Barnadesieae, Gochnatieae, Mutisieae and Nassauvieae (Asteraceae) in the Atlantic Forest of Caparaó National Park, Brazil

Abstract

Asteraceae is highly diverse in the Atlantic Forest, especially in montane environments. The early-diverging tribes within Asteraceae are of particular interest because of their great representation in Brazil and unique features within the family. Thus, a floristic inventory and systematic treatment of these basal grade in Caparaó National Park, an important remnant of Atlantic Forest that has been highlighted for its diversity and endemism, were undertaken. The study was conducted by examining specimens from representative collections of Caparaó and a field expedition. A total of 15 species were found, representing six genera and four tribes, with Mutisieae and Nassauvieae being the most representatives. Six of the species have restricted distributions, either to Brazil (3 spp.), the Atlantic Forest (2) or Caparaó mountain (Mutisia lutzii). Three new records were documented for the park, namely Chaptalia integerrima, Moquiniastrum polymorphum and Mutisia coccinea. Floristic inventories and systematic treatments can make great contributions to science by bringing a better understanding our biodiversity and providing relevant information about flora for the conservation of priority areas.

Key words
Compositae; collection; Mantiqueira Setentrional; Southeast Brazil; taxonomy

INTRODUCTION

Caparaó National Park (CNP; Parque Nacional do Caparaó) is an Atlantic Forest conservation unit that has already been highlighted for its richness and endemism (Moreira et al. 2020, Araújo et al. 2021). It is situated in the northern portion of Serra da Mantiqueira (Mantiqueira mountain range) straddling the border between the states of Minas Gerais and Espírito Santo (IBDF 1981). CNP holds remarkable plant diversity with roughly 1.791 species, 891 endemic species, representing 198 families (Moreira et al. 2020). Asteraceae is the second richest family in CNP with 139 species, of which 74 are endemic (Moreira et al. 2020).

Asteraceae is the largest family of vascular plants with 32.000 species distributed in 1.600–1.700 genera (Cheek et al. 2020), corresponding 10–12% of angiosperm (Mandel et al. 2017). Currently has a cosmopolitan distribution, but its greatest diversity remains in South America (Panero & Crozier 2016 , Munõz-Moreira et al. 2024).

One of the most intriguing complexes of the family are the early-diverging tribes, because they share several morphological characters and a complex evolutionary history (Funk et al. 2005, Katinas et al. 2008, Mandel et al. 2019). Barnadesieae, sister group to all other Asteraceae, a tribe of 91 species with axillary spines on branches, a feature not found elsewhere within the family (Jansen & Palmer 1987, Mandel et al. 2019). The following tribes diverged later, together Barnadesieae represent the basal grade, a non-monophyletic group with unique features within Asteraceae: Famatinantheae, Stifftieae, Hyalideae, Onoserideae, Nassauvieae, Mutisieae, Wunderlichieae, Gochnatieae, Hecastocleideae and Pertyeae (Susanna et al. 2020).

Brazil is a center of diversity of early-diverging tribes within Asteraceae, with eight tribes representing 73% of basal grade, 30 genera and 140 species; only Famatinantheae, Hecastocleideae and Pertyeae do not occur within the country (Hind 2011, Panero et al. 2014, Flora e Funga do Brasil 2023, SpeciesLink 2023). In Brazil, the Atlantic Forest is the richest phytogeographical domain for the early-diverging tribes holding 60 species, 42.8% of grade diversity (Sussana et al. 2020, Flora e Funga do Brasil 2023).

Moreira et al. (2020), in a first land plants checklist for CNP, registered 13 species of Asteraceae basal clades. This number is expected to be higher, because further exploration in CNP is needed. Although the region of Caparaó has high richness and endemism, there has been little research into its flora (Araújo et al. 2018, 2021, Moreira et al. 2020), and even less into Asteraceae (Lopes et al. 2021).

Whereas floristic inventories are fundamental to documenting the biodiversity, and thus promoting its conservation (Funk 2006, Cheek et al. 2020), more are needed for Caparaó, especially approaching Asteraceae, one of most richest families in the Atlantic Forest. Thus, this study aimed to perform a floristic inventory and systematic treatment for early-diverging tribes of Asteraceae in Caparaó National Park.

MATERIALS AND METHODS

Study area

Caparaó National Park (CNP; Parque Nacional do Caparaó) is a fully protected conservation unit with around 32.000 hectares of Atlantic Forest straddling the boundary between the states of Minas Gerais and Espírito Santo, Brazil (ICMBio 2023), around 41°51’– 41°43’W, 20°36’– 20°18’S (Fig. 1). The CNP is located in the northern portion of Serra da Mantiqueira (Mantiqueira mountain range; Araçuaí Orogen) (Heilbron et al. 2004, Pedrosa-Soares et al. 2007) one of the highest regions in Southeast Brazil (IBDF 1981). CNP is popularly known for “Pico da Bandeira”, the third highest peak in Brazil at 2.890 meters above sea level (m asl) (ICMBio 2015, IBGE 2023). Other peaks in CNP also are attractive, such as “Pico do Calçado” at 2.849 m asl and “Pico do Cristal” at 2.770 m asl (ICMBio 2023).

Figure 1
Study area. a. Caparaó National Park location; Brazil, Minas Gerais and Espírito Santo states. b. Boundaries of the Caparaó National Park and coordinates. c. Altitude in meters of the park. Coordinates in EPSG: 4674 – SIRGAS 2000. QGIS Development Team.

The climate of the region is classified as subtropical highland Cwb I, according Köeppen´s classification (IBDF 1981, Beck et al. 2018, Mindat 2023). The annual average temperature ranges 16°C–22°C, the elevation limits the daily maximum temperature in the summer, while the temperature in coldest months (June-August) can hit -4°C at highest peak (IBDF 1981, ICMBio 2015). Annual rainfall ranges 750–2.250 mm and relative air humidity usually above 70% (IBDF 1981, ICMBio 2015).

CNP experiences two seasons, a rainy-summer season from November to March and a dry-winter season from April to October which has a short rain declivity (IBDF 1981, ICMBio 2015). The vegetation in CNP is montane with Montane Semideciduous Forest (Fig. 2a, Montane Mixed Ombrophilous Forest (Fig. 2b), Upper Montane Mixed Ombrophilous Forest (700–1.780m), transitional vegetation between forest and High-altitude grassland, referred to here as “Candeal” (Fig. 2c) due the abundance of Eremanthus erythropappus (DC) MacLeish (1.800–2.000m), and High-altitude grassland (Fig. 2d-f). The high-altitude grassland can be “scrub” when there are scattered herbaceous and subshrub/shrub plants ≥60cm in height; Chusquea sp. grassland (Fig. 2e); or “clean” grassland characterized by rock outcrops (Fig. 2f) with few scattered herbaceous and small subshrub/shrub plants <60cm in height (2.000–2.800m) (IBGE 2012, ICMBio 2015, Moreira et al. 2020, information also from our expedition).

Figure 2
Phytophysiognomies of the Atlantic Forest in Caparaó National Park. a. Montane Semideciduous Seasonal Forest. b. Montane Ombrophilous Forest. c. Transitional vegetation between Montane Forest and High-altitude grassland, called here by “Candeal”. d. High-altitude grassland with scattered shrubs (scrub). e. Chusquea sp. grassland. f. High-altitude grassland. (Photos: P Renon).

The region of Caparaó has intensive coffee monoculture and the adjacent areas of CNP are highly fragmented with permitted anthropic activities (Silva et al. 2021). Another important degradation factor in the CNP is the tourist visitation, which can pressure population of species endangered and with restricted distribution.

Floristic inventory

Collections from CNP were studied at the main herbaria for the region: BHCB, ESA, HUFU, RB, HUEMG and via loans from CAP, GFJP, VIES, UEC (acronyms following Thiers 2023).

A field expedition was carried out for five days of collecting at the beginning of rainy season. The sampled specimens were georeferenced and data regarding habit, height, flower and involucre color, vegetation and altitude were recorded. The specimens were subsequently herborized according to traditional methodology (Rotta et al. 2008), with the support of HUEMG. The specimens were incorporated into the HUFU Herbarium and duplicates were sent to reference herbaria (HUEMG and VIES).

Distribution, phenology and conservation

Distribution data were checked through “Flora e Funga do Brasil” (http://reflora.jbrj.gov.br/), POWO (https://powo.science.kew.org/) and SpeciesLink (https://specieslink.net/). Phenological data were extracted from the specimen’s labels. Conservation status was checked by consulting the online databases CNCFlora (http://www.cncflora.jbrj.gov.br/) and IUCN Red List (https://www.iucnredlist.org/).

Systematic Treatment

Taxa were identified using specific references for basal clades of Asteraceae, consultation of descriptions, identification keys (Katinas et al. 2008, Monge et al. 2017, Pasini 2017, Saavedra & Roque 2017), monographs for genera (Katinas et al. 1996, Saavedra et al. 2018) and regional Floras (Monge & Semir 2012).

Comparisons with specimens of the HUFU herbarium, records from SpeciesLink (https://specieslink.net/) and Virtual Herbarium Reflora (http://reflora.jbrj.gov.br/) were also used during identification, and the type specimens were reviewed on Jstor Plant Sciences (https://plants.jstor.org/).

The species treatments herein included identification keys, descriptions, examined material, distribution, phenology, conservation data, and taxonomic comments. The descriptions were systematized within the genera, as those species belonging to different genus were not closely standardized. The descriptions and material examined sections were assembled using the MonographaR package from R software (The R Foundation for Statistical Computing 2021, Reginato 2016).

Taxonomic comments and identification keys covered all taxa within each genus and tribe to facilitate the recognition of new records. Thus, the taxonomic comments for tribes and genera, as well as the identification keys, relied on relevant literature, such as Funk et al. (2009) and Katinas et al. (2008), as well as original descriptions of species and genera. Images of specimens available on SpeciesLink and “Flora e Funga do Brasil” were also consulted.

Nomenclature follows IPNI (https://www.ipni.org/) and Tropicos (https://www.tropicos.org/). Terminology followed Ellis et al. (2009), Hickey (1973) and Radford (1986) for vegetative structures, and Funk et al. (2009) for reproductive structures.

RESULTS

Floristic inventory

A total of 15 species and six genera were recorded, representing Barnadesieae, Gochnatieae, Mutisieae and Nassauvieae (Supplementary Material - Table SI). Mutisieae (7 species, 2 genera) and Nassauvieae (5 species, 2 genera) were the richest tribes, while Chaptalia Vent. (4 spp.) and Trixis P. Browne (each with 4 spp.) were the richest genera.

