Lectotypification of Salvia bogotensis Benth. (Lamiaceae)

39 Lectotypification of Salvia bogotensis Benth. (Lamiaceae) Ángela María Morales-Trujillo1, Brenda Yudith Bedolla-García1, Patricia Hernández-Ledesma1, Jorge Gabriel Sánchez Ken1 1 Instituto de Ecología A.C. (INECOL), Centro Regional del Bajío, Red de Diversidad Biológica del Occidente Mexicano, Avenida Lázaro Cárdenas 253, 61600 Pátzcuaro, Michoacán, Mexico Corresponding author: Ángela María Morales-Trujillo (angela.mor[email protected]) Copyright: © Ángela María Morales-Trujillo et al. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International – CC BY 4.0). Research Article Abstract The species Salvia bogotensis (sect. Angulatae) was not adequately typified at the time of its description. The original material used by the author of the species corresponds to collections by Goudot s.n. carried out in Bogotá, Colombia. There are five specimens that match with this data distributed in three herbaria: F, K and P. Epling designated the material deposited in K as lectotype, however, these specimens rather correspond to two different collections, and he failed to choose one of them unambiguously. Because of this, it is necessary to clearly determine which of the two specimens corresponds to the lectotype. The specimen K000479515 (K!) is designated here as lectotype because it is the most complete showing the key morphological characters of the species (leaves, inflorescences, and flowers), in addition to the note made by Epling. Of the other specimens, one of those deposited at P(P00715001) is an isolectotype. Key words: Calosphace, Colombia, herbaria Introduction The sixth-largest angiosperm family is Lamiaceae Martinov (1820: 355). It has a cosmopolitan distribution and contains about 236 genera and 7200 species (Kubitzki 2004). Salvia L. (In Linnaeus 1753: 23) is the largest genus of Lamiaceae, comprising approximately 1000 species, and also has a cosmopolitan distribution (Drew et al. 2017; Drew 2020). It occurs in diverse biomes from sea level to 4000 m elevation (Claßen-Bockhoff et al. 2004; Kriebel et al. 2019), with centers of diversity in Eurasia, including the Mediterranean (ca. 250 spp.) and Central and Eastern Asia (ca. 90 spp.), as well as Eastern and Southern Africa (ca. 60 spp.), North America including Mexico (ca. 314 spp.), Central America (ca. 164 spp.), and South America (ca. 330 spp.) (Claßen-Bockhoff et al. 2004; Walker et al. 2004; Kriebel et al. 2019; González-Gallegos et al. 2020). The genus Salvia is classified worldwide into 11 subgenera, two of which are endemic in the American Continent: Audibertia J. B. Walker, B. T. Drew, & K. J. Sytsma (2015: 837) is restricted to the United States and northern Mexico and Calosphace (Benth.) Epling (Bentham 1833: 198; Epling 1939: 4) with approximately 582 species, has about half of the species richness of the genus, and it is distributed from the southern United States, Academic editor: Eberhard Fischer Received: 20 July 2025 Accepted: 6 September 2025 Published: 14 October 2025 Citation: Morales-Trujillo ÁngelaM, Bedolla-García BY, HernándezLedesma P, Sánchez Ken JG (2025) Lectotypification of Salvia bogotensis Benth. (Lamiaceae). PhytoKeys 265: 39–48. https://doi.org/10.3897/ phytokeys.265.165733 PhytoKeys 265: 39–48 (2025) DOI: 10.3897/phytokeys.265.165733 40 PhytoKeys 265: 39–48 (2025), DOI: 10.3897/phytokeys.265.165733 Ángela María Morales-Trujillo et al.: Lectotypification of Salvia bogotensis to Argentina, including the Caribbean islands, covering a total of 42 countries (González-Gallegos et al. 2020). Mexico, the Andes, southern Brazil and Argentina are a center of diversity of Calosphace (Walker and Sytsma 2006). Mexico is the primary center of diversification of the subgenus with 295 spp., of which 243 are endemic. This places the country as the richest, followed by Peru with 77 spp. (49 endemics) and Colombia with 60 spp. (37 endemics) (González-Gallegos et al. 2020). Based on floral morphology and geographic distribution, the subgenus Calosphace is divided (Walker et al. 2004) into 104 sections (Epling 1939; Epling and Mathias 1957). One of these sections, Angulatae (Epling 1939: 234), represents a major challenge due to its species richness (52) (Epling, 1937, 1939, 1940, 1941; Epling and Mathias 1957; Epling and Játiva 1966) and wide geographic range, extending from the southern United States to Argentina and the Caribbean islands (González-Gallegos et al. 2020). However, recent phylogenetic studies (Fragoso-Martínez et al. 2018) have confirmed that section Angulatae is polyphyletic, with species distributed across at least eight distinct clades. In Colombia, section Angulatae is represented by 14 species, primarily distributed in Cordillera Oriental (Boyacá and Cundinamarca departments) and Sierra Nevada de Santa Marta (Magdalena department), where several microendemisms occur. Wood and Harley (1989) included the section in their taxonomic revision of Salvia for Colombia, while Fernández-Alonso (2003) provided a synopsis of the Colombian species in the section; both works included Salvia bogotensis Benth. (1848: 312). After reviewing the nomenclature of this species, we found that the information regarding