy in the distal half, tufted; cream petals, 1.9-2 mm long, free, obovate, concave, acute apex, hairy abaxially in the distal half, tufted; gynoecium 2 mm long, 3 nectariferous branches, releasing from the column below the simple stigmatic branches.

Examined material: Serra do Caraça, 1906, A.T. Baeta-Neves (OUPR 13134); 15.XII.2000, R.C. Mota 1097 (BHCB); subida para o Pico do Inficcionado, 20°08’04”S, 43°27’48”W, 1,821 m, 8.XII.2008, C.T. Oliveira et al. 232 (BHCB).

Leiothrix mucronata is endemic to Minas Gerais, occurring in the Serra do Cipó, RPPN Santuário do Caraça, and Ouro Preto (Giulietti 2020). It was found in the following localities in the RPPN Santuário do Caraça: Campos de Fora and ascent to Pico do Inficcionado. It grows in quartzitic Campo Rupestre, over dry to moist sandy soils, among open vegetation. Specimens with flowers were recorded in December. As noted by Giulietti (1984), the peak of flowering occurs from January to March.

Leiothrix mucronata is remarkable by the presence of rhizome forming clumps, differing from Leiothrix curvifolia by its smaller stature and rosettes with linear leaves of mucronate apex, spathes castaneous and lax, larger than leaves, and by smaller scapes (Giulietti 1984, 2020b).

Leiothrix curvifolia var. setacea Ruhland was described based on the material from RPPN Santuário do Caraça (Glaziou 15542, B, NY, P), from Morro São João in Ouro Preto (Glaziou 15543, 24.VI.1884, P, NY) and from Morro São Sebastião in Ouro Preto (Magalhães Gomes 3886, BHCB) (Ruhland 1903). Giulietti (2020b) synonymized L. curvifolia var. setacea as L. mucronata and selected the material collected by Glaziou (15543, 14.II.1884, P) from the Itacolomi in Ouro Preto as the lectotype. We observed that the voucher Glaziou 15543 number is composed of more than one material from Ouro Preto: the one dated 24.VI.1884 is from Morro São Sebastião, which was cited in the protologue, and the other dated 14.II.1884 is from the Itacolomi.

9. Leiothrix vivipara (Bong.) Ruhland, Pflanzenr. 4(30): 238. 1903.Fig. 3o

Plants 5-30 cm tall. Rhizome present. Stem short, restricted to the leaf rosette, non-branched. Rosette leaves 0.7-2 cm × 0.2-0.5 mm, linear, mucronate apex, pilose to glabrescent on both surfaces, not ciliate. Inflorescences with scapes emerging from the stem at the leaf axil.Spathes 1.2-2.7 cm long, abaxially pilose to glabrescent, oblique opening. Scapes 1.5-35 cm long, free, 3-20 per rosette, sparsely pilose to glabrescent, presenting pseudoviviparity. Capitula 1.5-4 mm wide. Involucral bracts cream, 1-2 mm long, not surpassing the flower disc, 4-5 series, lanceolate, acuminate apex, abaxially pilose, ciliate. Floral bracts cream, 2-2.5 mm long, lanceolate, concave, acuminate apex, ciliate at the proximal half. Staminate flowers 2.5-3 mm long; cream sepals, 1.5-2 mm long, fused at base, lanceolate, concave, acuminate apex, abaxially pilose in the middle, ciliate at the proximal half; cream corolla, 1.5-1.7 mm long, infundibular, acute lobes, glabrous; stamens 3, anthers basifixed; 3 nectariferous carpellodes. Pistillate flowers 2.2 mm long; cream sepals, 2 mm long, free, lanceolate, concave, acuminate apex, abaxially pilose in the middle, ciliate in the middle; cream petals, 2 mm long, free, narrow-lanceolate, flat, acute apex, ciliate at proximal half; gynoecium 1.7-2 mm long, 3 nectariferous branches, releasing from the column below the simple stigmatic branches. .

Examined material: atrás da Capelinha, 2.X.1998, A.R. Marques et al. (BHCB 43451); 20°03’14”S, 43°17’07”W, 1,395 m, 3.XI.2020, A. Soldevila & P. Vanucci 304 (OUPR); 20°05’46”S, 43°28’45”W, 30.VIII.1997, J.R. Stehmann et al. 2268 (BHCB, CESJ); caminho para a Gruta do Padre Caio, 11.XII.1986, I.R. Andrade et al. (BHCB 9049); 11.IX.1990, J.R. Stehmann et al. (BHCB 18925); Campos de Fora, 20°06’33”S, 43°31’18”W, 1,503 m, 17.VII.2021, L. Echternacht & P. Mendes 2897 (OUPR); 20°04’13”S, 43°18’39”W, 1,497 m, 24.XI.2020, A. Soldevila & P. Vanucci 330 (OUPR); estrada para pico Canjerana, próximo ao platô, 20°04’31”S, 43°18’17”W, 1,767 m, 24.XI.2020, A. Soldevila & P. Vanucci 333 (OUPR); Parque do Caraça, 10.I.1996, V.C. Souza et al. 10007 (OUPR, BHCB); trilha entre Capelinha e Gruta de Lourdes, 20°03’13”S, 43°17’02”W, 1,471 m, 3.XI.2020, A. Soldevila & P. Vanucci 308 (OUPR); trilha para Pico da Conceição, 20°07’03”S, 43°31’08”W, 1,504 m, 17.VII.2021, L. Echternacht & P. Mendes 2892 (OUPR); 20°07’03”S, 43°31’08”W, 1,504 m, 17.VII.2021, L. Echternacht & P. Mendes 2893 (OUPR).

Leiothrix vivipara is endemic to the Espinhaço Range plus the Quadrilátero Ferrífero in Minas Gerais, occurring from Datas to Serra do Caraça (Giulietti 2020). It is widely distributed in RPPN Santuário do Caraça and was recorded from the following localities: trail to Capelinha, trail to Gruta de Lourdes, trail to Padre Caio cave, Pico do Inficcionado, Campos de Fora, ascent to Pico da Canjerana and ascent to Pico da Conceição. This species forms abundant populations due to its pseudoviviparity and clonal growth; rosettes stack on top of each other and may form piles of up to 50 cm high, sometimes covering the ground, in open areas, gullies, or forest edges, over dry to moist sandy soils (Giulietti 1984). Specimens were recorded flowering all over the year.

Leiothrix vivipara is mainly recognized by its pseudoviviparous habit, forming clonal clumps and tussocks; the capitula meristem sprouts into a new rosette, rooting when it touches the ground or prostrating over the vegetation; in addition to this habit, it can be recognized by small linear delicate leaves and scapes, lanceolate involucral bracts, with acuminate apex, ciliate and abaxially pilose.

Leiothrix vivipara var. angusta Ruhland was described based on the material from Serra do Caraça, collected by Mendonça (323, B) and Glaziou (15515, B, G, NY), and also based on the material collected by Claussen (1163, B) from an unknown locality (Ruhland 1903). Giulietti (2020b) synonymized L. vivipara var. angusta under L. vivipara and selected the material collected by Glaziou (15515, K) from Serra do Caraça as the lectotype.

10. Paepalanthus calvus Körn., Fl. bras. 3(1): 391. 1863.Fig. 4a-b

Plants 30-70 cm tall. Rhizome present. Stem reptant, non-branched. Rosette leaves 16-23 × 1.2-3.2 cm, lanceolate, acute to acuminate apex, glabrous on both surfaces, not ciliate. Inflorescences with scapes emerging from the stem at the leaf axil. Spathes 16-23 cm long, striate, glabrous, oblique opening. Scapes 50-70 cm long, free, 8 per rosette, glabrous. Capitula 14-16 mm wide. Involucral bracts brown, 3.5-4.5 mm long, not surpassing the flower disc, 5-6 series, ovate, obtuse apex, abaxially glabrous, ciliate with tiny hairs. Floral bracts castaneous, 3.5-4.5 mm long, spathulate, concave, rounded apex, abaxially pilose at top, ciliate, tufted. Staminate flowers 4-4.5 mm long; castaneous sepals, 2.5-3 mm long, free, spathulate, concave, obtuse apex, abaxially densely pilose, ciliate, tufted; reddish castaneous corolla, 3-3.2 mm long, tubular, acute lobes, glabrous; stamens 3, castaneous anthers, dorsifixed; 3 nectariferous carpellodes. Pistillate flowers 3.5-4 mm long; castaneous sepals at apex, light castaneous at base and middle, 2.5-3 mm long, free, linear, concave, obtuse apex, distally pilose on both surfaces, tufted; light castaneous petals, 2-2.2 mm long, free, linear, flat, truncate apex, ciliate at the distal half, tufted; gynoecium 2.5-3 mm long, 3 nectariferous branches, stigmatic branches bifid.

Examined material: trilha para Pico da Conceição, 20°06’18”S, 43°31’22”W, 1,507 m, 17.VII.2021, L. Echternacht & P. Mendes 2899 (OUPR).

Paepalanthus calvus was described from Minas Gerais, at 7,000 feet elevation (around 2,100 m), without precise locality. Records in SpeciesLink (<https://specieslink.net/search/>) are identified from the Atlantic Forest, in Minas Gerais and São Paulo, mainly in the region of the Serra da Mantiqueira, but with records also from Serra da Piedade. We expanded the species distribution to RPPN Santuário do Caraça with a new record.) During our fieldwork, a small population of Paepalanthus calvus (around 30 individuals) was found on the trail to Pico da Conceição, in a partly shaded environment at a bamboo grove understory (Chusquea nutans L.G.Clark), within a quartzitic Campo Rupestre. Individuals grow at the border of a small stream, over humid sandy soils, at around 1500 m of altitude. Specimens were recorded in July with old capitula.

Paepalanthus calvus is recognized by large and glabrous leaves on both surfaces, glabrous spathes, long scapes, large capitula, and involucral bracts dark brown, abaxially glabrous and ciliate with tiny hairs. The population found in RPPN Santuário do Caraça was notable by its rosette of appressed leaves, which are large and deep green. Individuals are perennial and robust, with elongated stems creeping above the ground covered by marcescent leaves.

11. Paepalanthus camptophyllus Ruhland, Pflanzenr. 4(30): 174. 1903.Fig. 4c-e

Plants 10-30 cm tall. Rhizome absent. Aerial stem elongated, 5-20 cm long, decumbent to erect, branched. Rosette absent, spiraled leaves 0.6-2 cm × 0.5-2 mm, lanceolate to linear, patent to recurved, acute apex, pilose to glabrescent on both surfaces, ciliate with long hairs. Inflorescences with lateral branches (paraclades) ca. 1-3 cm long, sparse throughout the stem, paracladial bracts 0.5-2 cm long, similar to the leaves in shape, length, and indumentum. Spathes 0.8-2.7 cm long, abaxially sparsely pilose to glabrescent, ciliate oblique opening. Scapes in lateral fascicles, 2.5-11 cm long, free, 1-25 per fascicle, densely pilose. Capitula 3-7 mm wide. Involucral bracts castaneous to brown, 2-3 mm long, not surpassing the flower disc, 2-3 series, ovate, acute apex, abaxially pilose, ciliate. Floral bracts brown, 3.5-4 mm long, elliptic to oblong, concave, rounded apex, abaxially pilose, ciliate, tufted. Staminate flowers 3-4 mm long; castaneous sepals, 2.5-3 mm long, free, obovate, concave, truncate to rounded apex, tufted; cream corolla, 1-1.5 mm long, tubular, glabrous; stamens 3, anther dorsifixed; 3 nectariferous carpellodes. Pistillate flowers 4-4.2 mm long; castaneous sepals, 3-3.2 mm long, free, oblong to wide-elliptic, concave, truncate apex, tufted; cream petals, 3-3.2 mm long, free, linear, flat, obtuse apex, tufted, distally ciliate; gynoecium 3 mm long, 3 nectariferous branches, stigmatic branches simple.

