Research Article |
Corresponding author: Itzi Fragoso-Martínez ( itzi.fragoso@inecol.mx ) Academic editor: Chun-Lei Xiang
© 2024 Itzi Fragoso-Martínez, Gerardo A. Salazar, Emmanuel Martínez-Ambriz, Martin Reith.
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:
Fragoso-Martínez I, Salazar GA, Martínez-Ambriz E, Reith M (2024) Two new species of Salvia (Lamiaceae) from the dry forests of Dominican Republic. PhytoKeys 249: 299-315. https://doi.org/10.3897/phytokeys.249.137556
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We describe two new species of Salvia from the Antillean dry forests, belonging to Salvia section Urbania. These species’ names honor two Latin American botanists who have advanced our understanding either of the Dominican flora or the mint family (Lamiaceae). Salvia claseana is found in the Sierra Martín García. It resembles S. calaminthifolia but differs in having strigose stems, rhombic to trullate leaves with a cuneate, decurrent base, and larger flowers. Salvia martineziana inhabits the Sierra de Bahoruco. It resembles Salvia brachyphylla but differs in having strigose stems with retrorse trichomes and flowers disposed in the axils of the distal leaves. We provide descriptions, photographs, a distribution map and an identification key for the species of Salvia section Urbania from the Dominican Republic. Additionally, we sequenced three molecular markers (nrITS, trnL-trnF, and trnH-psbA) for the new taxa and other Dominican and Haitian Salvia species to investigate their phylogenetic relationships.
Describimos dos nuevas especies de Salvia provenientes de los bosques secos antillanos, pertenecientes a Salvia sección Urbania. Estas especies honran a dos botánicos latinoamericanos que han ampliado nuestro conocimiento ya sea sobre la flora dominicana o la familia de la menta (Lamiaceae). Salvia claseana se encuentra en la Sierra Martín García. Se asemeja a S. calaminthifolia pero se diferencia de ella por tener tallos estrigosos, hojas rómbicas a truladas con una base cuneada, decurrente, y flores más grandes. Salvia martineziana habita en la Sierra de Bahoruco. Es semejante a Salvia brachyphylla pero difiere en tener tallos estrigosos con tricomas retrorsos y las flores dispuestas en las axilas de las hojas distales. Presentamos descripciones, fotografías, mapas de distribución y una clave de identificación para las especies de Salvia sección Urbania de República Dominicana. Además, secuenciamos tres marcadores moleculares (nrITS, trnL-trnF y trnH-psbA) para los nuevos taxones y otras especies de Salvia dominicanas y haitianas, con el fin de explorar sus relaciones filogenéticas.
Antilles, endemism, Hispaniola, phylogeny, sages
The genus Salvia L. (sage) comprises more than 1000 species, which renders it the most diverse group in the mint family or Lamiaceae (
Salvia subg. Calosphace section Urbania Epling is a group of 10 species endemic to Hispaniola, most of them distributed exclusively in Haiti (
Dominican Republic houses 21 species of Salvia (
Herbarium specimens were collected and processed, and the individuals were photographed in situ. Voucher specimens of all the collected taxa were deposited at the herbarium of the
Jardín Botánico Nacional Dr. Rafael M. Moscoso (JBSD)
and the duplicates (when available) were sent to MEXU and XAL herbaria in Mexico (
The identification of the specimens was based on the most comprehensive revision of Salvia subgenus Calosphace by
The distribution map was made with QGIS v. 3.30 (QGIS Development Team, 2023), using the distribution data from the herbarium specimen and iNaturalist observations from one of the authors, these occurrences and coordinates are not provided due to the species global conservation status of Critically Endangered. The species’ extent of occurrence (EOO) and species’ area of occupancy (AOO) were calculated using GeoCAT (