Six species have narrower distributions. Chaptalia denticulata (Baker) Zardini (Fig. 4a-e), Dasyphyllum flagellare (Casar.) Cabrera (Fig. 3a-d) and Trixis glaziovii Baker are restricted to Brazil, Holocheilus pinnatifidus (Less.) Cabrera (Fig. 3e-h) and Trixis verbascifolia (Gardner) Blake (Fig. 3i-l) are restricted to the Atlantic Forest, and Mutisia lutzii G.M.Barroso is endemic to the Caparaó mountain (Fig. 4l-n).

Figure 3
Barnadesieae. Dasyphyllum flagellare a. Habit. b. Capitulum discoid and axillary spines. c. Corolla bilabiate. d. Cypsela densely setose and pappus plumose. Nassauvieae. Holocheilus pinnatifidus e. Habit. f. Flower billabiate and cypsela ellipsoid with pappus barbellate. g. Basal leaf pinnatissecta. h. Co-inflorescence cymose and capitulum with involucral bracts 2–3-seriate. Trixis verbascifolia i. Habit. j. Co-inflorescence corymbiform. k. Peryphery flower with corolla bilabiate-liguliform. l. Disc flower with corolla bilabiate and cypsela cylindrical. (Photos: c-f, h, k, l: P Renon & GM Paranhos; G: M Pastore; a, b, i, j: M Monge).

Figure 4
Mutisieae. Chaptalia denticulata a. Habit. b. Capitulum. c. Leaves cordiform with runcinated margin, abaxial surface densely lanose. d. Ray flower. e. Cypsela not rostrate and pappus barbellate. Chaptalia piloselloides f. Habit. g. Capitulum. h. Front view of capitulum showing the ray and disc flowers. i. Ray flower. j. Disc flower. k. Cypsela rostrate. Mutisia lutzii. l. Habit. m. Disc flower and cypsela. n. Ray flower and anthers with apical appendage apiculate. (Photos: a-d, j, k, m, n: GM Paranhos; e, f, g, l: M Monge; h: P Renon).

Three new records were found for Caparaó, Chaptalia integerrima (Vell.) Burkart, Moquiniastrum polymorphum (Less.) G.Sancho and Mutisia coccinea A.St.-Hill. According to Moreira et al. (2020), the latest list compiled for the basal groups of Asteraceae in Caparaó, only Moquiniastrum densicephalum (Cabrera) G.Sancho was not found there. Consulting the specimen of M. densicephalum cited by Moreira et al. (2020) on SpeciesLink (UEC122970), was observed that it does not correspond to that species, and it indeed belongs Orchidaceae. Furthermore, the specimen was collected in Itacambira, state of Minas Gerais, located 650 km of Caparaó region. Thus, M. densicephalum does not occur in Caparaó or has yet to be found there.

Distribution, phenology and conservation

Data regarding distribution and phenology are presented in the systematic treatment. Some records relevant to the distribution of Chaptalia nutans, C. piloselloides (Fig. 4f-k), Holocheilus pinnatifidus, Mutisia campanulata, M. coccinea, Trixis lessingii and T. verbascifolia, are difficult to validate due to the absence of images to confirm the identification and the lack of consensus among platforms become even more challenging. In such cases the literature was reviewed, herbaria were contacted, and more online platforms were searched for a consensus between two or more, with improbable records ultimately being eliminated.

Only four of the 15 species had their conservation status evaluated previously, Dasyphyllum brasiliense, M. polymorphum and H. pinnatifidus as Least Concern (LC) and Trixis glaziovii as Vulnerable (VU). Of the eleven species not evaluated, five are restricted, two to Brazil (C. denticulata, D. flagellare), two to the Atlantic Forest (H. pinnatifidus, T. verbascifolia) and one to the Caparaó mountain (M. lutzii).

Systematic treatment

A total of 98 specimens were analyzed, of which 49 specimens (50%) were not identified or had their identification corrected. Fourteen specimens with names of non-valid synonyms (not updated), twelve with incomplete identifications (only name of genus), twelve incorrectly identified, six without identification and five had uncertain identifications added of “aff.” Or “cf.”.

Identification key for basal tribes of Asteraceae in the Caparaó National Park

1. Presence of spines in leaf-axils or stems; leaves and involucral bracts with apex mucronate or spinescent; corolla often pseudobilabiate (4+1); anthers with bidentate apical appendages (Dasyphyllum) …………......................….... Barnadesieae

1’. Lack of spines in leaf-axils or stems; leaves and involucral bracts without apex mucronate or spinescent; corolla never pseudobilabiate (4+1); anthers with entire apical appendages ……..…….. 2

2. Pappus 1–3-seriate; when 1-seriate bristles of pappus connate at base …….......… Gochnatieae

2’. Pappus 1-seriate; bristles of pappus not connate at base ................................................................ 3

4. Branches, leaves or involucral bracts with lanose indument; capitula heterogamous; style branches obtuse, acute or clavate, glabrous or papillose ………….........................…………….….... Mutisieae

4’. Branches, leaves or involucral bracts lacking lanose indument; capitula homogamous; style branches frequently truncate, penicellate ……..…………................................…………...…….. Nassauvieae

Barnadesieae D. Don, Trans. Linn. Soc. London 16(2): 273 (1830)

Barnadesieae is recognized by shrub or tree habit, rarely herbaceous, spines in leaf-axils or stems, leaves and involucral bracts with mucronate or spinescent apexes, venation acrodomous, actinodromous or paralellodromous, capitula discoid or radiate, flower 1–numerous, corolla tubulose, ligulate, bilabiate or pseudobilabiate, pappus plumose, rarely barbellate or scaled. Trichomes of corolla and cypsela “Barnadesioides type” (3-cells). Barnadesieae is represented in CNP by Dasyphyllum (2 spp.).

Dasyphyllum Kunth, Nov. Gen. Sp. [H.B.K.] 4(14): 13 (ed. fol.) (1818)

The genus Dasyphyllum is recognized by involucral bracts coriaceous and stiff, leaves with acrodomous venation, capitula discoid, receptacle pilose, anthers with bidentate apical appendage and pappus plumose.

Identification key to Dasyphyllum in the Caparaó National Park

1. Presence of lenticels in branches; axillary spines 2.5–3.3 cm long, straight; receptacle flat; anthers with sagittate basal appendages; cypselae cylindrical or ellipsoid …....................................................................................................... D. brasiliense

1’. Absent of lenticels in branches; axillary spines 0.3–0.8 cm long, recurvated; receptacle concave; anthers with obtuse basal appendages; cypselae obovoid ....................................... D. flagellare

Dasyphyllum brasiliense (Spreng.) Cabrera, Revista Mus. La Plata, Secc. Bot. 9: 72 (1959)

Scandent shrubs or trees, 2.0–5.0 m tall; branches densely tomentose-sericeous, glabrescent, trichomes bristly, yellow-brownish, lenticels present; axillary spines 2.5–3.3 cm long, straight. Leaves alternate, petiole 1.5–2.0 cm long; lamina 6.5–24.5×2.3–9.6 cm, elliptic or obovate, membranaceous, chartaceous or coriaceous, base acute, margin entire, apex acute, acuminate or mucronate, venation acrodomous basal or suprabasal, concolor, both surfaces tomentose-sericeous, glabrescent, trichomes bristly, brownish. Co-inflorescence corymb-paniculiform, capitula pedunculated; involucre 0.8–1.5×0.4–0.6 cm, campanulate or cylindrical, 8–12-seriate; involucral bracts imbricate, sericeous, glabrescent, outer bracts ovate or elliptical, apex acute or mucronate, not revolute, inner bracts linear, apex acute, revolute; receptacle flat. Flowers bisexual, corolla 0.7–0.9 cm long, tubulose, tube glabrous, lobes pilose; anthers with basal appendages sagittate, apical appendages bidentate; style branches rounded, papillose. Cypselae cylindrical or ellipsoid, densely setose, trichomes brownish, ribs not evident; carpopodium inconspicuous; pappus 1-seriate, beige.

Examined material: Brazil. Espírito Santo: Dores do Rio Preto, Caparaó National Park, “Macieira”, 13 August 2011 (vg.), M. Monge et al. 986 (UEC169879). Iúna, “Comunidade do Rio Claro”, Montane Semideciduous Forest, 20°22’09”S, 41°49’53”W, 19 March 2014 (vg.), M. Monge et al. 2723 (UEC193437). Minas Gerais: Alto Caparaó, Caparaó National Park, nearly “Córrego Caparaó”, 1.200 m, 07 August 1996 (fl., fr.), L.S. Leoni 3424 (ESA28079, HUFU81503).

Distribution, phenology and conservation: Dasyphyllum brasiliense was collected in Montane Semideciduous Forest at 1.200 m asl. The reproductive period is during August. The species occurs in Argentina, Bolivia, Brazil, Paraguay, Peru and Uruguay. In Brazil it occurs in Atlantic Forest, Cerrado (Brazilian savanna) and Pampa (southern grassland). The conservation status of D. brasiliense is Least Concern (LC).

Taxonomic comments: Dasyphyllum brasiliense is a scandent shrub or a tree of up 5.0 m in height. It is recognized by branches with lenticels, axillary spines straight, leaves elliptic or obovate with basal or suprabasal acrodomous venation, receptacle flat, anthers with sagittate basal appendages and cypsela cylindrical or ellipsoid.

Dasyphyllum brasiliense differs from Dasyphyllum flagellare by its larger axillary spines (2.5–3.3 cm vs. 0.2–0.6 cm long), larger leaves (6.5–24.5×2.3–9.6 cm vs. 0.5–4.0×0.5–1.6 cm), receptacle flat (vs. concave) and anthers with sagittate basal appendages (vs. obtuse).

Dasyphyllum flagellare (Casar.) Cabrera, Revista Mus. La Plata, Secc. Bot. 9(38): 60 (1959)

Fig. 3a-d

Scandent shrubs, 0.8–1.5 m tall; branches densely tomentose-villous, glabrescent, trichomes not bristly, yellow-brownish, lenticels absent; axillary spines 0.3–0.8 cm long, recurvated. Leaves: alternate, petiole 0.2–0.6 cm long; lamina 0.5–4.0×0.5–1.6 cm, elliptic or orbicular, chartaceous or subcoriaceous, base obtuse, margin entire, apex cuspidate or mucronate, venation basal acrodromous, concolor, abaxial surface tomentose-sericeous, glabrescent, trichomes not bristly, brownish, adaxial surface strigose, glabrescent, trichomes not bristly, brownish. Co-inflorescence corymbiform, capitula pedunculated; involucre 1.0–1.5×0.5–0.8 cm, campanulate, 8–10-seriate; involucral bracts imbricate, tomentose or villous, glabrescent, outer bracts ovate, apex mucronate, not revolute, inner bracts linear, apex acute, revolute; receptacle concave. Flowers bisexual, corolla 0.7–0.8 cm long, tubulose, tube glabrous, lobes pilose; anthers with basal appendage obtuse, apical appendage bidentate; style branches rounded, glabrous. Cypselae obovoid, densely setose, trichomes whitish, ribs not evident; carpopodium inconspicuous; pappus 1-seriate, beige.

