Lepanthes ernestii (Salazar & Soto Arenas) Solano & E.Licona, 2025

Solano, Rodolfo, Licona, Ethian, Carral, Rodrigo & Téllez-Baños, Bruno, 2025, Two new species, nomenclatural notes and an updated checklist in Lepanthes (Orchidaceae, Pleurothallidinae) from Mexico, Phytotaxa 687 (1), pp. 21-64 : 27-41

Lepanthes ernestii (Salazar & Soto Arenas) Solano & E.Licona stat. nov.

≡ Lepanthes acuminata Schlter subsp. ernestii Salazar & Soto Arenas (1996: 51, f. 6).

Type:— MEXICO. Chiapas: Berriozábal, ca. km 16 del camino Berriozabal-El Cairo, selva del Ocote, ca. 1050 m, selva baja perennifolia achaparrada, ventosa, collected at 17 abril 1989, dryed at 18 Febrero 1989, M. Soto & E. Martínez 5405 (holotype: AMO!).

Taxonomic discussion:— Lepanthes ernestii is recognized by its ovate leaves, almost as long as the stem, raceme shorter than the leaf, ovate and shortly apiculate sepals, strongly bilobed petals, with quadrangular and rounded lobes, the upper one larger, lanceolate and falcate blades of the lip, and a mammilated, pubescent, yellowish appendix ( Figures 1–3 ). Initially, this taxon was considered a form of Lepanthes acuminata with subspecific rank, geographically isolated in northern Chiapas ( Salazar & Soto 1996). However, the discovery of morphologically similar individuals in Chiapas, Veracruz, and Guatemala, indicates that this taxon is not a subspecies, but a species represented by several populations with a wider distribution than originally thought. Lepanthes acuminata ( Figure 4 ) differs in its larger habit (10–12 cm tall vs. up to 5.0 cm tall), stems longer than the leaf (vs. as long as the leaf), larger flowers (10.0–11.0 mm tall vs. 7.0- 8.0 mm tall), lanceolate and long attenuate-acuminate sepals (vs. ovate and acute), petals barely bilobed with similar lobes (vs. prominently bilobed with lobes different in size), and an appendix incurved (parallel and folded against the lip’s connectives vs. perpendicular in relation to the connectives).

Distribution and ecology: — Mexico (Chiapas, Oaxaca, Veracruz), Guatemala (Alta Verapaz). The first records of this taxon in the flora of the states of Oaxaca and Veracruz ( Mexico), as well as that of Guatemala, are documented here. The best distribution model for L. ernestii had an AUC = 0.902, where Bio_10 (Mean Temperature of Warmest Quarter) was the most contributing variable with 65.7%, followed by Bio_14 (Precipitation of Driest Month) with 27.2.%, and Bio_5 (Max Temperature of Warmest Month) with 6.6%. The MaxEnt Jackknife test showed that Bio_5 (Max Temperature of Warmest Month) was the most contributing variable when used in isolation, thus providing the most useful information. Meanwhile, Bio_14 (Precipitation of Driest Month) was the variable that, when omitted, caused the greatest reduction in contribution, suggesting it provides unique information absent in the other variables. The model predicts four zones with high probability for the distribution of L. ernestii : the southern portion of Sierra Madre Oriental, Sierras of northern Oaxaca, Altos of Chiapas, and the Sierra Los Cuchumatanes ( Guatemala) along with the southern portion of the Sierra Madre of Chiapas ( Figure 5 ). The adjusted model had a cutoff value of 0.397, generating a potential occupancy area of 103,337.56 km 2, of which 76,865.13 km 2 are in Mexico (3.91% of its territory) and 26,472.43 km 2 in Guatemala (24.31% of its territory).

Lepanthes ernestii grows as an epiphyte on Quercus trunks, in the cloud forest, ranging between 1050–1460 m elev. Until recently, this taxon was thought to be restricted to the Central Plateau of Chiapas. However, the records presented here expand its distribution area to the neighboring states of Oaxaca and Veracruz, as well as to Guatemala. Flowers from April to June.

Conservation status: —Not evaluated by the IUCN Red List or the Extinction Risk Evaluation Method for Mexican wild flora species ( SEMARNAT 2019). Additional field studies are required to assess the viability of its populations and the quality of its habitat. In Mexico, L. ernestii is known from four disjunct localities (except those found in Veracruz), with a potential distribution covering 3.91% of the country’s surface area. In Veracruz, the population of this species is within a protected area, the Cofre de Perote National Park, where favorable habitat still exists for it. The populations in Chiapas and Oaxaca persist in remnant forests, close to human settlements, and face anthropogenic effects such as the expansion of urban areas, opening of lands for agriculture, or extraction of wood and firewood.