Figure 4 -
a-p. Species of Eriocaulaceae - a-b. Paepalanthus calvus - a. rosette of wide and glabrous leaves; b. dark brown involucral bracts; c-e. P. camptophyllus - c. habit; d. densely pilose scapes, capitulum with castaneous involucral bracts, abaxially pilose; e. elongated stem with spiraled leaves and paraclade; f-h. P. ciliolatus - f. capitulum with brown involucral bracts; g. habit; h. habitat; i-k. P. conduplicatus - i. rosette with distichous and conduplicate leaves; j. wholly fused scapes and capitula; k. habitat and habit; l-n. P. dianthoides - l. habit; m. capitulum (frontal view) with involucral bracts adaxially pilose at apex; n. capitulum (lateral view) showing the brown involucral bracts; o. P. elongatus - recurvate involucral bracts; p. P. erectifolius - habit, stem elongated into a fertile inflorescence axis, scapes in a spherical umbel. Images: a-e,p. Livia Echternacht; f-o. Ambar Soldevila

Examined material: Pico do Inficcionado, 9.IX.2013, C.A. Ferreira Junior et al. 1232 (OUPR, BHZB); Pico do Inficcionado, Garganta do Diabo, entrada de fenda entre a gruta do Centenário e a gruta da Bocaina, 20°08’01”S, 43°26’53”W, 2,000 m, 26.VII.2009, L. Echternacht et al. 2076 (BHCB); trilha entre o Campo de Fora e o Pico da Canjerana, 20°08’24”S, 43°31’31”W, 1,641 m, 7.V.2008, L. Echternacht & C. Chaussidon 1643 (BHCB).

Paepalanthus camptophyllus is endemic to the Quadrilátero Ferrífero, occurring in Serra do Caraça, Serra de Capanema, and Serra da Piedade (Rodrigues 2022). It was found in the following localities in RPPN Santuário do Caraça: Pico da Carapuça, Pico do Inficcionado, and the trail to Pico da Canjerana. It grows in high altitudinal fields, in quartzitic Campo Rupestre, between 1,600 and 2,000 m elevation, and occurs over gravelly quartzitic soil. Individuals are perennial and rupicolous. Specimens were recorded in May with young flowers, and in June and July with mature flowers.

Paepalanthus camptophyllus differs from the species with fertile lateral branches at Serra do Caraça by its elongated flexible stem (vs. rigid and erect), with scapes in fascicles (vs. in spherical umbels). It differs from the Paepalanthus with an elongated aerial stem by the branched stem, scapes densely pilose, brown involucral bracts, ovate with acute apex, pilose abaxially, ciliate, and glabrous adaxially, trimerous flowers, and anthers cream.

Paepalanthus camptophyllus was described by Ruhland (1903) based on Glaziou (15520) from RPPN Santuário do Caraça, Morro da Carapuça. Andrino et al. (2021) mention specimens in the herbaria B, P, K, BR, G, IAN. Here we report having seen specimens also in herbaria F and NY.

12. Paepalanthus ciliolatus Ruhland, Pflanzenr. 4(30): 147. 1903.Fig. 4f-h

Plants 15-30 cm tall. Rhizome absent. Stem short, 0.4-0.5 mm long, restricted to the leaf rosette, non-branched. Rosette leaves 3.5-10 cm × 0.3-0.4 cm, linear, striate, acute apex, glabrous on both surfaces, ciliate at the base. Inflorescences with scapes emerging from the stem at the leaf axil. Spathes 1.5-5 cm long, lax, striate, abaxially sparsely pilose, adaxially pilose, oblique opening. Scapes 7-20 cm long, free, 3-12 per stem, glabrous. Capitula 4-6 mm wide. Involucral bracts brown, 1.5-2 mm long, not surpassing the flower disc, 4-5 series, ovate to widely ovate, obtuse, acuminate to rounded apex, abaxially glabrous, ciliate. Floral bracts brown, 1.5 mm long, oblong, concave, obtuse apex, abaxially pilose, tufted. Staminate flowers 1.7-2.5 mm long; castaneous sepals, 1-1.5 mm long, free, obovate, concave, obtuse apex, pilose on both surfaces, tufted; cream corolla, 1-1.5 mm long, tubular, acute lobes, glabrous; stamens 3, black anthers dorsifixed; 3 nectariferous carpellodes. Pistillate flowers 2 mm long; light castaneous sepals, 1.7 mm long, free, obovate, concave, obtuse apex, pilose abaxially, tufted; cream petals, 1.7 mm long, free, linear, concave, obtuse apex, adaxially pilose, tufted; gynoecium 1.5 mm long, 3 nectariferous branches releasing below the bifid stigmatic branches.

Examined material: Cachoeira Cascatona, 20°02’29”S, 43°17’30”W, 1,052 m, 26.XI.2020, A. Soldevila & P. Vanucci 339 (OUPR).

Paepalanthus ciliolatus is endemic to Minas Gerais, recorded in the southern Espinhaço Range (Santana do Pirapama and Itambé do Mato Dentro) and in the Quadrilátero Ferrífero (Ouro Preto, Ouro Branco, and Serra do Caraça) (Rodrigues 2022). In the RPPN Santuário do Caraça, one population was found at the base of Cascatona. The population of this species grows at the riverside, a constantly humid environment by the waterfall spray, shaded by bushes and trees, over moist sandy organic soil. Individuals are perennial, and specimens were recorded in November and June with young and mature flowers.

Paepalanthus ciliolatus is mainly recognized by its relatively small size and delicate habit (15-30 cm long), with a small rosette of markedly spirodistichous leaves (5-10 cm long), striate and glabrous, and by brown involucral bracts, abaxially glabrous.

Paepalanthus ciliolatus was described by Ruhland (1903), based on type material from RPPN Santuário do Caraça, “Cachoeira de Caraça” collected by Glaziou (15527, B, F, G, MO, NY, P).

13. Paepalanthus conduplicatus Körn., Fl. bras. 3(1): 414. 1863.Fig. 4i-k

Plants 30-80 cm tall. Rhizome present. Stem short, 0.5-2 mm long, restricted to the leaf rosette, non-branched. Rosette leaves distichous and conduplicate, 23-42 × 0.6-1.4 cm, linear, striate, acuminate to acute apex, glabrous on both surfaces, sparsely ciliate at base. Inflorescences with scapes emerging from the rhizome at the leaf axil. Spathes 20-28 cm long, striate, abaxially glabrous, truncate opening. Scapes 50-70 cm long, fused to the top, 2-5 sets per rosette, sparsely pilose to glabrescent. Capitula 10-17 mm wide. Involucral bracts brown, 2-5 mm long, not surpassing the flower disc, 3-4 series, lanceolate to ovate, obtuse apex, glabrous abaxially, ciliate. Floral bracts brown, 3 mm long, linear to ovate, concave, obtuse or acute apex, tufted. Staminate flowers 2.7-3.2 mm long; castaneous sepals, 1.7-2 mm long, fused from base to middle, free at top, obovate, concave, obtuse apex, tufted; cream corolla, 1.5-1.7 mm long, infundibular, acute lobes, glabrous; stamens 3, anthers dorsifixed; 3 nectariferous carpellodes. Pistillate flowers 2.2-3 mm long; brown sepals, 2.2-2.8 mm long, free, spathulate, concave, obtuse apex, tufted; light castaneous petals, 2.2-2.5 mm long, free, obovate, concave, rounded apex, adaxially pilose, tufted; gynoecium 2.5 mm long, 3 nectariferous branches, stigmatic branches bifid.

Examined material: estrada para pico Canjerana, platô antes de chegar no cume, 20°04’32”S, 43°18’15”W, 1,790 m, 24.XI.2020, A. Soldevila & P. Vanucci 336 (OUPR); próximo ao Pico do Inficcionado, 20°08’16”S, 43°27’33”W, 1,848 m, 9.XII.2008, C.T. Oliveira & J.R. Stehmann 296 (BHCB).

Paepalanthus conduplicatus is endemic to Minas Gerais, with records in SpeciesLink (<https://specieslink.net/search/>) from the Espinhaço Range and the Quadrilátero Ferrífero (Itabira, Ouro Preto, and Serra do Caraça). It was found in RPPN Santuário do Caraça on the trail to Pico da Canjerana and near the Pico do Inficcionado. Populations grow at riversides, over sandy to organic, and constantly or seasonally flooded soils, at altitudes up to 1,850 m. Individuals are perennial and recorded in November with young flowers and in December with mature flowers.

Paepalanthus conduplicatus is mainly recognized by its robust habit, with a distichous rosette of conduplicate and glabrous leaves, spathes opening truncate, scapes bearing numerous capitula, not splitted at the apex, and brown involucral bracts.

Paepalanthus conduplicatus was described by Koernicke (1863) based on type material from RPPN Santuário do Caraça collected by Sellow (s.n., B).

14. Paepalanthus dianthoides Mart., Fl. bras. 3(1): 339. 1863.Fig. 4l-n

Plants 20-35 cm tall. Rhizome absent. Aerial stem elongated, 12-20 cm long, erect or decumbent, branched or not. Rosette absent, leaves spiraled 3-6 cm × 1.5-4.5 mm, lanceolate, acute to cuspidate apex, pubescent to glabrescent on both surfaces, ciliate at base with long hairs. Inflorescences with scapes emerging from the stem at the leaf axil. Spathes 3.5-6 cm long, abaxially pubescent to glabrescent, ciliate, oblique opening. Scapes 12-28 cm long, free, 3-5 per stem branch, pubescent to glabrescent. Capitula 10-13 mm wide. Involucral bracts brown, 4-7 mm long, surpassing or not the flower disc up to 0.5-1 mm, 5-6 series, ovate to lanceolate, acuminate to long acuminate apex, abaxially densely pilose with patent and adpressed hairs, adaxially distally pilose, ciliate. Floral bracts brown, 5.5-6 mm long, apiculate, concave, acute apex, tufted, distally ciliate. Staminate flowers 5.5-6 mm long; light castaneous sepals, 5-6 mm long, fused at base, elliptic, concave, acute apex, tufted; cream corolla, 4-4.5 mm long, infundibular, acute to round lobes, adaxially pilose, ciliate; stamens 3, anthers dorsifixed; 3 nectariferous carpellodes. Pistillate flowers 5.5-6 mm long; light castaneous sepals, 5-5.5 mm long, free, elliptic, concave, acute apex, tufted; cream petals, 4-4.5 mm long, free, elliptic to oblong, concave, truncate to rounded apex, adaxially pilose, tufted; gynoecium 5 mm long, 3 nectariferous branches, stigmatic branches bifid.

Examined material: Serra do Caraça, VIII.1971, M.A. Lisboa (OUPR 13256); Tabuões lado direito, primeiro sumidouro subindo o rio, 20°02’44”S, 43°18’04”W, 1,180 m, 23.XI.2020, A. Soldevila & P. Vanucci 322 (OUPR); Tabuões lado direito, segundo sumidouro subindo o rio, 20°02’44”S, 44°18’04”W, 1,188 m, 23.XI.2020, A. Soldevila & P. Vanucci 324 (OUPR).

Paepalanthus dianthoides is endemic to Minas Gerais, occurring from the RPPN Santuário do Caraça at the south, throughout the adjacent Serra do Garimpo to the north, at the southern Serra do Cipó, and with a disjunct population in the Diamantina Plateau (Hensold 1988; Andrino & Costa 2013). In the RPPN Santuário do Caraça, it was found in the Pico da Conceição and Tabuões. The population found in Tabuões was over a river sinkhole, growing in quartzitic Campo Rupestre, over gravelly to sandy soil, among shrubby to herbaceous vegetations. Individuals are perennial, and specimens with flowers were recorded in March, June, August, and November, the latter two showing young and mature flowers.

