Research Article
Research Article
A new epilithic bromeliad (Stigmatodon, Tillandsioideae) from Pedra dos Três Pontões, Espírito Santo, Brazil, with an emended combination for the genus
expand article infoDayvid Rodrigues Couto, André Pavioti Fontana, Andrea Ferreira da Costa§, Claudio Nicoletti de Fraga|
‡ Instituto Nacional da Mata Atlântica, Santa Teresa, Brazil
§ Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil
| Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, Rio de Janeiro, Brazil
Open Access


Background and aimsStigmatodon (Bromeliaceae) is a monophyletic genus currently comprising 33 species, all endemic to rocky outcrops, mainly inselbergs, from the Atlantic Forest and campos rupestres of the Espinhaço range, Brazil.

Material and methods – Our fieldwork in the inselbergs of the Atlantic Forest of the state of Espírito Santo led to the discovery of a new Stigmatodon species with a tubo-laciniate stigma type. Standard herbarium taxonomy practices were adopted to study the novelty. Morphological data were obtained from herbarium specimens and fresh material collected in the field.

Key resultsStigmatodon lemeanus is here described and illustrated. Diagnostic characters and affinities of the new species are discussed, accompanied by notes on its ecology, geographic distribution, and conservation status. A key to the species of Stigmatodon from Espírito Santo state is provided. The novelty is similar to S. goniorachis from coastal inselbergs of Rio de Janeiro and S. attenuatoides, another inselberg endemic from southern Espírito Santo. Stigmatodon lemeanus is preliminarily assessed as Vulnerable (VU) according to the IUCN Red List criteria. Additionally, we present a new combination of Tillandsia oligantha in Stigmatodon, correcting a nomenclatural error.


Atlantic Forest, Brazilian endemics, campo rupestre, inselbergs, taxonomy, Vrieseinae


On isolated granitic and/or gneissic outcrops, known as inselbergs, plant communities with different ecological requirements inhabit different vegetational habitats, such as mats, shallow depressions, ephemeral flush vegetation, rock pools, and crevices (Porembski and Barthlott 2000; Porembski 2007; Paula et al. 2016). In Sugarloaf Land, in south-eastern Brazil, vertical rock walls of inselbergs are colonized by specialized epilithic bromeliads (mainly Alcantarea (É.Morren ex Mez) Harms, Stigmatodon Leme, G.K.Br. & Barfuss, and Tillandsia L.), forming impressive vertical mats (Couto et al. in press). These extremophile mat communities occur in one of the most extreme and neglected habitats in the tropics, where new species of bromeliads are constantly discovered, mainly related to the genus Stigmatodon, which is endemic to Brazil (Leme et al. 2010a, 2014, 2022; Couto et al. 2020a, 2020b).

Stigmatodon (Bromeliaceae, Tillandsioideae) was created to accommodate 18 lithophytic species that were formerly included in Vriesea Lindl. (Barfuss et al. 2016). The first broad phylogenetic analysis focussing on Stigmatodon, using two plastid markers (matK and rps16-trnK) and the nuclear gene PHYC, was recently published, including the reconstruction of ancestral states for ecological and morphological characters (Couto et al. 2022). Accordingly, the taxonomic limits of Stigmatodon proposed by Barfuss et al. (2016) were expanded to include the eight species of the “Vriesea limae L.B.Sm. complex” (sensu Leme and Siqueira Filho 2006). The genus is morphologically characterized by (1) the semi-xeromorphic to xeromorphic, coriaceous, and usually triangular leaves with truncate margins; (2) the stamens usually positioned in two groups of three on each lateral side of the corolla; (3) the stigma of the tubo-laciniate or convolute-blade type; (4) the stomata usually positioned above the ordinary epidermal cells; and (5) the water-storage parenchyma differentiated on both surfaces (Barfuss et al. 2016; Couto et al. 2022).

