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Research Article
A new dwarf palm from the campo rupestre: another piece of the Syagrus evansiana complex (Arecaceae)
expand article infoStephane da Silva Reis, Dayana Maria Teodoro Francino§, Bernardo Otávio Dias§, Ramon Martins Ferreira-Filho§, Lucas Giovanni Pinto, Elaine Lopes Pereira Nunes|, Bruno Francisco Sant’Anna-Santos
‡ Universidade Federal do Paraná, Curitiba, Brazil
§ Universidade Federal dos Vales do Jequitinhonha e Mucuri (UFVJM, Campus JK), Diamantina, Brazil
| National Coalition of Independent Scholars, Battleboro, United States of America
Open Access

Abstract

Background and aims – In the Espinhaço Range, the campos rupestres harbour many endemic acaulescent Syagrus. Populations assigned to Syagrus evansiana include geographically and ecologically distinct populations. We reassessed the two populations used in the original circumscription of S. evansiana and revealed that they represent distinct species.

Material and methods – We compared morphological and anatomical characters from field collections of both populations. The pinnae anatomy was studied using light microscopy of free-hand cross sections. A diagnostic comparison with other members of the S. evansiana complex is provided, together with a distribution map.

Key resultsSyagrus montana sp. nov. resembles S. evansiana but differs in leaf size, inflorescence architecture, and flower arrangement. Syagrus montana shows flowers only in triads, whereas S. evansiana frequently bears tetrads and pentads. Additional diagnostic traits include indumentum on the pistil of the flowers in pre-anthesis, staminode shape, fruit and endocarp size, and several pinnae anatomy characters. An accessory vascular bundle within the expansion tissue is recorded only in S. evansiana, first recorded for the genus.

Conclusion – Morphological and anatomical evidence, together with geographic isolation and habitat differences, supports recognition of Syagrus montana as a new species. Syagrus evansiana possess rare and exclusive morphological and anatomical characters. Our results highlight the need to reassess other poorly studied populations currently assigned as S. evansiana. In addition, the data presented here corroborate the Espinhaço Range and its disjunctions as one of the centres of diversity of Syagrus. The discovery of this new species reinforces the uniqueness of the local flora and its classification as a priority area for conservation.

Keywords

Meridional Espinhaço, Palmae, plant anatomy, taxonomy

Introduction

The Espinhaço Range is found in eastern Brazil and extends for approximately 1,200 km (Rapini et al. 2008; Silveira et al. 2016). It occurs in a transition region encompassing two biodiversity hotspots, the Atlantic Forest and the Cerrado, and due to its remarkable biological diversity the Southern Espinhaço Range was recognized as a Biosphere Reserve (UNESCO 2005). In this region, the campos rupestres stand out for their high rates of endemism (Silveira et al. 2016; Morellato and Silveira 2018). These environments harbour a high number of rare and endemic species, including acaulescent palm species of the genus Syagrus Mart. (Rabinowitz 1981; Noblick 2013).

The genus is monophyletic (Dransfield et al. 2005, 2008; Meerow et al. 2009), but Syagrus comprises several species whose circumscription is challenging due to strong morphological similarity—a taxonomic problem that is particularly severe in species with subterranean or short stems, commonly found in high-elevation environments such as the Southern Espinhaço Range (Noblick 2017a). Even experienced taxonomists have difficulty distinguishing acaulescent Syagrus (Marcato and Pirani 2001), both in the field and when analysing herbarium specimens, because diagnostic vegetative and reproductive characters used for identification—such as the arrangement and colour of pinnae—may be lost during pressing/drying or may be absent from specimen labels (Glassman 1972; Dransfield et al. 2008; Noblick 2009, 2013, 2017a, 2017b; Noblick and Lorenzi 2010; Firmo et al. 2021; Sant’Anna-Santos et al. 2023a, 2023b, 2023c, 2025).

Before Noblick and Lorenzi (2010), many acaulescent Syagrus species currently recognized were treated as a single species, Syagrus petraea (Mart.) Becc. However, analysis of pinnae anatomy, combined with increased field experience by Noblick and Lorenzi (2010), revealed that many populations formerly treated as S. petraea represented undescribed species, and that S. petraea probably does not occur in Brazil. Even species that are very similar may be anatomically distinct, making pinnae anatomy an important dataset for more detailed morphological studies of a given population, and a useful tool for resolving species complexes (Noblick 2017a).

More recently, Firmo et al. (2021) showed that new anatomical data are also useful for separating similar species, revealing that Syagrus evansiana Noblick represents a complex of acaulescent Syagrus. In Sant’Anna-Santos et al. (2023c), the S. evansiana complex began to be disentangled with the description of Syagrus aristeae Sant’Anna-Santos and, more recently, another population of the complex was recognized as Syagrus harenae Sant’Anna-Santos (Firmo et al. 2021; Sant’Anna-Santos et al. 2023c, 2025).