Paepalanthus dianthoides shares with P. mollis the elongated aerial stem, with spiraled leaves, brown involucral bracts, and ciliate and adaxially pilose corolla of the staminate flowers. The former differs from the latter by the acuminate to the long acuminate apex of the external series of involucral bracts (vs. obtuse to acute apex), the internal series sometimes surpassing the flower disc 0.5-1 mm long (vs. constantly surpassing, 1.5-3 mm long), adaxially pilose at the apex only (vs. adaxially densely pilose from middle to top), floral bracts apiculate (vs. lanciform), petals of pistillate flowers cream (vs. blackish at apex), and by non pigmented anthers (vs. pigmented).

15. Paepalanthus elongatus (Bong.) Körn., Fl. bras. 3(1): 312. 1863.Fig. 5o

Plants 45-75 cm tall. Rhizome present. Stem short, 1-3 cm long, restricted to the base of the leaves, branched. Rosette absent, leaves caespitose 9.5-20 cm × 1-3 mm, linear, obtuse to rounded apex, glabrous on both surfaces, ciliate or not. Inflorescences with scapes emerging from the stem at the leaf axil. Spathes 7-12 cm long, abaxially sparsely pilose to glabrescent, oblique opening. Scapes 28-70 cm long, free, 1-2 per stem branch, sericeous with adpressed hairs. Capitula 10-20 mm wide. Involucral bracts dark brown with light brown margins, 3-5 mm long, recurvate, surpassing the flower disc 1-2 mm long, 6-9 series, lanceolate to ovate, acute apex, abaxially glabrous, ciliate. Flowers dimerous. Floral bracts brown, 2-3 mm long, lanceolate, concave, acute apex, tufted with long hairs. Staminate flowers 2-2.5 mm long; castaneous sepals, 2-2.3 mm long, free, obovate to oblong, concave, truncate apex, tufted; cream corolla, 2 mm long, tubular, glabrous; stamens 2; dorsifixed anthers; 3 nectariferous carpellodes. Pistillate flowers 2.2-2.5 mm long; brown sepals, 2.5 mm long, free, obovate, concave, rounded apex, tufted; light castaneous petals, cream in the apex, 2-2.2 mm long, free, elliptic, flat, obtuse apex, tufted; gynoecium 2.5 mm long, 3 nectariferous branches, stigmatic branches bifid.

Examined material: Campos de Fora, 29.V.2002, J. Ordones 873 (BHZB, OUPR); 20°04’13”S, 43°18’39”W, 1,497 m, 24.XI.2020, A. Soldevila & P. Vanucci 329 (OUPR); Serra do Caraça, 17.IV.1933, H.L.M. Barreto 2579 (BHCB, NYBG); 19.VIII.2000, R.C. Mota 880 (BHCB).

Paepalanthus elongatus is widely distributed in Brazil, occurring in Tocantins, Bahia, Góias, Minas Gerais, and São Paulo, in the Domains of the Caatinga, Cerrado, and Atlantic Forest (Andrino et al. 2024). It was found in the following localities in RPPN Santuário do Caraça: Pico da Conceição, Campos de Fora, Pico da Canjerana, and the trail to Mirante. The population found in Campos de Fora was growing in quartzitic Campo Rupestre, over humid sandy soil, among open herbaceous vegetation. Individuals are perennial, and specimens with flowers were recorded in February, March, April, May, August, and November. In May, specimens have young flowers; in April and August, mature flowers, and in November, old flowers.

Paepalanthus elongatus is mainly recognized by its caespitose habit, with linear leaves, recurvate involucral bracts, dark brown in the middle and light brown in the margins, remarkably ciliate, slightly surpassing the flower disc (1-2 mm long), and by dimerous flowers.

16. Paepalanthus erectifolius Silveira, Flora e Serras Mineiras: 51. 1908.Fig. 5p

Plants 45-120 cm tall. Rhizome present. Stem short, 1-2.5 cm long, restricted to the leaf rosette in sterile plants, non-branched, elongating into an inflorescence (fertile) axis. Rosette leaves 10-12 × 0.2-0.5 cm, lanceolate, acute apex, sparsely pilose to glabrescent on both surfaces, ciliate or not. Inflorescences axis 45-100 cm long, erect, with scapes at the apex; axis bracts 3-6 cm long, lanceolate, patent, acute apex, glabrescent on both surfaces, sparsely ciliate to glabrescent. Spathes 3-6 cm long, abaxially glabrous, oblique opening. Scapes 21-32 cm long, free, 25-100 in spherical umbels, glabrous. Capitula 7-9 mm wide. Involucral bracts brown, 1.5-2 mm long, not surpassing the flower disc, 3-5 series, obovate, acute to obtuse apex, abaxially glabrous, ciliate. Flowers dimerous. Floral bracts ligth castaneous, 2.5-3 mm long, linear, flat, acute apex, ciliate at the apex. Staminate flowers 2.5-3 mm long; light castaneous sepals, 2 mm long, free, obovate, concave, acute apex, tufted; corolla cream, 2 mm long, tubular, glabrous; stamens 2; dorsifixed anthers; 3 nectariferous carpellodes. Pistillate flowers 2-2.5 mm long; light castaneous sepals, 2.5 mm long, free, dolabriform, concave, mucronate apex, abaxially densely pilose; cream petals, 2.2 mm long, free, spathulate, flat, obtuse apex, abaxially densely pilose, tufted; gynoecium 1.8 mm long, 3 nectariferous branches, stigmatic branches bifid.

Examined material: trilha para a cachoeira da Bocaina, ca. 500 m antes da cachoeira, 20°07’25”S, 43°28’12”W, 1,272 m, 3.VI.2012, LA Echternacht, A Diaz 2261 (BHCB, SPF); trilha para Cachoeira da Bocaina, depois da Pedra da Paciência, 20°04’20”S, 43°16’52”W, 1,271 m, 4.XI.2020, A. Soldevila & P. Vanucci 315 (OUPR).

Paepalanthus erectifolius is distributed in Minas Gerais and Bahia, and presents disjunctive populations in quartzitic Campo Rupestre, occurring in the Diamantina Plateau, Serra do Cabral, Serra do Cipó, and Chapada Diamantina; it occurs in the Domains of the Caatinga and Cerrado (Trovó et al. 2013). One population only is recorded in RPPN Santuário do Caraça, on the trail to Bocaina waterfall, with about ten individuals growing in quartzitic Campo Rupestre, over sandy soil, among shrubby to herbaceous vegetation. Individuals are monocarpic; specimens were recorded in June with young and mature flowers and in November with old and dry flowers.

Paepalanthus erectifolius is mainly recognized by its robust habit, the stem developing into an erect elongated fertile axis, with patent bract, scapes in spherical umbels at the apex of the fertile axis, and dimerous flowers.

17. Paepalanthus flaccidus (Bong.) Kunth, Enum. Pl. 3: 511. 1841.Fig. 5a

Perennial plants 8-20 cm tall. Rhizome absent. Aerial stem elongated, 10-30 cm long, decumbent, branched or not, bearing an umbel of inflorescences at the apex. Rosette absent, leaves spiraled 8-16 × 0.5-1.2 mm, lanceolate, acute apex, glabrous on both surfaces, ciliate with long hairs. Inflorescences with scapes emerging from the stem at the leaf axil. Spathes 1.5-2 cm long, glabrous abaxially, oblique opening. Scapes 8-22 cm long, free, 5-20 in umbels per stem branch, sparsely pilose to gabrescent. Capitula 2-5 mm wide. Involucral bracts cream to golden, 1-1.5 mm long, not surpassing the flower disc, 3-4 series, ovate, obtuse to rounded apex, abaxially glabrous, not ciliate. Flowers dimerous. Floral bracts brown, 1.5 mm long, linear, flat, acute apex, tufted. Staminate flowers 1.2-1.5 mm long; brown sepals, 1.2 mm long, free, obovate, concave, rounded apex, tufted; cream corolla, 0.5 mm long, tubular, glabrous; stamens 2, dorsifixed anthers; 3 nectariferous carpellodes. Pistillate flowers 1.5-1.7 mm long; brown sepals, 1.5-1.7 mm long, free, dolabriform, concave, rounded apex, tufted; cream petals, 1.5-1.7 mm long, free, obovate, flat, rounded apex, tufted; gynoecium 1.7 mm long, 3 nectariferous branches, stigmatic branches bifid.

Local records: Caraça, Plantes des environs de Rio de Janeiro et d’Ouro Preto, 14.VI.1884, A.F.M. Glaziou 15518 (NYBG)

Examined material: Catas Altas, RPPN Santuário do Caraça, 14.IV.1933, HLM Barreto 2528 (BHCB). Itabirito, Serra de Capanema, 27.VI.1975, M. Lisboa (OUPR 12614). Ouro Preto, Bico de Pedra, 21.II.1975, M. Lisboa (OUPR 12613); Campo Grande, 6.XI.1980, J. Badini (OUPR 12605).

Paepalanthus flaccidus is distributed in Goiás, Minas Gerais, and São Paulo (Andrino et al. 2024). Two records only were registered for this species in RPPN Santuário do Caraça, with no precise locality. Label notes mention that it was collected in a marshy site, being infrequent. Individuals are perennial, and specimens were recorded in April and June with immature inflorescences.

Paepalanthus flaccidus is mainly recognized by its decumbent and delicate habit, aerial stem elongate, with small, linear, and spiraled leaves, involucral bracts cream to gold, and dimerous flowers.

Two records were found in RPPN Santuário do Caraça, but only one was available at the herbarium (BHCB) for analysis, and it has immature flowers (Barreto, HLM 2528). Therefore, the description of fertile parts was based on records from Serra do Capanema and the municipality of Ouro Preto, the closest available sites.

18. Paepalanthus freyreissii (Thunb.) Körn., Fl. bras. 3(1): 370. 1863.Fig. 5b

Plants 10-50 cm tall. Rhizome present or not. Aerial stem short to elongated, 4-15 cm long, erect, branched. Rosette leaves spiraled 1.8-7 cm × 0.5-1.5 mm, striate, linear, acute apex, glabrous on both surfaces, not ciliate. Inflorescences with scapes emerging from the stem at the leaf axil. Spathes 1-2.2 cm long, abaxially glabrous, oblique opening. Scapes 4-20 cm long, free, 2-30 per rosette, glabrous. Capitula 4-6 mm wide. Involucral bracts brown, 5-2 mm long, not surpassing the flower disc, 3-4 series, elliptic, obtuse apex, abaxially glabrous, not ciliate. Floral bracts brown, 2-2.5 mm long, oblong, concave, mucronate apex, abaxially densely pilsose, tufted with long hairs. Staminate flowers 3.5-4 mm long; brown sepals, 1.5-2 mm long, free, oblanceolate, concave, rounded apex, abaxially pilose, tufted with long hairs; cream corolla, 1.5 mm long, infundibular, obtuse to round lobes, glabrous; stamens 3, black anthers, dorsifixed; 3 nectariferous carpellodes. Pistillate flowers 2-2.5 mm long; castaneous to light castaneous sepals, 1.5-1.7 mm long, free, obovate, concave, rounded apex, abaxially pilose, tufted with long hairs; cream petals, 1.7 mm long, free, linear, flat, rounded apex, adaxially pilose, tufted; gynoecium 2 mm long, 3 nectariferous branches, stigmatic branches bifid.

Examined material: Fraldas da Serra do Caraça, próximo a Santa Rita, 4.VIII.1971, J. Badini (OUPR 12590); próximo ao Pico do Inficcionado, 20°08’16”S, 43°27’33”W, 1,848 m, 9.XII.2008, C.T. Oliveira & J.R. Stehmann 298 (BHCB).

Paepalanthus freyreissii is endemic to Minas Gerais, distributed in disjunct populations among the Serra da Mantiqueira, Espinhaço Range, and Quadrilátero Ferrífero (Ferreira et al. 2011; Andrino et al. 2024). Two records only were recorded for this species in RPPN Santuário do Caraça, one with no precise locality and the other near the Pico do Inficcionado. Population grows in quartzitic Campo Rupestre, at altitudes up to 1,850 m, occurring in shady and humid environments, over riverside or near caves (Ferreira et al. 2011). Specimens were recorded in August and December with mature flowers.