Currently, Stigmatodon comprises 33 species (Gouda et al. 2023), 15 of them recently added to the genus (Couto et al. 2020a, 2020b, 2022, 2023; Leme et al. 2022). They occur on inselbergs within the Atlantic Forest of the states of Alagoas (AL), Bahia (BA), Espírito Santo (ES), Minas Gerais (MG), Pernambuco (PE), and Rio de Janeiro (RJ), and in campos rupestres of the Espinhaço mountain range from BA and MG (Couto et al. 2022). Species in Stigmatodon s.s. (sensu Barfuss et al. 2016), with tubo-laciniate stigma type I—with the exception of S. plurifolius (Leme) Leme, G.K.Br. & Barfuss (Leme 1987: 314; Barfuss et al. 2016: 58), which has tubo-laciniate stigma type II (Leme et al. 2022)—are ‘rock-climber bromeliads’, growing exclusively on vertical rock walls on dome-shaped inselbergs, in south-eastern Brazil (BA, ES, MG, and RJ). According to Leme et al. (2022), the Stigmatodon limae group includes species with three different stigma types: (1) tubo-laciniate stigma type II, observed in S. ilhanus Leme & D.R.Couto (Leme et al. 2022: 239) and S. rosulatulus (Leme) Leme, G.K.Br. & Barfuss (Leme 2012: 10; Barfuss et al. 2016: 58); (2) convolute blade type III (stigmatodontoid type), observed in S. andaraiensis (Leme) D.R.Couto & A.F.Costa (Leme 2012: 16; Couto et al. 2022: 352), S. limae (L.B.Sm.) D.R.Couto & A.F.Costa (Smith 1970: 181; Couto et al. 2022: 354), and S. zonatus (Leme & J.A.Siqueira) D.R.Couto & A.F.Costa (Leme and Siqueira Filho 2006: 374; Couto et al. 2022: 354); and (3) convolute blade type II (vrieseoid type), observed in S. vellozicolus (Leme & J.A.Siqueira) D.R.Couto & A.F.Costa (Leme and Siqueira Filho 2006: 406; Couto et al. 2022: 354), S. freicanecanus (J.A.Siqueira & Leme) D.R.Couto & A.F.Costa (Leme and Siqueira Filho 2006: 377; Couto et al. 2022: 352), S. vexatus (Leme) Leme & D.R.Couto (Leme 2017: 137; Leme et al. 2022: 245), and S. oliganthus (Baker) D.R.Couto & A.F.Costa, whose name was not validly published. These species grow on different rock types (granite, gneiss, and quartzite outcrops), on different degrees of rock inclination, including vertical to slightly inclined rock slopes, flat outcrops, or more rarely as epiphytes on stems of Vellozia J.Agardh (Leme and Siqueira Filho 2006; Cacossi et al. 2019; Couto et al. 2022).

A total of 16 species of Stigmatodon occur in the State of Espírito Santo, which encompasses several mountain ranges. In the central region of the State rises the Serra do Castelo, or Serra Capixaba (Chiron and Bolsanello 2015), consisting of imposing granitic mountains such as the Pico do Forno Grande (2,039 m) in the Castelo County, the Pedra Azul (1,822 m) in Domingos Martins, and the Pedra dos Três Pontões (1,968 m) in Afonso Cláudio, all of which harbour a diverse vascular flora (Pinto-Junior et al. 2021; Marcusso et al. 2022). In the last decades, several new plant species were described from the Pedra dos Três Pontões, most being narrow endemics, namely Mandevilla declinata J.F.Morales, A.P.Fontana & L.Kollmann (Apocynaceae, Morales et al. 2022: 1081), Pleroma subsessilis F.S.Mey. & L.Kollmann (Melastomataceae, Meyer et al. 2016: 202), and the bromeliads Alcantarea glaucifolia Leme & L.Kollmann (Leme and Kollmann 2016: 156), Orthophytum boudetianum Leme & L.Kollmann (Leme and Kollmann 2007: 149), and Pitcairnia diversifolia Leme & A.P.Fontana (Leme et al. 2010b: 491).

During field expeditions, a new rock-climbing species of Stigmatodon was found in this locality, growing exclusively on large inselberg vertical walls. This new taxon is described and illustrated in this work, its conservation status assessed, and its taxonomic affinities discussed. In addition, this paper provides an emended key for the species of Stigmatodon from Espírito Santo and a nomenclatural note for Stigmatodon oliganthus.

Material and Methods

Morphological data were obtained by studying herbarium specimens and fresh material collected randomly in pre-selected sites during fieldwork in Bahia and Espírito Santo. Descriptions and illustrations are based on both living and dried material, which were studied using a stereomicroscope Motic K700; the descriptive terminology follows Stearn (1973), Radford (1986), Smith and Downs (1977), with the modifications suggested by Scharf and Gouda (2008), and Leme et al. (2022) for stigma types. Voucher specimens were dried and pressed according to Fidalgo and Bononi (1984) and deposited at MBML, RB, and R (acronyms following Thiers 2023). The nomenclature follows the Shenzhen Code (Turland et al. 2018).