Currently, 70 Syagrus species are recognized, of which 35 are acaulescent (Noblick 2009, 2013, 2017a, 2018; Noblick et al. 2014; Firmo et al. 2021; Sant’Anna-Santos et al. 2023a, 2023b, 2023c, 2025). In the Southern Espinhaço Range, S. evansiana has the highest number of collection records, including populations geographically isolated (Noblick 2017a; Sant’Anna-Santos et al. 2025). Some of these populations are known from only one or a few collections, reinforcing the need to revisit them and reassess their morphology and pinnae anatomy (Firmo et al. 2021; Sant’Anna-Santos et al. 2025). When described by Noblick (2009), in addition to the type population in an area adjacent to the Espinhaço Range, data from another geographically isolated population in the Espinhaço Range were also used to define the morphological spectrum of S. evansiana.

Therefore, considering prior evidence (Firmo et al. 2021; Sant’Anna-Santos et al. 2023a, 2025), we evaluate, separately, the morphology and anatomy of these two populations used by Noblick (2009) in the description of S. evansiana. We hypothesize that the two populations represent distinct species, which would further increase the need to reassess other populations treated as S. evansiana since the revision of the genus by Noblick (2017a). If confirmed, this new taxon will increase the diversity of endemic Syagrus in campos rupestres and reinforce the need to protect and conserve these areas, which are under severe anthropogenic pressure.

Material and methods

This study was based on field observations in two different areas of the Minas Gerais State, Brazil (Fig. 1). The first area lies outside the Espinhaço Range and corresponds to the type population of S. evansiana, in a Cerrado sensu stricto area in the municipality of Jequitaí. The second area lies within the Espinhaço Range and corresponds to a campo rupestre in the municipality of Itacambira. These two populations were used in the protologue of Syagrus evansiana for the description of the species (Noblick 2009) We conducted bibliographic surveys of available publications on the anatomy and taxonomy of the S. evansiana complex (Glassman 1967, 1968, 1970, 1972, 1979, 1987; Henderson et al. 1995; Noblick 2009, 2010, 2013, 2017a, 2017b; Noblick and Lorenzi 2010; Soares and Guimarães 2019; Firmo et al. 2021; Sant’Anna-Santos et al. 2023a, 2023b, 2023c, 2025) and examined herbarium material (DIAM, RB, SP, SPF, HPL, MBM, MCMG, UPCB) and online databases (UB, ESA, NY, K).

Figure 1. 

Geographic distribution of species of the Syagrus evansiana complex in Minas Gerais State, Brazil. The coloured lines indicate the municipalities of the type populations of S. montana, S. evansiana, and S. aristeae.

Morphological and anatomical terminology follows Glassman (1972), Dransfield et al. (2008), Tomlinson et al. (2011), Noblick (2017a, 2017b), and Firmo et al. (2021). The circumscription adopted for Syagrus follows Noblick (2017a) and Sant’Anna-Santos et al. (2023a, 2023b, 2023c, 2025). Morphobiometric data and photographs were recorded in situ for both populations previously treated as S. evansiana since Noblick (2009), but here treated as different species: S. evansiana and S. montana sp. nov. Data were obtained from 15 reproductive individuals (bearing inflorescences and/or fruits) of each species, from which two botanical illustrations were prepared. The illustration of S. evansiana in Noblick (2009) was prepared using specimens from different populations (Fig. 2; Noblick 2009); therefore, we provide illustrations for both S. evansiana and S. montana. For S. aristeae, considered by Sant’Anna-Santos et al. (2023c) as part of the S. evansiana complex, data were taken from Firmo et al. (2021) and Sant’Anna-Santos et al. (2023c) and used to prepare Tables 1 and 2. Flower samples from both populations were analysed fresh and immediately stored in ethanol for examination under a Bioptika L60T stereomicroscope coupled with a CMOS 12mp PLUS camera.

Figure 2. 

Syagrus evansiana. A. Acaulescent habit. B. Asymmetric pinnae with a long tapering tip. C. Unbranched inflorescence. D. Staminate flower. E. Sepals connate at the base. F. Filament bases connate. G. Lateral view of the stamen, sagittate anther base. H. Stamen, dorsal view. I. Trifid pistillode. J. Pistillate flower, sepals. K. Pistillate flower, petals. L. Pistil, glabrous ovary and dentate staminodes. M. Fruit covered by dense indumentum. N. Ellipsoid epicarp, lateral view. O. Basal view of the epicarp, pores. Based on Sant’Anna-Santos 405 (DIAM). Illustration by Gustavo Surlo.

Table 1.

Differences in habitat and morphology among Syagrus montana, S. evansiana, and S. aristeae.

