Sauvagesia (Ochnaceae) in the campos rupestres of the Espinhaço Range, Brazil

Abstract The campos rupestres (rupestrian grasslands) of the Brazilian Espinhaço Range are renowned for harboring extraordinary plant diversity and endemism, yet many groups representative of this habitat remain taxonomically understudied. Sauvagesia, a predominantly neotropical genus of Sauvagesieae (Ochnaceae), is a classic example. In this study, we refine the long-neglected and problematic taxonomy of the Espinhaço-Range-inhabiting species of Sauvagesia. Our taxonomic treatment includes 24 species of which seven are derived from recently described taxa or new combinations presented herein. For each of the three new combinations from variety into species status, we also provide new lectotypifications. The studied species are remarkably variable in habit, leaf, and floral architectures, and their distribution is often narrowly restricted to particular mountains across the Espinhaço Range, a pattern especially observed in the highly endemic S. subsect. Vellozianae. We provide an updated identification key, distribution maps, photographs, and a discussion of diagnostic morphological features.


Introduction
The ca. 1,000 km long S-shaped Espinhaço Range occupies an area of ca.120,000 km 2 in the center of Minas Gerais and Bahia states and is well known for the high rates of endemism of its species-rich and unique flora (Harley 1995;Giulietti et al. 1997;Rapini et al. 2008).Whereas the northern Espinhaço Range is nested in the Rodriguésia 74: e00992022.2023 Caatinga seasonally dry woodland, the southern part lies between two South American biodiversity hotspots (Myers et al. 2000): the Atlantic tropical rain forest to the east and the Cerrado (savanna vegetation) to the west.Most of the remarkably scenic landscape of the Espinhaço Range is, however, covered by the campo rupestre, a savanna-like vegetation closely associated with quartzite and ironstone outcrops on mountain tops at altitudes above 900 m a.s.l.isolated by vallies and major depressions (Silveira et al. 2015).Soil conditions, temperature and wind variation, water stress, and fire regimes have favored the convergent evolution of scleromorphic and ericoid plants in different angiosperm families (Giulietti et al. 1997;Silveira et al. 2015).
Although the Espinhaço Range represents less than 1.5% of the Brazilian territory, it harbors ca.10% of the entire flowering plant species reported for the country (Ribeiro et al. 2012), of which 30% are endemic to the campos rupestres (Giulietti et al. 1987).Endemism is concentrated in particular plant families, such as Asteraceae, Eriocaulaceae, Melastomataceae, and Velloziaceae.However, other taxonomically neglected families such as Ochnaceae have begun to catch the attention by the speed at which new species have been discovered over the last decade especially in the genus Sauvagesia L.: the focus of this treatment.
Sauvagesia has a pantropical distribution and ca.50 species, and it is the largest and morphologically most heterogeneous genus in Sauvagesieae (Schneider et al. 2014) with a complex taxonomic history.Recently, Schneider et al. (2021a,b) shed light on the phylogenetic understanding of Ochnaceae based on nuclear and plastid phylogenomic analyses, respectively.In addition to unraveling the internal relationships within Ochnaceae, these studies showed with maximum support the non-monophyletic nature of Sauvagesia.Strikingly, Sastre's (1981) subsect.Vellozianae appeared as a separate lineage with respect to the remaining Sauvagesia species.Interestingly, most species of subsect.Vellozianae are geographically localized, being mostly endemic to the campos rupestres vegetation in the ridges of the mountains across the Espinhaço Range in the Brazilian states of Minas Gerais and Bahia.A future revision of the entire Sauvagesia s.l. in a molecular phylogenetic context including a more comprehensive sampling would help to clarify the generic delimitation in the group.
In Brazil, Sauvagesia comprises 36 species, ca.80% of these species occur in the campos rupestres and cerrado vegetation (Cardoso et al. 2022).The significant diversity and endemism of Sauvagesia in such formations is evidenced by the numerous new species that have already been described from Bahia and Minas Gerais only in the last decade (Cardoso & Conceição 2008;Cardoso 2011;Cardoso & Harley 2015;Harley et al. 2005;Queiroz-Lima et al. 2017a,b, 2018;Zappi & Lucas 2002).Such recent discoveries clearly reinforce the need for a thorough floristic and taxonomic assessment of the genus involving a review of herbarium collections and extensive fieldwork, especially in the poorly-known campos rupestres areas that still remain understudied throughout the Espinhaço Range.