Examined material: Brazil. Espírito Santo: Dores do Rio Preto, Caparaó National Park, “Farofa” waterfall, 20°27’58”S, 41°49’01”W, 2.223 m, 16 March 2004 (bd.), M. Monge et al. 2621 (UEC 193282). Pedra Menina, Caparaó National Park, “Farofa” waterfall trail, Montane Semideciduous Forest, 20°28’21”S, 41°49’40”W, 1.937 m, 12 October 2022 (fr.), P. Renon 535 (HUFU83170). Minas Gerais: Alto Caparaó, Caparaó National Park, High-altitude grassland, 2.200 m, 05 May 2001 (fl., fr.), L.S. Leoni 4658 (HUFU27020); ibidem, “Macieira”, 20.4333°S, 41.8683°W, 21 June 2013 (fl., fr.), V.C. Manhães et al. 345 (UEC193958); ibidem, “Tronqueira-Terreirão” trail, 20°26’00”S, 41°53’00”W, 21 September 2012 (fl., fr.), M. Monge et al. 1278 (UEC197756); ibidem, High-altitude grassland, 20°24’49”S, 41°49’58”W, 2.086 m, 09 October 2022 (fr.), P. Renon 515 (HUFU83151); ibidem, “Terreirão”, High-altitude grassland, 23 November 2006 (fl., fr.), A.M. Teles et al. 293 (BHCB103990, RB468834).

Distribution, phenology and conservation: Dasyphyllum flagellare was collected in High-altitude grassland and Montane Semideciduous Forest at 1.930–2.220 m asl. The reproductive period spends from March to November. The species is restricted to Brazil and spread through Atlantic Forest and Cerrado domains. Dasyphyllum flagellare is accessed as Least Concern (LC).

Taxonomic comments: Dasyphyllum flagellare is recognized by the scadent shrubby habit, axillary spines recurvated, leaves elliptic or orbicular, apex cuspidate or mucronate, venation basal acrodomous, receptacle concave, anthers with obtuse basal appendages and cypsela densely setose, obovoid.

Gochnatieae Panero & V.A.Funk, Proc. Biol. Soc. Washington 115(4): 913 (2002)

Gochnatieae can be shrub, subshrub, herb or tree, monoecious or rarely gynodioecious or polygamo-dioecious, leaves alternate or rosulate, flowers 3–300, corolla tubulose, bilabiate or pseudobilabiate, anthers with apical appendage acuminate or apiculate, cypsela villous or sericeous, pappus 1–3-seriate, barbellate, sometimes plumose at the apex, rarely paleaceous, bristles equal or unequal in length, connate or not at the base. Moquiniastrum (1 sp.) represents the tribe in CNP.

Moquiniastrum (Cabrera) G.Sancho, Phytotaxa 147(1): 29 (2013)

The genus Moquiniastrum is recognized by shrub or tree habit, trichomes 2–3-armed in branches, leaves and involucral bracts, leaves discolor, co-inflorescence paniculiform, corymbiform or racemiform, style branches glabrous, pappus 2–3-seriate, unequal in length.

Moquiniastrum polymorphum (Less.) G.Sancho, Phytotaxa 147(1): 32 (2013)

Shrub or tree, 2.0–5.0 m tall; branches densely tomentose-incanous, persistent, trichomes 2-armed, whitish. Leaves alternate, petiole 0.4–2.0 cm long; lamina 3.5–16.0×1.5–8.0 cm long, elliptic or obovate, chartaceous or coriaceous, base acute or obtuse, margin irregular entire or serrate, apex acute or obcordate, venation eucamptodromous, abaxial surface densely tomentose-incanous, persistent, trichomes 2-armed (Y-shaped), persistent, adaxial surface tomentose, glabrescent, trichomes 2-armed (Y-shaped), whitish. Co-inflorescence corymb-racemiform or paniculiform; capitula discoid, penduculated; involucre, 0.4–0.6×0.3–0.5 cm, campanulate, 6-seriate; involucral bracts imbricate, membranaceous, tomentose-villous, glabrescent, trichomes 2-armed, whitish, outer bracts ovate, apex acute, inner bracts elliptic or narrowly elliptic, apex acute; receptacle convex, glabrous. Flowers bisexual, corolla 0.6–0.7 cm long, tubulose, tube glabrous, lobes glabrous; anthers with basal appendage calcarate, apical appendage acuminate; style branches obtuse, glabrous. Cypselae ellipsoid, setose, trichomes whitish, 5–6-ribs; carpopodium inconspicuous; pappus barbellate, stramineous.

Examined material: Brazil. Espírito Santo: Iúna, “Comunidade do Rio Claro”, Ciliar Forest, 20°22’09’’S,41°49’53”W, 1.079 m, 19 March 2014 (vg.), M. Monge et al. 2713 (UEC193691). Minas Gerais: Alto Caparaó, Caparaó National Park, “Macieira”, 20°49’81”S, 41°97’14”W, 18 June 2013 (fr.), J.P.F. Zorzanelli et al. 751 (VIES31989). Espera Feliz, Caparaó National Park, “Macieira” trail, 20°28’48”S, 41°49’57”W, 1.841 m, 05 March 2010 (fr.), M.O. Bünger et al. 367 (BHCB136323); ibidem, “Macieira”, 1.870 m, July 2005 (fl., fr.), L.S. Leoni & M.J. Rocha 6232 (HUFU81498).

Supplementary material: Brazil. Minas Gerais: Lima Duarte, Conceição do Ibitipoca, road to “Moreiras”, 12 January 2002 (fl., fr.), D.S., Pifano & A. Valente 285 (HUFU26481).

Distribution, phenology and conservation: Moquiniastrum polymorphum was collected in Montane Semideciduous Forest at 1.000–1.900 me asl. Its reproductive period is around January to March. The species occurs in Argentina, Brazil, Paraguay and Uruguay. In Brazil can be found in Atlantic Forest, Cerrado and Pampa. Regarding the conservation status, M. polymorphum is accessed as Least Concern (LC).

Taxonomic comments: Moquiniastrum polymorphum is easily recognized by indument of branches, abaxial leaf surface and involucral bracts densely tomentose-incanous with Y-shaped trichomes, leaves petiolate, coriaceous or chartaceous, elliptic or obovate, co-inflorescence corymb-racemiform or paniculiform and capitula pedunculated. Moquiniastrum polymorphum has been mistakenly identified as Critoniopsis Sch.Bip. or Eremanthus Less., both genera belonging to the later-diverging tribe Vernonieae Cass. Although M. polymorphum has features that also occur in Vernonieae such as, tree habit, 2–3-seriate pappus and discolor leaves with dense incanus trichomes. It can be distinguished from Vernonieae by its obtuse (vs. acute or acuminate), briefly bifurcated (vs. long bifurcated) and glabrous (vs. pilose) style branches.

Mutisieae Cass., J. Phys. Chim. Hist. Nat. Arts 88: 199-200 (1819)

Mutisieae is recognized by herbaceous and climbing habit, lanose indument in branches, leaves, involucral bracts or corolla, leaves alternate, opposite or rosulate, entire or lobed, capitula heterogamous, radiate or disciform, flowers dimorphic or trimorphic, corolla liguliform, bilabiate, filiform or tubulose, anthers with basal appendage calcarate or sagittate, pappus plumose, barbellate or scabrid. Chaptalia (4 spp.) and Mutisia (3 spp.) represents the tribe in the CNP.

Identification key to Mutisieae in the Caparaó National Park

1. Herbs; leaves rosulate; tendrils absent in leaf-apex; flowers ≤ 2.0 cm long, trimorphic; disc flowers tubulose or filiform; anthers with apical appendage obtuse or acute; cypselae generally papillose, rostrate (except in C. denticulata); pappus barbellate ............................…......... Chaptalia

1’. Scandent woody plant; leaves alternate, opposite or subopposite; tendrils in leaf-apex; flowers >2.0 cm long, dimorphic; disc flowers bilabiate; anthers with apical appendage apiculate; cypselae glabrous, not rostrate; pappus plumose …………..........…....................... Mutisia

Chaptalia Vent., Descr. Pl. Nouv. ad t. 61 (1802)

The species from Chaptalia are small herbs, leaves arranged in rosette, flowers trimorphic, ray flowers concolor, disc flowers tubulose or filiform, anthers with apical appendage obtuse or acute, cypselae generally papillose, rostrate (except in C. denticulata), pappus barbellate.

Identification key to Chaptalia genus in the Caparaó National Park

1. Older leaves twisted, leaves narrowly oblong or oblanceolate; scape bracteate; style branches clavate ….................................……………... C. piloselloides

1’. Older leaves not twisted, leaves cordiform, elliptic or spatulate; scape not bracteate; style branches obtuse……..…………………………………...…...……. 2

2. Leaves cordiform, petiole not winged; involucral bracts elliptic; ray flowers with 4-nerves; cypselae cylindrical, not rostrate, 8-ribs …………….……...............................…… C. denticulata

2’. Leaves elliptic or spatulate, petiole winged; involucral bracts linear; ray flowers with 3-nerves; cypselae fusiform, rostrate, 4–5-ribs ………….........................................................................………….. 3

3. Leaves elliptic, entire; receptacle flat; disc flowers 1.3–1.4 cm long, anthers with apical appendage acute; carpopodium inconspicuous ………………………................................…........... C. integerrima

3’. Leaves spatulate, lobed; receptacle convex; disc flowers 1.0–1.2 cm long; anthers with apical appendage obtuse, carpopodium conspicuous ...……………...............………………... C. nutans

Chaptalia denticulata (Baker) Zardini, Darwiniana 19: 728 (1975)

Fig. 4a-e

Herbs, ca. 0.1–0.3 m tall; stem not evident. Leaves rosulate, petiole 2.5–13.0 cm long, not winged, sheaths absent; lamina 2.0–3.5×1.5–2.5 cm, cordiform, membranaceous, base cordate, margin runcinate, revolute, apex obtuse or rarely acute, venation eucamptodromous, discolor, abaxial surface densely lanose, persistent, trichomes beige or yellowish, adaxial surface lanose, glabrescent, trichomes beige or yellowish, older leaves not twisted. Capitula radiate, solitary; scape 5.0–28.0 cm long, not bracteate, densely lanate, glabrescent; involucre 1.0–1.3×0.6–0.8 cm, campanulate, 4–5-seriate; involucral bracts subimbricate, membranaceous, lanose, narrowly elliptic or elliptic, apex acuminate or acute; receptacle flat, glabrous. Flowers trimorphic; ray flowers pistillate, corolla 1.3–1.5 cm long, liguliform, 3-lobed, 4-nerves, white, glabrous; disc flowers bisexual or pistillate, corolla 0.5–0.7 cm long, tubulose or filiform, whitish, tube glabrous, lobes glabrous; anthers with basal appendage calcarate, apical appendage obtuse; style branches obtuse, papillose. Cypselae cylindrical, 0.1–0.2 cm long, papillose, glabrescent, trichomes apparently not glandular, 8-ribs; rostrum absent; carpopodium inconspicuous; pappus 1-seriate, yellowish.