Additional specimens examined: — MEXICO. Chiapas: ca. km 16 del camino Berriozabal-El Cairo, selva del Ocote, 1050 m, selva baja perennifolia achaparrada, ventosa, 17 April 1989, M. Soto & E. Martínez 5408 ( AMO!). Veracruz: Atzacán, barranca de Metlac, 1460 m, 15 December 2019, E. Licona 95 ( MEXU!).

Other records: — MEXICO. Chiapas: Cacahoatán, 22 March 2023, P. Gómez s.n. (https://mexico.inaturalist. org/observations/183330816). Oaxaca: San Felipe Usila , November 2022, S. Avila (https://mexico.inaturalist. org/observations/145085412). Veracruz: Chocaman , Tenejapa , 1790 m, March 2016, A. Fernández (https://www. naturalista.mx/observations/2783859) . GUATEMALA. Alta Verapaz: San Pedro Carcha, Sencaal , 1245 m, E. Mó (photo!) ; Cobán, San Luis Chicoyou , 1420 m, E. Mo (photo!) .

Lepanthes luicitae Solano , sp. nov. (figs. 6, 7).

Type:— MEXICO. Oaxaca: Totontepec Villa de Morelos, Santa María Huitepec, en la colindancia con Ocotepec, 2073 m, 17º 12’ 22.11’’ N, 96º 0’ 33.07’’ W, 01 October 2009, R. Solano 2982 & A. Martínez (holotype: OAX-27192!).

Similar to Lepanthes thurstoniorum Salazar, Soto Arenas & O. Suarez (in Salazar & Soto, 1996: 203, f. 70), from which differs by its flexuous rachis (vs. zigzag), dorsal sepal as wide as the lateral ones (vs. lateral sepals almost 2/3 the width of the dorsal one), lateral sepals obliquely ovate and diverging from each other towards their apices (vs. obliquely triangular-ovate and subparallel to each other towards their apices), and petals forming butterfly wing-like blades (vs. forming oblong and oblique blades).

Description:— Epiphyte, caespitous herb, up to 4.5 cm tall, including the inflorescence. Roots 1.0 mm in diameter, flexuous, cylindrical, whitish. Stems 2.0–3.0 cm long, 0.4 mm in diameter, erect, straight, formed by 3–5 internodes, with an annulus 0.5–0.6 mm beneath the apex; completely covered by imbricate, funnel-shape, shortly apiculate, scarious, brownish sheaths, thickened and papillose-ciliate along the veins and apical margin. Leaf 10.0–14.0 × 6.0– 8.5 mm, fleshy, lenticular, ovate to elliptic-ovate, obtuse to rounded, shortly 2-lobed mucronate at the apex, green bright on adaxial surface, green dark and purple suffused on the abaxial one, attenuate at the base into a petiole 1.0– 1.6 mm long. Inflorescence as long as the leaf at the beginning of flowering, but eventually surpassing its length, from the annulus, erect, racemose; peduncle 3.0–5.0 mm long, 0.2 mm in diameter, straight, filiform, purple, with 1–2 funnel-shaped, obtuse, shortly apiculate, purple-translucent bracts, papillose-ciliate along the vein and the apical margin, 1.0– 1.3 mm long; rachis flexuous, producing up to 9 successive flowers. Floral bracts 0.8–1.2 mm long, obliquely funnel-shaped, obtuse, shortly apiculate, hyaline, carinate, and muriculate along the vein. Flowers extended; sepals yellow to yellowish translucent, red striped along the veins, with reddish cilia, petals yellowish orange to dark yellow, red-spotted toward their margins, lip red, column reddish, anther whitish and red suffused. Dorsal sepal 6.2–7.2 × 3.6–4.0 mm, ovate, acute, ciliate, 3-veined; lateral sepals 5.3–6.2 × 3.3–3.5 mm, united each other near the middle of their length, obliquely ovate, acute, ciliate, slightly divergent at their apices, 2-veined. Petals 4.2–4.6 × 1.0 mm, asymmetric, transversally 2-lobed, minutely apiculate, 1-veined, minutely glandular-papillose; upper lobes 2.1–2.3 × 0.7–1.0 mm, obliquely triangular-lanceolate, subrounded, divergent each other; basal lobes 1.9–2.0 × 0.4– 0–7 mm, narrowly triangular, acute, parallel each other. Lip 3-lobed; blades 2.2–2.3 × 0.4 mm, slightly sigmoid, narrowly lanceolate, glabrous, recurved along the external margins, obliquely rounded at the proximal end, acuminate and incurved at the distal end, united by trapezoidal connectives, 0.5 mm long and wide; appendix 0.2–0.3 × 0.2 mm, oblong, truncate, recurved, pubescent at the base. Column 1.8 mm long, 0.7 mm wide, claviform, widened toward the apex; clinandrium truncate. Stigma subapical, horseshoe-shape, concave; rostellum apical, transversally laminar, ending into a minute, erect, laminar projection. Anther 0.8 × 0.6 mm, dorsal, ovate-subquadrate. Pollinarium 0.6 mm long, with 2 claviform, yellow pollinia, united by a drop of viscidium. Ovary 1.1 mm long, 0.4–0.5 mm diameter, arching; pedicel 0.6–1.3 mm long, 0.25 mm diameter. Capsule not seen.