Material and Methods
The taxonomic treatment involves diagnostic morphological descriptions, an updated identification key, and comments on flowering phenology, geographical distribution, and diagnostic features which help to differentiate morphologically similar species.We reviewed the following herbaria known to archive important collections for the flora of the campos rupestres of the Espinhaço Range (acronyms are according to Thiers, continuously updated): Brazilian herbaria (ALCB, BHCB, BHZB, CEN, CEPEC, CTES, DIAM, ESA, FURB, HRB, HUEFS, HUFSJ, MBML, SP, SPF, UB, UEC, UFG, and UNB), and foreign herbaria (BR, F, FI, G-DC, K, MO, NHN, NY, P, and W).
The terminology for the morphological descriptions is based primarily on Radford et al. (1976) and Stearn (1992), while the other vegetative and reproductive structures follow Rizzini (1977), Weberling (1992), Harris & Harris (1994).Whenever specific terminologies are used on Sauvagesia morphology, these were based primarily on Sastre (1970Sastre ( , 1971aSastre ( ,b, 1973Sastre ( , 1978Sastre ( , 1981) ) and Harley et al. (2005).We also carried out extensive field expeditions to remote and underexplored areas in the Espinhaço Range, some of which were known to inhabit poorly collected taxa or promising sites to discover new species or to reveal new records.Photographs were taken for most species, and we followed the usual recommendations of drying and collecting plant specimens (Mori 2011) prior to being incorporated into the HUEFS herbarium.Duplicate specimens were also shared among different herbaria.
Leaves and flowers obtained during our fieldwork were fixed in 70% alcohol to enable an accurate examination of the specimens for description and illustration.In addition, silica-gel dried leaf samples were separated for future molecular phylogenetic investigation.
Currently, phylogenetic analyses of molecular data have left no doubt that Sauvagesia is a member of the Ochnaceae (Schneider et al. 2014(Schneider et al. , 2021a,b),b), where it appears as most closely related to the genera Luxemburgia A.St.-Hil., Philacra Dwyer, and Testulea Pellegr.
The pantropically-distributed genus Sauvagesia is represented by endemic species in China (one sp.), Malaysia (two spp.), and Africa (one sp.), however its species diversity is greatly concentrated in the Neotropics, mainly in Brazil, where 36 species have been recorded thus far (Amaral & Bittrich 2014;Cardoso et al. 2020).Sauvagesia species inhabit the understory of tropical rain forests or drier environments such as savannas and campos rupestres (rupestrian grasslands).The remarkable diversity of the genus in the scenic mountain tops of the Espinhaço Range , which is home to more than half of the species known in the genus, is perhaps another example of species radiation in the evolutionary theater of the campos rupestres (Ribeiro et al. 2012;Bitencourt & Rapini 2013;Conceição et al. 2016;Alcântara et al. 2018;Vasconcelos et al. 2020;Rapini et al. 2021).Subshrub up to 40 cm tall, slender, cylindrical, ericoid, the leaves shedding at the base of the branchlets.Flowers in this species open widely and have pale pink petals, while the corona-like internal staminodal whorl is pink, becoming white in the apex.Sauvagesia alpestris is vegetatively very similar to S. ericoides and for a long time they were treated as a single species (e.g., Sastre 1971aSastre , 1981)).Both species occur sympatrically in the campos rupestres of Serra do Caraça in Minas Gerais, however, the type of inflorescence in these species is of great importance to differentiate between them.We agree with Eichler (1871) and Zappi & Lucas (2002) who have considered S. alpestris as distinct species.The inflorescence in S. alpestris is a pedunculate lax panicle and the sepals ovate 2:1, while S. ericoides bears flowers on a terminal sessile fascicle and sepals lanceolate 3:1.Sauvagesia alpestris is currently known by only a few collections from the Santuário do Caraça Natural Heritage Private Reserve (RPPN in Portuguese) in Minas Gerais.It occupies the campo rupestre on the sandstone summit at ca. 1,400 m a.s.l.It has been found flowering from April to August, and fruiting from October to December.Shrub up to 1.7 m tall, erect, single-stemmed, branched mainly towards the apex, the branches 2-4 mm thick, slender, cylindrical; leaves and stipules densely grouped on rosette-like short fascicules, the whole plant body looks like it was entirely colonized by mosses; fruit-like galls scattered along the branches.Flowers in this species open widely and have intense pink petals turning white proximally and distally, while the corona-like internal staminodal whorl is pink, becoming white in the apex.