Data on the distribution of the new and related species, gathered from field notes and herbarium labels, were plotted on a map using DIVA-GIS v.5.2 (Hijmans et al. 2005), over a layer of inselbergs prepared by IEMA (2016). The conservation assessment complies with the criteria of the IUCN (2022), with the extent of occurrence (EOO) and the area of occupancy (AOO) estimated with GeoCAT and the AOO based on a user-defined cell size of 2 km2 (Bachman et al. 2011).

Taxonomic treatment

Stigmatodon lemeanus D.R.Couto, A.P.Fontana & Fraga, sp. nov.

Figs 1, 2, 3, 4


BRAZIL – Espírito Santo • Afonso Cláudio, Pedra dos Três Pontões, inselberg, epilithic on vertical rock wall; 20°04’33”S, 41°02’39”W; 1078 m; 21 Oct. 2019; fl.; D.R. Couto & C.N. Fraga 4926; holotype: MBML [MBML055811]; isotype: RB.


Similar to Stigmatodon goniorachis (Baker) Leme, G.K.Br. & Barfuss but distinguished by the suberect to patent peduncle (vs erect to suberect) that is more robust with a diameter of 6.5–7.8 mm (vs 3.5–6.2 mm), the main axis with shorter internodes (8.5–9.5 mm vs 15–25 mm long) that supports fewer flowers (11–13 vs 14–32), the longer flowers (56–60 mm vs 35–40 mm long), the larger petals (38–39 × 17–21 mm vs 22–28 × 13–15 mm), the basal appendages with irregularly acute-dentate apex (vs irregularly bidentate), and the shorter stamens (19–22 mm vs 26–28 mm long).

Figure 1. 

Stigmatodon lemeanus. A. Habit. B. Leaf. C. Peduncle bract. D. Inflorescence before anthesis. E. Flower, frontal view with detail of the stamens arranged in two groups of three on each lateral side of the corolla. F. Flower, lateral view. G. Floral bract, adaxial side. H. Floral bract, abaxial side. I. Floral bract, detail of the thickest part at the base. J. Petal with two basal appendages. K. Basal appendages, details. L. Stamen. M. Pistil. From the holotype D.R. Couto & C.N. Fraga 4926 (MBML). Drawn by Joelcio Freitas.


Lithophytic herb, 50–67 cm tall when flowering, propagating by basal shoots. Stem long, creeping, 7–20 cm in length. Leaves 12–20 in number, rosulate, thickly coriaceous, forming a funnelform rosette; sheath elliptic, 10–14 × 6–8.5 cm, densely and minutely castaneous-lepidote on both sides; blade narrowly triangular, apex acuminate-caudate, recurved, flat, suberect to arcuate, (11–)17–20 cm long, 2–2.8 cm wide at the base, green on both sides and usually with purplish-vinaceous spots on the adaxial side, usually vinaceous along margins, densely white-lepidote on both sides, with trichomes forming white crossbands adaxially, not obscuring the green colour of the blade, margins truncate, ca 0.5 mm thick. Inflorescence simple, ascending, 14–17 cm long, apex obtuse before and at anthesis, 11–13 flowered; peduncle suberect to patent, 35–42 cm long, 6.5–7.8 mm in diameter, green, glabrous, smooth at anthesis, sulcate in sicco; peduncle bracts subfoliaceous at the base, ovate above, apex obtuse-acuminate to apiculate, 2.5–3.5 × 2.4–2.8 cm, erect, imbricate, exceeding the internodes, green and usually with purplish-vinaceous spots on the adaxial side, densely lepidote abaxially, sparsely white-lepidote becoming denser towards the apex adaxially, smooth at anthesis; main axis smooth at anthesis, sulcate in sicco, green, glabrous, internodes 8.5–9.5 mm long; floral bracts broadly ovate, apex obtuse, 27–30 × 40–42 mm, shorter than sepals, green at anthesis, densely lepidote abaxially, adaxially glabrous, ecarinate, secund with the flowers before and during anthesis, coriaceous, thicker toward the base, smooth at anthesis, corrugate-sulcate in sicco. Flowers 56–60 mm long, night-blooming, with garlic scent and producing translucent mucilage, distichous, densely disposed, secund at anthesis; pedicel 9.5–11 mm long, 9–11 mm in diameter at distal end, stout, green, glabrous; sepals obovate, apex obtuse to rounded, (22–) 30–35 × (15–)21–23.5 mm, green, glabrous on adaxial side, densely lepidote abaxially, thick and coriaceous near the base, margins membranaceous; corolla 33–37 mm in diameter; petals obovate, apex rounded to emarginate, recurved near the apex at anthesis, 38–39 × 17–21 mm, greenish-white, strongly thickened with membranous margins, bearing 2 appendages at the base; appendages 9.5–11.6 × 3–3.8 mm, spathulate, basally adnate to the petals for 4.5–6.5 mm, apex irregularly acute-dentate; filaments free, complanate, 19–22 × 1.7–2 mm wide, white; anthers 8.5–9.5 mm long, dorsifixed near the base, base bilobed, apex obtuse, arranged in 2 groups of three on each lateral side of the corolla at anthesis; pistil equalling or exceeding the stamens; ovary superior, 5.6–7.4 mm long; ovules caudate; style 24.5–27 mm long, green; stigma tubo-laciniate type I, denticulate at the margins, 1.7–1.8 mm in diameter, green. Fruits a capsule, ellipsoid, apex apiculate, 32–45 mm long, 9.5–10.5 mm diameter, brown. Seeds fusiform, 3–4 mm long, brown; basal coma 15–21 mm long, beige; apical coma 1–1.6 mm, beige.