Distribution and ecology: —Endemic to Mexico, in Oaxaca. The best distribution model for L. luicitae had an AUC = 0.975, where Bio_8 (Mean Temperature of Wettest Quarter) was the most contributing variable with 42.4%, followed by Bio_3 (Isothermality) with 22.3%, and Bio_15 (Precipitation Seasonality) with 19.2%. The MaxEnt Jackknife test showed that Bio_5 (Max Temperature of the Warmest Month) was the most contributing variable when used in isolation, thus providing the most useful information. On the other hand, Bio_3 (Isothermality) was the variable that, when omitted, caused the greatest reduction in contribution, suggesting it provides unique information absent in the other variables. The model predicts a single region with high probability for the distribution of L. luicitae , which would be a micro-endemism restricted to Sierras of northern Oaxaca ( Figure 8 ). The adjusted model had a cutoff value of 0.286 and generated a potential occupancy area of 4,013.10 km 2, which corresponds to 0.20% of the Mexican territory.

Lepanthes luicitae ranges between 1850–2400 m, where it grows as an epiphyte on small to medium tree trunks, in shaded, humid, and well-ventilated sites within the cloud forest ( Figure 7 ). The climate is semi-warm humid, with average annual temperature of 18º–20º C, and an annual precipitation higher than 1700 mm, much of it in the form of fog, which is frequent throughout the year.

It has been seen flowering from November to April.

Etymology: —The specific name of this taxon is dedicated to Luicita Lagunez, in recognition of her contribution to the ethnopharmacological study of Mexican orchids.

Additional specimens examined (paratypes): — MEXICO. Oaxaca: San Juan Juquila Vijanos, Tarántulas, 1850 m, bosque mesofilo de montaña, 7 December 2004, R. Solano s.n., R. Rivera & A. Ríos (OAX[in spirit and drawing]!). San Juan Juquila Vijanos, 1.5 km antes de la desviación de la carretera Juquila Vijanos a Talea de Castro, entre los parajes El Mirador y La Virgen, 2000 m, 10 August 2022, C. Tomás-López (OAX[photo!]). Santiago Xiacui, bosque mesofilo de montaña, 2000 m, February–May 2013, L. Jiménez-Bautista s.n. (OAX[photo]!). Totontepec Villa de Morelos, Santa María Huitepec, en la colindancia con Ocotepec, 2073 m, 12 March 2009, R. Solano 2926 & A. Martínez (OAX-27127!). Totontepec Villa de Morelos, Santa María Huitepec, ca. 1 km sobre el camino a Huitepec partiendo de la carretera Mitla-Totontepec, 2427 m, 14 March 2009, R. Solano 2949 (OAX-27150!), R. Solano 2994 & A. Martínez (OAX-27183!).

Other records: — MEXICO. Oaxaca, Villa Talea de Castro, 23 Sepember 2024, C. Tomás-López (https://mexico. inaturalist.org/observations/244400266).