Sauvagesia bryoclada can be differentiated from S. elegantissima and S. spicata by its singlestemmed shrubby habit densely branched towards the apex (vs.poorly-branched, single-stemmed shrub in S. elegantissima and multi-stemmed shrub densely branched from the base in S. spicata), absence of glands in the leaf and sepal margins (vs.glands rare in S. elegantissima and often present in the sepals and leaves of branch apices in S. spicata), inflorescence a lax, 20-50-flowered, 1.2-3.8cm long bostryx (vs.congested, 10-20-flowered, 1.3-2.2cm long bostryx in S. elegantissima), pedicel 5−10 mm long (vs.3-6 mm long in S. elegantissima), reddish-pink and widely open petals resulting in a star-like flower shape (vs.nonstar-like flowers with pale-pink, strongly reflexed petals in S. elegantissima and star-like flowers with white petals in S. spicata).
Sauvagesia bryoclada has restricted distribution in the campo rupestre of the municipalities of Botumirim and Grão Mogol in Minas Gerais state, Brazil.It grows directly on rocks or in damp sandy soil at 1,000−1,300 m a.s.l.Flowering specimens have been collected from March to November and fruit specimens from July to November.(A.St.-Hil.)Sastre, Sellowia 23: 13. 1971.

Sauvagesia capillaris
Fig. 2a-b Shrub up to 1 m tall, branched, the branches, cylindrical, glabrous; leaves with different sizes in the same branch.Flowers in this species have revolute pale pink petals, while the corona-like internal staminodal whorl is pink, becoming white proximally and distally.Sauvagesia capillaris is morphologically very similar to S. glandulosopubescens.Indeed, both species were considered as a single species thus far (Saint-Hilaire 1823).Although they share a similar habit, S. capillaris readily differs from S. glanduloso-pubescens by the glabrous branches (vs.fimbriate-laminar glandular branches in S. glanduloso-pubescens), stipules with ca. 10 fimbriae (vs.ca.3), and a relatively smaller, 1.8-4 cm long inflorescence (vs.3.5-9 cm long).Sauvagesia capillaris occurs only in Minas Gerais state (Fig. 3c), where it dwells in sandstone campo rupestre, between cracks and depressions, and steep valleys in more humid shady areas, where streams are usually common.It occurs at 1,300-1,700 m a.s.l.It has been recorded flowering and fruiting all over the year.Cardoso, comb. nov. et  Fig. 1e-f Shrub 20-90 cm tall with erect, slender, cylindrical, and glabrous branches without leaves at the base.Flowers have revolute white petals, while the corona-like internal staminodal whorl is pink, becoming white in the apex.It has been historically related and confounded to the herein called glandular species complex (S.glandulosa and S. rubra), but it is readily distinguished by the combination of elliptic 2:1 leaves measuring 0.9-1.6 × 0.5-1.2cm (vs.leaves 1.6-4 × 0.8-2.1 cm and obovate, in S. glandulosa and 2-6 × 1-2.5 cm and obovate in S. rubra), sepals with transparent, conical, sessile glands along the margin (vs.sepals with stalked glands in S. rubra), petals 4-7 × 2.5-3 mm and pale pink to white (vs.petals 6-8 × 3-4 mm and white in S. glandulosa and 9-10 × 3-4 mm and deep-pink in S. rubra).Sauvagesia congesta also appears morphologically akin to S. lagevianae, but it differs on its elliptic leaves with acute apex, slightly serrate margin, and inconspicuous secondary veins (vs.leaves obovate with obtuse apex, sharply serrate margin, and distinct secondary veins), as well as the subcylindrical panicles (vs.umbellate inflorescences borne above a distinct peduncle on terminal branches) with flowers star-like in shape, the petals opening widely and reflexed (vs.campanulate-like flowers).

Sauvagesia congesta
In the original description of Lavradia glandulosa var.congesta Eichl., three syntype specimens were cited: Saint-Hilaire s.n., Riedel s.n.(Serra São José prope São João d'El Rey), and Lund s.n.(Serra Itabira do campo).One hundred years later, Sastre (1971a) synonymized this variety under a broad concept of Lavradia glandulosa A.St.-Hil., yet he erroneously assigned the same type collection as that of Lavradia glandulosa var.rubra A.St.-Hil.We were able to locate only Lund's duplicate specimen that is deposited at NY herbarium (no.NY00888225) and thus have we have chosen it for the lectotype.