Figure 2. 

Stigmatodon lemeanus. AB. Plants in situ at the type locality. C. Flowering individual in cultivation. D. Lateral view of the flower and floral bract. E. Frontal view of the flower. F. Details of the tubo-laciniate stigma and anthers. Photographs: A, C–F by Dayvid R. Couto, B by André P. Fontana.


Stigmatodon lemeanus is endemic to Pedra dos Três Pontões, from the county of Afonso Cláudio in the central region of Espírito Santo, Brazil (Fig. 3), growing on bare and vertical walls of inselbergs, 1,000 m a.s.l. (Fig. 4). The Municipality of Afonso Cláudio recently created the Conservation Unit “Monumento Natural Municipal Pedra dos Três Pontões” to protect the local fauna and flora.

Figure 3. 

Geographic distribution of the 16 species of Stigmatodon on the inselbergs of Espírito Santo. A. S. amadoi, S. apparicianus, S. attenuatoides, S. belloi, S. bifidus. B. S. euclidianus, S. fontellanus, S. funebris, S. harrylutheri, S. magnibracteatus. C. S. multifoliatus, S. plurifolius, S. sanctateresensis, S. vellozicolus, S. viridibracteatus. D. S. lemeanus.

Figure 4. 

Stigmatodon lemeanus. AB. General view of the landscape at the type locality, highlighting the large and vertical pointed peaks of Pedra dos Três Pontões. C. Detail of the population on a vertical rock wall alongside sparse individuals of Pitcairnia diversifolia; some individuals grow close to the base of the slope, which facilitated the collection of some specimens for the study and description of the new species. Photographs: A by André P. Fontana, B–C by Claudio N. Fraga.

Habitat and ecology

The plant is epilithic, heliophilic, living on bare and vertical rock walls of inselbergs, where it forms large populations (Figs 2A, 4C), surrounded by Atlantic Forest fragments and farmland. In the type locality, Stigmatodon lemeanus grows on vertical rock walls (Fig. 2A–B), including on the large and pointed peaks of the mountain, from which the name “Pedra dos Três Pontões” is derived (Fig. 4B). In some places, they grow alongside sparse individuals of Pitcairnia diversifolia (Bromeliaceae), Mandevilla declinata (Apocynaceae), and the orchids Bulbophyllum weddellii Rchb.f., Bifrenaria harrisoniae (Hook.) Rchb.f., and Pseudolaelia dutrae Ruschi.


Flowers were recorded in October and fruits from May to October.


The specific epithet honours our friend Dr Elton Martinez Carvalho Leme (1960−), from Rio de Janeiro, Brazil, whose personal efforts have substantially contributed to the knowledge of the diversity, taxonomy, and systematics of Brazilian bromeliads. In addition, Dr Elton Leme is one of the authors of the genus Stigmatodon and has greatly contributed to the knowledge of these plants.