Taxonomic discussion: — Lepanthes luicitae is distinguished by its elliptical leaves, a raceme progressively elongating when new flowers are produced to be longer than the leaf, ovate, ciliated sepals, with the lateral ones diverging each other at their apices, narrowly lanceolate blades of the lip, and an oblong and pubescent appendix ( Figures 6–7 ). The more similar species is Lepanthes thurstoniorum ( Figure 9A ), which becomes sympatric in some localities. However, it differs by its zigzag rachis (vs. flexuous rachis), lateral sepals almost 2/3 the width of the dorsal one (vs. dorsal sepal as wide as the lateral ones), lateral sepals obliquely triangular-ovate and subparallel to each other towards their apices (vs. obliquely ovate and diverging from each other towards their apices), and petals forming oblong and oblique blades (vs. forming butterfly wing-like blades).

Lepanthes gabriellae Salazar & Soto (1996: 97) is another similar species ( Figure 9B ) and becomes sympatric with L. luicitae . However, the first differs in having an inflorescence shorter than the leaf (vs. longer than the leaf), zigzag rachis (vs. flexuous), prominently ciliated sepals (vs. shortly ciliate), oblong and truncate to rounded lobes of the petals (vs. obliquely triangular, acute to subrounded), and obliquely oblong blades of the lip (vs. narrowly lanceolate).

Another similar species is L. schultesii Salazar & Soto (1996: 183) ( Figure 9C ), also endemic in northern Oaxaca and sympatric with L. luicitae , which differs by its longer peduncle (10.0–15.0 mm vs. 3.0–5.0 mm), smaller flowers (7.2–8.5 mm × 2.2. mm vs. 8.0– 13.2 mm × 5.0– 5.5 mm), narrowly ovate sepals with prominent cilia (vs. ovate, shortly ciliate), inward apices of the lateral sepals (vs. divergent apices), smaller petals (1.6–2.0 mm vs. 4.2–4.6 mm) with oblong lobes (vs. triangular lobes), and ovate-lanceolate blades of the lip (vs. narrowly lanceolate).

Lepanthes schiedei is somewhat similar too ( Figure 9D ), but it differs by its narrower flowers (2.0– 2.5 mm in wide vs. 55.5 mm in wide), triangular-lanceolate not ciliate sepals (vs. ovate and ciliate), oblong lobes of the petals (vs. triangular), the upper ones overlapping at their apices (vs. non-overlapping), and lanceolate blades of the lip (vs. narrowly lanceolate).

Conservation status. Not evaluated by the IUCN Red List nor the Extinction Risk Evaluation Method for Mexican wild flora species ( SEMARNAT 2019). Additional field studies are required to assess the viability of its populations and the quality of its habitat. So far, the species is known from five very close localities in a restricted region, with a potential distribution covering 0.20% of the country’s surface area. These localities occur within or near natural protected areas by local communities in Oaxaca, where the forest has a management program compatible with the conservation. However, beyond these areas, the forest is being altered by agricultural practices of itinerant or permanent crops (Pérez-García & del Castillo-Sánchez 2016, 2017). The increase in human settlements and the border effects associated with the aperture of new roads in the region is an additional risk for the species.

Plants of Lepanthes luicitae grow in the richest Mexican region for Lepanthes , the northern highlands of Oaxaca. So far, 27 species have been recorded, representing 39.7 % of the genus’s richness in Mexico.

Lepanthes machorroi Salazar & Soto Arenas (1996: 121, f. 35).

Type: — MEXICO. Oaxaca: 2 km de Totontepec [Villa de Morelos] Mixes hacia [San Pedro y San Pablo] Ayutla, 2000 m, colectado 15 Sep 1991, prensado en cultivo 20 Feb 1993, R. Jiménez 1524 & O. Suárez (holotype: AMO!).

Lepanthes machorroi is distinguished among the species of the genus in Mexico by its white flowers, abruptly acuminate-caudate and ciliate sepals, symmetric, apiculate, and papillose petals, oblong laminae of the lip, emarginate at the apex, with sharp margins, and prominently papillose, connectives of the lip with a hemispheric ventral projection, and a filiform and glabrous appendix ( Figure 10 ). This species was previously only known from three localities in Northern Highlands of Oaxaca, Mexico ( Salazar & Soto 1996). Thus, the distribution of the species extends its range to a more northern portion in the neighboring state of Veracruz.