Examined material: Brazil. Minas Gerais, Caparaó mountain, 2.500 m, 1941 (fr.), A.C. Brade 17003 (HUFU81285). Alto Caparaó, “Pico do Cristal” trail, humid area, 23 November 2006 (fl., fr.), A.M. Teles et al. 284 (BHCB64241).

Distribution, phenology and conservation: Chaptalia denticulata was found in High-altitude grassland at a humid area, around 2.500 m asl. The reproductive period is around November. It is restricted to Brazil occurring in Atlantic Forest, Cerrado and Caatinga domains. Regarding conservation status, Chaptalia denticulata is not evaluated.

Taxonomic comments: Chaptalia denticulata is easily recognized by leaves cordiform, margin runcinated, petiole prolonged reaching nearly 13.0 cm long, not winged, ray flowers with 4-nerves and cypselae not rostrate with 8-ribs.

Chaptalia integerrima (Vell.) Burkart, Darwiniana 6: 576 (1944)

Herbs, ca. 0.3–0.6 m tall; stem not evident. Leaves rosulate, petiole 1.5–2.5 cm long, winged, sheaths present; lamina 3.5–21.5×1.5–3.0 cm, elliptic, membranaceous, base atenuate or decurrent, margin dentate, slightly revolute, apex acute, venation eucamptodromous, discolor, abaxial surface densely lanose, persistent, trichomes yellowish, adaxial surface lanose, glabrescent, trichomes yellowish, older leaves not twisted. Capitula radiate, solitary; scape 30.0–46.0 cm long, not bracteate, lanose, glabrescent; involucre 1.7–2.4×1.5–2.4 cm, campanulate, 4–8-seriate; involucral bracts imbricate, membranaceous, lanose, linear, apex acuminate; receptacle flat, glabrous. Flowers trimorphic; ray flowers pistillate, corolla 1.4–1.6 cm long, liguliform, 3–4-lobed, 3-nerves, white, glabrous; disc flowers bisexual or pistillate, corolla 1.3–1.4 cm long, tubulose or filiform, whitish or yellowish, tube glabrous, lobes glabrous; anthers with basal appendage calcarate, apical appendage acute; style branches obtuse, glabrous. Cypselae fusiform, 0.3–0.4 cm long, papillose, persistent, trichomes apparently glandular, 4-ribs; rostrum 0.5–1.0 cm long; carpopodium inconspicuous; pappus 1-seriate, white-yellowish.

Examined material: Brazil. Minas Gerais: Alto Caparaó, Caparaó National Park, High-altitude grassland, 2.200 m, 24 November 1998 (fl., fr.), L.S. Leoni 4063 (GFJP).

Supplementary material: Brazil. Minas Gerais: Diamantina, Biribiri State Park, 18°10’59.3”S, 43°46’34.5”W, 03 December 2012 (fl., fr.), I.M. Franco et al. 1141 (HUFU66421).

Distribution, phenology and conservation: Chaptalia integerrima was collected in High-altitude grassland at 2.200 m asl. Its reproductive period is around November and December. The species occurs in Argentina, Bolivia, Guyana, Paraguay, Peru, Venezuela and Uruguay. In Brazil, it is spread through Atlantic Forest, Cerrado, Caatinga, and Pampa. Chaptalia integerrima is not evaluated regarding conservation status.

Taxonomic comments: Chaptalia integerrima is recognized by leaves elliptic, entire, anthers with apical appendage acute, cypselae with 4-ribs and carpopodium inconspicuous. The species is similar to C. nutans, but can be distinguished mainly by leaves elliptic and entire (vs. spatulate and lobed), receptacle flat (vs. receptacle convex), ray flowers white (vs. lilac or rosaceous), anthers with apical appendage acute (vs. obtuse) and carpopodium inconspicuous (vs, conspicuous).

Chaptalia nutans (L.) Pol., Linnaea 41(7): 582 (1878)

Herbs, ca. 0.3–0.6 m tall; stem not evident. Leaves rosulate, petiole 0.5–3.0 cm long or absent, winged, sheaths present; lamina 3.0–25.0×1.2–7.5 cm, spatulate, lobed at base, membranaceous, base decurrent, margin dentate, serrate or crenate, slightly revolute, apex acute, venation eucamptodromous, discolor, abaxial surface densely lanose, glabrescent, trichomes white or beige, adaxial surface lanose, glabrescent, trichomes white or beige, older leaves not twisted. Capitula radiate, solitary; scape 8.3–72.2 cm long, not bracteate, lanose, glabrescent; involucre 1.3–2.0×0.8–1.5 cm, campanulate, 5–8-seriate; involucral bracts imbricate, membranaceous, lanose, linear, apex acute or acuminate; receptacle convex, glabrous. Flowers trimorphic; ray flowers pistillate, corolla 1.3–1.4 cm long, liguliform, 3-lobed, 3-nerves, lilac or rosaceous, glabrous; disc flowers bisexual or pistillate, corolla 1.0–1.2 cm long, tubulose or filiform, cream or albescent, tube glabrous, lobes papillose; anthers with basal appendage calcarate, apical appendage obtuse; style branches obtuse, glabrous. Cypselae fusiform, 1.4–1.6 cm long, papillose, persistent, trichomes apparently glandular, 4–5-ribs; rostrum ca. 1.0 cm long; carpopodium conspicuous; pappus 1-seriate, whitish or brownish.

Examined material: Brazil. Espírito Santo: Patrimônio da Penha, Caparaó mountain, 18 May 2007 (fl., fr.), T. Chimalli s.n. (HUFU83413). Minas Gerais: Alto Caparaó, Caparaó National Park, “Tronqueira” trail, 20.2600°S, 41.5260°W, 18 September 2012 (fl., fr.), M. Monge et al. 1181/1184 (UEC204308, UEC204305); ibidem, “Vale Verde” trail, Montane Semideciduous Forest, 20°25’11”S, 41°50’41”W, 1.352 m, 06 October 2022 (fl., fr.), P. Renon 490 (HUFU83179); ibidem, “Tronqueira” trail, Montane Semideciduous Forest, 20°24’30”S,41°50’36”W, 1.616 m, 07 October 2022 (fl., fr.) P. Renon 500 (HUFU83134).

Distribution, pheonology and conservation: Chaptalia nutans was found in Montane Semideciduous Forest at 1.300–1.600 m asl. Its reproductive period is during September to May. The species is widely spread through New World occurring in Argentina, Belize, Bolivia, Brazil, Colombia, Costa Rica, Cuba, El Salvador, Ecuador, Guatemala, Guiana, Haiti, Honduras, Jamaica, Leeward Island, Mexico, Nicaragua, Panama, Paraguay, Peru, Puerto Rico, Dominican Republic, Suriname, Trinidad-Tobago, Uruguay, Venezuela and Windward Island. In Brazil, Chaptalia nutans is distributed across all phytogeography domains. The species is not evaluated regarding conservation status.

Taxonomic comments: Chaptalia nutans is easily recognized by leaves spatulate and lobed, scape very prolonged reaching nearly 72.0 cm long, ray flowers lilac or rosaceous, receptacle convex and cypselae with carpopodium conspicuous.

Chaptalia piloselloides (Vahl) Baker, Fl. Bras. (Martius) 6(3): 378-379 (1884)

Fig. 4f-k

Herbs, ca. 0.1–0.4 m tall; stem not evident. Leaves rosulate, petiole 2.0–7.0 cm long, winged, sheaths present; lamina (1.5)3.0–19.0×0.3–1.2 cm, narrowly oblong or oblanceolate, membranaceous, margin entire or serrate, revolute, base attenuated or decurrent, apex acute, venation eucamptodromous, discolor, abaxial surface densely lanose, glabrescent, trichomes whitish, adaxial surface lanose, glabrescent, trichomes whitish, oldest leaves twisted. Capitula radiate, solitary; scape 8.5–32.5 cm long, bracteate, lanose, glabrescent; involucre 1.5–1.9×0.8–2.0 cm, campanulate, 4–6-seriate; involucral bracts imbricate, membranaceous, lanate, glabrescent, lanceolate or linear, apex acute or acuminate; receptacle flat, glabrous. Flowers trimorphic; ray flowers pistillate, corolla 0.7–1.0 cm long, liguliform, 3-lobed, 4-nerves, white, glabrous; disc flowers bisexual or pistillate, 0.5–0.7 cm long, tubulose or filiform, white, tube glabrous, lobes glabrous; anthers with basal appendage calcarate, apical appendage obtuse; style branches clavate, papillose. Cypselae cylindrical or elipsoid, 0.3–0.5 cm long, papillose, persistent, trichomes apparently glandular, 5–6-ribs; rostrum 0.2–0.4 cm long; carpopodium inconspicuous; pappus 1-seriate, yellowish.