Preliminary IUCN conservation assessment

Vulnerable: VU D2. The new species is known from only one very small population within a protected area, with an AOO of 4 km2. These figures fall within the limits for Critically Endangered (CR), based on criterion B2 (area of occupancy), taking into account only the number of locations or severely fragmented areas, which can lead to a continuous decline in the extent of occurrence, area of occupancy and quality of habitat of the new species. However, population isolation and fragmentation are natural conditions on inselbergs, thus evaluations of plant distribution on inselbergs probably yield overestimated values of AOO and EOO (Leme et al. 2022). We do not have reliable information about this species’ continuing decline in plant numbers or extreme population fluctuations to enable us to apply criterion B. The Pedra dos Três Pontões inselberg is included in the Monumento Natural Municipal Pedra dos Três Pontões, an integral protection unit with the basic objective of preserving nature, with only the indirect use of its natural resources being permitted. Therefore, we estimate the human impact to be neither high nor continuous on the bare granitic and vertical walls of the inselbergs where it occurs. Due to the apparent rarity and very narrow distribution (a single location and AOO < 20 km2), this species is prone to the effects of stochastic events in the near future, such as fire or impact from climbers. Therefore, it seems prudent to include it in the Vulnerable category: VU D2.

Additional material examined

BRAZIL – Espírito Santo • Afonso Cláudio, Pedra dos Três Pontões, inselberg, epilithic on vertical rock wall; 20°04’33”S, 41°02’39”W; 1078 m; 21 Oct. 2019; fr.; D.R. Couto & C.N. Fraga 4921; R • Afonso Cláudio, Pedra dos Três Pontões, rupícola no sol em paredão vertical; 18 May 2007; fr.; A.P. Fontana et al. 3386; MBML [MBML00029444].


Stigmatodon lemeanus is morphologically related to S. goniorachis, differing by the fewer number of leaves per rosette up to 20 (vs up to 30), acuminate-caudate leaf blade apex (vs acute apex), green, and usually with purplish-wine spots in the adaxial surface, usually vinaceous along margins (vs green); ovate peduncle bracts (vs elliptic); the main axis densely flowered at anthesis (vs laxly flowered), broadly ovate floral bracts (vs elliptic-ovate to suborbiculate) that are wider (40–43 mm vs 13–20 mm long); and the larger sepals (30–35 × 21–23.5 mm vs 21–30 × 15–19 mm), in addition to the characters cited in the diagnosis. Furthermore, while S. lemeanus is endemic to inselbergs in the mountainous region of Espírito Santo (> 1,000 m elevation), S. goniorachis is endemic to vertical walls of several coastal inselbergs by the sea, in the city of Rio de Janeiro, Brazil. When compared with S. attenuatoides D.R.Couto, Manhães & A.F.Costa (Couto et al. 2020a: 144), another pachypetalous species endemic to the inselbergs in the south of Espírito Santo, S. lemeanus can be distinguished by the distinctly smaller size when flowering (50–67 cm vs 90–105 cm tall); the acuminate-caudate apex of leaf blades (vs acute or acuminate), which are smaller (17–20 × 2–2.8 cm vs 20–25(–36) × 3–5.5 cm), green, and usually with purplish-wine spots in the adaxial surface, usually vinaceous along the margins, without epicuticular wax (vs green concolorous, covered by a thick layer of white epicuticular wax); the simple inflorescence (vs compound) with fewer flowers (11–13 vs 20–40); the larger flowers (56–60 mm vs 50–55 mm long) with obovate sepals (vs elliptic), which are longer and wider (30–35 × 21–23.5 mm vs 22–24 × 16–17 mm), and with larger corolla (33–37 mm vs 27–28 mm in diameter).

Stigmatodon oliganthus (Baker) D.R.Couto & A.F.Costa, comb. nov.

Tillandsia oligantha Baker, The Journal of Botany, British and Foreign 25: 345. 1887. (Baker 1887)

Vriesea oligantha (Baker) Mez (Mez 1894: 544).

Stigmatodon oliganthus (Baker) D.R.Couto & A.F.Costa, nom. inval. (Couto et al. 2022: 354).


BRAZIL – Minas Gerais • Serra d’Ouro Branco; 10 Nov. 1884; fl.; Glaziou 15472; holotype: K [K000322056]; isotype: P [P00753328].


Couto et al. (2022) proposed eight new combinations in Stigmatodon. However, in the combination of Tillandsia oligantha Baker, the protologue of the basionym was erroneously cited. To address this uncorrectable error under Art. 41.8 of the Shenzhen Code (Turland et al. 2018), the name is here validly published with the correct citation of the protologue as indicated by Art. 41.5 (see Ex. 14) of the same Code.