Figure 5 -
a-l. Species of Eriocaulaceae - a. Paepalanthus flaccidus - stem elongated, branched, scapes in an umbel at the apex; b. P. freyreissii - habitat and habit; c. P. hilairei - habit, elongated fertile axis bearing verticillate paraclades; d-e. P. leucoblepharus - d. detail of blackish involucral bracts; e. detail of adaxially glabrous leaves conspicuously ciliate, abaxially densely pilose spathes, with striate indumentum; f-g. P. melaleucus - f. habitat and habit; g. fused scapes, splitting at the top, castaneous involucral bracts; h-i. P. mollis var. mollis - h. habit; i. capitulum with involucral bracts adaxially densely pilose from middle to the top; j-k. P. planifolious - j. capitula and scapes wholly fused; k. habitat and habit; l. P. polyanthus - habit, elongated fertile axis bearing spiraled paraclades. Images: a. from Serra de Capanema; b. from Serra de Ibitipoca; c-l. from Serra do Caraça. a-c,h,i,l. Livia Echternacht; d-g,j,k. Ambar Soldevila

Paepalanthus freyreissii is mainly recognized by its delicate habit of dark green leaves and rosettes at the apex of short to elongated, branched aerial stems; fresh leaves often show horizontal stria, are linear, and glabrous; spathes, scapes, and the brown involucral bracts are also glabrous.

19. Paepalanthus hilairei Körn. Fl. bras. 3(1): 332. 1863.Fig. 5c

Plants 100-200 cm tall. Rhizome absent. Stem restricted to the leaf rosette when sterile, non-branched, elongating into an inflorescence (fertile) axis. Rosette leaves 17-31.5 × 1.2-2.5 cm, lanceolate, acute to mucronate apex, pilose to glabrescent on both surfaces, ciliate. Inflorescences bearing a fertile axis and lateral branches (paraclades) 14-33 cm long, these verticillate, paracladial bracts 3-7 cm long, lanceolate, acute apex, pilose to glabrescent on both surfaces, ciliate. Spathes 1.4-2 cm long, lax, abaxially pilose to glabrescent, oblique opening. Scapes 5.8-15.5 cm long, free, 60-220 in spherical umbels, glabrescent, or pilose when young. Capitula 3-4 mm wide. Involucral bracts castaneous, 1-1.8 mm long, not surpassing the flower disc, obovate, obtuse apex, abaxially pilose to glabrescent, ciliate. Floral bracts light castaneous, 1.8-2 mm long, narrow-elliptic, concave, cuspidate apex, abaxially densely pilose, tufted. Staminate flowers 2 mm long; light castaneous sepals, 1.6-1.8 mm long, free, obovate, concave, obtuse to rounded apex, tufted; cream corolla, 1.6-1.8 mm long, infundibular, glabrous; stamens 3, anther dorsifixed; 3 nectariferous carpellodes. Pistillate flowers 2.2 mm long; light castaneous sepals, 1.8-2.2 mm long, free, obovate, obtuse apex, tufted; cream petals, 1.8-2 mm long, free, elliptic, acute apex, abaxially densely pilose from middle to apex; gynoecium 1.8-2.2 mm long, 3 nectariferous branches, stigmatic branches simple, longer than the nectariferous branches.

Examined material: Caminho da Verruguinha, 1.V.1980, T.S.M. Grandi & Tales 79 (BHCB); Campos de Fora, 20°06’54”S, 43°31’13”W, 1,493 m, 17.VII.2021, L. Echternacht & P. Mendes 2889 (OUPR); Serra do Caraça, 19.VI.2002, T.M.A. Alves & M. Sobral 56 (BHCB); 19.VI.1974, J. Badini (OUPR 13231); 19.VI.1974, J. Badini (OUPR 13234); 16.IV.1933, H.L.M. Barreto 699 (BHCB); 14.IV.1933, H.L.M. Barreto 700 (BHCB, US); VI.1907, L.B. Damazio (OUPR 13221); L.B. Damazio (OUPR 13229); 2.IV.2000, R.C. Mota 762 (BHCB); 6.IV.1976, M.A. Zurlo (OUPR 13208); trilha para Capelinha, 20°03’16”S, 43°17’26”W, 1,325 m, 3.XI.2020, A. Soldevila & P. Vanucci 306 (OUPR); trilha para o Pico do Inficcionado, 15.VI.2000, M.F. Vasconcelos (BHCB 52713).

Paepalanthus hilairei is widely distributed in Brazil, occurring from Piauí to São Paulo (= Actinocephalus bongardii (A.St.-Hil.) Sano; Sano et al. 2010; Costa & Sano 2024). It is also widely distributed in RPPN Santuário do Caraça, being the most abundant species of the family, recorded in the following localities: the trail to Cascatona, the trail to Campos de Fora, the trail to Capelinha, Piscina, the trail to Padre Caio cave, Cascudo, Mirante da Piscina, and the trail to Tabuões and Bocaina. Populations occur in quartzitic Campo Rupestre, mainly in open herbaceous vegetation, over sandy to organic soil. Individuals are monocarpic, with one to two cohorts; specimens were recorded from October to July, with the flowering peak from January to March.

Paepalanthus hilairei is recognized by compound inflorescence, with an elongated fertile axis bearing lateral branches (paraclades), these verticillate along the axis and with scapes in spherical umbels; scapes are pilose when young, and the rosette dries out and eventually disappears in fertile individuals. The most similar species, Paepalanthus polyanthus, has spiraled paraclades and scapes villous when young.

20. Paepalanthus leucoblepharus Körn., Fl. bras. 3(1): 388. 1863.

= Paepalanthus globosusRuhland, Pflanzenr. 4(30): 143. 1903. Typus: BRAZIL. MINAS GERAIS: “Serra de Capanema, juin de Caraça”, 20.II.1884, A.F.M. Glaziou 15537 (lectotype B!, designated by Moldenke (1975: 483); isolectotypes C!, F “photo!”, LL!, P “photo!”), syn.nov.Fig. 5d-e

Perennial plants 27-70 cm tall. Rhizome absent. Stem short, 0.5-3 cm long, restricted to the leaf rosette, branched. Rosette leaves 3-17 cm × 3-12 mm, lanceolate, striate, obtuse to acute apex, pilose to glabrous on both surfaces, densely ciliate. Inflorescences with scapes emerging from the stem at the leaf axil. Spathes 6.5-15 cm long, striate, abaxially densely pilose in the strias, adaxially pilose at apex, oblique opening. Scapes 22-60 cm long, free, 1-10 per rosette, pilose to glabrous. Capitula 9-16 mm wide. Involucral bracts blackish, 2-3 mm long, not surpassing the flower disc, 4-5 series, oblong to orbicular, round, obtuse, or mucronate apex, abaxially glabrous, ciliate. Floral bracts brown, 3-3.2 mm long, linear, flat, obtuse apex, abaxially densely pilose, tufted with long hairs. Staminate flowers 3-3.2 mm long; castaneous sepals, 2 mm long, fused at base, linear to oblanceolate, flat, rounded apex, abaxially densely pilose, tufted with long hairs; cream corolla, 2 mm long, infundibular, acute lobes, glabrous; stamens 3, blackish anthers, dorsifixed; 3 nectariferous carpellodes. Pistillate flowers 3-3.2 mm long; castaneous sepals, 2 mm long, free, linear to oblanceolate, flat, rounded apex, abaxially densely pilose, tufted with long hairs; cream petals, 1.8 mm long, free, linear, flat, acute apex, ciliate in the middle and top with long hairs, tufted; gynoecium 1.7-2.2 mm long, 3 nectariferous branches, stigmatic branches bifid.

Examined material: caminho para o Pico do Sol, 20°06’34”S, 43°27’06”W, 1,837 m, 28.II.2009, L. Echternacht et al. 1940 (BHCB); 3.III.2010, L.N. Perillo 464 (BHCB); estrada para pico Canjerana, subida para o pico, 20°04’29”S, 43°18’19”W, 1,696 m, 24.XI.2020, A. Soldevila & P. Vanucci 335 (OUPR); Pico da Carapuça, 20°05’27”S, 43°28’17”W, 1,848 m, 17.II.2009, C.T. Oliveira & A.J. Arruda 328 (BHCB, RB); Pico da Conceição, 20°05’39”S, 43°31’33”W, 1,778 m, 17.VII.2021, L. Echternacht & P. Mendes 2898 (OUPR); 20°05’39”S, 43°31’37”W, 1,783 m, 26.VIII.2008, C.T. Oliveira & L.L. Giacomin 52 (BHCB); Pico do Inficcionado, 20°08’05”S, 43°27’05”W, 2,075 m, 8.III.2008, L. Echternacht et al. 1614 (BHCB); 9.IX.2013, C.A. Ferreira Junior et al. 1225 (BHZB, OUPR); Pico do Sol, 20°06’34”S, 43°27’03”W, 1,827 m, 25.IV.2009, C.T. Oliveira et al. 427 (BHCB); trilha entre o Campo de Fora e o Pico da Canjerana, 20°08’24”S, 43°31’31”W, 1,641 m, 7.V.2008, L. Echternacht & C. Chaussidon 1641 (BHCB); Serra do Caraça, 1905, A. Silveira (OUPR 13257).

Paepalanthus leucoblepharus is endemic to the southeastern Quadrilátero Ferrífero, occurring in the Serra do Caraça, Ouro Preto, and Serra de Capanema (Rodrigues 2022). It is widely distributed in the RPPN Santuário do Caraça, forming abundant populations at most local peaks. This species was recorded in the following localities: Pico do Sol, Pico da Carapuça, ascent to Pico da Canjerana, Pico do Inficcionado, and Pico da Conceição. It grows in quartzitic Campo Rupestre, at altitudes above 1,600 m, up to 2,068 m, over sandy to gravelly soil, among outcrops and a shrubby to herbaceous vegetation. Specimens were recorded with flowers in February, March, April, May, June, July, August, and November, indicating that it flourishes throughout the year.

Paepalanthus leucoblepharus is mainly recognized by leaves conspicuously ciliated, spathes with striate indumentum, densely pilose abaxially, adaxially tufted at apex, and blackish involucral bracts and anthers.

Paepalanthus leucoblepharus was described by Körnicke (1863) based on a material collected by Riedel s.n. without a specific location. In the LE herbarium, the second author found the Riedel s.n. material identified as P. leucoblepharus with handwriting that probably belongs to Köernicke. Although this specimen has no locality reference, it shows the diagnostic characteristics defined in the protologue: leaves conspicuously ciliate and spathes adaxially tufted at the apex. Therefore, we interpret it as the holotype since no other Riedel’s collection was found assigned to this species.

Paepalanthus globosus was described by Ruhland (1903) and has the type material from RPPN Santuário do Caraça (Glaziou 15537). The lectotype at B is quite fragmented but brings Ruhland’s notes and drawings. Dr. Echternacht analyzed it physically, as well as the isolectotype at C, which matches the diagnostic features of P. leucoblepharus, notably the ciliate leaves, the striate and adaxially tufted spathe, a rare feature in Eriocaulaceae. We conclude that these two names correspond to the same species that flourishes in the high mountains of Serra do Caraça, and we consider P. globosus synonymous with P. leucoblepharus. These features also allow the differentiation of this species from Paepalanthus falcatus, a close morphological species, as reported by Andrino et al. (2022).

21. Paepalanthus manicatus Poulsen ex Malme, Bih. Kongl. Svenska Vetensk.-Akad. Handl. 27(3, 11): 28-29. pl. 2. f. 3. 1901.

= Paepalanthus manicatus f. robustaRuhland, Pflanzenr. 4(30): 158. 1903. Typus: BRAZIL. MINAS GERAIS: “Na Serra de Caraça em Minas Geraes”, III.1892, E. Ule 2715 (type HGB “photo!”, R!), syn.nov.