Examined material: Brazil. Espírito Santo: Dores do Rio Preto, Caparaó National Park, “Macieira”, 1.897 m, 20 June 2005 (fl., fr.), L.S. Leoni & M.J.R Rocha 6196 (HUFU81504, RB747197); ibidem, “Casa Queimada” trail, Upper Montane Forest, 13 August 2011 (fl., fr.), M. Monge et al. 971 (UEC169879). Iúna, Caparaó National Park, High-altitude grassland, 20°24’36,1”S, 41°48’29,6”W, 18 February 2000 (fl., fr.), V.C. Souza et al. 23347 (RB738985); ibidem, nearly “Rancho dos Cabritos”, High-altitude grassland, 18 February 2000 (fl., fr.), V.C. Souza et al. 23392 (RB739018). Minas Gerais: Caparaó mountain, “Luiz Ignácio” peak, 2.300 m, 09 September 1941 (fl., fr.), A.C. Brade 16890 (RB45888, UEC155984). Alto Caparaó: Caparaó National Park, High-altitude grassland, 2.100 m, 29 February 1995 (fl., fr.), L.S. Leoni et al. 3090 (HUFU81470, RB739000); ibidem, High-altitude grassland, 2.200 m, 14 October 1998 (fr.), L.S. Leoni 4037 (HUFU27019); ibidem, “Pico da Bandeira” trail, 20.4333°S, 41.8683°W, 19 September 2013 (fl., fr.), V.C. Manhães et al. 435 (UEC193964); ibidem, “José Pedro” watercourse, Montane Semideciduous Forest, 20.2600°S, 41.5260°W, 18 September 2012 (fr.), M. Monge et al. 1209 (UEC197481); ibidem, “Vale Encantado”, 20.2600°S, 41.5260°W, 19 September 2012 (fl., fr.), M. Monge et al. 1218 (UEC204309); ibidem, “Tronqueira” trail, High-altitude grassland, 20°26’00”S, 41°53’00”W, 21 September 2012 (vg.), M. Monge et al. 1282 (UEC197843); ibidem, “Tronqueira-Terreirão”, High-altitude grassland, 20°26’00”S, 41°53’00”W, 22 September 2012 (fl., fr.), M. Monge et al. 1290 (UEC197850); ibidem, “Pico da Bandeira” trail, rock outcrops, humid area, 20°24’57”S, 41°53’37”W, 2.145 m, 17 March 2014 (fl., fr.), M. Monge et al. 2633 (UEC193509). Espera Feliz: Caparaó National Park, High-altitude grassland, 2.200 m, 21 April 2004 (fr.), L.S. Leoni et al. 5710 (HUFU81496).

Distribution, phenology and conservation: Chaptalia piloselloides was collected in high-altitude grassland at 1.900–2.300 m asl. Its reproductive period is during February to September. The species occurs in Argentina, Brazil, Bolivia, Paraguay, Uruguay and Venezuela. In Brazil is spread through Atlantic Forest and Pampa domains. Chaptalia piloselloides is not evaluated regarding its conservation status.

Taxonomic comments: Chaptalia piloselloides differs from others species within genus due to its leaves narrowly oblong or oblanceolate, the oldest leaves twisted, scape bracteate and style branches clavate.

Mutisia L.f., Suppl. Pl. 57: 373 (1782)

The genus Mutisia is recognized by its climbing habit, branches frequently quadrangular, leaves alternate, opposite or subopposite, lobed, venation acrodromous or actinodromous, tendrils in leaf-apex, ray flowers discolor, disc flowers bilabiate, anthers with apical appendage apiculate, cypselae cylindrical, glabrous and pappus plumose.

Identification key to genus Mutisia in the Caparaó National Park

1. Leaves with 9–13 pairs of lobes; lobes opposite or subopposite, base truncate; involucre 5–6-seriate; outer involucral bracts not recurved; receptacle flat; pappus 2.0–2.5 cm long ……………………………………………………………………………………….... M. lutzii.

1’. Leaves with 4–6 pairs of lobes; lobes alternate or rarely subopposite, base attenuate, obtuse or cuneate; involucre 8–10-seriate; outer involucral bracts recurved; receptacle concave; pappus 3.0–3.5 cm long ……….......................…...……….. 2

2. Branches winged; leaves with abaxial surface lanose-tomentose; involucre campanulate; corolla lobes of disc flowers without glandular-dots; cypsela with 4 ribs …………………………………………….……….... M. campanulata

2’. Branches not winged; leaves with abaxial surface lanose-villous; involucre cylindrical; corolla lobes of disc flowers with glandular-dots; cypsela with ca. 6 ribs ……...……………… M. coccinea

Mutisia campanulata Less., Linnaea 5(2): 269 (1830)

Scandent woody plant; branches quadrangular, winged, lanose, glabrescent, trichomes whitish to brownish. Leaves: alternate, sessile; lamina 11.0–20.0×5.5–12.0 cm, lobed, 4–6 pair of lobes; lobes alternate or rarely subopposite; lobes lamina 2.5–6.5×1.0–2.2 cm, elliptic or ovate, membranaceous, margin irregular entire, base attenuate, obtuse or cuneate, apex acute or obtuse, venation basal acrodromous, discolor, abaxial surface densely lanose-tomentose, persistent, trichomes whitish or brownish, adaxial surface lanose, glabrescent, trichomes whitish or brownish; tendrils in leaf-apex. Capitula radiate, solitary; involucre 3.5–4.0×2.0–2.5 cm, campanulate, 8–10-seriate; involucral bracts imbricate, chartaceous or membranaceous, densely lanose-tomentose, glabrescent, trichomes beige or brownish, outer bracts lanceolate, apex caudate, recurved, inner bracts oblong to ovate, apex acuminate, apiculate or obtuse, not recurved; receptacle concave, glabrous. Flowers dimorphic; ray flowers pistillate, 14–16 flowers per capitulum; corolla 4.5–5.8 cm long, liguliform, 2-lobed, 14–16-nerves, red or vinaceous, limb and lobes densely lanose-tomentose in abaxial surface; disc flowers bisexual, ca. 60 flowers per capitulum; corolla 3.8–4.0 cm long, bilabiate, 5-lobed, yellowish or orange, tube glabrous, lobes lanose; anthers with basal appendages calcarate; style branches obtuse, papillose. Cypselae ca. 0.2 cm long, 4-ribs; carpopodium inconspicuous; pappus 1-seriate, 3–3.5 cm long, brownish.

Examined material: Brazil. Espírito Santo: Dores do Rio Preto, Caparaó National Park, 13 August 2011 (vg.), M. Monge et al. 982 (UEC169875). Minas Gerais: Caparaó mountain, Transitional vegetation, 1.700 m, 10 August 1979 (vg.), A.F.M. Albuquerque s.n. (RB296782); Caparaó National Park, 1.400–1.600 m, 31 October 1922 (fl., fr.), P. Campos Porto 1174 (RB4986); ibidem, 01 May 1996 (vg.), L.S. Leoni 3321 (RB662572); ibidem, “Pico da Bandeira”, 1.500–1.600 m, 06 September 1977 (fl., fr.), G.J. Shepherd et al. 5753 (UEC2888). Alto Caparaó, Caparaó National Park, 06 January 1999 (fl., fr.), L.S. Leoni 4078 (RB748166); ibidem, 21 February 2000 (fl., fr.), V.C. Souza et al. 23631 (ESA64779). Espera Feliz, Caparaó National Park, “Macieira” trail, 1.400–1.870 m, 20 September 2016 (bd), C.D.M. Ferreira et al. 357 (RB665775).

Distribution, phenology and conservation: Mutisia campanulata has been found in Montane Semideciduous Forest and “Candeal” (transitional vegetation), at 1.400–1.900 meters asl. Its reproductive period is around September to February. The species occurs in Argentina, Brazil, Chile and Paraguay. In Brazil it is distributed in Atlantic Forest and Pampa. The conservation status of Mutisia campanulata has not been evaluated.

Taxonomic comments: Mutisia campanulata is easily recognized by winged branches, involucre campanulate and cypselae 4-ribbed. Despite the similarity between M. campanulata and M. coccinea, due to their leaves with the same number of lobes (4–6 pairs), lamina of lobes elliptic to ovate with obtuse base or cuneate, involucre 8–10-seriate, outer involucral bracts recurved and receptacle concave, both species can be distinguished mainly by branches never winged in M. coccinea (vs. winged), involucre cylindrical (vs. campanulate) and cypselae ca. 6-ribbed (vs. 4-ribbed).

Mutisia coccinea A.St.-Hil., Voy. Distr. Diam. 1: 386 (1833)

Scandent woody plant; branches quadrangular, not winged, lanose, glabrescent, trichomes whitish or yellowish. Leaves: alternate, sessile; lamina 9.0–10.5×5.0–6.5 cm, lobed, 5–6 pair of lobes; lobes alternate; lobes lamina 1.5–3.0×0.5–1.0 cm, elliptic to ovate, membranaceous, base obtuse or slightly cuneate, margin irregular entire, apex acute, venation basal acrodromous, discolor, abaxial surface densely lanose-villous, persistent, trichomes whitish or yellowish, adaxial surface lanose, glabrescent, trichomes whitish or yellowish; tendrils in leaf-apex. Capitula radiate, solitary; involucre ca. 3.5×2.5 cm, cylindrical, ca. 10-seriate; involucral bracts imbricate, membranaceous or chartaceous, densely lanose-villous, glabrescent, trichomes whitish or yellowish, outer bracts lanceolate, apex caudate, recurved, inner bracts elliptic to ovate, apex acute or acuminate; receptacle concave, glabrous. Flowers dimorphic; ray flowers pistillate, ca. 13 flowers per capitulum; corolla ca. 5.5 cm long, liguliform, 3-lobed, ca. 13-nerves, red with yellow, limb and lobes densely lanose-tomentose in abaxial surface; disc flowers bisexual, ca. 60 flowers per capitulum; corolla 3.5–3.8 cm long, bilabiate, yellowish, tube glabrous, lobes lanose with glandular-dots; anthers with basal appendages calcarate; style branches obtuse, papillose. Cypselae ca. 0.2 cm long, ca. 6-ribs; carpopodium inconspicuous; pappus 1-seriate, ca. 3.5 cm long, brownish.

Examined material: Brazil. Minas Gerais: Espera Feliz, Caparaó National Park, “Bonita” waterfall, Ombrophilous Forest, 41°83’73”S, 20°40’65”W, 1.818 m, 16 September 2016 (fl., fr.), J.A.M. Paiva et al. 1067 (BHCB182961).

Distribution, phenology and conservation: Mutisia coccinea was collected in Montane Ombrophilous Forest at 1.820 m asl, with flowers and fruits in September. The species occurs in Argentina, Brazil, Paraguay and Uruguay. In Brazil, it is distributed in Atlantic Forest and Pampa. Mutisia coccinea is not evaluated regarding conservation status.