Key to the species of Stigmatodon in Espírito Santo state, Brazil

1. Stigma of the convolute-blade type II (vrieseoid type II), epiphyte on Vellozia in inselberg vegetation S. vellozicolus (Leme & J.A.Siqueira) D.R.Couto & A.F.Costa
Stigma of the tubo-laciniate type, epilithic on inselberg vertical rock wall 2
2. Simple inflorescence 3
Compound inflorescence 14
3. Leaf blades at least 10 cm long 4
Leaf blades less than 10 cm long 13
4. Leaf blade less than 1 cm wide at the base, utricular rosette, forming non-impounding rosettes S. plurifolius (Leme) Leme, G.K.Br. & Barfuss
Leaf blade at least 1 cm wide at the base, infundibuliform rosette, forming impounding rosettes 5
5. Floral bracts smooth at anthesis 6
Floral bracts strongly nervate-sulcate at anthesis 7
6. Leaf blade with acute apex, flowers ca 4 cm long, sepals with emarginate apex, petals slightly thickened S. fontellanus (Leme & G.K.Br.) Leme, G.K.Br. & Barfuss
Leaf blade with acuminate-caudate apex, flowers 5.5–6 cm long, sepals with obtuse apex, petals strongly thickened S. lemeanus D.R.Couto, A.P.Fontana & Fraga
7. Leaves, 28–45 in number, blade up to 2.5 cm wide, with a truncate margin (ending very abruptly as if cut straight across) less than 1 mm thick, peduncle bracts smooth at anthesis S. multifoliatus (Leme & G.K.Br) Leme, G.K.Br. & Barfuss
Leaves, 15–25 in number, blade more than 2.5 cm wide, margins distinctly truncate thicker than 1 mm, peduncle bracts strongly nervate-sulcate at anthesis 8
8. Greyish-green leaf blades S. apparicianus (E.Pereira & Reitz) Leme, G.K.Br. & Barfuss
Cinereous leaf blades 9
9. Peduncle bracts ovate-lanceolate, more than 45 cm long S. magnibracteatus (Leme & L.Kollmann) Leme, G.K.Br. & Barfuss
Peduncle bracts ovate, up to 45 cm long 10
10. Floral bracts green, smooth, inconspicuously corrugate-sulcate near the apex at anthesis S. viridibracteatus D.R.Couto, Fraga & Leme
Floral bracts castaneous, stramineous, strongly nervate-sulcate at anthesis 11
11. Leaf sheaths ovate-elliptic to elliptic, 9–12 cm wide, peduncle erect to suberect, flowers 4.7–5.1 cm long S. euclidianus (Leme & G.K.Br.) Leme, G.K.Br. & Barfuss
Leaf sheaths obovate to broadly elliptic, 7–8.5 cm wide, peduncle suberect, flowers 4–4.5 cm long 12
12. Leaf blade canaliculate toward the base, with margins 0.9–1 mm thick, peduncle bracts lax, sepals elliptic with obtuse-emarginate apex, appendages with bifid-caudate apex S. bifidus (Leme & L.Kollmann) Leme, G.K.Br. & Barfuss
Leaf blade flat toward the base, with margins 1.5–2 mm thick, peduncle bracts imbricate, sepal oblong to ovate with rounded apex, appendages with irregularly long-dentate to acuminate apex S. harrylutheri (Leme & G.K.Br.) Leme, G.K.Br. & Barfuss
13. Leaf sheath 8.5–10 cm long (longer than the leaf blade), leaf blade ligulate, 5.8–7.3 cm long, flowers 2–3 cm long S. amadoi (Leme) Leme, G.K.Br. & Barfuss
Leaf sheath 7–9.8 cm long (equaling to the leaf blade length), leaf blade triangular, 7–10 cm long, flowers 3.4–4 cm long S. sanctateresensis (Leme & L.Kollmann) Leme, G.K.Br. & Barfuss
14. Sublinear-triangular leaf blade, erect to the suberect peduncle, dark-wine floral bracts S. funebris (L.B.Sm.) Leme, G.K.Br. & Barfuss
Narrowly triangular leaf blade, peduncle strongly curved, vinaceous-green floral bracts 15
15. Leaf blades vinaceous-green, peduncle pending, 35–50 cm long S. belloi (Leme) Leme, G.K.Br. & Barfuss
Leaf blades green, concolorous, peduncle patent, 57–60 cm long S. attenuatoides D.R.Couto, Manhães & A.F.Costa