Sauvagesia elegantissima
Shrub 1-1.5 m tall, erect, branched only toward the apex, the branches 2.5-4 mm thick, slender, cylindrical, covered with densely-grouped leaves and stipules in rosette-like short fascicles.Flowers in this species have revolute pale pink petals, while the corona-like internal staminodal whorl is pink, becoming white in the apex.Sauvagesia elegantissima is morphologically related to S. bryoclada and S. spicata.The taxonomic distinction of S. elegantissima is presented in the discussion of the other species.Sauvagesia elegantissima is narrowly distributed in the campo rupestre vegetation of the municipalities of Buenópolis and Diamantina in Minas Gerais state, Brazil, where it occurs in cracks between rocks at 1,200−1,300 m a.s.l.Flowering and fruiting individuals were observed from November to January.
Herb to subshrub 0.1-1 m tall, well branched, green to vinaceous, cylindrical.Flowers have flat or revolute white petals, with numerous free pink external staminodes around and internal whorl of five free pink petaloid staminodes, becoming greenish-white in the apex.The morphology of S. erecta is widely variable, probably as a result of its pantropical distribution, occupying many different environments, from the understory of tropical rain forests to the drier mountain tops of the Espinhaço Range.As such, the distribution of its leaves along the branches can be well spread or very close, sometimes becoming verticillate.Leaf size is also highly variable, as well as the habit.For example, we have seen delicate, herbaceous individuals measuring 10 cm tall to more robust, well-branched plants up to 1 m tall.In view of this polymorphism, the taxonomic circumscription of S. erecta has always been difficult, as expressed in the fact that it currently involves more than ten synonyms (Sastre 1968(Sastre , 1971b)).Regardless, S. erecta is easily differentiated from the remaining species in the campos rupestres by the combination of axillary solitary flowers with numerous external staminodes and five, free, petaloid internal staminodes.
Rodriguésia 74: e00992022.2023 7. Sauvagesia ericoides (A.St.-Hil.)Sastre, Sellowia 23: 15. 1971. Fig. 2e Subshrub up to 40 cm tall, densely branched from the base, branches erect.The flowers are pendent with petals opening widely and pale pink, the corona-like internal staminodal whorl pink, becoming white in the apex.The diagnostic features of S. ericoides with respect to its morphologically related S. alpestris were mentioned in the taxonomic comments about this species.Sauvagesia ericoides is likely narrowly endemic to the RPPN Santuário Serra do Caraça in the state of Minas Gerais, where it inhabits the sandstone campos rupestres in the mountain slopes in shady places at 1,750-2,030 m a.s.l.Flowering and fruiting specimens have been collected all over the year.(A.St.-Hil.)Sastre, Sellowia 23: 16. 1971.

Sauvagesia glandulosa
Fig. 1a-b Shrub up to 1 m tall, branches erect.The flowers are erect and star-like in shape, where the white petals open widely; the corona-like internal staminodal whorl becomes pink in the base.Sauvagesia glandulosa had a broad circumscription that encompassed the morphologically related S. rubra and S. congesta.Indeed, these species were earlier considered as varieties under S. glandulosa.Specific taxonomic considerations that help to differentiate S. glandulosa are given in the discussion of S. rubra and S. congesta.
Representative specimens examined: Belo Horizonte, Serra da Mutuca, 1,400 m a.s.l., 16.XI.1938, fl Sauvagesia glandulosa occurs only in the mountains of Minas Gerais state (Fig. 3d), on supergenic concretions of iron oxide and ferruginous concretions (campo cerrado ferruginoso locally known as canga), sandy soil between quartzite outcrops (campo rupestre) and sandy slopes, at 1,100-1,500 m a.s.l.It has been found in flower from October to February, and in fruit from December to April.Fig. 2c Subshrub up to 40 cm tall, well-branched.It was initially described as a variety of S. capillaris (Saint-Hilaire 1823).However, the cylindrical stems with glandular laminar indument, often fimbriate covering all branches in S. glandulosopubescens is the striking feature unknown in any species of the genus that has made us to recognize its species status.The flowers are pendent, with relatively small, pink petals opening widely, while the corona-like internal staminodal is reddishpink, becoming white in the base.A detailed morphological comparison of S. glanduloso-We found that the type collection of S. glanduloso-pubescens is composed of four gatherings made by A. Saint-Hilaire (three at P and one at MPU).We chose the gathering at P no.P02441344 for lectotype because it clearly refers to the same locality and taxon name as provided in the original publications (Saint-Hilaire 1823, 1829).