S. montana S. evansiana S. aristeae
Habitat Campo rupestre Cerrado Campo rupestre
Plant height 37–80 cm 62–100 cm 100–165 cm
Leaf rachis 21–57 cm 79–97 cm 94–145 cm
Leaves 3–6(–10) 6–11 3–6
Sheathing leaf base ca. 12–20 cm long 1.5–12 cm long 13–23 cm long
Pseudopetiole 10–23 cm long 15–33 cm long 15–34 cm long
True petiole 0–39.5 × 0.6–1.1 cm 17.5–32 × 1.0–1.5 cm 10–32.5 × 0.8–1.4 cm
Pinnae number 18–43 33–48 38–67
Pinna arrangement 2–5(–6) 2–4(–5) 2–3(–4)
Apical pinnae 3–8.5 × 0.1–0.9 cm 9–12 × 0.5–0.7 cm 7–18 × 0.6–1.0 cm
Middle pinnae 12–18 × 1.3–2.0 cm 19–30 × 1.5–3.0 cm 22–33 × 1.3–3.0 cm
Basal pinnae 11–18 × 0.5–0.9 cm 15.5–16 × 0.6–0.7 cm 20–32 × 0.5–1.2 cm
Prophyll size 5–14 × 1.0–1.8 cm 6–16 × 1.3–2.6 cm 7–23 × 1.3–4.8 cm
Inflorescence Usually branched Usually spicate Spicate or branched
Inflorescence length 8–26 cm 16.5–34 cm 36–71.5 cm
Peduncle 8–16 cm 8–27 cm 18–42 × 0.4–0.9 cm
Axis of the inflorescence 8–15 cm 9.5–20 cm 18–29.5 cm
Rachis of the inflorescence 0–2.5 cm 0–13 cm 0–5(–10) cm
Rachillae 1–6 1–8 1–9
Apical rachillae 10.5–11 cm 4.5–10 cm 11–21 cm
Basal rachillae (2–)4–9.5 cm 5–8 cm 11–21 cm
Flower arrangement Triad Triads, tetrads, and pentads Triad
Peduncular bract 12–31 cm 18.5–48 cm 32–67 cm
Beak 0.4–0.8 cm 0.3–1.5 cm 1–2 cm
Inflated portion 7–16.5 × 1.5–4.8 cm 13–22 × 3–7 cm 14–31 × 3.3–6.5 cm
Peduncular bract perimeter and thickness 2.5–6.2 cm× 1–2.5 mm 4–11 cm × 1.5–3 mm 4–7 cm × 2.5–3 mm
Staminate flowers
Flowers Pedicellate on the basal portion Sessile Sessile
Size at apex 8–14 × 3–6 mm 10–14.5 × 4–7 mm 10–12.5 mm
Size at base 13–15 × 4–6 mm 13–20 × 5–7 mm 11.2–15.9 mm
Sepal size 0.5–4 × 0.5–2 mm 1.5–5 × 1–1.5 mm 1.3–3.6 × 0.7–2.6 mm
Petal size 7–13 × 2–5 mm 9–16 × 2–5 mm 9–14.8 × 2.1–4.5 mm
Stamen 4–8 mm 4–9 mm 5.9–9.0 mm
Anther 3.5–6 mm 4–5.5 mm 4.4–7.7 mm
Base of the anther Cordate Sagittate Cordate
Filament 1–3 mm 1.5–4 mm 1.3–3.2 mm
Pistillode 0.5–1.5(–3) mm 0.5–1.0 mm 0.5–1.5 mm
Pistillate flowers
Size at apex 8–15 × 6–7 mm 15–18 × 4–7 mm 12.1–16 mm
Size at base 11–16 × 3–6 mm 17–20 × 5–10 mm 12.1–16 mm
Number of sepals 3 2–3 3
Sepal 11–16 × 3–7 mm 12–19 × 5–8 mm 11.1–14.6 × 3.9–8.2 mm
Number of petals 3 or occasionally 4 3 3
Petal 8–14 × 3.5–5 mm 14–18 × 8–7 mm 7–12 × 1.6–4.7 mm
Petal tip 2/3 of the petal length 1/3 to 1/2 of the petal length 2/5 to 1/2 of the petal length
Pistil 6–8 × 2–4 mm 9–13 × 2–4 mm 5.8–7.7 × 2.1–3.9 mm
Pistil indumentum Present Absent Absent
Stigma 2–5 mm 3–5 mm 2.8–3.4 mm
Staminodial ring 1–1.5 mm 1–3 mm 1 mm
Staminodes Undulate Dentate Dentate
Fruit Nearly globose Ellipsoid Nearly globose
Fruit sized 1.4–1.6 × 1.1–1.4 cm 2.0–2.3 × 1.4–1.6 cm 1.7–2.5 × 1.1–1.9 cm
Epicarp indumentum Thick brownish Thick brownish Cracked plates
Mesocarp ca 0.5 mm ca 1.0 mm 1.0–2.5 mm
Endocarp size 1.3–1.4 × 1.0–1.2 cm 1.6–2.0 × 0.9–1.3 cm 1.3–1.8 × 1.1–1.4 cm
Table 2.