Plants 6-8.5 cm tall. Rhizome absent. Aerial stem elongated, 0.5-3 cm long, non-branched. Rosette absent, leaves spiraled 2-15 × 0.5-1 mm, linear, parallel-veined, three-nerved, acute apex, glabrous on both surfaces, sparsely ciliate with long hairs. Inflorescences with scapes emerging from the stem, fasciculate at the apex. Spathes 2-5 mm long, abaxially glabrous, oblique opening. Scapes 3.5-7.5 mm long, free, 3-10 per fascicle, glabrous. Capitula 1-1.5 mm wide. Involucral bracts brown, 0.7-1 mm long, not surpassing the flower disc, 2-3 series, elliptic, obtuse to acute apex, abaxially glabrous, not ciliate. Floral bracts castaneous, 1 mm long, obovate to oblong, concave, obtuse apex, abaxially pilose. Staminate flowers 1 mm long; castaneous sepals, 0.7 mm long, fused at base, oblong to obovate, flat, obtuse apex, glabrous; cream corolla, 0.7 mm long, infundibular, obtuse lobes, glabrous; stamens 3, dorsifixed anthers; 3 nectariferous carpellodes. Pistillate flowers 1 mm long; castaneous sepals, 0.7 mm long, free, ovate, concave, obtuse apex, glabrous; light castaneous petals, 0.7 mm long, free, concave, elliptic, flat, acute apex, tufted; gynoecium 0.7-1 mm long, 3 nectariferous branches, stigmatic branches bifid.

Examined material: Serra do Caraça, 8.X.1974, J. Badini (OUPR 12569); 1904, C.A.W. Schwacke (BHCB 547).

Paepalanthus manicatus is widely distributed in Brazil, occurring in Bahia, Piauí, Goiás, Minas Gerais, and São Paulo (Andrino et al. 2024). Records from RPPN Santuário do Caraça have no precise locality. This species usually grows over humid sandy soil, often in shadowy places. Specimens were recorded in March with young and mature flowers and in October with mature and old flowers.

Paepalanthus manicatus is mainly recognized by its small size (6-8.5 cm tall), a delicate herb that forms clustered populations. It can be recognized also by its elongated stem, with spiraled leaves, which are three-nerved, membranaceous, glabrous on both surfaces, and sparsely ciliate with long hairs, in addition to the fasciculate scapes at the stem apex and brown involucral bracts.

Ruhland (1903) described Paepalanthus manicatus f. robusta with type material from RPPN Santuário do Caraça (voucher Ule 2715, HBG), and describes that this form differs from the typical by the following characteristics: more robust habit, thicker stem, slightly longer leaves, more numerous scapes and slightly larger capitula. Andrino et al. (2024) do not recognize infraspecific taxa for this species, but do not cite P. manicatus f. robusta. Here follow a broad concept of the species and consider this form synonymous with the autonym.

22. Paepalanthus melaleucus (Bong.) Kunth, Enum. Pl. 3: 510. 1841.Fig. 5f-g

Perennial plants 30-50 cm tall. Rhizome absent. Stem reptant, elongated, 7-20 cm long, erect, branched or not. Rosette leaves 15-20 × 0.5-0.6 cm, lanceolate to linear, acuminate apex, glabrous on both surfaces, sparsely ciliate with long hairs. Inflorescences with scapes emerging from the stem at the leaf axil. Spathes 2.2-5 cm long, glabrous abaxially, truncate opening. Scapes 12-22 cm long, fused, splitting at the top, 10-40 sets per rosette, pilose to glabrous. Capitula 6-18 mm wide. Involucral bracts brown, 2.5-3 mm long, not surpassing the flower disc, 3-4 series, ovate, acute apex, glabrous abaxially, not ciliate. Floral bracts brown, 3 mm long, oblong to obovate, concave, acute apex, tufted. Staminate flowers 3-3.2 mm long; castaneous sepals, 2.2 mm long, fused at base and middle, obovate, concave, round to truncate apex, tufted; cream corolla, 1.7 mm long, infundibular, acute lobes, glabrous; stamens 3, dorsifixed anthers; 3 nectariferous carpellodes. Pistillate flowers 3.5-4 mm long; castaneous sepals, 2.5-3 mm long, free, obovate, concave, rounded apex, tufted; cream petals, 2.5-3 mm long, oblong to obovate, flat, obtuse apex, tufted; gynoecium 2.5-3 mm long, 3 nectariferous branches, stigmatic branches bifid.

Examined material: Cachoeira da Bocaina, 20°04’20”S, 43°16’32”W, 1,264 m, 4.XI.2020, A. Soldevila & P. Vanucci 318 (OUPR).

Paepalanthus melaleucus has one record from Bahia, and all the others are from Minas Gerais. In the RPPN Santuário do Caraça, one population was found on the margins of waterfall Bocaina, with about 60 individuals. It grows at a riverside, on a slope, in a constantly humid environment, shaded by shrubs and trees, over humid, organic to sandy soil. Individuals are perennial, and specimens were recorded with flowers in February, March, June, August, and November, indicating that it blooms all over the year.

Some confusion pervades the taxonomic history of this taxon. It was described by Bongard (1831) from a Riedel collection from Minas Gerais. Dr. Echternacht found only one specimen assigned from Minas Gerais, with Riedel’s label and handwriting, which is in OXF. All the other collections from Riedel (in LE, G, and UPS) are identified by Körnicke, assigned by him from Bahia, and cited at the Flora brasiliensis (Körnicke 1863). However, all these specimens from Riedel seem very similar, and no other collection of this species from Bahia was made to the present day [Tissot-Squalli 1997; SpeciesLink (<https:// specieslink.net/search/>)]. We wonder whether there was a mislabeling from Körnicke, who possibly assigned the specimen from Bahia by mistake. In this case, the species should be considered endemic to Minas Gerais. In addition, Paepalanthus corymboides is currently under synonymy with P. melaleucus, and its type is from Serra do Caraça (Ule 2725, B).

Pepalanthus melaleucus is mainly recognized by its elongated stem, becoming reptant with age, spathe opening truncate and fused scapes, splitting distally. It can also be recognized by glabrous leaves on both surfaces, sparsely ciliate with long hairs, and castaneous involucral bracts.

23. Paepalanthus mollis Kunth var. mollis, Enum. Pl. 3: 507. 1841.Fig. 5h-i

Perennial plants 15-30 cm tall. Rhizome absent. Aerial stem usually elongate, sometimes short, 1-15 cm, erect or decumbent, branched or not. Rosette absent, leaves spiraled 2-7 × 0.1-0.3 cm, linear, acute apex, pilose to glabrescent on both surfaces, ciliate. Inflorescences with scapes emerging from the stem at the leaf axil. Spathes 3.5-6 cm long, abaxially pilose, ciliate with long hairs, oblique opening. Scapes 10-30 cm long, free, 1-10 per stem branch, pilose to glabrescent. Capitula 7-17 mm wide. Involucral bracts blackish, 5-6 series, ovate, abaxially sparsely pilose, with adpressed hairs, adaxially densely pilose from middle to top, ciliate, external series 1.5-3 mm long, obtuse to acute apex, internal series 4-5 mm long, sometimes recurvate, surpassing the flower disc 1.5-3 mm long, acute apex. Floral bracts blackish, 3.5 mm long, lanciform, flat, acute apex, ciliate at the distal half, tufted with long hairs. Staminate flowers 3-3.8 mm long; light castaneous sepals at the proximal half, blackish at the distal half, 2-2.2 mm long, fused at base, obovate, concave, obtuse apex, tufted with long hairs; castaneous corolla, 2 mm long, infundibular, obtuse lobes, adaxially pilose; stamens 3, pigmented anthers, dorsifixed; 3 nectariferous carpellodes. Pistillate flowers 3-3.5 mm long; light castaneous sepals at the base and middle, blackish at apex, 3-3.5 mm long, free, linear, flat, obtuse to rounded apex, tufted with long hairs; light castaneous petals at the base and middle, blackish at apex, 2.5-3 mm long, free, oblong to obovate, concave, rounded apex, adaxially pilose, tufted; gynoecium 3-3.2 mm long, 3 nectariferous branches, stigmatic branches bifid.

Examined material: caminho para o Pico do Sol, 20°06’34”S, 43°27’06”W, 1,837 m, 28.II.2009, L. Echternacht et al. 1939 (BHCB, HUFU); Pico do Inficcionado, 20°08’05”S, 43°27’05”W, 2,075 m, 9.III.2008, L. Echternacht et al. 1617 (BHCB); 7.VIII.2002, R.C. Mota 3554 (BHCB); 20°08’05”S, 43°27’05”W, 2,075 m, 1.X.2008, C.T. Oliveira & P.L. Viana 138 (BHCB); 20°08’05”S, 43°27’05”W, 2,075 m, 10.XII.2008, C.T. Oliveira & J.R. Stehmann 299 (BHCB); 7.VIII.2002, J. Ordones 923 (BHZB, OUPR); 2.IX.1999, M.F. Vasconcelos (BHCB 48905); 4.IX.1999, M.F. Vasconcelos (BHCB 48902); 11.X.1999, M.F. Vasconcelos (BHCB 52695); 9.I.2000, M.F. Vasconcelos (BHCB 52684); subida para Pico do Inficcionado, 8.IX.2013, C.A. Ferreira Junior & J. Ordones 1215 (BHZB, OUPR, SPF); trilha para o Pico do Sol, 14.XI.2009, L.N. Perillo 443 (BHCB).

Paepalanthus mollis var. mollis is endemic to Minas Gerais, with disjunct populations in Ouro Preto and Serra do Caraça (Hensold 1988), Serra da Bocaina and Serra de Carrancas (Mourão et al. 2017). The population from RPPN Santuário do Caraça is abundant and frequent at some peaks, being recorded in the Pico do Inficcionado, Pico do Sol, and Pico da Carapuça. It grows in altitudes from 1,800 up to 2,068 m a.s.l., in quartzitic Campo Rupestre, at riversides or outcrops, at the borders of crevices, in shaded or sunny places, over sandy soil. Individuals are perennial, and specimens with flowers were recorded throughout the year.

Paepalanthus mollis var. mollis is distinct mainly by the elongated stem with spiraled leaves, and blackish involucral bracts, which are pilose adaxially and surpass the flower disc. These features are shared with P. dianthoides, from which it differs by the external series of the involucral bracts with obtuse to acute apex (vs. acuminate to long acuminate), the internal series densely pilose adaxially from middle to top (vs. at the top only), surpassing the flower disc 1.5-3 mm long (vs. 0.5-1 mm), in addition to lanciform floral bracts (vs. apiculate), petals of pistillate flowers blackish at the apex (vs. cream), and by pigmented anthers (vs. cream).

24. Paepalanthus planifolius (Bong.) Körn., Fl. bras. 3(1): 413. 1863.Fig. 5j-k

Perennial plants 20-60 cm tall. Rhizome present. Stem restricted to the leaf rosette, non-branched. Rosette leaves 7-29 × 0.9-3 cm, lanceolate, apex acuminate, glabrous to pilose on both surfaces, not ciliate. Inflorescences with scapes emerging from the stem at the leaf axil. Spathes 8-13 cm long, abaxially glabrescent to glabrous, truncate opening. Scapes 15-35 cm long, fused to the top, 10-50 sets per rosette, sparsely pilose to glabrescent, with adpressed hairs. Capitula 9-19 mm wide. Involucral bracts castaneous, 3-4 mm long, not surpassing the flower disc, 2-3 series, ovate, acute apex, abaxially densely pilose, not ciliate. Floral bracts castaneous, 2.5 mm long, oblong, concave, acuminate apex, tufted. Staminate flowers 2.2-2.5 mm long; castaneous sepals, 2.2 mm long, fused at base, oblong to obovate, concave, mucronate apex, tufted; cream corolla, 1.2 mm long, infundibular, mucronate lobes, glabrous; stamens 3, castaneous anthers, dorsifixed; 3 nectariferous carpellodes. Pistillate flowers 2.2-2.7 mm long; castaneous sepals, 2.2 mm long, free, obovate, concave, rounded apex, tufted; cream petals, 2 mm long, free, obovate, concave, obtuse apex, adaxially pilose; gynoecium 1.5 mm long, 3 nectariferous branches, stigmatic branches bifid.