Taxonomic comments: Mutisia coccinea is recognized by leaves and involucral bracts with lanose-villous indument, involucre cylindrical with recurved outer bracts, corolla of disc flowers glandular-dotted and cypselae ca. 6-ribbed.

Mutisia lutzii G.M.Barroso, Mem. Inst. Oswaldo Cruz 53: [357], fig (1955)

Fig. 4l-n

Scandent woody plant; branches quadrangular, not winged, lanose, glabrescent, trichomes whitish or beige. Leaves alternate, sessile; lamina 8.0–14.0×2.0–4.5 cm, lobed, 9–13 pair of lobes; lobes opposite or subopposite; lobes lamina 0.3–2.6×0.4–1.1 cm, elliptic to ovate, membranaceous, base truncate, margin irregular entire, apex acute or obtuse, venation basal acrodromous, discolor, abaxial surface densely lanose-tomentose, persistent, trichomes beige or whitish, adaxial surface lanose, glabrescent, trichomes whitish to beige; tendrils in leaf-apex. Capitula radiate, solitary; involucre 2.3–2.5×1.4–1.8 cm, cylindrical, 5–6-seriate; involucral bracts imbricate, membranaceous or chartaceous, densely lanose-tomentose, glabrescent, trichomes whitish or beige, outer bracts lanceolate, apex caudate or acuminate, not recurved, inner bracts oblong or elliptic, apex acute or acuminate; receptacle flat, glabrous. Flowers dimorphic; ray flowers pistillate, ca. 12 flowers per capitulum; corolla 5.0–6.1 cm long, liguliform, 3-lobed, ca. 12-nerves, red or vinaceous, limb and lobes densely lanose in abaxial surface; disc flowers bisexual, ca. 40 flowers per capitulum; corolla 2.9–3.2 cm long, bilabiate, cream, lobes lanose; anthers with basal appendage calcarate; style branches obtuse, papillose. Cypselae ca. 0.2 cm long, 8–10-ribs; carpopodium inconspicuous; pappus 1-seriate, 2–2.5 cm long, brownish.

Examined material: Brazil. Espírito Santo: Dores do Rio Preto, Caparaó National Park, “Macieira” trail, 22 March 2012 (fl., fr.), J. Kuntz et al. 598 (RB649284); ibidem, “Farofa” waterfall, 20°29’00’’S, 41°49’37”W, 16 March 2014 (fl., fr.), M. Monge et al. 2576 (UEC194236); ibidem, “Lajão”, 20°27’58”S, 41°49’01”W, 2.223 m, 16 March 2014 (fl., fr.), M. Monge et al. 2580 (UEC194143). Iúna, Caparaó National Park, nearly “Rancho dos Cabritos”, High-altitude grassland, 18 February 2000 (fl., fr.), V.C. Souza et al. 23399 (HUFU56659, RB570526, RB662503, UEC178502). Minas Gerais: Alto Caparaó, nearly “Casa Queimada”, High-altitude grassland, 20°27’32”S, 41°48’34”W, 02 May 2019 (fl., fr.), J.P.F Zorzanelli & L. Batista 1901 (HUFU83414). Espera Feliz: Caparaó National Park, nearly “Lajão”, Transitional vegetation, 14 May 2016 (fl., fr.), L.S. Leoni. & I.T.F.V. Lopes 10148 (HUFU72640).

Distribution, phenology and conservation: Mutisia lutzii is endemic to Caparaó, where it has been found at High-altitude grassland and “Candeal” (transitional vegetation) around 2.200 meters asl. Its reproductive period occurs in February to May approximately. The species is not evaluated regarding conservation status.

Taxonomic comments: Mutisia lutzii is easily distinguished from its congeners by its leaves with 9–13 pairs of lobes, lobes opposite or subopposite with truncate base, involucral bracts non-recurved, receptacle flat and cypselae with 8–10-ribs.

Nassauvieae Cass., J. Phys. Chim. Hist. Nat. Arts 88: 198-199 (1819)

Nassauvieae is recognized by herbaceous, shrubby or climbing habit, leaves alternate or rosulate, entire, palmatifid, pinnatifid or pinnatisect, capitula homogamous, flowers isomorphic or dimorphic, bilabiate, bilabiate-liguliform or rarely tubulose, anthers with basal appendage calcarate or caudate, apical appendage acute, apiculate or falciform, style branches truncate, rarely rounded, panicellate, rarely papillose, cypselae with pappus uni-multisseriate, barbellate, paleaceous, plumose or absent. Holocheilus (1 sp.) and Trixis (4 spp.) represent the tribe in CNP.

Identification key to Nassauvieae in the Caparaó National Park

1. Involucre up to 3-seriate; receptacle glabrous; corolla glabrous in inner portion; anthers blackned ………...……………....………………....… Holocheilus

1’. Involucre up to 5-seriate; receptacle pilose; corolla pilose in inner portion; anthers brownish, yellowish or orange .......................... Trixis

Holocheilus Cass., Bull. Sci. Soc. Philom. Paris 1818: 73 (1818)

Holocheilus is recognized by herbaceous habit, leaves heteromorphic, basal rosulate, petiolate, entire, pinnatifid or pinnatisect, apical alternate, sessile, entire generally with semi-aplexicaul base, co-inflorescence corymbiform, involucre up to 3-seriate, receptacle glabrous, corolla isomorphic, glabrous in inner portion and anthers blackned.

Holocheilus pinnatifidus (Less.) Cabrera, Revista Mus. La Plata, Secc. Bot. 11(50): 4 (1968)

Fig. 3e-h

Herbs, ca. 0.6 m tall; branches fistulous, strigose-villous, glabrescent, trichomes multicellular, whitish. Leaves basal rosulate, petiole 6.0–10.0 cm long; lamina 3.5–26.2×1.3–5.5 cm, pinnatisect, base atenuate, apex acute; leaves apical alternate, sessile; lamina 4.0–11.0×0.5–2.0 cm, lanceolate, base hastate or semi-amplexicaul, apex acuminate; both basal and apical leaves membranaceous, margin denteate, venation cladodromous, concolor, both surfaces strigose, glabrescent, trichomes whitish. Co-inflorescence with corymbs aphyllous; capitula discoid, pedunculated; peduncle papillose with trichomes apparently glandular; involucre 0.4–0.6×0.3–0.5 cm, campanulate, 3-seriate; involucral bracts imbricate, membranaceous or chartaceous, strigose, trichomes whitish, glabrescent, outer and inner bracts obovate or oblong, apex acute; receptacle flat or convex. Flowers isomorphic, ca. 10 per capitulum, bisexual; corolla 0.5–0.8 cm long, bilabiate-liguliform, white, glabrous in inner portion, papillose in outer portion, trichomes apparently glandular; lobes glabrous; anthers with basal appendage calcarate, apical appendage apiculate; style branches truncate, penicellate. Cypselae ellipsoid or obovate, strigose, tector trichomes recurved, apex acute, glandular trichomes cylindrical, apex capitate, 4-ribs or absent; carpopodium conspicuous; pappus 1-seriate, barbellate, stramineous.

Examined material: Brazil. Espírito Santo: Caparaó National Park, “Macieira”, 18 January 2019 (fl., fr.), L. H. Silva & L.J. Lieven 270 (HUFU83635). Minas Gerais: Caparaó National Park, High-altitude grassland, humid area, 19 November 1988 (fl., fr.), L. Krieger et al. 555 (HUFU83313). Alto Caparaó, Caparaó Natinal Park, “Vale Encantado”, 2.100 m, 01 December 2000 (fl., fr.), L.S. Leoni 4536 (GFJP6028, GFJP s.n.); ibidem, “Morro da Jumenta”, 20°25’18,1”S, 41°49’16,1”W, 2.330 m, 20 February 2000 (fl., fr.), V.C. Souza et al. 23564 (RB748136); ibidem, “Macieira” trail, 20°49’46.9”S, 41°49’46.9”W, 1.854 m, 23 January 2008 (fl., fr.), B.V. Tinti et al. 442 (HUFU73532).

Distribution, phenology and conservation: Holocheilus pinnatifidus was found in “Candeal” (transitional vegetation) and High-altitude grassland, mainly in humid areas, at 1.850–2.330 m asl. Its reproductive period is during November to February. The species is restricted to Brazilian Atlantic Forest. The conversation status of Holocheilus pinnatifidus is Least Concern (LC).

Taxonomic comments: Holocheilus pinnatifidus is easily distinguished from its congeners due to its basal leaves long and pinnatisect, and co-inflorescence lax with numerous capitula. The pinnatisect basal leaves are found solely in H. pinnatifidus.

Trixis P. Browne, Civ. Nat. Hist. Jamaica 312: (1756)

The genus Trixis is recognized by herbaceous and shrubby habitat, glandular trichomes in branches, leaves, involucral bracts, corolla or cypselae, leaves homomorphic or heteromorphic, entire, receptacle pilose, epaleaceous, involucre up to 5-seriate, corolla dimorphic or isomorphic, pilose in inner portion and anthers brownish, yellowish or orange.

Identification key to genus Trixis in the Caparaó National Park

1. Basal leaves alternate; co-inflorescence pluricephala corymbiform or racemiform ...…….. 2

1’. Basal leaves in rosette; capitulum solitary or co-inflorescence paucicephala in dichasial cyme ……………………………........……………………………………… 3

2. Branches not winged; leaves petiolate, lamina 2.3–16.5×0.5–3.6 cm, membranaceous; ca. 10 flowers per capitulum; periphery flowers with 2-nerves; cypselae with 5-ribs ………………….............................................................…………… T. antimenorrhoea

2’. Branches winged; leaves sessile, lamina 2.5–34.5×1.0–11.0 cm, chartaceous; 60–80 flowers per capitulum; periphery flowers with 4-nerves; cypselae with 8–10 ribs ......................................................................................................................... T. verbascifolia

3. Branches not fistulous, strigose-pubescent; trichomes in branches and leaves with base expanded apparently glandular; leaf-margin dentate or serrate, venation broquidodromous; receptacle flat; foliaceous subinvolucral bracts absent; carpopodium conspicuous …….................................................................................................... T. glaziovii

3’. Branches fistulous, sericeous or tomentose; trichomes in branches and leaves without base expanded; leaf-margin entire, venation semicraspedodromous or paralellodromous; receptacle convex, foliaceous subinvolucral bracts present, carpopodium inconspicuous ............................................... T. lessingii

Trixis antimenorrhoea (Schrank) Mart. ex Baker, Fl. Bras. (Martius) 6(3): 385 (1884)