Differences in pinnae anatomy between S. montana, S. evansiana, and S. aristeae.

Syagrus montana Syagrus evansiana Syagrus aristeae
Expansion tissue Continuous Interrupted Interrupted
Stomata on adaxial surface Absent Present Present
Raphides Frequent Rare Absent
Mesophyll Homogeneous Heterogeneous Heterogeneous
Adaxial fibre bundles Very narrow Narrow Narrow
Number of accessory vascular bundles around the main vascular system of the midrib 0–1 0–2 2
Number of collateral bundles in the vascular system of the midrib 1 2–4 3
Fibrous ring reaching the abaxial hypodermis Absent Absent Present
Group of fibres in the abaxial hypodermis Present, small Present, large Present, small
Large first adaxial fibre bundle at the margin Present Absent Absent
Adaxial hypodermis walls Thick Thin Thin
Midrib hypodermis stratification Biseriate Uniseriate Uniseriate
Midrib format Triangular Truncate Truncate
Fibre bundles around the fibrous ring Up to 5 10–15 10–15
Small accessory bundle between the expansion tissue Absent Present Absent
Fibres in the expansion tissue arrangement In a line Dispersed In a line

A distribution map showing all species of the S. evansiana complex was produced using QGIS v.3.40.3 (QGIS Development Team 2024) based on the following sources: state and municipal boundaries (IBGE 2020), elevation (EMBRAPA 2005), and occurrence localities (data from the current species). Distribution data for S. aristeae were taken from Sant’Anna-Santos et al. (2023c). Anatomical analyses were based on 15 samples from each population, taken from the central region of a median pinna, from herbarium specimens from our own collections and from plants in the field, following Firmo et al. (2021). After rehydration (Meira and Martins 2003), freehand cross sections were prepared. The samples were cleared using sodium hypochlorite and distilled water (1:1) and stained with Safrablau (1% Astra blue + 1% Safranin) (Bukatsch 1972, modified). Temporary slides were mounted, and photographs were taken with a light photomicroscope (Bioptika B20+) with an attached digital camera CMOS 12mp PLUS.

Taxonomic treatment

Syagrus montana B.F.Sant’Anna-Santos, sp. nov.

Figs 1, 37; Tables 1, 2

Type

BRAZIL – Minas Gerais • Itacambira; 16°57’29.28”S, 43°25’4.98”W; 1240 m; 30 Jul. 2024; fl., fr.; Sant’Anna–Santos 425; holotype: DIAM; isotypes: UPCB, IBGE, HCF.

Figure 3. 

Syagrus montana. A. Acaulescent habit. B. Asymmetric pinnae with a long tapering tip. C. Branched inflorescence. D. Unbranched inflorescence. E. Staminate flower. F. Stamen, lateral view showing the cordate base of the anther. G. Stamen, dorsal view. H. Lateral view of the stamen, cordate anther base. I. Trifid pistillode. J. Pistillate flower, sepals. K. Pistillate flower, petals. L. Pistil, ovary covered by indumentum and undulate staminodes. M. Fruit, epicarp densely covered by indumentum. N. Nearly globose epicarp, lateral view. O. Basal view of the epicarp. Based on the holotype Sant’Anna-Santos 425 (DIAM). Illustration by Gustavo Surlo.

Figure 4. 

Habitat and vegetative morphological aspects of Syagrus montana (A, C, E, G, I, K) and S. evansiana (B, D, F, H, J, L). A. Campo rupestre, landscape view. B. Cerrado sensu stricto, landscape view. CD. Solitary habit (white circles). EF. Shorter leaf rachis in S. montana (E, two white arrowheads) compared with S. evansiana (F, two white arrowheads). GH. Fibrous leaf sheath (two white arrowheads). IJ. Discolorous pinnae, adaxial (ad) and abaxial (ab) surfaces. KL. Pinnae insertion on the leaf rachis (ra) without ramenta. Photographs by Bruno F. Sant’Anna-Santos.

Figure 5. 

Morphological features of inflorescences, flowers, and fruits of Syagrus montana (A–C, G–H, K) and Syagrus evansiana (D–F, I–J, L). A. Branched inflorescence. B. Unbranched inflorescence. C. Striate peduncular bract. D. Unbranched inflorescence. E. Branched inflorescence. F. Striate peduncular bract. G. Triad: one pistillate flower (white dot) flanked by two staminate flowers (black dots). H. Pedicellate staminate flowers (pe): sepals (se) connate at the base and petal (pt). I. Tetrad: two pistillate flowers (white dots), each flanked by a staminate flower (black dots). J. Pentad: two pistillate flowers (white dots) flanked by three staminate flowers (black dots). K. Nearly globose fruit: epicarp covered by brown indumentum. L. Ellipsoid fruit: epicarp covered by brown indumentum. Photographs by Bruno F. Sant’Anna-Santos.