In Sastre's (1971a) taxonomic revision, when he lectotypified S. insignis he cited the wrong collection number "7899".This typo is corrected here so as the correct corresponding Ule's type material numbered "7099".
Sauvagesia insignis occurs in the campos rupestres of the Chapada Diamantina in Bahia state (Fig. 3b) at ca. 1,200 m a.s.l.Flowering specimens were collected from July to November, and fruiting set from September to May.Shrub up to 1.5 m tall, branches erect, without leaves at the base.The flowers pendent, opening widely, sometimes the petals reflexed distally, with reddish-pink petals and internal staminodal whorl; free external staminodes eventually present in some flowers.Sauvagesia insolita is morphologically related to S. paganuccii.However, S. insolita are single-stemmed, few-branched, erect shrubs (vs.multi-stemmed subshrubs that are densely branched from base in S. paganuccii) and has leaves not exceeding 45° to the branch (vs.leaves ca.90° to the branch), broadly elliptic to ovate with length/width ratio 2:1 (vs.narrowly elliptic leaves with length/ width ratio 3:1), and the margin homogeneously crenulate (vs.margin irregularly crenulate, the crenulae more spaced basally and closer distally).Moreover, based on the number of pedicel scars along the inflorescence axes, we estimate that S. insolita has a bloom of less than 20 flowers (vs.bloom of ca.70 flowers).
Sauvagesia lagevianae occurs in the Diamantina Plateau in Minas Gerais state (Fig. 3d), where it grows on quartzite-sandstone outcrops and sandy soils at ca. 1,500 m a.s.l.It has been found flowering from September to November and in April, and fruiting in October.A.St.-Hil., Bull. Soc. Phil. Paris 174. 1823. Fig. 2i Herb to shrub 15-60 cm tall, erect, mostly poorly branched at the apex.Flowers erect, the petals opening widely, white to pale-pink; the internal staminodia purplish.Sauvagesia linearifolia is morphologically similar to S. tenella.Both occur in periodically flooded environments, but their leaves are somewhat different.Sauvagesia linearifolia has obovate petals (vs.slightly unguiculate in S. tenella) that remain revolute in the upper half (vs.flat).Also, S. tenella has never been recorded in the Espinhaço Range.Amaral et al. 1973 (HUFU).

Sauvagesia linearifolia
Despite being a relatively widespread species in the savannas of Central Brazil, S. linearifolia is only known from a few collections in the campos rupestres of Bahia and Minas Gerais states.It usually grows on sandy and flooded soils at 380-767 m a.s.l.Flowering and fruiting specimens have been found during the whole year.Zappi & E. Lucas, Kew Bull. 57(3): 711. 2002. Fig. 2f Shrub to small tree up to 2.5 m tall, branches erect and without leaves at the base.Flowers erect with white petals opening widely and the internal staminodal whorl pink.Sauvagesia nitida is morphologically similar to S. insignis, but was also long confounded with S. glandulosa (Zappi & Lucas 2002).The morphological differences of S. nitida from S. insignis are presented in the taxonomic comments under this species.The main features that are helpful to distinguish S. nitida from S. glandulosa are the 6-9 mm long stipules (vs.9-13 mm long in S. glandulosa), leaf apex nonmucronate or only shortly mucronate (vs.distinctly mucronate), and the pink corona (vs.white with the base pink).
Sauvagesia oliveirae has been encountered thus far only in the Gobira peak of the Chapada Diamantina region (Fig. 3a), where it grows on sandy soil between rocks at ca. 1,500 m a.s.l.Flowering and fruiting specimens have been recorded from September to March.Shrub up to 1.2 m tall, densely branched from the base, the branches erect and leafless below.Flowers erect with petals reddish-pink and opening widely, and the internal staminodia reddish-pink.We have provided a morphological discussion for S. paganuccii in the taxonomic section of its morphologically closely related S. insolita.