Distribution and ecology: —Endemic to Mexico, in Oaxaca and Veracruz. The first record of the species for the flora of Veracruz state is reported here. The best distribution model for L. machorroi had an AUC = 0.986, where Bio_5 (Max Temperature of Warmest Month) was the most contributing variable with 31.0%, followed by Bio_7 (Temperature Annual Range) with 29.5%, and Bio_2 (Mean Diurnal Temperature Range) with 12.9%. The MaxEnt Jackknife test showed that Bio_7 (Temperature Annual Range) was the most contributing variable when used in isolation, thus providing the most useful information. On the other hand, Bio_4 (Temperature Seasonality) was the variable that, when omitted, caused the greatest reduction in contribution, suggesting it provides unique information absent in the other variables. The model predicts two areas with a high probability for the distribution of L. machorroi , one in the southern part of the Sierra Madre Oriental and the other in Sierras of northern Oaxaca ( Figure 11 ). The adjusted model had a cutoff value of 0.301 and generated a potential occupancy area of 10,198.23 km 2, which corresponds to 0.52% of the Mexican territory.

Lepanthes machorroi grows as an epiphyte in the cloud forest, ranging between 2000–2250 m elev. In Veracruz the species was found growing on Quercus delgadoana Valencia, Nixon & Kelly (2001: 275) , together with Lepanthes avis , L. calopetala Salazar & Soto (1996: 77) , L. salvaterrae , and L. totontepecensis Salazar & Soto (1996: 206) . It flowers between June and November.

Conservation status: —Not evaluated by the IUCN Red List or the Extinction Risk Evaluation Method for Mexican wild flora species ( SEMARNAT 2019). Additional field studies are required to assess the viability of its populations and the quality of its habitat. The species is known from five localities, one in Veracruz and four more in Oaxaca, which are very close to each other, with a potential distribution covering 0.52% of the country’s surface area. In Veracruz, the known populations of this species are within a protected area. the Salvaterra Ecological Reserve. It is also very close to the Cofre de Perote National Park, which could provide additional favorable habitat. The localities in Oaxaca occur within or near natural protected areas by local communities, where the forest has a management program compatible with the conservation. However, beyond these areas, the forest is being altered by agricultural practices of itinerant or permanent crops (Pérez-García & del Castillo-Sánchez 2016, 2017).

Additional specimen examined: — MEXICO. Veracruz: Xico, Rancho Salvaterra, localidad Buenavista, 2137 m, 08 July 2023, R. Carral et al. RCD-876 (OAX!).

Lepanthes papilionacea Salazar, Soto Arenas & O. Sua ́rez (in Salazar & Soto 1996: 161, f. 53).

Type: — MEXICO. Oaxaca: [Santiago Comaltepec] carretera Oaxaca-Tuxtepec, km 115.9, 2400 m, 1 October 1978, O. Suárez 315-1 & E. W. Greenwood (holotype: AMO!, isotype: AMO [in spirit]!).

Lepanthes papilionacea is distinguished from its congeners by its sepals with long, rigid, and spaced-out cilia, ascending on the dorsal sepal and descending on the lateral ones, with some cilia along the veins on the abaxial surface; the petals are strongly bilobed, with highly dissimilar lobes; the appendix of the lip is formed by a triangular, glabrous, deflexed blade surrounded by stiff hairs ( Figure 12 ). This species was previously only known from three widely separated localities, one in the mountains of northern Oaxaca ( Salazar & Soto 1996), other two in region known as Huasteca, in Queretaro (García-Cruz et al. 2003) and San Luis Potosi ( Fortanelli-Martínez et al 2021). The new locality in Veracruz shows continuity among widely dispersed populations known for the taxon.

Distribution and ecology: —Endemic to Mexico, in Oaxaca, Queretaro, San Luis Potosi, and Veracruz. The first record of the species for the flora of Veracruz state is reported here. The best distribution model for L. papilionacea had an AUC = 0.944, where Bio_2 (Mean Diurnal Temperature Range) was the most contributing variable with 43.8%, followed by Bio_5 (Max Temperature of Warmest Month) with 28.5%, and Bio_DEM (Mexican Digital Elevation Model) with 26.5%. The MaxEnt Jackknife test showed that Bio_5 (Max Temperature of Warmest Month) was the most contributing variable when used in isolation, thus providing the most useful information. Meanwhile, Bio_2 (Mean Diurnal Temperature Range) was the variable that, when omitted, caused the greatest reduction in contribution, suggesting it provides unique information absent in the other variables. The model predicts three areas with high probability for the distribution of L. papilionacea : two of them in the Sierra Madre Oriental, one in its central portion, known as the Huastec region (Queretaro and San Luis Potosi), the other in the southern part of it; Sierras of northern Oaxaca ( Figure 13 ). The adjusted model had a cutoff value of 0.396 and generated a potential occupancy area of 23,842.53 km 2, which corresponds to 1.21% of the Mexican territory.