To test the phylogenetic position of the new taxa, as well as other sage species from the Caribbean, we sampled most of the species from Dominican Republic and a few taxa from Haiti, mainly belonging to section Urbania (Table
Voucher information and GenBank accession codes for the sampled sage species in this study. Detailed specimen information can be found in Suppl. material
Species | Voucher information | GenBank accession numbers | ||
---|---|---|---|---|
ITS | trnH-psbA | trnL-trnF | ||
S. arborescens Urb. & Ekman | I. Fragoso-Martínez et al. 459 (JBSD, MEXU) | PP905439 | PP907529 | PP907549 |
S. arduinervis Urb. & Ekman | E. Ekman 3168 (TEX) | PP905440 | PP907530 | PP907550 |
S. bahorucona Urb. & Ekman | I. Fragoso-Martínez et al. 517 (JBSD, MEXU) | PP905441 | PP907531 | PP907551 |
S. brachyloba Urb. | I. Fragoso-Martínez et al. 509 (JBSD, MEXU) | PP905442 | PP907532 | PP907552 |
*S. claseana Fragoso & Salazar | I. Fragoso-Martínez et al. 529 (JBSD, MEXU) | PP905459 | PP907548 | PP907568 |
*S. calaminthifolia Vahl | E. Ekman 9443 (TEX) | PP905443 | PP907533 | PP907553 |
S. caymanensis Millsp. & Uline | I. Fragoso-Martínez & R. Middleton 309, cultivated | PP905444 | PP907534 | PP907554 |
S. decumbens Alain | I. Fragoso-Martínez et al. 519 (JBSD, MEXU) | PP905445 | PP907535 | PP907555 |
S. foveolata Urb. & Ekman | I. Fragoso-Martínez et al. 508 (JBSD, MEXU) | PP905446 | PP907536 | PP907556 |
S. lachnaioclada Briq. | I. Fragoso-Martínez et al. 608 (JBSD, MEXU) | PP905447 | PP907537 | PP907557 |
S. lavendula Alain | T. Clase et al. 1059 (JBSD) | PP905448 | PP907538 | PP907558 |
*S. martineziana Fragoso & Martínez-Ambr. | I. Fragoso-Martínez et al. 497 (JBSD, MEXU) | PP905458 | PP907547 | PP907567 |
*S. montecristina Urb. & Ekman | I. Fragoso-Martínez et al. 527 (JBSD, MEXU) | PP905449 | PP907539 | PP907559 |
*S. praeterita Epling | I. Fragoso-Martínez et al. 607 (JBSD, MEXU) | PP905450 | PP907540 | PP907560 |
S. selleana Urb. | I. Fragoso-Martínez et al. 503 (JBSD, MEXU) | PP905451 | PP907541 | PP907561 |
S. serotina L. | I. Fragoso-Martínez et al. 506 (JBSD, MEXU) | PP905452 | PP907542 | PP907562 |
*S. subaequalis Epling | GenBank BioSample: SAMN22547053 | PP905453 | – | – |
S. tenella Sw. | T. Clase et al. 8266 (JBSD) | PP905454 | PP907543 | PP907563 |
S. thormanii Urb. | T. Clase et al. 8059 (JBSD) | PP905455 | PP907544 | PP907564 |
S. tuerckheimii Urb. | I. Fragoso-Martínez et al. 607 (JBSD, MEXU) | PP905456 | PP907545 | PP907565 |
S. uncinata Urb. | I. Fragoso-Martínez et al. 575 (JBSD, MEXU) | PP905457 | PP907546 | PP907566 |
The sequences were edited and assembled using Geneious v.10.2.6 (http://www.geneious.com,
The matrix comprising the three molecular markers included 288 taxa, of which 269 belong to Salvia subgenus Calosphace. The length of this matrix was 2,197 bp: 722 bp from the nrITS region, 505 bp from the trnH-psbA intergenic spacer and 970 bp from the trnL-trnF region. Based on the results of the congruence test between plastid and nuclear data performed in previous phylogenetic studies of Salvia subgenus Calosphace (
The concatenated matrix included three partitions corresponding to each molecular marker. For each partition, we evaluated 88 molecular substitution models using ModelFinder (
The Maximum likelihood analysis of the concatenated matrix was performed using the IQ-TREE algorithm (
Similar to Salvia calaminthifolia, but with the stems strigose (vs. cinereous); the leaves rhombic to trullate with the base cuneate, decurrent, the margin crenate-serrate (vs. deltoid-ovate, base truncate, margin subentire); the flowers bigger, with the calyx 5.5–8 mm long and the corolla tube 7.7–9 mm long (vs. 5–5.5 mm long and 5–6 mm long, respectively), and the lower lip reclinate (vs. reflexed).