Figure 6. 

Anatomy of the margin and intermediate region of pinnae of Syagrus montana (A–D) and Syagrus evansiana (E–J) using LM in transverse sections. A. Empty raphide idioblasts (eight white arrowheads) in stained sample; narrow adaxial fibres (fd); stomata on the abaxial surface (black arrowheads); large first adaxial fibre bundle at the margin (la); and homogeneous mesophyll. B. Idioblast containing raphides (ra) in an unstained sample. C. Biseriate adaxial hypodermis (hy). D. Detail of the abaxial surface: subsidiary cells (two yellow dots), guard cells (two white dots) and fibres (fi) in the hypodermis. E. Stomata on both surfaces (black arrowheads); large adaxial fibres (fd); and dorsiventral mesophyll. F. Raphides (ra). G. Biseriate adaxial hypodermis (hy). HI. Stomata on the adaxial (H) and abaxial (I) surfaces: subsidiary cells (two yellow dots), guard cells (two white dots). J. Fibre bundle on the abaxial surface (fi). Photographs by Bruno F. Sant’Anna-Santos.

Figure 7. 

Midrib anatomy of Syagrus montana (A–E) and S. evansiana (F–J) using LM in transverse sections. A. Triangular midrib: collateral bundle (white circle), continuous expansion tissue (ex) with linearly arranged fibres (two yellow circles), accessory vascular bundle (white arrowhead), and non-vascular fibres (four black arrowheads). BC. Detail of the expansion tissue: only non-vascular fibre bundles (black circle). D. Biseriate hypodermis (hy) with thickened walls. E. Detail of the collateral bundle: phloem poles (four white arrowheads). FG. Truncate midrib: collateral bundles (three white circles), interrupted expansion tissue (ex) with non-linearly arranged fibres (two yellow circles), small accessory vascular bundle within the expansion-tissue caps (green arrowhead), small vascular bundle (white arrowhead), and non-vascular fibres (11 black arrowheads). H. Detail of the accessory vascular bundle: vascular tissue (white circle). I. Uniseriate hypodermis (hy) with thin walls. J. Detail of the collateral bundles: phloem poles (six white arrowheads). Photographs by Bruno F. Sant’Anna-Santos.

Diagnosis

Syagrus montana is similar to Syagrus evansiana Noblick, from which it differs in leaf rachis length (21–57 vs 79–97 cm); leaf sheath length (12–20 vs 1.5–12 cm); apical pinnae length (3–8.5 vs 9–12 cm), middle pinnae length (12–18 vs 19–30 cm); inflorescence rachis length (0–2.5 vs 0–13 cm); apical rachillae length (10.5–11 vs 4.5–10 cm); flowers always arranged in triads (vs triads, tetrads, and pentads); basal staminate flowers pedicellate (vs sessile); anther bases cordate (vs sagittate); apical pistillate flower length (8–15 vs 15–18 mm), basal pistillate flower length (11–16 vs 17–20 mm); pistillate flowers with 3 sepals (vs 2 to 3); pistillate flowers with 3 petals or occasionally 4 (vs 3), with valvate tips reaching 2/3 of the petal length (vs 1/3–1/2); pistil length (6–8 vs 9–13 mm), pistil with indumentum (vs glabrous flowers in pre-anthesis), staminodes undulate (vs dentate); fruit nearly globose (vs ellipsoid); endocarp length (1.3–1.4 vs 1.6–2.0 mm).