Shrubs, 2.0–4.0 m tall; branches not fistulous, not winged, strigose-pubescent, glandular, glabrescent, tector trichomes recurved, base expanded, apex acute, whitish, glandular trichomes cylindrical, not recurved, base not expanded, apex capitate, brownish. Leaves homomorphic, alternate, petiole 0.3–1.0 cm long; lamina 2.3–16.5×0.5–3.6 cm, elliptic or lanceolate, membranaceous, base acute, margin entire with uncinate trichomes, apex acuminate, venation broquidodromous, concolor, abaxial surface densely strigose-sericeous, glabrescent, trichomes whitish, adaxial surface strigose-pubescent, glabrescent, trichomes whitish. Co-inflorescence pluricephala, corymbiform-racemiform; capitula discoid, pedunculated; involucre 0.8–1.2×0.5–1.0 cm, campanulate, 2–3-seriate; involucral bracts subimbricate, chartaceous or membranaceous, strigose-pubescent, tector trichomes cylindrical, apex acute, whitish, glandular trichomes cylindrical, apex capitate, brownish, bracts linear or elliptic, apex acute; receptacle flat. Flowers dimorphic, ca. 10 per capitulum, periphery flowers bisexual; corolla 1.1–1.2 cm, bilabilate-liguliform, 2-nerves; disc flowers bisexual; corolla 1.0–1.2 cm long, bilabiate; both periphery and disc flowers yellowish or cream; corolla-limb glandular in outer portion, trichomes cylindrical, apex capitate, pubescent in inner portion, trichomes cylindrical, apex acute; lobes strigose, glandular in outer portion, glabrous in inner portion; anthers basal appendage calcarate, apical appendage apiculate or falciform; style branches truncate, penicellate. Cypselae narrowly cylindrical or ellipsoid, densely glandular, trichomes cylindrical, apex capitate, 5-ribs; carpopodium conspicuous; pappus 1-seriate, barbellate, stramineous.

Examined material: Brazil. Espírito Santo: Dores do Rio Preto, Caparaó National Park, “Macieira” road, 20°29’33.8”S, 41°49’18.2”W, 1.546 m, 07 June 2008 (fl., fr.), B.V. Tinti et al. 579 (HUFU73468). Iúna, “São João do Príncipe” farm, 20°20’30”S, 41°49’14”W, 1.107 m, 14 July 2021 (fr.), M.I.A. Horst et al. 160 (HUFU83412). Minas Gerais: Caparaó National Park, “Vale Verde”, 15 June 1991 (fl., fr.), G. Hatschbach et al. 55546 (HUFU26417).

Distribution, phenology and conservation: Trixis antimenorrhoea was found in Montane Semideciduous Forest at 1.100–1.550 m asl. The reproductive period is around July and June. The species is spread through Argentina, Brazil, Bolivia, Colombia, Ecuador, Paraguay, Peru and Venezuela. In Brazil occurs in Atlantic Forest and Cerrado domains. Regarding the conservation status, Trixis antimenorrhoea is not evaluated.

Taxonomic comments: Trixis antimenorrhoea can be distinguished from others species due to its leaves homomorphic, petiolate, elliptic or lanceolate, co-inflorescence laxa, pluricephalus, corymbiform-racemiform, capitula with ca. 10 flowers and periphery flowers with 2-nerves.

Trixis glaziovii Baker, Fl. Bras. (Martius) 6(3): 391 (1884)

Herbs or subshrubs, ca. 0.4 m tall; branches not fistulous, slightly winged, strigose-pubescent, glabrescent, trichomes recurved, base expanded apparently glandular, apex acute, whitish or brownish. Leaves heteromorphic; basal leaves rosulate, sessile; lamina 3.5–12.5×1.4–5.0 cm; apical leaves alternate, sessile; lamina 1.3–11.0×0.5–2.5 cm; both elliptic or obovate, chartaceous, margin dentate or serrate, spinescent, base attenuate or decurrent, apex obtuse or acute, venation broquidodromous, concolor, adaxial and abaxial surfaces strigose-pubescent, glabrescent, trichomes with base apparently glandular, whitish or brownish. Co-inflorescence paucicephala, dichasial cyme or solitary capitulum; capitula discoid, pedunculated; involucre 1.0–1.3×1.8–2.5 cm, campanulate, 2–3-seriate; involucral bracts subimbricate, chartaceous or membranaceous, densely strigose-pubescent, trichomes with base apparently glandular, brownish, bracts linear or elliptic, apex acute or acuminate; receptacle flat. Flowers dimorphic, 20–85 per capitulum; periphery flowers bisexual; corolla 1.6–1.8 cm long, bilabiate-liguliform, 4-nerves; disc flowers bisexual; corolla 1.3–1.5 cm long, bilabiate; both periphery and disc flowers yellowish or orangish; corolla-limb glabrous in outer portion, strigose-pubescent in inner portion; lobes strigose-setose in outer portion, glabrous in inner portion; anthers basal appendage calcarate, apical appendage apiculate; style branches truncate, panicellate. Cypselae narrowly cylindrical, densely strigose-pubescent, glandular, tector trichomes slender, bristly, apex acute, glandular trichomes cylindrical, apex capitate, 10-ribs; carpopodium conspicuous; pappus 1-seriate, barbellate, stramineous.

Examined material: Brazil. Minas Gerais: Alto Caparaó, Caparaó National Park, “Pico da Bandeira” trail, High-altitude grassland, humid area, 20°25’32”S,41°48’05”W, 2.581 m, 17 March 2014 (fr.), M. Monge et al. 2654 (UEC194055); ibidem, nearly “Terreirão”, High-altitude grassland, 2.300 m, 20°25’S, 41°49’W, 17 February 2000 (fl., fr.), V.C. Souza et al. 23302 (HUFU56670, UEC204176); ibidem, High-altitude grassland, 12 February 1996 (fl., fr.), L.S. Leoni 3198 (HUFU27029, RB736216, RB736222).

Distribution, phenology and conservation: Trixis glaziovii was found in High-altitude grassland and can be associated with humid locations, at 2.300–2.600 m asl. Its reproductive period is around February and March. The species is restricted to Brazil, where it occurs in Atlantic Forest and Cerrado domains. Trixis glaziovii is accessed as Vulnerable (VU).

Taxonomic comments: Trixis glaziovii is recognized by branches slightly winged, leaves basally rosulate, sessile, elliptic or obovate with attenuate to decurrent base, capitulum solitary or dichasium paucicephalus. Trixis glaziovii and T. lessingii have often been confused, both are species very similar, but can be differentiated mainly by branches, leaves, subinvolucral bracts and carpopodium. Trixis glaziovii has non-fistulosus branches (vs. fistulosus), dentate or serrate leaf margin (vs. entire), absence of subinvolucral bracts (vs. foliaceous and covering the capitulum) and conspicuous carpopodium (vs. inconspicuous).

Trixis lessingii DC., Prodr. [A. P. de Candolle] 7(1): 70 (1838)

Herbs or subshrubs, ca. 0.8 m tall; branches fistulous, not winged, densely sericeous or tomentose, glabrescent, trichomes slender, base not expanded, apex acute, whitish or brownish. Leaves heteromorphic; basal leaves rosulate, sessile; lamina ca. 45.0×3.2 cm, oblong or lanceolate, base attenuate, apex cuspidate or acute, venation semicraspedodromous; apical leaves alternate, sessile; lamina 3.2–38.0×1.6–2.0 cm, elliptic, base acute or obtuse, apex cuspidate or acute, venation parallelodromous; both basal and apical leaves chartaceous, margin irregular entire, spinescent, concolor, abaxial and adaxial surfaces strigose or strigose-sericeous, glabrescent, tector trichomes slender, whitish or brownish. Co-inflorescence paucicephala, dichasial cyme; capitula discoid, pedunculated; involucre 1.2–1.5×0.8–1.3 cm, campanulate, 1–2-seriate; involucral bracts subimbricate, chartaceous or membranaceous, densely strigose-sericeous, trichomes slender, whitish or brownish, bracts linear or lanceolate, apex acute or acuminate; receptacle convex. Flowers dimorphic, 25–95 per capitulum; periphery flowers bisexual; corolla 1.3–1.5 cm long, bilabiate-liguliform, 4-nerves; disc flowers bisexual; corolla 1.2–1.3 cm long, bilabiate; both periphery and disc flowers yellowish or orangish; corolla-limb glandular in outer portion, trichomes cylindrical, apex rounded, pubescent in inner portion, trichomes cylindrical, apex acute; lobes strigose in outer portion, trichomes apparently with glandular base, glabrous in inner portion; anthers basal appendage calcarate, apical appendage apiculate; style branches truncate, penicellate. Cypselae narrowly cylindrical, densely pubescent-tomentose, glandular, tector trichomes cylindrical, slender, apex acute, whitish or stramineous, glandular trichomes cylindrical, apex clavate, brownish, ribs not evident; carpopodium inconspicuous; pappus 1-seriate, barbellate, stramineous.

Examined material: Brazil. Minas Gerais: Alto Caparaó, Caparaó National Park, “Aurélio” waterfall trail, 20°28’S, 41°50’W, 1.834 m, 01 December 2010 (fl., fr.), T.M. Machado et al. 253 (RB540281); ibidem, “Macieira”, 20°28’54.2”S, 41°50’11.3”W, 1.833 m, 22 January 2008 (fl., fr.), B.V. Tinti et al. 433 (HUFU73457).

Distribution, phenology and conservation: Trixis lessingii was found propably in “Candeal” (transitional vegetation) at 1.830 meters asl. The species is spread through Brazil, Paraguay and Uruguay. Regarding the conservation status, Trixis lessingii is not evaluated.

Taxonomic comments: Trixis lessingii is recognized mainly by fistulous branches, leaves heteromorphic, basal leaves rosulate, lamina reaching ca. 45 cm long, oblong, lanceolate or elliptic, venation semicraspedodromous or parallelodromous, subinvolucral bracts foliaceous, involucre 1–2-seriate and co-inflorescence paucicephalus in dichasial cyme.

