Research Article |
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Corresponding author: Bruno Francisco Sant’Anna-Santos ( bsantannaufmg@gmail.com ) Academic editor: Igor Kessous
© 2026 Stephane 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.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Reis SdaS, Francino DMT, Dias BO, Ferreira-Filho R, Pinto LG, Nunes ELP, Sant’Anna-Santos BF (2026) A new dwarf palm from the campo rupestre: another piece of the Syagrus evansiana complex (Arecaceae). Plant Ecology and Evolution 159(2): 356-369. https://doi.org/10.5091/plecevo.188286
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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 results – Syagrus 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.
Meridional Espinhaço, Palmae, plant anatomy, taxonomy
The Espinhaço Range is found in eastern Brazil and extends for approximately 1,200 km (
The genus is monophyletic (
Before
More recently,
Currently, 70 Syagrus species are recognized, of which 35 are acaulescent (
Therefore, considering prior evidence (
This study was based on field observations in two different areas of the Minas Gerais State, Brazil (Fig.
Morphological and anatomical terminology follows
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 (
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 |
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 (
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:
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 (
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. C–D. Solitary habit (white circles). E–F. Shorter leaf rachis in S. montana (E, two white arrowheads) compared with S. evansiana (F, two white arrowheads). G–H. Fibrous leaf sheath (two white arrowheads). I–J. Discolorous pinnae, adaxial (ad) and abaxial (ab) surfaces. K–L. Pinnae insertion on the leaf rachis (ra) without ramenta. Photographs by Bruno F. Sant’Anna-Santos.
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.
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). H–I. 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.
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). B–C. 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). F–G. 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.
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).
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.
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.
Syagrus montana was observed with flowers and fruits from November to December and from June to July in 2019 and 2024.
The specific epithet, montana, means “mountain” and refers to the high-elevation plateau where the species occurs.
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 (
BRAZIL – Minas Gerais • Itacambira, à beira da rodovia Montes Claros-Itacambira (MG-308); 19 Nov. 2013; fl.; Medeiros & Fonseca 84;
In S. montana, stomata occur only on the abaxial surface, whereas in S. evansiana stomata are present on both surfaces (Fig.
Adaxial fibre bundles are connected to the adaxial hypodermis and reach nearly half of the mesophyll in both species (Fig.
The midrib is triangular in S. montana and the expansion tissue is continuous (Fig.
When described, Syagrus evansiana was characterized as a miniature of Syagrus glaucescens Becc. and Syagrus duartei Glassman due to strong similarity in leaf morphology (
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 (
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 (
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 (
In Syagrus species delimitation, the size of leaf parts is an important diagnostic aspect (
Anatomically, S. montana is surprisingly different from S. evansiana (Table
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.
This study was financed in part by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - Brasil (CAPES) - Finance Code 001.