Dominican Republic. Azua, Las Charcas • Paraje Boquerón, cañada subiendo por el puente Juana Guayacán, a 4 km de la desviación de la carretera Sánchez (Bani-Azua); 18°21'0.9138"N, 70°31'51.8118"W; 307 m; 28 Nov 2016; I. Fragoso-Martínez, G.A. Salazar & T. Clase 529 (holotype: JBSD 129667; isotypes: MEXU 1512146, XAL 0154233).
Suffruticose herbs , ca. 0.5 m tall; stems strigose with antrorsely appressed trichomes, internodes 1.5–4 cm long. Leaves rhombic to trullate, 1.3–3.6 × 1–3 cm, base cuneate, decurrent, apex acute, margin crenate-serrate, ciliate; upper leaf surface bullate, glabrescent; lower leaf surface pale, densely pubescent, simple trichomes minute, white, with amber spherical glands; petioles 0.5–1.2 cm long. Flowers axillary, 4(–8) per node; pedicels 2.5–6 mm long, trichomes simple, adpressed. Calyx green, tubular-campanulate, 5.5–8 mm long, strigose, with simple and glandular trichomes; tube 4.7–6.5 × 3 mm; lobes deltate, apex apiculate, upper lobe 1.3–1.8 mm long, straight, 3-veined, margin ciliate, lower lobes 1.3–2.5 mm long, straight. Corolla violet with white nectar guides in the lower lobe, ca. 1 cm long, tube 7.7–9 × 2.6–3 mm, ventricose, invaginate, internally epapillate; lobes unequal in length, upper lobe galeate, 3.8–6 mm long, densely pubescent, lower lobe 5–8 × 7.8–8.4 mm, tetralobate, reclinate. Stamens included in the upper corolla lobe, fused close to the corolla opening; filaments 1.5–2 mm long; connective 7–8 mm long, sparsely pilose, with a bilobed tooth close to the insertion with the filament, straight; upper arm of the connective shorter than the lower arm, 3–3.5 mm long, thecae 1–1.5 mm long; lower arm 4.5–5 mm long. Style 9–11 mm long, densely pubescent near the branches, upper branch longer than the lower one, lower branch spathulate. Nectary disc surface with spherical glands near the base of the mericarps, nectary horn ca. 2 mm long, oblong, laterally compressed. Mericarps ovoid, 1.6–2 × 0.7–0.9 mm, smooth.
Flowers were documented from November to May. Fruits have been observed after this period.
The epithet “claseana” honors the Dominican botanist Teodoro Clase, head of the Botany Department of the Jardín Botánico Nacional “Dr. Rafael M. Moscoso”. His botanical expeditions have resulted in ca. 12,000 collected specimens and he has described ca. 12 new species of angiosperms from Hispaniola. These contributions are undoubtedly crucial to the knowledge of the flora of Dominican Republic.
Endemic to the dry forests with limestone soils from the Sierra Martín García in Azua, Dominican Republic (Fig.
Phylogenetically, this species is closely related to Salvia praeterita (Fig.