Trixis verbascifolia S.F.Blake, Contr. U.S. Natl. Herb. 22: 655 (1924)

Fig. 3i-l

Herbs or subshrubs, 0.5–2.0 m tall; branches not fistulous, winged, densely tomentose-strigose, glandular, glabrescent, tector trichomes recurved, base expanded apparently glandular, apex acute, whitish, glandular trichomes cylindrical, apex capitate, brownish. Leaves homomorphic, alternate, sessile; lamina 2.5–34.5×1.0–11.0 cm, elliptic or lanceolate, chartaceous, base decurrent, margin dentate or serreate, spinescent, apex acute or acuminate, venation broquidodromous, concolor, abaxial surface densely villous, glabrescent, tector trichomes cylindrical, yellowish, glandular, persistent, glandular trichomes rounded, brownish, adaxial surface strigose or strigose-pubescent, glabrescent, trichomes with expanded base apparently glandular. Co-inflorescence pluricephala, corymbiform; capitula discoid, pedunculated; involucre 1.2–1.6×1.5–2.0 cm, campanulate, 3-seriate; involucral bracts subimbricate, chartaceous or membranaceous, densely strigose or strigose-sericeous, glandular, tector trichomes cylindrical, apex acute, glandular trichomes rounded or cylindrical with apex capitate; receptacle convex. Flowers dimorphic, 60–80 flowers per capitulum; periphery flowers bisexual; corolla 1.6–2.0 cm long, bilabiate-liguliform, 4-nerves; disc flowers bisexual; corolla 1.3–1.5 cm long, bilabiate; both periphery and disc flowers yellowish or orangish; corolla-limb glabrous in outer portion, strigose-pubescent in inner portion; lobes glabrous in outer and inner portion; anthers basal appendage calcarate, apical appendage apiculate; style branches truncate, penicellate. Cypselae narrowly cylindrical, densely glandular, trichomes cylindrical, apex capitate, 8–10-ribs; carpopodium inconspicuous; pappus 1-seriate, barbellate, stramineous.

Examined material: Brazil: Caparaó National Park, 1.800 m, 18 May 1971 (fl., fr.), A.P. Duarte 13972 (RB 149003); ibidem, “Pico da Bandeira”, 1.500-1.600 m, 06 September 1977 (fl., fr.), G.J. Shepherd et al. 5764 (UEC2904). Espírito Santo: Caparaó National Park, nearly “Pico da Bandeira”, 2.600-2.800 m, 05 February 1985 (fl., fr.), P.E. Berry et al. 4526 (RB363245). Dores do Rio Preto, Caparaó National Park, 20 June 2005 (fl., fr.), L.S. Leoni 6198 (HUFU83415). Ibitirama, Caparaó National Park, “Casa Queimada” trail, High-altitude grassland, 20°26’17”S, 41°47’56”W, 2.810 m, 10 May 2014 (fl., fr.), A. Salino et al. s.n. (BHCB169855). Minas Gerais: Caparaó mountain, 2.200 m, September 1941 (fr.), A.C. Brade 17063 (RB45908); Caparaó National Park, path to “Pico da Bandeira”, 1.500-1.600 m, 06 September 1977 (fl., fr.), G.J. Shepherd et al. 5764 (UEC2904). Alto Caparaó, Caparaó National Park, High-altitude grassland, 21 March 2012 (fl., fr.), J. Kuntz et al. 582 (RB674530); ibidem, “Terreirão”, High-altitude grassland, 2.370 m, 27 March 1996 (fl., fr.), L.S. Leoni 3242 (GFJP3512); ibidem, nearly “Tronqueira”, 1.500 m, June 1996 (fl., fr.), L.S. Leoni 3389 (GFJP3724); ibidem, “Aurélio” waterfall trail, 20.4333°S, 41.8683°W, 30 August 2013 (fl., fr.), V.C. Manhães et al. 342 (UEC194010); ibidem, “Pico da Bandeira” trail, High-altitude grassland, humid area, 20°25’47”S, 41°48’03”W, 2.656 m, 17 March 2014 (fl., fr.), M. Monge et al. 2656 (UEC193483); ibidem, trail between “Tronqueira” and “Pico da Bandeira”, High-altitude grassland, 20°25’15”S, 41°51’12.6”W, 2.000-2.890 m, 11 April 2010 (fl., fr.), G.O. Romão et al. 2729 (HUFU74962, RB575037); ibidem, “Pico da Bandeira” trail, High-altitude grassland, 20°25’S, 41°48’W, 1.790-2.400 m, 12 February 1998 (fl., fr.), J.P. Souza et al. 2130 (HUFU56669); ibidem, “Aurélio” waterfall trail, 20°28’49”S, 41°49’59”W, 1.848 m, 2013 (fr.), J.P.F. Zorzanelli 1471 (HUFU83602); ibidem, “Aurélio” waterfall trail, 18 June 2013 (fl., fr.), J.P.F. Zorzanelli et al. 752 (HUFU83416). Espera Feliz, Caparaó National Park, 20.2733°S, 41.4836°W, 13 August 2011 (fl., fr.), M. Monge et al. 981Y37 (UEC201208).

Distribution, phenology and conservation: Trixis verbascifolia was collected in Montane Semideciduous Forest and High-altitude grassland, in humid or dry locations, at 1.500–2.890 m asl. Its reproductive period is during March to September. The species is restricted to Brazilian Atlantic Forest. Regarding conservation status, Trixis verbascifolia is not evaluated.

Taxonomic comments: Trixis verbascifolia is easily recognized by its branches winged, leaves homomorphic, sessile, elliptic or lanceolate with decurrent base, 60–80 flowers per capitulum, involucre 3-seriate and co-inflorescence laxa, pluricephalous, corymbiform.

DISCUSSION

Floristic inventory

Mutisieae and Nassauvieae stood out as the most diverse among the basal grade of Asteraceae in Caparaó National Park. These tribes also exhibit the greatest species richness, overall, compared to the other early-diverging tribes within the family (Susanna et al. 2020). Mutisieae and Nassauvieae contain a total of 254 species and 313 species, respectively, both primarily distributed throughout South America (Katinas et al. 2008, Funk et al. 2009, Susanna et al. 2020). In Brazil, the Atlantic Forest has the greatest representation of Mutisieae (4 genera, 16 species) and Nassauvieae (6 genera, 27 species), with Trixis (9 spp.) and Chaptalia (7 spp.) being the most diverse genera for the tribes in the domain (Flora e Funga do Brasil 2023) and the richest genera of basal clades in CNP. Based on the total number of species in the Atlantic Forest, CNP contain 57% of the species of Chaptalia and 44% of the species of Trixis.

The Caparaó National Park protects populations of six species (40% of total) with narrow distributions, including Mutisia lutzii, which is endemic to the Caparaó mountain. This highlights the importance of preserving natural environments, especially biodiversity hotspots. Since such hotspots harbor a large diversity of species with restricted distributions, the failure to preserve them would drive many species to extinction. It is important to consider that distributions can be extremely limited, such as with M. lutzii, which only occurs at a single specific location.

This study documented Chaptalia integerrima, Moquiniastrum polymorphum and Mutisia coccinea in CNP for the first time, expanding the latest list compiled by Moreira et al. (2020). Floristic inventories based on collections and field expeditions are an effective way to enhance our understanding of biodiversity (Funk 2006, 2018). In addition, accurate species recognition is crucial to establish a reliable listing.

Distribution, phenology and conservation

Data regarding the geographic distribution of seven species includes doubtful records that are difficult to validate. A shortfall in determining species distributions using databases has become apparent due to the lack of images of specimens in records, inconsistencies in data across online platforms, deficiencies in linking herbaria specimens to online databases, and incorrect or missing identifications. Consequently, it has become harder or even impossible to corroborate numerous records.

There is also a lack of information in online databases regarding the conservation status of the species found here. The majority of the species have not been evaluated or had equivocal data in CNCFlora, IUCN Red List or “Flora e Funga do Brasil”.

Considering ongoing environmental degradation, an effort must be made to evaluate the threat level of uncategorized species, particularly those with restricted distributions. Furthermore, there needs to be periodic reviews for species that have already been evaluated.

Systematic treatment

Half of the analyzed specimens (49 of 98 specimens, 50%) had their identifications corrected during the present study, highlighting the importance of species identification and the relevance of taxonomic works. Taxonomic studies enhance morphological knowledge of species and provide a review of specimens in herbaria, thus enabling the identification and/or correction of material. According to Funk et al. (2006), biodiversity studies will be better driven when biological collections and taxonomists are closely linked.

Taxonomic research is not only important for filling identification gaps in collections, but also plays a crucial role in supporting other biodiversity studies that require the recognition of taxa, consultation regarding plant morphology or the gathering of biogeographical and phenological data. According to Lagomarsino & Frost (2020), taxonomic studies are increasingly used to understand the ecological and evolutionary patterns of species, encompassing a wide range of scientific knowledge areas.

Although taxonomic studies are seen as less relevant by the scientific community, whereby the publication of studies becomes challenging due to page limitations of journals and the lack of acceptance in high-impact journals, they can contribute greatly to overall scientific progress and support species conservation planning.

Through taxonomy, science can recognize existing species and discover new ones (Joppa et al. 2011b). When there is knowledge about a species, it is possible to incorporate them into conservation planning, thereby increasing the probability of their protection. On the other hand, if a species remains unknown, the capacity for their protection decreases (Joppa et al. 2011a, Lagomarsino & Frost 2020).

CONCLUSIONS

Caparaó National Park harbors a diverse flora of basal clades of Asteraceae, including species with restricted distributions and endemic species, such as Mutisia lutzii. An intensive floristic survey encompassing all seasons of the year should be undertaken in CNP and its surrounding areas. It is likely that additional unrecorded and potentially endemic species occur in the Caparaó region.

The western portion of CNP has been little investigated due to difficult access, and the western side of the mountain peak remains unexplored. The management of CNP should consider the creation of a trail for access to the peak of the west, to allow collections that can contribute to understanding the Caparaó flora. The surrounding areas of the park also require attention, as they include mountain ranges located within a biodiversity hotspot, and probably harbor a high level of endemism. Meanwhile, these areas are currently being disturbed by extensive coffee monocultures.

A notable shortfall for online databases regarding the conservation status of species reveals the need for further studies to evaluate the threat level of plant species. The presence of dubious records for seven species highlights the importance of having images accompany records and accurate identifications of specimens.

Floristic inventories associated with taxonomic studies play a significant role in science. Collections and taxonomy can lead to a better understanding of biodiversity and its distribution, as well as the identification of species to mitigate gaps caused by erroneous or missing identifications.

ACKNOWLEDGMENTS

The authors are grateful to Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for financial support, to the Herbário da Universidade do Estado de Minas Gerais – HUEMG for assistance during the field expedition and to Gabriel Merola Paranhos for supporting the macro photographs.

SUPPLEMENTARY MATERIAL

Table SI.

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Publication Dates

• Publication in this collection
  21 Oct 2024
• Date of issue
  2024

History

• Received
  06 Oct 2023
• Accepted
  16 May 2024