Examined material: caminho para Belchior, 12.XII.1986, M.B. Horta et al. 11 (BHCB); Pico da Canjerana, 20°08’06”S, 43°30’49”W, 1,904 m, 27.VIII.2008, C.T. Oliveira & L.L. Giacomin 83 (BHCB); Pico do Inficcionado, 5.IX.1999, M.F. Vasconcelos (BHCB 48910); Serra do Caraça, 19.VIII.2000, R.C. Mota 881 (BHCB); 11.X.1986, J.A. Paula 42 (BHCB, MBM); 11.IX.1990, J.R. Stehmann et al. (BHCB 18962); 26.IX.1998, M.F. Vasconcelos & P.N.M. Oliveira (BHCB 44046); trilha para Bocaina, 10.I.2000, M.F. Vasconcelos (BHCB 52683); trilha para Cachoeira da Bocaina, depois da Pedra da Paciência, 20°04’20”S, 43°16’49”W, 1,272 m, 4.XI.2020, A. Soldevila & P. Vanucci 316 (OUPR); trilha para Tabuões lado direito, 20°02’43”S, 43°18’07”W, 1,181 m, 23.XI.2020, A. Soldevila & P. Vanucci 325 (OUPR).

Paepalanthus planifolius is widely distributed in Brazil, occurring in Distrito Federal, Goiás, and in all states of Southeast and South Brazil (Andrino et al. 2024). It is widely distributed in RPPN Santuário do Caraça, being one of the most abundant species of the family. It was found in the following localities: Campos de Fora, Casa do Pesquisador, the trail to Tabuões, the trail to Bocaina, the trail to Cascatinha, the trail to Banho do Belchior, Pico do Inficcionado, Pico da Canjerana, Cascudo, and Mirante da Piscina. It grows in quartzitic Campo Rupestre, over moist, sandy to organic soil, among shrubby to herbaceous vegetation, sometimes in marshes, and most frequently at stream edges. Individuals are perennial, and specimens with flowers were recorded from August to January, with the flowering peak in the rainy season.

Paepalanthus planifolius is mainly recognized by its completely fused scapes, the stem restricted to the leaf rosette, which is robust and forms tanks, in addition to the truncate spathes. It differs from P. conduplicatus by the flat leaves (vs. conduplicate), involucral bracts abaxially densely pilose and not ciliate (vs. glabrous and ciliate), and castaneous anthers (vs. cream).

25. Paepalanthus polyanthus (Bong.) Kunth, Enum. Pl. 3: 516. 1841.Fig. 5l

Plants 60-150 cm tall. Rhizome absent. Stem restricted to the leaf rosette when sterile, non-branched, elongating into an inflorescence (fertile) axis. Rosette leaves 13.5-21 × 0.5-3.3 cm, lanceolate, acute apex, pilose to glabrescent on both surfaces, ciliate. Inflorescences bearing a fertile axis and lateral branches (paraclades) 7-28.5 cm long, these alternate spiraled, paracladial bracts 1.5-3.5 cm long, lanceolate, acute apex, pilose to glabrescent on both surfaces, ciliate. Spathes 0.7-1.5 cm long, lax, abaxially pilose to glabrescent, oblique opening. Scapes 2-3.5 cm long, free, 80-150 in spherical umbels, villous when young. Capitula 3 mm wide. Involucral bracts castaneous, 2-3 mm long, not surpassing the flower disc, obovate, obtuse apex, abaxially pilose to glabrescent, ciliate. Floral bracts castaneous, 1.8-2 mm long, obovate-elliptic, concave, obtuse apex, tufted. Staminate flowers 2 mm long; light castaneous sepals, 1.6-1.8 mm long, free, obovate, concave, obtuse to rounded apex, tufted; cream corolla, 1.6-1.8 mm long, infundibular, glabrous; stamens 3, anthers dorsifixed; 3 nectariferous carpellodes. Pistillate flowers 2.2 mm long; light castaneous sepals, 1.8-2.2 mm long, free, obovate, obtuse apex, tufted; cream petals, 1.8-2 mm long, free, elliptic, acute apex, densely pilose from middle to apex; gynoecium 1.8-2.2 mm long, 3 nectariferous branches, stigmatic branches simple.

Examined material: Pico do Inficcionado, 10.I.2000, M.F. Vasconcelos (BHCB 52571); subida para o Pico do Inficcionado, 20°08’05”S, 43°27’49”W, 1,829 m, 9.III.2008, L. Echternacht et al. 1619 (BHCB); subida para Pico da Canjerana, 20°08’04”S, 43°30’46”W, 1,859 m, 27.VIII.2008, C.T. Oliveira & L.L. Giacomin 101 (BHCB).

Paepalanthus polyanthus is widely distributed in Brazil (= Actinocephalus polyanthus (Bong.) Sano; Costa & Sano 2024). It was found in the following localities in RPPN Santuário do Caraça: ascent to Pico do Inficcionado, Pico do Inficcionado, and ascent to Pico da Canjerana, above 1,800 m elevation. Abundant populations occur in quartzitic Campo Rupestre, mainly in open sandy areas. Individuals are monocarpic, flowering between November and March.

Paepalanthus polyanthus is recognized by compound inflorescence, with an elongated fertile axis bearing spiraled paraclades along the axis and with scapes in spherical umbels; scapes are villous when young, and the rosette dries out, but leaves are persistent. The most similar species is A. bongardii, which has verticillate paraclades and pilose scapes when young.

26. Paepalanthus pubescens Körn., Fl. bras. 3(1): 384. 1863.Fig. 6a-b

Plants 15-30 cm tall, forming clumps. Rhizome present. Aerial stem short, restricted to the leaf rosette, branched. Rosette leaves 1.5-6.5 × 0.15-0.3 cm, linear, acute to acuminate apex, pilose to glabrescent on both surfaces, ciliate. Inflorescences with scapes emerging from the stem at the leaf axil. Spathes 3.5-6 cm long, striate, pilose abaxially, oblique opening. Scapes 10-25 cm long, free, 1-6 per rosette, pilose to glabrescent with patent or adpressed hairs. Capitula 8-11 mm wide. Involucral bracts castaneous, 2-3.5 mm long, not surpassing the flower disc, 4-5 series, oblong to ovate, obtuse to rounded apex, abaxially densely pilose at apex, ciliate. Floral bracts light castaneous, 3 mm long, narrow-linear, flat, acute apex, abaxially pilose, ciliate. Staminate flowers 3-4 mm long; light castaneous to brown sepals, 2.2 mm long, fused at base, obovate, concave, obtuse apex, glabrous; cream corolla, 3 mm long, infundibular, round lobes, glabrous; stamens 3, dorsifixed anthers; 3 nectariferous carpellodes. Pistillate flowers 3 mm long; light castaneous sepals, 2.7 mm long, free, oblanceolate, concave, acuminate apex, glabrous; light castaneous petals, 2.5 mm long, free, linear, concave, acute apex, tufted; gynoecium 3 mm long, 3 nectariferous branches, stigmatic branches bifid.

Examined material: trilha para Pico da Conceição, 20°06’37”S, 43°31’17”W, 1,501 m, 17.VII.2021, L. Echternacht & P. Mendes 2894 (OUPR).

Paepalanthus pubescens is distributed in Minas Gerais and São Paulo (Andrino et al. 2024), with disjunct populations in the Espinhaço Range and Quadrilátero Ferrífero . One population only was found in RPPN Santuário do Caraça, on the trail to Pico da Conceição. It occurs in quartzitic Campo Rupestre, over sandy soil, among herbaceous vegetation. Individuals were collected in July with mature flowers.

Paepalanthus pubescens is recognized by the pilose to glabrescent leaves on both surfaces, ciliate, spathes pilose with patent and adpressed hairs, pilose to glabrescent scapes with adpressed hairs at the distal half, and by the castaneous involucral bracts, ciliate and densely pilose abaxially at the distal half.

27. Paepalanthus scleranthus Ruhland, Pflanzenr. 4(30): 199. 1903.

Plants 2.5-5 cm tall. Rhizome absent. Aerial stem elongate, 2.5-4 cm long, erect, sometimes branched. Rosette absent, leaves spiraled 5-10 × 0.5-0.8 mm, linear, striate, acute to acuminate apex, sparsely pilose to glabrescent on both surfaces, not ciliate. Inflorescences congest at the stem apex. Spathes absent. Scapes absent. Capitula 1.5-3 mm wide. Involucral bracts green, 2-6 mm long, surpassing the flower disc 1-4 mm long, 1 series, linear, acute apex, glabrous, not ciliate. Flowers dimerous. Floral bracts cream at base and middle, blackish at apex, 1.5 mm long, obovate, concave, long acuminate apex, glabrous. Staminate flowers 1.2 mm long; cream sepals, 1 mm long, fused at base, oblong to obovate, flat, round to truncate apex, tufted; cream corolla, 0.5 mm long, tubular, obtuse lobes, glabrous; stamens 2, anthers dorsifixed; 3 nectariferous carpellodes. Pistillate flowers 1.7 mm long; cream sepals, blackish at apex, 1.5 mm long, free, elliptic, concave, obtuse apex, distally ciliate; cream sepals, blackish at the apex, 1.5 mm long, free, oblong, flat, rounded apex, distally ciliate; gynoecium 1 mm long, 3 nectariferous branches, stigmatic branches simple.

Local records: Minas Geraës, auf Felsen der Serra de Caraca, III.1892, E.H.G. Ule 2717 (B, HBG, NY).

Examined material: Pico do Sol, 20°06’34”S, 43°27’06”W, 1,828 m, 24.IV.2009, C.T. Oliveira et al. 399 (BHCB).

Paepalanthus scleranthus is distributed in Bahia and Minas Gerais (Andrino et al. 2024), where it occurs in the Espinhaço Range and Quadrilátero Ferrífero. We found two records from RPPN Santuário do Caraça, one belonging to the type collection (Ule 2717), with no precise locality, and another more recent record from Pico do Sol. As a very small species, it is frequently overlooked. It grows in quartzitic Campo Rupestre, over sandy to organic, humid soil. Specimens were recorded with flowers in March and April.

Paepalanthus scleranthus is recognized mainly as tiny plants (2.5-5 cm tall), lacking spathes and scapes, with elongated and branched aerial stems, spiraled leaves, green involucral bracts, and dimerous flowers.

Figure 6 -
a-n. Species of Eriocaulaceae - a-b. Paepalanthus pubescens - a. involucral bracts castaneous; b. detail of the leaf rosette; c-e. P. suffruticans - c. stem elongated, leaves spiraled showing morphological variation in the stem length; d. blackish involucral bracts; e. habit; f-h. P. vellozioides - f. habit, castaneous leaves at base; g. light castaneous involucral bracts, slightly surpassing the flower disc, long acuminate; h. detail of the rosette and fused scapes, splitting at the top; i-k. Syngonanthus anthemiflorus - i. detail of synflorescence, the primary axis with whorled bracts at the apex and one spathe and scape; j. habit; k. involucral bracts surpassing the flower disc; l. S. caulescens - stem short to elongate, branched or not, leaves spiraled scapes in umbels; m-n. S. gracilis - m. recurved rosette leaves; n. delicate capitula. Images: a-b,i-k,m,n. Livia Echternacht; c-h,l. Ambar Soldevila.

Paepalanthus scleranthus was described by Ruhland (1903) based on type material from RPPN Santuário do Caraça, collected by Ule (2717, B, HBG, NY, R), also on material type from Serra de Ouro Preto (Ule 2714, B, HBG, R), from Serra das Camarinhas (Schwacke 11987, B), and Diamantina (Glaziou 19990, B, F, G, P). Andrade et al. (2022) selected Glaziou 19990 at B as lectotype; consequently, the specimen from Caraça is considered a paralectotype.