Morphological comparison among the new taxa and their more similar species, based data from type specimens,
S. claseana Fragoso & Salazar | S. calaminthifolia Vahl | S. praeterita Epling | S. martineziana Fragoso & Martínez-Ambr. | S. brachyphylla Urb. | |
---|---|---|---|---|---|
STEM | |||||
Pubescence | Strigose, trichomes antrorse | Cinereous, trichomes straight | Strigose, trichomes antrorse | Strigose, trichomes retrorse | Hispid, trichomes straight |
LEAF | |||||
Shape | Rhombic to trullate | Deltoid-ovate | Ovate to deltoid-ovate | Obovate to flabellate | Obovate to flabellate |
Size (cm) | 1.3–3.6 × 1–3 | 0.5–1.2 × 0.7–1.5 | 1–2 × 0.5–1 | 1–1.5 × 0.5–1 | 1.5–3 × 1–2.5 |
Base | Cuneate, decurrent | Truncate | Cuneate, decurrent | Cuneate, decurrent | Cuneate, decurrent |
Margin | Crenate-serrate | Subentire | Crenate-serrate | Crenate-serrate | Crenate-serrate |
Upper surface | Bullate | Smooth | Bullate | Bullate | Bullate |
Lower surface | Densely pubescent | Cinereous, trichomes adpressed | Densely pubescent | Tomentulose | Hispid |
INFLORESCENCE | |||||
Presence | Absent (axillary flowers) | Absent (axillary flowers) | Present, terminal racemes | Absent (axillary flowers) | Present, axillary and terminal racemes |
Flowers per node | 4(–8) | 2–6 | 2–6 | 2 | 3 |
CALYX | |||||
Shape | Tubular-campanulate | Tubular-campanulate | Tubular-campanulate | Campanulate | Tubular-campanulate |
Length (mm) | 5.5–8 | 5–5.5 | 5–7 | 5–6.8 | 5.5–6.5 |
COROLLA | |||||
Tube length (mm) | 7.7–9 | 5–6 | 8.5–9.5 | 7–9 | 7.5–9.5 |
Upper lip length (mm) | 3.8–6 | 3–4.7 | 4–4.6 | 4.5–6.5 | 5.6–6 |
Lower lip length (mm) | 5–8 | 3.5–5.5 | 5–6 | 7–7.5 | ca. 6.6 |
Lower lip position | Reclinate | Reflexed | Reclinate | Reflexed | Reclinate |
Dominican Republic. Azua, Barreras • Sierra Martín García, en los alrededores de Barreras; 170 m; 11 Sep 1984; M. Mejía et al. 1180 (JBSD). Las Charcas • Parque Nacional Francisco Alberto Caamaño, paraje Boquerón; 18° 21'38.4366"N, 70°31'51.24"W; 298 m; 9 May 2014; T. Clase & R. Ovidio S. 8645 (JBSD, MEXU).
Similar to Salvia brachyphylla, but differing from it by the pubescence of the plant (strigose with retrorse trichomes vs. hispid with erect trichomes); having smaller leaves (1–1.5 × 0.5–1 cm vs. 1.5–3 × 1–2.5) with a tomentulose lower leaf side (vs. hispid); flowers axillary (vs. forming racemes), calyx campanulate (vs. tubular-campanulate) and lower lobe of the corolla reflexed (vs. reclinate).
Dominican Republic. Independencia, Duvergé • Parque Nacional Sierra de Bahoruco, 6.8 km al S de Puerto Escondido por el camino a la Caseta 1; 18°17'10.4994"N, 71°34'6.3978"W; 965 m; 24 Nov 2016; I. Fragoso-Martínez, G.A. Salazar & T. Clase 497 (holotype: JBSD 129573; isotypes: MEXU 1512142, XAL 0154234).