the type specimen is ambiguous, as it is only noted that the type is at K, where there are two specimens, in addition to three more in other herbaria. This study arises from the need to clarify the typification of this species as an essential step in the taxonomic research of the genus. Materials and methods All information regarding Salvia bogotensis was gathered from Tropicos.org (https://www.tropicos.org/home), and the protologue was consulted in the Botanicus Digital Library (2024) (http://www.botanicus.org/). Other online resources such as type images in JSTOR Global Plants 2025 (https://plants.jstor. org/), the Mid-Atlantic Herbaria Consortium (https://midatlanticherbaria.org/), and the Kew Data Portal (2025) (https://species.data.kew.org/) were examined. Typification of names follows the International Code of Nomenclature for Algae, Fungi, and Plants (ICN) (Turland et al. 2018). Herbarium acronyms are according to Thiers (2017 onwards). Typification of the name Salvia bogotensis was described by the English botanist George Bentham in 1848. The specific epithet refers to the collection locality in Bogotá, Cundinamarca, Colombia, an area extensively explored in the early 1800s by the French naturalist Justin Goudot (JSTOR 2025). In the protologue, Bentham (In De Candolle 1848: 312) states: “prope Bogotá, Goudot! (v. in h. Hook.)”, indicating that he examined the specimens of the Hooker’s herbarium that was incorporated 41 PhytoKeys 265: 39–48 (2025), DOI: 10.3897/phytokeys.265.165733 Ángela María Morales-Trujillo et al.: Lectotypification of Salvia bogotensis at Kew Herbarium, where Bentham worked and where most of his type specimens are deposited (O’Leary and Moroni 2016). Two Goudot’s specimens with original labels were deposited at K, K000479514, collected in “Bogotá” in April (as avril), and K000479515, in “Bogotá” locality of “Boquerón” in March (as martii (Latin)) both correspond to “Nelle. [Nouvelle] Grenade”. At the time of collection, Colombian territory was officially known as the Republic of New Granada, a name written on the labels. The locality “Boquerón” likely refers to the páramo of the same name surrounding Bogotá. Epling (1939), in his monograph on American Salvias, only mentioned that he saw the type of S. bogotensis at K (Goudot’s collection). However, upon reviewing the online specimens, we found that there are currently five sheets and a photo of a “type”: two in K, two in P, one in F and another in PH (photo). Epling left a note on the specimen K000479515 in 1927 stating: ‘This and the second Goudot specimen may be taken together as the type.’ However, in Kew both correspond to different specimens collected at different times and places K000479514 and K000479515. Meanwhile, a photo of a “type” (PH00024448, Fig. 1) was also located at PH (Thiers 2017), containing photographs taken by Epling between 1927 and 1928 of the specimens from K, along with images of specimens collected at the type locality by Isaac F. Holton 479 (23 November 1852, PH00024446) and J. Triana 3597 (11 May 1855, PH00024447), both collected at the same location. Additionally, other Goudot’s specimens collected in Bogotá were located: two in the P herbarium, P00715001 collected in Bogotá, locality of Boquerón, in March of 1844. This specimen is a duplicate of K000479515. And the specimen P00715002 collected in Bogotá in April of 1844 is a duplicate of K000479514. Finally, one in F (F0061175F - Catalog Number: 976885), without collection date. The photos of types reveal that the Kew specimens were originally mounted onto two mixed sheets: the first included a Goudot specimen alongside one collected by Holton, while the second combined a Goudot specimen with one collected by Triana. This means that both K sheets contained four different collections that later were separated and are now cataloged as individual specimens: Goudot’s (K000479514, Fig. 2; K000479515, Fig. 3), Holton’s (K004937578), and Triana’s (K004937579). Mixed-sheet mounting was a common practice implemented in the 1930s during the Great Depression (Forman and Bridson 1989). This practice was due to limited resources during the 19th and early 20th century, as mounting materials were costly and scarce. This forced institutions and collectors to group multiple specimens on the same sheet to save costs and storage space; however, there were misidentification errors, where collectors inadvertently grouped different species under the same collection number, a situation detected later during taxonomic revisions (Forman and Bridson 1989). By leaving a note on the specimen, Epling indicated that the material from the two sheets at K should be together as the type; they belong to two different collections and therefore cannot be considered one type specimen. For this reason, it is likely that the K staff mounted each plant separately without cross-referencing labels indicating they constituted a single specimen (ICN, Art. 8.3). Epling later reviewed additional type materials of Salvia bogotensis in other herbaria, wrongly designating P00715001 and P00715002 (P) as isotypes in 1928, and possibly F0061175F (F) as an isotype, though the 42 PhytoKeys 265: 39–48 (2025), DOI: 10.3897/phytokeys.265.165733 