Sauvagesia paganuccii is apparently endemic to Serra do Orobó in the municipality of Ruy Barbosa, Bahia, where it grows in more humid and shady rocky cliffs at ca. 1,000 m a.s.l.This campo rupestre vegetation is disjunct with the core area of the Serra do Sincorá in Chapada Diamantina.Flowering from August to January and fruiting from November to January.17. Sauvagesia paniculata D.B.O.S. Cardoso & A.A. Conc., Brittonia 60(4): 306. 2008.
Sauvagesia paniculata has been collected only in the Serra do Esbarrancado of the Chapada Diamantina in Bahia state (Fig. 3a) at ca. 1,500 m a.s.l., where it often occurs associated with rocky outcrops.Flowering and fruiting specimens were collected from January to September.pale-pink petals, external staminodes numerous, filamentous, and the petaloid internal staminodes completely free, pink, and with white apex.Sauvagesia racemosa is similar to the Amazonian species S. ramosa (Gleason) Sastre, from which it has usually been confused in herbarium collections.However, S. racemosa differs by the leaves scattered along the branches (vs.densely arranged in S. ramosa), acute or obtuse at the apex (vs.acuminate), the margin callose-crenate (vs.revolute), inflorescence a simple raceme 5-11 cm long (vs. compound  Sauvagesia racemosa has a wide distribution throughout the Brazilian savannas (Cerrado domain).In Brazil, this species has been recorded in the states of Bahia, Goiás, Mato Grosso, Mato Grosso do Sul, Minas Gerais, Paraná, Rio de Janeiro, São Paulo, and Tocantins.It is usually associated with sandy and flooded soils at 380-774 m a.s.l.Flowering and fruiting individuals were recorded during the whole year.Harley & Giul., Kew Bull. 60(4): 576. 2005.Shrub 0.5-1 m tall, erect, stems leafless below, well-branched to the apex, scars of the insertion of stipules very evident in branches.Flowers opening widely, the petals pink and the fused internal staminodes pink with white apex.Sauvagesia ribeiroi was largely misidentified among the herbarium collections as the morphologically very similar, acicular-leaved species S. semicylindrifolia.In addition to sharing needle-like leaves, both species have flowers with pink, star-like shaped corolla.However, S. ribeiroi differs by having a multi-stemmed subshrubby habit with the leaves never shedding at the base of the branches (vs.mostly single-stemmed, ericoid shrubs, the leaves more clustered at branchlet apices in S. semicylindrifolia), leaf lamina crenulate-glandular at the margin and measuring less than 15 mm long (vs.leaves entire and 20 cm long), and flowers with glandular sepals (vs.sepals without glands).TYPE: BRAZIL.Minas Gerais: "Ponte Alta, à la Serra d'Itambé, cap des Mines", 1817, A. Saint-Hilaire 409 (lectotype, here designated: P no.P02441378).

Sauvagesia ribeiroi
Fig. 1c-d Shrub up to 1 m tall, branched, erect, and without leaves at the base.The flowers are erect and with pink petals opening widely whereas the fused internal staminodes are pink, becoming white in the apex.Sauvagesia rubra is readily distinguished from its morphologically related S. glandulosa by the stipules and leaves loosely imbricate (vs. the branches densely covered by stipules and imbricate leaves in S. glandulosa), the leaf mucron at the apex measuring ca. 1 mm long (vs.ca. 5 mm long), larger, ca.22 cm long inflorescences (vs.ca. 12 cm long), stalked glands along the sepals margin (vs.conical glands), pink petals (vs.white), corona entirely pink (vs.white becoming pink in the base).This species was originally described as a variety under the name Lauradia glandulosa var.rubra (Saint-Hilaire, 1823, 1829).Three different locations ("Itambé", "Ponte Alta", and "Candonga") were cited in the original publications, for which we were able to locate seven gatherings at P and K herbaria.We chose the specimen P no.P02441378 for lectotype, because it is the most complete and has the associated labels clearly matching with the original locality "Ponte Alta".