Description

Small palm, solitary palm, 37–80 cm tall. Apparently acaulescent. Leaves pinnate, 3–6(–10) in number; leaf sheath ca 12–20 cm long; pseudopetiole 10–23 cm long; true petiole absent to 39.5 × 0.6–1.1 cm and 0.3–0.6 cm thick, adaxially grooved and abaxially rounded; abaxial surface of the petiole and leaf rachis with white tomentum; leaf rachis 21–57 cm long; pinnae medium to dark-green, discolorous, abaxial surface glaucous, linear, rigid-coriaceous, with a more or less asymmetric apex and an elongated, tapering midrib, 18–43 pairs, arranged in 2–5(–6) along the leaf rachis and inserted in divergent planes; ramenta or tomentum absent at the insertion of pinnae on the leaf rachis and along the abaxial surface of the pinna midrib; apical pinnae 3–8.5 × 0.1–0.9 cm; median pinnae 12–18 × 1.3–2.0 cm; basal pinnae 11–18 × 0.5–0.9 cm; prophyll 5–14 × 1.0–1.8 cm; peduncle indumentum glabrous. Inflorescence erect, spicate or spirally branched, but usually spirally branched; prophyll 5–14 × 1.0–1.8 cm; peduncular bract ca 12–31 cm long, inflated portion 7–16.5 × 1.5–4.8 cm, including a beak 0.4–0.8 cm long, perimeter 2.5–6.2 cm, 1–2.5 mm thick, woody, striate, exterior glabrous; peduncle 8–16 cm long, 1.6–5 × 1.5–4 mm wide, elliptic in transverse section, glabrous; inflorescence axis 8–15 cm long; rachis 0–2.5 cm long; rachillae 1–6, 10.5–11 cm long at the apex, (2–)4–9.5 cm long at the base, glabrous. Staminate flowers 8–14 × 3–6 mm at the apex, 13–15 × 4–6 mm at the base, shortly pedicellate at the base of the inflorescence; pedicels ca 1 mm long, yellow; sepals 3, 0.5–4 × 0.5–2 mm, glabrous, without evident nerves, briefly connate at the base; petals 3, 7–13 × 2.0–5.0 mm, with acute apices, nerves inconspicuous; stamens 4–8 mm long, anthers 3.5–6 mm long; filaments 1–3 mm long, briefly connate at the base; pistillode trifid, ca 0.5–1.5(–3) mm long. Pistillate flowers elongate-pyramidal, 8–15 × 6–7 mm at the apex, 11–16 × 3–6 mm at the base, glabrous; sepals 3, 11–16 × 4–6.5 mm, yellow, without visible venation, imbricate; petals 3 or occasionally 4, 8–14 × 3.5–5 mm, with valvate apices reaching 2/3 of the petal length; pistil 6–8 × 2–4 mm, with lepidote indumentum from the base of the outer ovary wall to approximately the base of the stigmas; stigmas 3, 2–5 mm long; staminodial ring ca 1–1.5 mm tall; staminodes undulate. Fruits nearly globose, 1.4–1.6 × 1.1–1.4 cm, brown when mature; epicarp less than 1 mm thick, covered with dense brownish tomentum; mesocarp ca 0.5 mm thick, succulent and fibrous; endocarp ca 1.3–1.4 × 1.0–1.2 cm, ca 1 mm thick, with 3 pores in the basal portion. Seed nearly globose, endosperm homogeneous. Germination remote-tubular.

Distribution and habitat

Syagrus montana is endemic to Minas Gerais State, Brazil, in a region known as the “Northern Mountains Complex”, in the southern portion of the Espinhaço Range (Fig. 1). The species occurs in campos rupestres, where it may be locally dominant, forming part of the graminoid stratum on high-elevation plateausat around 1240 m altitude, in the municipality of Itacambira and adjacent areas (Noblick 2017a). Campos rupestres are characterized by shallow, stony, nutrient-poor soils and are subject to severe fires (Almada et al. 2016; Fernandes 2016).

Phenology

Syagrus montana was observed with flowers and fruits from November to December and from June to July in 2019 and 2024.

Etymology

The specific epithet, montana, means “mountain” and refers to the high-elevation plateau where the species occurs.

Preliminary IUCN conservation assessment

The population of the new species is known only from the municipality of Itacambira, where no protected areas exist. Near the type population, there is a highway and Eucalyptus plantations. Considering the area of occupancy (AOO = 32 km2) and extent of occurrence (EOO = 42.998 km2) and following the IUCN Categories and Criteria (IUCN 2022), S. montana should be considered Critically Endangered: CR B1ab(i,iii).

Additional specimens examined

BRAZIL – Minas Gerais • Itacambira, à beira da rodovia Montes Claros-Itacambira (MG-308); 19 Nov. 2013; fl.; Medeiros & Fonseca 84; SPF • same data as for preceding; Medeiros & Fonseca 86; SPF • Itacambira, estrada para Montes Claros; 9 Jan. 1986; fl.; Mello-Silva et al. 9158; SPF • Itacambira, fazenda da plantar siderúrgica, em área de plantação de eucalipto, na rodovia entre Juramento e Itacambira; 13 Dec. 2019; fl., fr.; Sant’Anna-Santos & Firmo 184; UPCB • same data as for preceding; Sant’Anna-Santos & Firmo 185; UPCB • same data as for preceding; Sant’Anna-Santos & Firmo 189; UPCB • same data as for preceding; Sant’Anna-Santos & Firmo 195; UPCB • same data as for preceding; Sant’Anna-Santos & Firmo 196; UPCB • same data as for preceding; Sant’Anna-Santos & Firmo 197; UPCB.