28. Paepalanthus suffruticans Ruhland, Pflanzenr. 4(30): 138. 1903. Typus: BRAZIL. MINAS GERAIS: “Minas Geraes: Caraça, au Morro de Carapuça”, 11.VI.1884, A.F.M. Glaziou 15529 (lectotype B!, designated by Moldenke (1976: 115); isolectotypes LE!, B!, C!, P “photo !”, LL!).

= Paepalanthus chloroblepharusRuhland, Pflanzenr. 4(30): 139. 1903. Typus: BRAZIL. MINAS GERAIS: “Minas Gerais; in Sümpfen bei Caraça”, III.1892, E.H.G. Ule s.n. (lectotype B!, designated by Moldenke (1973b: 198); isolectotypes LL!, R!), syn.nov.

= Paepalanthus langsdorffii var. caracensisMoldenke, Phytologia 25: 430. 1973. Typus: BRAZIL. MINAS GERAIS: “Sandstone summit of Serra da Caraça”, 25.I.1971, H.S. Irwin 29110 (holotype LL!, isotypes C!, F!, NY!, P “photo !”, US “photo !”), syn.nov.

= P. suffruticans var. angustifolius Silveira exMoldenke, Phytologia 35: 115. 1976. Typus: BRAZIL: MINAS GERAIS: “In campis ad Verruguinha, Serra de Caraça, Minas”, IV.1905, A. Silveira 331 (holotype R!), syn.nov.Fig. 6c-e

Perennial plants 30-100 cm tall. Rhizome absent. Aerial stems usually elongate, sometimes short in young individuals, 1.5-65 cm long, erect to decumbent, branched or not. Rosette absent, leaves spiraled 6.5-18 cm × 1.8-6 mm, linear to lanceolate, acute to acuminate apex, pilose to glabrescent or glabrous on both surfaces, ciliate or not. Inflorescences with scapes emerging from the stem at the leaf axil. Spathes 5.5-11.5 cm long, densely to sparsely pilose, glabrescent or glabrous, oblique opening. Scapes 16-60 cm long, free, 3-20 per stem branch, glabrous. Capitula 6-11 mm wide. Involucral bracts blackish, 2-3 mm long, not surpassing the flower disc, 6-8 series, oblong, wide orbiculate or ovate, round, obtuse or acuminate apex, abaxially pilose with adpressed hairs, ciliate. Floral bracts cream at the base and middle, blackish at the apex, 1.8-2.2 mm long, linear to spathulate, flat, acuminate to rounded apex, densely pilose abaxially, tufted. Staminate flowers 2.5-3 mm long; light castaneous sepals, 2 mm long, fused at base, obovate, concave, rounded apex, densely pilose on both surfaces, tufted; reddish castaneous corollas, 1.8-2 mm long, tubular, obtuse lobes, glabrous; stamens 3, dorsifixed anthers; 3 nectariferous carpellodes. Pistillate flowers 2.7-3 mm long; castaneous to light castaneous sepals, 1.7-2 mm long, free, linear to obovate, concave, obtuse to rounded apex, abaxially densely pilose, tufted; cream to castaneous petals, 1.8-2 mm long, free, linear, concave, obtuse apex, adaxially pilose, tufted; gynoecium 2-2.2 mm long, 3 nectariferous branches, nectariferous branches bifid.

Examined material: Pico da Carapuça, 20°05’27”S, 43°28’17”W, 1,848 m, 17.II.2009, C.T. Oliveira & A.J. Arruda 329 (BHCB, RB); Pico do Sol, 20°06’38”S, 43°26’52”W, 1,885 m, 29.VI.2009, C.T. Oliveira et al. 513 (BHCB, RB); Serra do Caraça, VI.1907, L.B. Damazio (OUPR 12633); trilha para Capelinha e Gruta de Lourdes, 9.I.1982, N. Hensold et al. (BHCB 70446); 20°03’14”S, 43°17’06”W, 1,413 m, 3.XI.2020, A. Soldevila & P. Vanucci 305 (OUPR); trilha para a Gruta de Lourdes, próximo a gruta, 20°03’13”S, 43°17’01”W, 1,434 m, 3.XI.2020, A. Soldevila & P. Vanucci 309 (OUPR).

Paepalanthus suffruticans is micro-endemic to the RPPN Santuário do Caraça. It was recorded in the following localities: the trail to Capelinha, the trail to Gruta de Lourdes, Pico da Carapuça, Pico do Sol, and Pico da Verruguinha. The population observed in the trail to Capelinha and Gruta de Lourdes has approximately 10-20 individuals growing in shaded and humid slopes, along the edge of riparian forests, among shrubs and trees, in moist sandy soils. It was also recorded in quartzitic Campo Rupestre at altitudes above 1,800 m, in Pico da Carapuça and Pico do Sol. Individuals are perennial, and specimens with flowers were recorded throughout the year.

Paepalanthus suffruticans presents a morphological variation in the stem length, leaf indumentum, and morphology of the involucral bracts. It usually has elongated aerial stem, with spiraled leaves, glabrous to pilose on both surfaces, ciliate or not, and involucral bracts are blackish, oblong to wide orbiculate. Its habitat in shaded understory is unusual for Eriocaulaceae in the RPPN Santuário do Caraça, which occurs mainly at open, sunny sites.

Paepalanthus suffruticans and Paepalanthus chloroblepharus have both type materials from RPPN Santuário do Caraça and were described in the same work by Ruhland (1903). Analyzing the protologue and later descriptions of the taxa, and the type specimens, we observed that types differ in the following characters: in P. chloroblepharus stem is briefly elongated (vs. quite elongated), leaves are glabrous or puberulous and ciliate (vs. glabrous or hirsute and ciliate with long hairs), involucral bracts are ovate to obovate, almost orbiculate (vs. wide orbiculate). During fieldwork, the observed within populations a morphological variation in these features, with individuals presenting short to elongated stems, the leaf indumentum varying from glabrous to pilose, and involucral bracts from oblong to wide orbiculate (even in the same capitulum). We also noticed a variation in vegetative organs in individuals growing above 1,800 m elevation, which have thinner leaves and denser indumentum, evidencing the plasticity of the species. Therefore, the characteristics raised by Ruhland (1903) to differentiate the two taxa are considered here as part of a gradient of the morphological variation of the species. Considering that both names were published together (Ruhland 1903) and have no priority over each other, we chose P. suffruticans as the accepted name and considered P. chloroblepharus synonymous with it.

Paepalanthus langsdorffii var. caracensis has also type material from RPPN Santuário do Caraça (Moldenke 1973). Analyzing the type collection, we consider it synonymous with P. suffruticans. Hensold (1988) revised P. subg. Xeractis, which includes P. langsdorffii, and excluded this variety from the subgenus.

The name Paepalanthus suffruticans var. angustifolia was cited by Silveira in Floralia Montium in the list of specimens of the “Herbarii Silveira” (Silveira 1928, page 414), referring to specimen Silveira 331 collected in Caraça in 1906. In the same work, Silveira (1928, page 106) highlighted that the specimen collected at Morro da Verruguinha (Silveira 331, R) has narrower leaves than the specimen collected at Morro da Carapuça (1.5-3 vs 5-10 mm wide). We assume that Silveira (1928) did not intend to publish a new name, and to that moment, it should be treated as a name in the herbarium. However, Moldenke (1976, page 115) published the name P. suffruticans var. angustifolius based on Silveira (1928). Moldenke validly publishes the name and makes reference to the work that contains a diagnosis. Therefore it can not be considered as a nomen nudum. We consider it following the International Code of Nomenclature for Algae, Fungi and Plants (Turland et al. 2018). Nevertheless, this taxon is not included in Flora and Funga do Brasil (Andrino et al. 2020) nor in the International Plan Names Index (IPNI, continuously updated). As previously described, P. suffruticans has a wide vegetative morphological variation, and individuals with different leaves widths can be found even in the same subpopulation. We do not recognize infraspecific taxa, proposing here to synonymize P. suffruticans var. angustifolius.

29. Paepalanthus vaginatus Körn., Fl. bras. 3(1): 313. 1863.

Plants 30-70 cm tall. Rhizome absent. Stem restricted to the leaf rosette, non-branched. Rosette leaves 3-6 cm × 1-4 mm, linear, acute apex, pilose to glabrescent with adpressed hairs on both surfaces, ciliate or not. Inflorescences with scapes emerging from the stem at the leaf axil. Spathes 6-13 cm long, sparsely pilose to glabrescent, or glabrous, oblique opening. Scapes 30-70 cm long, free, 1-6 per rosette, sparsely pilose to glabrescent, or glabrous. Capitula 5-10 mm wide. Involucral bracts dark brown, 1.5-2.5 mm long, not surpassing the flower disc, 5-6 series, ovate, obtuse to acute apex, abaxially densely pilose with appressed hairs, ciliate. Flowers dimerous. Floral bracts castaneous, 2-2.5 mm long, oblong, concave, acuminate apex, tufted. Staminate flowers 2-2.5 mm long; castaneous sepals, 2-2.2 mm long, fused at base and middle, oblong, concave, obtuse apex, abaxially pilose, tufted; cream corolla, 1.5 mm long, tubular, glabrous; stamens 2, dorsifixed anthers; 2 nectariferous carpellodes. Pistillate flowers 2-2.5 mm long; castaneous sepals, fused at base, oblong, concave, acute apex, abaxially pilose, tufted; cream petals, 1.5-2 mm long, free, linear, flat, rounded apex, tufted, gynoecium 1.5 mm long, 2 nectariferous branches, stigmatic branches bifid.

Local records: S. da Carassa, 1.I.1825, L. Riedel 1462 (B, LE, UPS).

Examined material: Mariana, Bento Rodrigues, estrada para Santa Rita do Durão, 26.III.2008, M.C.T.B. Messias et al. 1861 (OUPR); 29.III.2008, M.C.T.B. Messias et al. 1829 (OUPR).

Paepalanthus vaginatus is endemic to Minas Gerais (Andrino et al. 2024) and widely distributed in the Espinhaço Range, also occurring in the Quadrilátero Ferrífero and the Serra da Canastra. In the RPPN Santuário do Caraça, there is one record only, which is the type material (Riedel 1462). It grows in quartzitic Campo Rupestre, among shrubby to herbaceous vegetation, in humid sandy soils. The fertile specimen was recorded in January.

Description of Paepalanthus vaginatus was based on the type material and materials from the municipality of Mariana, from the closest available sites. P. vaginatus is mainly recognized by the spathes much longer than the leaves, the short stem, restricted to leaf rosette, long scapes, involucral bracts dark brown, and dimerous flowers.

30. Paepalanthus vellozioides Körn., Fl. bras. 3(1): 401-402. 1863.Fig. 6f-h

Perennial plants 20-35 cm tall. Rhizome present. Stem restricted to the leaf rosette, non-branched. Rosette leaves 13-32 × 0.4-1 cm, lanceolate, brown at base adaxially, acute to acuminate apex, glabrous on both surfaces, distally ciliate. Inflorescences with scapes emerging from the stem at the leaf axil. Spathes 5.5-9 cm long, glabrous, truncate opening. Scapes 19-32 cm long, fused, splitting at top, 6-15 sets per rosette, pilose to sparsely pilose. Capitula 7-9 mm wide. Involucral bracts light castaneous to golden, 3-5 mm long, surpassing the flower disc 0.5-1.5 mm long, 4-5 series, ovate, apex long acuminate, abaxially glabrous, ciliate. Floral bracts light brown at base and middle, dark brown to blackish at apex 3.5-4 mm long, elliptic to oblong, concave, acute apex, tufted, ciliate. Staminate flowers 3.5-4.5 mm long; light castaneous sepals at base and middle, brown to blackish at apex, 2.5-3 mm long, fused at base and middle, obovate, concave, obtuse to rounded apex, tufted; cream corolla, 2-2.5 mm long, infundibular, long acuminate lobes, glabrous; stamens 3, dorsifixed anthers; 3 nectariferous carpellodes. Pistillate flowers 3-3.2 mm long; castaneous sepals, 2.5-2.8 mm long, free, oblong, concave, obtuse apex, tufted; cream petals, 2-2.5 mm long, free, linear, concave, rounded apex, adaxially densely pilose, tufted; gynoecium 1.5 mm long, 3 nectariferous branches, stigmatic branches bifid.