Ángela María Morales-Trujillo et al.: Lectotypification of Salvia bogotensis Figure 1. Photographs of a mixed collection of Salvia bogotensis Benth. (Lamiaceae) a photo of “type” from the Academy of Natural Sciences [PH (PH00024448, PH00024446, PH00024447)]. The arrows refer to the collections of Goudot s.n. authorship of this annotation is unclear. Regardless, duplicates of this collection have been found in multiple herbaria, all considered part of the same original collection (ICN, Art. 8.2). After the name Salvia bogotensis was published, in all subsequent studies the specimens deposited at K were considered the type collection (Wood and 43 PhytoKeys 265: 39–48 (2025), DOI: 10.3897/phytokeys.265.165733 Ángela María Morales-Trujillo et al.: Lectotypification of Salvia bogotensis Figure 2. Specimen of Salvia bogotensis Benth. (Lamiaceae) Goudot s.n. (K000479514), collected in April. Harley 1989; Fernández-Alonso 2003), without mentioning clearly which of the two specimen represents the type. Following the ICN Arts. 9.3, 9.14, and 9.17, a lectotype must be designated from the original material. If the lectotype designation refers to a single gathering but multiple specimens, a second-step lectotypification must restrict it to one specimen. Therefore, here we designate K000479515 as the lectotype that has a more complete specimen (leaves, inflorescences, and flowers) and its inclusion of an original note by Epling supporting its selection. Only one of the others Goudot’s collection, those of P (P00715001) correspond to an isolectotype (Fig. 4) since it has the same collection date of the lectotype. 44 PhytoKeys 265: 39–48 (2025), DOI: 10.3897/phytokeys.265.165733 Ángela María Morales-Trujillo et al.: Lectotypification of Salvia bogotensis Figure 3. Lectotype of Salvia bogotensis Benth. (Lamiaceae) Goudot s.n. (K000479515), collected in March. Salvia bogotensis Benth., Prodr. [A. P. De Candolle] 12: 312. 1848. Lectotype. Colombia • Dep. Cundinamarca, Boquerón, Bogotá, N. [Nouvelle] Granade, M. J. Goudot s.n. 1844 (K! first-step designed by Epling 1939:254), Second-step designated here: — K [barcode K000479515 image!]; isolectotype: P [barcode P00715001 image!]. Note. In a handwritten annotation on specimen K000479515 (Goudot s.n.), in 1927, Epling stated that this specimen should together be taken with K000479514 to jointly constitute type material. However, this last one corresponds to another collection with another date and locality collection. 45 PhytoKeys 265: 39–48 (2025), DOI: 10.3897/phytokeys.265.165733 Ángela María Morales-Trujillo et al.: Lectotypification of Salvia bogotensis Further remarks Salvia bogotensis is an endemic species of Colombia that grows in high-altitude regions (2500–4000 m) in El Cocuy (Boyacá), located at the Northern end of the Eastern Mountain range. It is characterized as a small to medium-sized shrub with narrow, bullate leaves, moderately short inflorescences, and an intense blue corolla (Fernández-Alonso 2003). On the other hand, Wood and Harley (1989) recognized three subspecies of Salvia bogotensis Benth. complex: subsp. aratocensis, subsp. sochensis, Figure 4. Isolectotype of Salvia bogotensis Benth. (Lamiaceae) Goudot s.n. (P00715001), collected in March. 46 PhytoKeys 265: 39–48 (2025), DOI: 10.3897/phytokeys.265.165733 Ángela María Morales-Trujillo et al.: Lectotypification of Salvia bogotensis and subsp. bogotensis, which exhibit overlapping distributions in Boyacá and Santander. Fernández-Alonso (2003) later elevated the first two to species rank (Salvia aratocensis and S. sochensis), while reclassifying S. bogotensis subsp. suratensis as S. aratocensis subsp. suratensis due to its close morphological affinity and restricted distribution to the Chicamocha Valley (Santander, department) (Fernández-Alonso 2003). Acknowledgments ÁMMT thanks the Instituto de Ecología A.C., Centro Regional del Bajío, the postgraduate course “Nomenclatura botánica y herramientas digitales en estudios taxonómicos,” and Dr. Juan José Ancona Aragón for his valuable contributions. Finally, we acknowledge the editor and two anonymous reviewers for their comments that enriched this manuscript. Additional information Conflict of interest The authors have declared that no competing interests exist. Ethical statement No ethical statement was reported. Use of AI No use of AI was reported. Funding This work was supported by Secretaría de Ciencia, Humanidades, Tecnología e Innovación - Secihti. Gobierno de México, Convocatoria Becas Nacionales para Estudios de Posgrado 2023. Author contributions Writing - review and editing: PHL, BYBG, ÁMMT, JGSK. Author ORCIDs Ángela María Morales-Trujillo https://orcid.org/0000-0002-5723-0160 Brenda Yudith Bedolla-García https://orcid.org/0000-0002-1560-1090 Patricia Hernández-Ledesma https://orcid.org/0000-0002-0071-4201 Jorge Gabriel Sánchez Ken https://orcid.org/0000-0001-9623-7245 Data availability All of the data that support the findings of this study are available in the main text. References Bentham G (1833) Labiatarum genera et species. Fasc. 3. James Ridgway & Sons, London. Botanicus Digital Library (2024) Missouri Botanical Garden. 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