Suffruticose herbs , 0.3–0.5 m tall; stems strigose with retrorsely appressed trichomes, internodes 0.8–2.4 cm long. Leaves obovate to flabellate, 1–1.5 × 0.5–1 cm, base cuneate, decurrent, apex acute to obtuse, margin crenate-serrate, ciliate; upper leaf surface bullate, densely hirsute; lower leaf surface whitish, tomentulose simple trichomes minute, curved, white, with yellow spherical glands; petioles 4–8 mm long. Flowers axillary, 2 per node; pedicels 1.5–4.5 mm long, hirsute. Calyx green, campanulate, 5–6.8 mm long, densely hirsute, with simple and glandular trichomes; tube 3–5 × 3 mm; lobes ovate-deltate, apex apiculate, upper lobe 2.5–4 mm long, curved backwards, 3-veined, margin ciliate, lower lobes 1.6–2.5 mm long, straight. Corolla violet with white nectar guides in the lower lobe, 1.2–1.4 cm long, tube 7–9 × 2.9 mm, ventricose, invaginate, internally epapillate, lobes unequal in length, upper lobe galeate, 4.5–6.5 mm long, densely pubescent, lower lobe 7–7.5 × 8.4 mm, tetralobate, reflexed. Stamens included in the upper corolla lobe, fused close to the corolla opening; filaments 1.5–2 mm long; connective 7.5–8.5 mm long, sparsely pilose, with an entire tooth close to the insertion with the filament, retrorse; upper arm of the connective slightly longer than the lower arm, 4–4.5 mm long, thecae 1–1.5 mm long; lower arm 3.5–4 mm long. Style 13–15 mm long, densely pubescent near the branches, with simple and capitate glandular trichomes, upper branch longer than the lower one, lower branch spathulate. Nectary disc surface with spherical glands near the base of the mericarps, nectary horn ca. 1 mm long, oblong, laterally compressed. Mericarps ovoid, 1.5–2 × 0.4–0.6 mm, smooth.
Flowers were documented from November to May. Fruits have been observed after this period.
The epithet “martineziana” honors the Mexican botanist Martha Martínez Gordillo, specialist of the Euphorbiaceae and Lamiaceae families. Dr. Martínez works at the FCME herbarium at the Universidad Nacional Autónoma de México. She has conducted fieldwork mainly in the states of Chiapas, Guerrero, Mexico and Oaxaca. Her study of the Mexican flora, particularly that of Guerrero, has led to the description of more than 30 species of angiosperms, many of them from the genus Salvia. Dr. Martínez has taught botany to numerous generations of Mexican biologists, and her exemplary professional and academic ethics, determination, kindness and generosity are an inspiration to her students.
Endemic to the dry forests with limestone soils from Sierra de Bahoruco in Independencia, Dominical Republic (Fig.
This species is sister to a clade formed by two other Dominican species of Salvia section Urbania (Fig.
Phylogenetic relationships of the Angulatae clade based on three molecular markers (nrITS, trnH-psbA and trnL-trnF) with an increased taxon sampling including the new species and other Hispaniolan taxa. Bootstrap values ≥80% are shown above the branches. The names of the new species are written in purple, while the names in gray correspond to the sections of the taxa from the Hispaniolan subclade. An extended version of the phylogenetic tree is provided in Suppl. material
Dominican Republic. Independencia, Duvergé • 5.2 km al S de Puerto Escondido en camino a Acetillar, Sierra de Bahoruco; 170 m; 9 May 1985; T. Zanoni et al. 34648 (JBSD).
1a | Flowers in racemes | 2 |
2a | Leaves and calyces densely hispid | S. brachyphylla |
2b | Leaves and calyces strigose to glabrate | 3 |
3a | Bracts persistent | 4 |
4a | Bracts flabellate, surrounding the flowers almost completely | S. saccifera |
4b | Bracts ovate, not surrounding the flowers completely | S. subglabra |
3b | Bracts deciduous | 5 |
5a | Lower leaf surface greenish, glabrous; flowers 6–12 per node | S. hotteana |
5b | Lower leaf surface whitish, strigose; flowers 2–6 per node | S. praeterita |
1b | Flowers axillary, not forming racemes | 6 |
6a | Leaf margin subentire, entire or sinuate | 7 |
7a | Leaves coriaceous | S. montecristina |
7b | Leaves membranaceus | 8 |
8a | Lower leaf surface incanous, grayish | S. mornicola |
8b | Lower leaf surface glabrate, greenish | 9 |
9a | Leaves oblong-elliptical | S. tortuensis |
9b | Leaves deltoid-ovate | S. subaequalis |
6b | Leaf margin crenate-serrate | 10 |
10a | Leaf base truncate | S. calaminthifolia |
10b | Leaf base cuneate | 11 |
11a | Leaves rhombic to trullate, 1–3 cm wide, lower leaf surface densely pubescent | S. claseana |
11b | Leaves obovate to flabellate, 0.5–1 cm wide, lower leaf surface tomentulose | S. martineziana |
The Hispaniolan species included in the phylogenetic analysis are distributed in four different clades. The species from Salvia section Micranthae (Benth.) Epling — Salvia serotina L. and S. tenella Sw.—, are part of a clade that also contains taxa from sections Bracteata Epling and Subrotundae (Epling) Epling (99% BS; Suppl. material
Epling’s sections sampled | Taxa sampled from the total |
---|---|
Ekmania Epling | 8 spp. (100%) |
*Gardoquiiflorae Epling | 2 spp. (66%) |
Micranthae | 4 spp. (57%) |
*Urbania Epling | 6 spp. (50%) |
Wrightiana Epling | 2 spp. (66%) |
The new species described in the present study belong to the monophyletic section Urbania. Within this clade Salvia montecristina Urb. & Ekman, endemic to Dominican Republic, is sister to the remaining species (Fig.