Sauvagesia rubra occurs in Minas Gerais state (Fig. 3d), on sandy soil between quartzite outcrops (campo rupestre), swampy fields.It is always associated with places where water accumulates.It occurs at 680-1,300 m a.s.l.Flowering recorded from April to February and fruiting set from November to February.Sastre, Bull. Jard. Bot. Natl. Belg. 51(3-4): 398. 1981.Shrub 0.4-1.2m tall, erect, stems leafless below, well-branched to the apex, and scars of the insertion of stipules very evident in branches.Flowers, opening widely with pink petals and fused internal staminodes pink, with the apex white.Sauvagesia semicylindrifolia is the most widespread of the acicular-leaved Sauvagesia species, all of which are endemic of the campos rupestres in Chapada Diamantina of Bahia state (Fig. 3a).A detailed morphological comparison of S. semicylindrifolia is given in the taxonomic notes of S. ribeiroi.
Subshrub up to 40 cm tall, branched from the base, branches erect, ericoid, the leaves clustered at branch apices and caducous at the base.Flowers nearly campanulate-like, nodding, petals white, and the fused internal staminodes reddish-pink, becoming white in the apex.Sauvagesia setulosa is unlike among all species of Sauvagesia because of its dwarf subshrubby habit with a unique combination of morphological features: the remarkable bristle-bearing leaves and flowers with white petals that are borne on terminal, umbel-like fascicules of dichasial cymes.It can be tentatively associated to the species of subsect.Vellozianae that also have umbel-like inflorescences: S. ericoides, S. oliveirae, S. ribeiroi, and S. semicylindrifolia.However, most of these species grow mostly as shrubs up to 1.3 m tall, the leaves never bearing marginal stalked glands, and the flowers have pink petals.A notable, non-overlapping ecological feature of S. setulosa helps to distinguish it from the previously mentioned species: it often inhabits open grassy campo rupestre on sandy moist soils, whereas the others are always directly associated with rocks or sandy hillsides amongst rocks.1,194 m a.s.l., 16.XII.2011, fl. and fr., D.J.P. Gonçalves et al. 343 (DIAM, UEC); Parque Nacional das Sempre-Vivas, estrada para o alojamento, campo rupestre, solo arenoso úmido, 17°57'07 ''S, 43°46'44''W, 1,258 m a.s.l., 9.I.2016, fl. and fr., A. Queiroz-Lima et al. 183   Shrub up to 1-2 m tall, erect, branched mainly at the apex, branches conspicuously covered with densely-imbricate leaves and stipules on short rosette-like fascicles.Flowers erect, petals white, opening widely, resulting in a star-like shape, and the corona-like fused internal staminodia reddishpink.Sauvagesia spicata had long been treated as a synonym of the morphologically similar S. elegantissima (Sastre 1971), but it was recently re-established (Queiroz-Lima et al. 2018).This species is distinct within the S. elegantissima species complex (S.bryoclada, S. elegantissima, and S. spicata) by the larger flowers, the petals measuring 5-7 × 3-3.5 mm and staminodal whorl 3-6 mm long, the stalked-glands on the sepal margin, as well as the heteromorphic leaves, where the leaves towards the inflorescences are often glandular on the margin.Furthermore, S. spicata can be separated from S. elegantissima by its multistemmed shrubby habit that is densely branched from the base (vs.poorly-branched, single-stemmed shrub), the 4-8 mm thick branchlets (vs.slender, 2.5-4 mm thick), the lax, 20-30-flowered, 2.3-10 cm long bostryx inflorescences (vs.congested, 10-20-flowered, 1.3-2.2cm long bostryx), flowers with 4-10 cm long pedicels (vs.flowers with shorter, 3-6 cm long pedicels).We have also observed that the vegetative branches of S. spicata are often infested with galls that are deceptively similar in shape to the fruits.

Figure 3
Figure 3 -a-d.Geographical distribution of Sauvagesia species in the Brazilian Espinhaço Range -a. the acicularleaved species Sauvagesia oliveirae, S. paniculata, S. ribeiroi, and S. semicylindrifolia; b. the morphologically closely related species S. insignis and S. nitida; c. the S. capillaris species complex comprised of S. capillaris and S. glanduloso-pubescens; d. the closely related species S. congesta, S. glandulosa, S. rubra, and S. lagevianae, all of which largely marked by the gland-bearing leaves with expanded lamina.
raceme or panicles 10-17 cm long).With respect to the Sauvagesia species found in the Espinhaço Range, S. racemosa may only be confounded with some robust individuals of S. erecta that bear coriaceous leaves.Nevertheless, S. racemosa never has solitary flowers at leaf axils, which is a diagnostic feature for S. erecta.