Lepanthes papilionacea grows as an epiphyte in the cloud forest and Quercus -Pinus forest, ranging between 2000– 2400 m elevation. In Veracruz it was found growing on Quercus delgadoana , together with Lepanthes salvaterrae , and L. totontepecensis . Flowers between April to November.

Conservation status: —Not evaluated by the IUCN Red List or the Extinction Risk Evaluation Method for Mexican wild flora species ( SEMARNAT 2019). Additional field studies are required to assess the viability of its populations and the quality of its habitat. Lepanthes papilionacea is known from six disjunct localities (except the two between the border of Queretaro and San Luis Potosi, and the two in northern Oaxaca), with a potential distribution covering 1.21% of the country’s surface area. Only three populations are protected, one in Oaxaca in forests under community protection, and the other in Veracruz, in the Salvaterra Ecological Reserve. It is also very close to the Cofre de Perote National Park, which could provide additional favorable habitat.

Additional specimens examined: — MEXICO. Veracruz: Xico, Rancho Salvaterra, localidad Buenavista, 2059 m, 26 September 2021, R. Carral et al. RCD-871 ( OAX!). Xico, Rancho Salvaterra, localidad Buenavista, 2059 m, 08 July 2023, R. Carral RCD-874 ( OAX!), R. Carral RCD-875 ( OAX!), R. Carral RCD-875.

Lepanthes salvaterrae Solano, J.R.Carral & Téllez , sp. nov. (figs 14, 15)

Type: — MEXICO. Veracruz: Municipio Xico, Rancho Salvaterra, localidad Buenavista, bosque mesófilo de montaña, 2000 m, 07 August 2021, R. Carral, B. Téllez-Baños, I. Escamilla-Gómez & L. Islas-Tello RCD-863 (holotype: OAX!).

Similar to L. gabriellae from which differs by its peduncle shorter than the leaf (vs. equal or longer than the leaf), orbicular to orbicular-elliptic leaves (vs. narrowly elliptic leaves), sepals marginally glabrous (vs. with apparent reddish cilia), and petals with upper and lower lobes different in shape and size (vs. similar to each other).

Description:— Epiphyte, caespitous herb, 3.2–6.4 cm tall including the inflorescence. Roots 0.4–0.8 mm diameter, flexuous, whitish. Stems 9.0–27.0 mm long, 0.2–0.4 mm diameter, erect, straight, formed by 3–5 internodes, with an annulus close to the apex; completely covered by imbricate, funnel-shape, apiculate, scarious, brownish sheaths, papillose-ciliate along the veins and the apical margin. Leaf 10.0–16.5 × 5.0– 8.5 mm, fleshy, orbicular to orbicular-elliptic, rounded, shortly 2-lobed and mucronate at the apex, green bright on both surface, attenuate at the base into a petiole 1.0–2.0 mm long. Inflorescence up to 26.0 mm long, from the annulus, erect, racemose, longer than the leaf; peduncle 7.0– 25.5 mm long, 0.12–0.16 mm in diameter, straight, filiform, greenish, with 1 funnel-shaped, obtuse, purple-translucent bract, ca. 1.0 mm long; rachis flexuous, producing up to 3 successive flowers. Floral bracts 0.8–1.3 × 1.0– 1.2 mm when spread out, obliquely funnel-shaped, obtuse, apiculate, brownish. Flowers 7.0– 11.2 mm tall, 2.0– 3.7 mm wide, extended; sepals cream or yellow pale, translucent, densely suffused with red-purple, petals yellowish orange, with the upper lobe purple along the inner margin, lip, column, and anther cap purple. Dorsal sepal 4.0–5.0 × 1.6–2.8 mm, ovate-lanceolate, acute-acuminate, 3-veined, glabrous; lateral sepals 3.0–5.0 × 1.3–2.0 mm, united each other almost one-third of their length, obliquely ovate-lanceolate, acuminate, marginally papillose, 2-veined. Petals 2.8 × 1.0 mm, oblong-oblanceolate, asymmetric, transversally 2-lobed, 1-veined, minutely glandular-papillose; upper lobes 1.2–1.5 × 0.8–1.0 mm, quadrangular, rounded to subtruncate, slightly overlapping at their apices; basal lobes 1.1–1.3 × 0.4–0.7 mm, obovate-triangular, obtuse, divergent each other at their apices. Lip 1.5–1.6 × 0.8–1.0 mm, 3- lobed; blades 1.5–1.6 × 0.4 mm, obliquely lanceolate-falcate, rounded at the proximal end, acuminate at the distal end, glabrous, united by trapezoidal connectives; appendix 0.1 × 0.8 mm, oblong, recurved, yellowish, densely covered with whitish hairs. Column 1.3 mm long, 0.6 mm wide, claviform, widened toward the apex; clinandrium truncate. Stigma subapical, concave; rostellum apical, transversally laminar. Anther 0.5 × 0.4 mm, dorsal, ovate. Pollinarium 0.47 mm long, with 2 ovate-claviform, pollinia, yellowish toward their bases, united by an adhesive viscidium. Ovary 1.5–1.7 mm long, 0.4–0.5 mm diameter; pedicel 0.4–2.7 mm long, 0.15 mm diameter. Capsule 2.3 mm long, 3.0 mm diameter, ellipsoid, longitudinally thickened along the ribs and suture lines.