Pinnae anatomy

In S. montana, stomata occur only on the abaxial surface, whereas in S. evansiana stomata are present on both surfaces (Fig. 6D–E, H–I). In both species, subsidiary cells are arcuate and located entirely below the level of the cuticle (Fig. 6D, H–I). Adaxial non-vascular fibre bundles are narrower in S. montana than in S. evansiana (Fig. 6A and 6E, respectively). On the abaxial surface, groups of fibres and isolated fibres are rarer in S. montana than in S. evansiana (Fig. 6A–D, E–J). In S. montana, raphide-containing idioblasts are frequent and occur both at the margin and in the intermediate region of the pinnae (Fig. 6A–B). In S. evansiana, raphides are rare and restricted to the marginal region (Fig. 6E–F).

Adaxial fibre bundles are connected to the adaxial hypodermis and reach nearly half of the mesophyll in both species (Fig. 6A–E). Only primary vascular bundles are connected to the hypodermis on both surfaces and are always completely surrounded by fibres in both species (Fig. 6A–E). In both species, primary vascular bundles always have a larger diameter, distinct phloem poles, and conspicuous protoxylem and metaxylem elements (Fig. 6A–E). Secondary and tertiary vascular bundles are surrounded abaxially only by a sclerenchymatous sheath (Fig. 6A–E). Whereas secondary vascular bundles are connected only to the abaxial hypodermis, tertiary vascular bundles may or may not be connected to the abaxial hypodermis (Fig. 6A–E). The mesophyll is homogeneous in S. montana (Fig. 6A) and dorsiventral in S. evansiana (Fig. 6E). At the margin, a large first adaxial non-vascular fibre bundle is present in the new species, whereas in S. evansiana this bundle is not always present (Fig. 6A, E).

The midrib is triangular in S. montana and the expansion tissue is continuous (Fig. 7A–B). In S. evansiana, the midrib is truncate and the expansion tissue is interrupted (Fig. 7F). In both species, the midrib is adaxially projected, and the expansion tissue contains immersed fibre groups arranged in a line in the new species (Fig. 7A–B) and dispersed in S. evansiana (Fig. 7F–G). The main vascular system of the midrib consists of a single collateral bundle in S. montana (Fig. 7A–E) and 2–4 in S. evansiana (Fig. 7F–J). In both species, the collateral bundles are surrounded by a fibrous ring with a reinforced sheath that does not connect to the adaxial or abaxial hypodermis (Fig. 7B–G). There are 0–5 and 10–14 non-vascular fibre bundles around the fibrous ring in S. montana and S. evansiana, respectively (Fig. 7A, F). The presence of a small accessory vascular bundle within the expansion-tissue caps is observed only in S. evansiana (Fig. 7G), representing the first record for the genus. The midrib hypodermis is biseriate in S. montana (Fig. 7D) and uniseriate in S. evansiana (Fig. 7I). Table 2 compares the leaf anatomy of S. evansiana, S. montana, and S. aristeae.

Notes

When described, Syagrus evansiana was characterized as a miniature of Syagrus glaucescens Becc. and Syagrus duartei Glassman due to strong similarity in leaf morphology (Noblick 2009). However, the acaulescent habit and other important morphological characters, such as the occurrence of both branched and unbranched inflorescences, strongly supported its recognition as a distinct species (Noblick 2009). In the following year, additional acaulescent Syagrus species were revealed as the result of an extensive effort to study these palms in their natural habitats combined with leaflet anatomy data (Noblick and Lorenzi 2010).

Since then, it has become clear that most dwarf Syagrus species do not occur over large geographic ranges and/or occupy different, geographically isolated areas (Noblick 2017a; Firmo et al. 2021; Sant’Anna-Santos et al. 2023a, 2023b, 2023c). However, some acaulescent Syagrus still show broader geographic distributions, such as S. evansiana (Noblick 2017a; Reflora 2026; SpeciesLink Network 2026), because they represent unresolved species complexes (Sant’Anna-Santos et al. 2025). In Noblick (2009), populations from two distinct regions were used in the characterization of the species: a cerrado sensu stricto population in the municipality of Jequitaí and a campo rupestre population in the municipality of Itacambira and adjacent areas. These two regions are more than 100 km apart in a straight line (Fig. 1), are geographically isolated, and occupy habitats with distinct conditions. Over subsequent years, additional areas were added to the distribution map of the species, such as Serra do Cabral and Serra do Ambrósio mountains (Noblick et al. 2014; Noblick 2017a).

Firmo et. al. (2021) provided the first step towards disentangling the Syagrus evansiana complex. The population treated as Syagrus evansiana from Serra do Cabral mountain, a disjunction in the southern portion of the Espinhaço Range, was formally described by Sant’Anna-Santos et al. (2023a) as Syagrus aristeae. In Noblick (2017a), another disjunction in the Espinhaço Range was indicated for the occurrence of S. evansiana: Serra do Ambrósio mountain. Recognized as an area of high endemism, Serra do Ambrósio harbours unique high-elevation environments known as carrascos, characterized by coarse sandy substrates and rare, microendemic species (Meguro et al. 1994; Pirani et al. 1994; Oliveira et al. 2014; Costa et al. 2016, 2018; Sant’Anna-Santos et al. 2025). Accordingly, Sant’Anna-Santos et al. (2025) described Syagrus harenae, representing yet another step in resolving the complex.