Examined material: caminho para Gruta do Padre Caio, 23.V.1987, J.A. Trindade 37 (BHCB); estrada entre o Seminário e a cancela de entrada do parque, 11.I.1996, V.C. Souza et al. 10030 (BHCB-ESA); estrada para Piscina, 20°03’16”S, 43°17’43”W, 1,307 m, 23.XI.2020, A. Soldevila & P. Vanucci 326 (OUPR); Serra do Caraça, 31.VIII.1973, J. Badini (OUPR 12652); Tabuões lado direito, segundo sumidouro subindo o rio, 20°02’44”S, 43°18’04”W, 1,188 m, 23.XI.2020, A. Soldevila & P. Vanucci 323 (OUPR); trilha para Cascatona, 20°05’02”S, 43°30’07”W, 1,130 m, 6.V.2008, L. Echternacht & C. Chaussidon 1638 (BHCB).

Paepalanthus vellozioides is endemic to Minas Gerais (Andrino et al. 2024), occurring in the Espinhaço Range and Quadrilátero Ferrífero, forming abundant populations. It has been recorded in the RPPN Santuário do Caraça in the following localities: Tabuões, along the Park road, on the road to Piscina, the trail to Cascatona, and the trail to Padre Caio cave. It grows in quartzitic Campo Rupestre, over gravelly to sandy soil, among shrubby to herbaceous vegetation. Individuals are perennial, and specimens with flowers were recorded in January, February, May, June, August, and November. In August, individuals have old flowers, and in November, young flowers, indicating that the flowering peak is in the rainy season.

Paepalanthus vellozioides is mainly recognized by fused scapes, distally splitting, and the golden involucral bracts with long acuminate apex. Moreover, the stem is restricted to the rosette, leaves are glabrous and brown at the adaxial base, and the spathe opening is truncated.

Paepalanthus vellozioides was described by Koernicke (1863) based on type material type collected by Martius (874, F, NY), without specific locality, and by materials from RPPN Santuário do Caraça collected by Sellow (s.n., B) and Clausen (96, B).

31. Syngonanthus anthemiflorus (Bong.) Ruhland, Pflanzenr. 4(30): 258. 1903.Fig. 6i-k

Plants 22-40 cm tall. Rhizome present, 3-15 cm long, densely branched. Aerial stem restricted to the rosette. Rosette leaves 4-11 × 0.5-0.9 mm, linear, recurved, acute apex, abaxially pilose, adaxially sparsely pilose to glabrescent, not ciliate. Inflorescences with capes emerging from a reproductive axis. Axis 2-2.5 cm long, 1 per rosette, pilose; bracts restricted to the apex, 1.5-5 mm long, lanceolate, acuminate apex, pilose on both surfaces, ciliate. Spathes 2.7-3.2 cm long, abaxially pilose to glabrescent, oblique opening. Scapes 18-21 cm long, free, 1 per reproductive axis, densely pilose. Capitula 6-10 mm wide. Involucral bracts 4-6 series, the external series light brown, 1.5-3 mm long, ovate to oblong, acuminate apex, abaxially glabrescent, internal series castaneous at base and cream from middle to the top, 4-6 mm long, spathulate, acute apex, abaxially pilose in the middle, ciliate or not, surpassing the flower disc 1.5-2.5 mm long. Floral bracts cream, 3-3.2 mm long, oblanceolate, acuminate apex, abaxially pilose. Staminate flowers 3-3.2 mm long; cream sepals, 2.5-2.7 mm long, fused at base, elliptic, concave, acute apex, pilose to glabrescent on both surfaces; cream corolla, 2-2.2 mm long, infundibular, acuminate lobes, glabrous; stamens 3, dorsifixed anthers; 3 nectariferous carpellodes. Pistillate flowers 2.9-3 mm long; cream sepals, 2.2-3 mm long, free, elliptic, concave, acute apex, abaxially pilose; cream petals, 1.5-2 mm long, free, obovate, acuminate apex, abaxially glabrous, adaxially pilose; gynoecium 1.5 mm long, 3 nectariferous branches, stigmatic branches simple.

Examined material: Campos de Fora, 20°06’29”S, 43°31’21”W, 1,503 m, 17.VII.2021, L. Echternacht & P. Mendes 2895 (OUPR); Serra do Caraça, 19.VIII.2000, R.C. Mota 885 (BHCB).

Syngonanthus anthemiflorus is endemic to the Espinhaço Range in Minas Gerais, distributed from the municipalities of Ouro Preto to Grão Mogol (Echternacht 2012). In RPPN Santuário do Caraça, it was found in the locality of the Campos de Fora only, growing in quartzitic Campo Rupestre, among herbaceous vegetation, over sandy soil. Specimens with flowers were recorded in August and July.

Syngonanthus anthemiflorus is recognized by one scape emerging from a repoductive axis, one per rosette, with a whorl of bracts at the axis apex; capitula are very characteristic, with castaneous involucral bracts in the external series, cream and spathulate in the internal series, long surpassing the flower disc.

32. Syngonanthus caulescens (Poir.) Ruhland, Pflanzenr. 4(30): 267, f. 38. 1903.Fig. 6l

Plants 4-23 cm tall. Rhizome absent. Aerial stem elongated, 0.5-9 cm long, occasionally branching at the base, with white wooly indumentum. Rosette absent, leaves spiraled 0.5-2.3 × 0.1-0.3 cm, lanceolate, acute apex, pilose to glabrescent on both surfaces with adpressed hairs, not ciliate. Inflorescences with scapes emerging at the stem apex. Spathes 1.5-4 cm long, abaxially pilose with adpressed hairs, oblique opening. Scapes in umbel at the stem apex, 2.5-21 cm long, free, 5-40 per umbel, pilose to glabrescent with adpressed hairs. Capitula 1-4 mm wide. Involucral bracts cream, 1.5-3 mm long, surpassing the flower disc 0.5-1 mm long, ovate, 3-4 series, acute apex, glabrous abaxially, not ciliate. Floral bracts absent. Staminate flowers 1.7-2 mm long; cream sepals, 1.7-1.9 mm long, free, elliptic, cymbiform, acuminate apex, glabrous; cream corolla, 1-1.2 mm long, fleshy, urceolate, acute lobes, glabrous; stamens 3, dorsifixed anthers; 3 nectariferous carpellodes. Pistillate flowers 1.7-1.9 mm long; cream sepals, 1.1-1.2 mm long, free, elliptic, cymbiform, acuminate apex, glabrous; cream petals, 1.2-1.5 mm long, free at base, fused distaly with short acute lobes, abaxially pilose; gynoecium 1.2-1.4 mm long, 3 nectariferous branches, stigmatic branches simple.

Examined material: Serra do Caraça, 15.IV.1933, H.L.M. Barreto 2584 (BHCB); 14.XII.2000, R.C. Mota 1059 (BHCB); J.R. Stehmann et al. (BHCB 28397); trilha da Cascatinha, 14.XII.2000, J. Ordones et al. 558 (BHZB, OUPR); 20°03’44”S, 43°16’59”W, 1,245 m, 4.XI.2020, A. Soldevila & P. Vanucci 311 (OUPR).

Syngonanthus caulescens is the most widespread species in the genus and is widely distributed in South America (Watanabe 2015). It was found in the following localities in RPPN Santuário do Caraça: the trail to Cascatinha and Pico do Inficcionado. It occurs in moist sandy and organic soils, marshes or river banks. Specimens with flowers were recorded in January, April, June, September, November, and December.

Syngonanthus caulescens can be recognized by its elongated stem with a white wooly indumentum, and spiraled leaves, by cream and acute involucral bracts, slightly surpassing the flower disc 0.5-1 mm long, floral bracts absent, and by fleshy (also described sometimes as spongy) petals in staminate flowers. In some individuals, the aerial stem is very short, and leaves seem to form rosettes. This plant is always in humid environments.

33. Syngonanthus gracilis (Bong.) Ruhland, Pflanzenr. 4(30): 249. 1903.Fig. 6m-n

Perennial plants, 14-21 cm tall. Rhizome present or not. Aerial stem short, 0.5-1 mm long, restricted to the leaf rosette, branched or not. Rosette leaves 5-12 × 0.2-1 mm, linear, recurved, acute apex, abaxially pilose, adaxially densely pilose, not ciliate. Inflorescences with scapes emerging from the stem at the center of the rosette. Spathes 1-2.6 mm long, abaxially pilose to glabrescent with capitate patent hairs or simple filamentous adpressed hairs, oblique opening. Scapes 5.5-21 cm long, free, 1-6 per rosette, pilose to glabrescent with capitate patent hairs or simple adpressed hairs. Capitula 3-5 mm wide. Involucral bracts straw-colored to golden, 1-2.5 mm long, not surpassing the flower disc, 5-6 series, obtuse to rounded apex, abaxially glabrous, not ciliate, external series elliptic, internal series oblong. Floral bracts absent. Staminate flowers 1.5-2 mm long; cream sepals, 1.2-1.3 mm long, fused at base, elliptic, concave, acute apex, glabrous; cream corolla 0.7-1 mm long, infundibular, acute lobes, glabrous; stamens 3, dorsifixed anthers; 3 nectariferous carpellodes. Pistillate flowers 2.5-3 mm long; cream sepals, 1.5-2 mm long, free, elliptic, concave, acuminate apex, glabrous; petals cream, 1.2-1.5 mm long, free at base, fused at top, with short acuminate lobes, oblanceolate, glabrous; gynoecium 1.5-2 mm long, 3 nectariferous branches, stigmatic branches simple.

Examined material: Campos de Fora, 20°07’03”S, 43°31’08”W, 1,504 m, 17.VII.2021, L. Echternacht & P. Mendes 2890 (OUPR); 20°04’13”S, 43°18’39”W, 1,497 m, 24.XI.2020, A. Soldevila & P. Vanucci 331 (OUPR); Pico do Inficcionado, 9.I.2000, M.F. Vasconcelos (BHCB 52694); Serra do Caraça, 14.IV.1933, H.L.M. Barreto 2551 (BHCB).

Syngonanthus gracilis is distributed in the Domains of Caatinga, Cerrado, and Atlantic Forest, occurring in Bahia, Paraíba, Espiríto Santo, Minas Gerais, and Rio de Janeiro (Echternacht 2012; Echternacht & Watanabe 2024). In RPPN Santuário do Caraça, it was found in the Campos de Fora and Pico do Inficcionado. It grows in quartzitic Campo Rupestre, over sandy to humid organic soils, among open herbaceous vegetation. Specimens were recorded in April and July with mature flowers and in November and January with old capitula.

Syngonanthus gracilis is recognized by its delicate habit, aerial stem restricted to the leaf rosette, with small leaves (5-12 mm long), recurved, involucral bracts straw-colored to golden, and floral bracts absent. In some young individuals, we observed capitate patent hairs in spathes and scapes.

Acknowledgements

We are grateful to the Universidade Federal de Ouro Preto (UFOP), especially the Laboratório de Taxonomia and Herbarium OUPR, for laboratory structure and staff support. We also thank RPPN Santuário do Caraça staff, for field support, collecting permit and logistical provision; and the curators and staff of the visited herbaria (BHCB and OUPR). The master’s scholarship was funded by Coordenação de Aperfeiçoamento de Pessoal de Nível Superior-CAPES (CAPES 88887.185914/2019-00); part of the research was financed throughout the researcher’s own resources.

Data availability statement

In accordance with Open Science communication practices, the authors inform that all data are available within the manuscript.

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Edited by

Area Editor:

Dr. Marcelo Trovó

Publication Dates

• Publication in this collection
  22 July 2024
• Date of issue
  2024

History

• Received
  16 Feb 2023
• Accepted
  19 Jan 2024