Distribution and ecology: —Currently, this species is a Mexican micro-endemism restricted to the southern part of the Sierra Madre Oriental in Veracruz ( Figure 16 ). Due to the limited number of known localities, all located in a very small area, it was impossible to create a distribution model for this species. However, it is possible that the species may also inhabit nearby municipalities close to Xico.

Lepanthes salvaterrae is an inhabitant of the cloud forest, ranging between 2000–2160 m elev. It grows as an epiphyte on trunks of Quercus delgadoana , Liquidambar styraciflua Linnaeaus (1753: 999) , Vaccinium leucanthum Schlechtendal (1834: 524) , and Quercus corrugata Hooker (1841: 403) , together with Lepanthes acuminata , L. calopetala , L. moorei , L. papilionacea , and L. totontepecensis . It flowers from August to November.

Etymology: —The specific name comes from Salvaterra, the name of a privately owned forested land ranch that preserve fragments of cloud forest. Located in Veracruz, this land is dedicated by its owners to the conservation of this habitat, where the species was discovered.

Additional specimens examined (paratypes): — MEXICO. Veracruz: Xico, Rancho Salvaterra, localidad Buenavista , 2000 m, 07 August 2021, R. Carral, B. Téllez-Baños, I. Escamilla-Gómez & L. Islas-Tello RCD-860 ( OAX!) ; R. Carral et al. RCD-861 ( OAX!), R. Carral et al. RCD-862 ( OAX!), R. Carral et al. RCD-865 ( OAX!), R. Carral et al. RCD-866 ( OAX!). Xico , Rancho Salvaterra , localidad Buenavista , 2066 m, 07 Aug 2021, R. Carral et al. RCD-864 ( OAX!). Xico, Rancho Salvaterra, localidad Buenavista, 2051 m, 07 Aug 2021, R. Carral et al. RCD-867 ( OAX!), R. Carral et al. RCD-868 ( OAX!) ; R. Carral et al. RCD-869 (OAX!).

Taxonomic discussion: — Lepanthes salvaterrae is recognized among its Mexican congeners by its orbicular to orbicular-elliptic leaves, flowers almost as long as the leaf, cream or pale yellow in color and densely spotted with red-purple, with ovate-lanceolate, acuminate, and glabrous sepals ( Figures 14–15 ). The species is florally similar in shape and color to L. gabriellae ( Figure 9A ), but it is distinguished by sepals with reddish cilia apparent on their margins (vs. entire sepals), petals with upper and lower lobes similar to each other (vs. lobes different in shape and size), and its narrowly elliptic leaves (vs. orbicular to orbicular-elliptic).

Lepanthes brachystele Salazar & Soto (1996: 71 , f. 14) is somewhat similar ( Figure 17A ), but it is distinguished by its narrowly elliptic to elliptic-lanceolate leaves (vs. orbicular to orbicular-elliptic), inflorescence shorter than the leaf (vs. longer than the leaf), rachis with floral bracts subimbricated each other (vs. separated from each other), and ovate and acute sepals (vs. ovate-lanceolate and acuminate).