Therefore, targeting previously known localities of S. evansiana for fieldwork is an effective strategy to document Arecaceae diversity in the southern Espinhaço Range and adjacent areas (Firmo et al. 2021; Sant’Anna-Santos et al. 2023a, 2025). Many of these areas have experienced increased anthropogenic pressure in recent years, and localities formerly considered less suitable for agriculture—such as rocky outcrops along the Espinhaço Range—have become strongly impacted by mining, cattle raising, and silviculture (Costa et al. 2018; Carvalho et al. 2024), increasing the likelihood that new species will go extinct before being documented.

However, an obstacle impedes studies of the Syagrus evansiana complex: it is necessary to take a step back and reassess the different populations used in the species’ original circumscription, as they may represent distinct species. Therefore, establishing the true morphological spectrum of S. evansiana is crucial for further work on the complex. To date, populations treated as S. evansiana in the Espinhaço Range and adjacent areas still lack field-based study and detailed morphological and anatomical analyses (Reflora 2026; SpeciesLink Network 2026).

In Syagrus species delimitation, the size of leaf parts is an important diagnostic aspect (Glassman 1987; Henderson et al. 1995; Noblick 2009, 2017a; Firmo et al. 2021; Sant’Anna-Santos et al. 2023a, 2023b, 2023c, 2025). In S. montana, the length of the leaf rachis and sheath, as well as the length of apical and middle pinnae, are useful to distinguish it from S. evansiana (Table 1). Regarding reproductive traits, both inflorescence architectures (unbranched and branched) occur in both species (Figs 2C, 3C–D, 5A–B, 5D–E). However, branched inflorescences are more common in S. montana, whereas unbranched inflorescences are more common in S. evansiana (Table 1). In S. montana, flowers are always arranged in triads (Fig. 5G), whereas in S. evansiana we observed triads, tetrads, and pentads at similar frequencies (Fig. 5I–J). Pentads were first described for S. harenae (Sant’Anna-Santos et al. 2025); therefore, this is the second record for the genus. We also observed differences in the length of apical rachillae (Table 1). The length of pistillate flowers, as well as the number of sepals and petals, are also useful for distinguishing the two species, in addition to the valvate petal apices, which reach 2/3 of the petal length in S. montana (Fig. 3K) and 1/3–1/2 in S. evansiana (Fig. 2K). The pistil is smaller in S. montana (Table 1), bears indumentum and has an undulate staminodial ring (Fig. 3L), in contrast to S. evansiana, where the pistil is glabrous and the staminodial ring is dentate (Fig. 2L). The fruit is smaller and nearly globose in S. montana (Fig. 3M, 5K) and ellipsoid in S. evansiana (Fig. 2M, 5L).

Anatomically, S. montana is surprisingly different from S. evansiana (Table 2; Figs 6, 7). Notable differences include the frequency and location of raphides, the alignment of fibre groups in the expansion tissue, stomatal position, mesophyll differentiation, and an exclusive feature in S. evansiana: the presence of a small accessory vascular bundle within the expansion tissue. Our results confirm that knowledge of the plant in its natural habitat, combined with leaf anatomical studies, is essential for circumscribing these dwarf palms. In Noblick and Lorenzi (2010), these two factors supported the reinstatement of species previously synonymized and revealed multiple dwarf species that were unknown or treated as a single species. Since Glassman (1972), pinnae anatomy has been known to be useful not only for distinguishing morphologically very distinct species, but also morphologically very similar species, and distinct populations treated as a single species (Glassman 1972, 1987; Noblick and Lorenzi 2010; Noblick 2013, 2017a, 2017b; Noblick and Sant’Anna-Santos 2021; Sant’Anna-Santos 2021, 2023; Sant’Anna-Santos et al. 2015, 2018), as in the case of S. montana.

Syagrus montana shows marked morphological and anatomical differences from S. evansiana; together with geographic isolation and significant habitat differences, these data support its recognition as a distinct species. Syagrus evansiana exhibits rare characters for the genus, such as flowers arranged in tetrads and pentads, and an exclusive trait: an accessory vascular bundle within the expansion tissue. These results reinforce the need to revise poorly studied populations of acaulescent Syagrus that are still treated as S. evansiana. In addition, the data presented here corroborate the southern Espinhaço Range and its disjunctions as one of the centres of diversity of Syagrus. The discovery of this new species highlights the singularity of the local flora and supports its classification as a priority area for conservation.

Acknowledgements

This study was financed in part by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - Brasil (CAPES) - Finance Code 001.

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