We thank Joelcio Freitas for the line drawing, the National Institute of the Atlantic Forest (INMA), in particular the staff of the herbarium MBML, and the Instituto de Pesquisas Jardim Botânico do Rio de Janeiro for sponsoring fieldwork in the state of Espírito Santo. DRC is supported by a research grant from the Nacional Council for Scientific and Technological Development (CNPq) in the Programa de Capacitação Institucional of the National Institute of the Atlantic Forest – INMA (PCI/INMA), from the Brazilian Ministry of Science, Technology, and Innovation (MCTI: grant no. 301141/2022-3). CNF’s biodiversity research is supported by grants from Fundação Carlos Chagas Filho de Amparo à Pesquisa do Estado do Rio de Janeiro (FAPERJ) through a Jovem Cientista do Nosso Estado Fellowship (grant no. 201280/2022) and CNPq in the Edital Universal (grant no. 422191/2021-3). AFC thanks to CNPq for the productivity grant (grant no. 311111/2021-1). Finally, we thank the reviewers and editor João Farminhão for their valuable suggestions that improved our manuscript.


  • Bachman S, Moat J, Hill AW, la Torre J, Scott B (2011) Supporting Red List threat assessments with GeoCAT: geospatial conservation assessment tool. ZooKeys 150: 117–126.
  • Baker JG (1887) A synopsis of Tillandsieae. The Journal of Botany, British and Foreign 25: 344–347.
  • Barfuss MHJ, Till W, Leme EMC, Pinzón JP, Manzanares JM, Halbritter H, Samuel R, Brown GK (2016) Taxonomic revision of Bromeliaceae subfam. Tillandsioideae based on a multi-locus DNA sequence phylogeny and morphology. Phytotaxa 279: 1–97.
  • Cacossi T, Dantas-Queiroz MV, Palma-Silva C (2019) Transferability of nuclear microsatellites markers to Vriesea oligantha (Bromeliaceae), an endemic species from Espinhaço Range, Brazil. Brazilian Journal of Botany 42: 727–733.
  • Chiron GR, Bolsanello RX (2015) Orchidées du Brésil - as orquídeas da Serra do Castelo (Espírito Santo - Brasil), vol. 1. Tropicalia, Voreppe, 1–371.
  • Couto DR, Manhães VC, Costa AF (2020a) Stigmatodon attenuatoides (Tillandsioideae, Bromeliaceae): a new mat-forming species on inselbergs of southeastern Brazil. Phytotaxa 461: 143–149.
  • Couto DR, Manhães VC, Costa AF (2020b) Stigmatodon francae (Tillandsioideae, Bromeliaceae): a new species from cliff ecosystems on inselbergs in southeastern Brazil. Novon 28: 275–280.
  • Couto DR, Kessous IM, Neves B, Paixão-Souza B, Faria CG, Barfuss MHJ, Salgueiro F, Sa-Haiad B, Costa AF (2022) Molecular phylogenetics and trait evolution in Stigmatodon (Bromeliaceae, Tillandsioideae), an endemic genus to Brazilian rocky outcrops. Systematic Botany 47: 347–362.
  • Couto DR, Gonella PM, Costa AF (2023) Stigmatodon enigmaticus (Bromeliaceae, Tillandsioideae), a new lithophytic species from the campos rupestres within the Brazilian Atlantic Forest. Phytotaxa 584: 207–215.
  • Couto DR, Porembski S, Barthlott W, Paula LFA (in press) Hyperepilithics – an overlooked life form of vascular plants on tropical vertical rock walls. Austral Ecology.
  • Fidalgo O, Bononi VLR (1984) Técnicas de coleta, preservação e herborização de material botânico. Manual n° 4. São Paulo, Instituto de Botânica, 1–61.
  • Gouda EJ, Butcher D, Gouda CS (2023) Encyclopaedia of Bromeliads, version 4. University Botanical Garden, Utrecht. [accessed 20.02.2023]
  • Hijmans RJ, Guarino L, Jarvis A, O’Brien R, Mathur P, Bussink C, Cruz M, Barrantes I, Rojas E (2005) DIVA-GIS: Version 5.2. Manual. California, Lizard Tech, Inc. & University of California. [accessed 26.04.2023]