The new taxa described here are an addition to the sage species of Dominican Republic, increasing its diversity to 23 species (
Both species described here fit the circumscription of Salvia section Urbania provided by
Despite a recent surge in the discovery of new Neotropical sages, the Dominican Republic has remained relatively unexplored in this regard. The last new sage species from the country was described almost 40 years ago (
These newly discovered taxa from Hispaniola, which are critically endangered, are the first species of Salvia section Urbania to be evaluated under the IUCN criteria. However, they mirror a pattern seen in other native sages on the island, where some species are vulnerable (Salvia arborescens Urb. & Ekman, and S. decumbens Alain) and others are critically endangered (Salvia buchii Urb., S. haitiensis Urb., S. lachnaioclada Briq. and S. paryskii Skean & Judd). These results highlight the need for a comprehensive assessment of all Hispaniolan sage species, to develop effective strategies for their protection.
IFM is grateful for the support from the Posgrado en Ciencias Biológicas, Universidad Nacional Autónoma de México during her doctoral studies and the scholarship for doctoral studies (grant 324065) from Consejo Nacional de Ciencia y Tecnología (CONACyT, Mexico). The authors thank the Ministerio de Medio Ambiente y Recursos Naturales from the Dominican Republic for the collection permit; the staff of the JBSD herbarium, for managing the exchange and donation of the specimens; Teodoro Clase by providing information on the localities of the Dominican sages and conducting fieldwork with IFM and GAS. We also acknowledge María del Rosario García Peña of the MEXU herbarium, Elizabeth Séptimo from the JBSD herbarium and Israel Acosta Rosado of the XAL herbarium, for the facilities provided to process the type specimens. Finally, the authors are grateful to A.I. Bieler Antolín for the photographs of the floral structures; L.I. Cabrera for guidance with lab work.
The authors have declared that no competing interests exist.
No ethical statement was reported.
The Society of Systematic Biologists (SSB) provided a grant to IFM to conduct fieldwork in the Dominican Republic. Additional financial support for the laboratory work and sequencing was provided by the Programa Fronteras de la Ciencia FC-2016 (Project 1867). Publication fees were covered by the SNII-CONAHCYT grant (83797) provided to IFM.
IFM wrote the manuscript, EMA, GAS and MR provided feedback on the draft and made insightful suggestions to improve it. IFM and GAS conducted the field expedition in 2016 and MR provided them with information about the localities of the surveyed taxa. EMA extracted the DNA samples, amplified the molecular markers, made the map, assessed the conservation status of the new taxa and assembled all the figures of the manuscript. IFM processed the molecular data and performed phylogenetic analyses.
Itzi Fragoso-Martínez https://orcid.org/0000-0003-3661-1076
Gerardo A. Salazar https://orcid.org/0000-0002-5203-5374
Emmanuel Martínez-Ambriz https://orcid.org/0000-0002-4770-4174
Martin Reith https://orcid.org/0000-0001-7830-9161
All the newly sequenced data was uploaded to the NCBI with the accession numbers provided in Table
Additional information of the sampled sage species in the study
Data type: pdf
Complete phylogenetic trees of the plastid, nuclear and combined dataset
Data type: pdf