  • Leme EMC (1987) Novas Bromeliáceas nativas do Brasil – IV. Bradea 39(4): 309–326.
  • Leme EMC (2012) Revision of the lithophytic Vriesea species from Minas Gerais (and vicinities), Brazil – Part V: two tubolaciniate stigma-type new species. Journal of the Bromeliad Society 62: 10–19.
  • Leme EMC (2017) Revision of the lithophytic Vriesea species of Minas Gerais, Brazil – Part VII: a new species from the summit of Pico da Aliança. Journal of the Bromeliad Society 66: 137–148.
  • Leme EMC, Siqueira Filho JA (2006) Taxonomia das bromélias dos fragmentos de mata atlântica de Pernambuco e Alagoas. In: Siqueira Filho JA, Leme EMC (Eds) Fragmentos de Mata Atlântica do Nordeste, Biodiversidade, Conservação e suas Bromélias. Andrea Jakobsson Estúdio, Rio de Janeiro, 191–381.
  • Leme EMC, Kollmann LJC (2016) A new Alcantarea species from Espírito Santo, Brazil. Journal of the Bromeliad Society 65(3): 154–165.
  • Leme EMC, Kollmann LJC (2007) Studies on Orthophytum – Part VI: three new species from Espírito Santo, Brazil. Journal of the Bromeliad Society 57: 149–158.
  • Leme EMC, Fraga CN, Kollmann LJC, Brown GK, Till W, Ribeiro OBC, Machado MC, Monteiro FJS, Fontana AP (2010a) Miscellaneous new species in the Brazilian Bromeliaceae. Rodriguésia 61: 21–67.
  • Leme EMC, Fontana AP, Halbritter H (2010b) Three new Pitcairnia species (Bromeliaceae) from the inselbergs of Espírito Santo, Brazil. Systematic Botany 35: 487–496.
  • Leme EMC, Couto DR, Kollmann LJC, Fraga CN (2022) Novelties in Stigmatodon (Bromeliaceae, Tillandsioideae), a genus endemic to Brazil: three new species, one new combination, and two new stigma types. Phytotaxa 576: 233–249.
  • Marcusso GM, Menini Neto L, Lombardi JA (2022) Plants in the clouds: vascular epiphytes of Pedra Azul, a mountain top in Espírito Santo, Southeastern Brazil. Rodriguésia 73: e02322020.
  • Mez C (1894) Bromeliaceae. In: von Martius CFP, Eichler AW, Urban I (Eds) Flora Brasiliensis, vol. 3(3). F. Fleischer, Leipzig, 425–816.
  • Paula LFA, Forzza RC, Neri AV, Bueno ML, Porembski S (2016) Sugar Loaf Land in south-eastern Brazil: a centre of diversity form at-forming bromeliads on inselbergs. Botanical Journal of the Linnean Society 181: 459–476.
  • Pinto-Junior HV, Villa PM, Pereira MCA, Menezes LFTD (2021) The pattern of high plant diversity of Neotropical inselbergs: highlighting endemic, threatened and unique species. Acta Botanica Brasilica 34: 645–661.
  • Porembski S, Barthlott W (2000) Granitic and gneissic outcrops (inselbergs) as centers of diversity for desiccation-tolerant vascular plants. Plant Ecology 151: 19–28.
  • Radford AE (1986) Fundamentals of Plant Systematics. Harper & Row Publ. Inc, New York, 1–512.
  • Scharf U, Gouda EJ (2008) Bringing Bromeliaceae back to homeland botany. Journal of the Bromeliad Society 58: 123–129.
  • Smith LB (1970) Notes on Bromeliaceae XXXI. Phytologia 20(3): 121–183.
  • Smith LB, Downs RJ (1977) Bromeliaceae (Tillandsioideae). Flora Neotropica Monograph, vol. 14. Hafner Press, New York, 663–1492.
  • Stearn WT (1973) Botanical Latin, Second Edition. David & Charles Newton Abbot, London, 1–592.
  • Turland NJ, Wiersema JH, Barrie FR, Greuter W, Hawksworth DL, Herendeen PS, Knapp S, Kusber W-H, Li D-Z, Marhold K, May TW, McNeill J, Monro AM, Prado J, Price MJ, Smith GF (2018) International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum Vegetabile 159. Koeltz Botanical Books, Glashütten, 1–254.
login to comment