Another similar taxon is L. calopetala ( Figure 17B ) which differs by its lanceolate leaves (vs. orbicular to orbicular-elliptic), inflorescence shorter than the leaf (vs. longer than the leaf), ovate and obtuse sepals (vs. ovate-lanceolate and acuminate), boomerang-shaped petals (vs. no boomerang shaped), and ovate laminae of the lip (vs. lanceolate-falcate).

Conservation status: —Not evaluated by the IUCN Red List nor the Extinction Risk Evaluation Method for Mexican wild flora species SEMARNAT 2019). Although L. salvaterrae is known from various sites, all of them are in the same locality, within a privately owned reserve that protects its habitat. This locality is located close to Xico, a tourist city in Veracruz, where the development of tourism infrastructure could impact the remaining fragments of cloud forest and the habitat of the species. Additionally, this region is affected by the conversion of forests into coffee plantations and increasing urbanization. However, nature tourism activities are offered there, and some coffee producers strive for their products to be organic, which represents conservation-friendly actions. In Veracruz, Rancho Salvaterra ecological reserve includes one of the richest localities for Lepanthes in Mexico, where eight species have been recorded so far: L. acuminata , L. avis , L. calopetala , L. machorroi , L. moorei , L. papilionacea , L. salvaterrae , and L. totontepecensis .

Figures

FIGURE 1. Lepanthes ernestii. A) habit. B) Close-up of stems with inflorescences. C) Flower, frontal view. D) Flower, lateral view. E) Floral dissection. F) Lip and column, frontal and lateral views. Plate by E. Licona, based on E. Licona 95.

FIGURE 2. Lepanthes ernestii. A) Habit. B) Inflorescence with one open flower. C) Flower, frontal view. D) Sepals dissected. E) Lip, column, and ovary, lateral view. F) Petals. G) lip. H) Column, ventral, lateral, and dorsal view. I) Anther cap. J) Pollinarium. Drawing form R. Jiménez, reproduced with the permission of AMO Herbarium.

FIGURE 3. Lepanthes ernestii. A) specimen from Veracruz, Mexico, photograph by Adrian Fernández. B) specimen from Alta Verapáz, Guatemala, photograph by E. Mó.

FIGURE 4. Lepanthes acuminata. A) specimen from Las Margaritas, Chiapas, photograph by E. Licona. B) specimen from Veracruz, photograph by E. Hágsater based on E. Hágsater 4445.

FIGURE 5. Potential distribution of Lepanthes ernestii. The points correspond to the known localities of the species. Map by E. Licona.

FIGURE 6. Lepanthes luicitae. A) habit.B) caulinar sheet, detail.C) Inflorescence, segment. D) Flower, frontal view.E) Flower dissected. F) Ovary, lip, and column, lateral view. G) lip and appendix. H) Column, ventral and dorsal views. I) Anther cap and pollinarium. Line drawing by R. Solano based on the type.

FIGURE 7. Lepanthes luicitae. A) Specimen from Santa Maria Huitepec, Oaxaca, photograph by Rodolfo Solano. B) Specimen from San Juan Juquila Vijanos, Oaxaca, photograph by C. Tomás-López.

FIGURE 8. Potential distribution of Lepanthes luicitae. The points correspond to the known localities of the species. Map by E. Licona.

FIGURE 9. A) Lepanthes thurstoniorum, photograph by Edward W. Greenwood, based on O. Suárez 294-1. B) Lepanthes gabriellae, photograph by Gerardo Salazar, based on G. Salazar 5140. C) Lepanthes schiedei, photograph by Rodolfo Solano, based on G. Salazar 5150. D) Lepanthes schultesii, photograph by Edward W. Greenwood, based on O. Suárez 294-4.

FIGURE 10. Lepanthes machorroi. Specimen from Xico, Veracruz, photograph by B. Téllez-Baños, based on R. Carral-Dominguez et al. 876.

FIGURE 11. Potential distribution of Lepanthes machorroi. The points correspond to the known localities of the species. Map by E. Licona.

FIGURE 12. Lepanthes papilionacea. Specimen from Xico, Veracruz, photographs by B. Téllez-Baños, based on R. Carral-Domínguez 874.

FIGURE 13. Potential distribution of Lepanthes papilionacea. The points correspond to the known localities of the species. Map by E. Licona.

FIGURE 14. Lepanthes salvaterrae. A) Habit. B) Leaf. C) Stem with lepanthiform sheaths. D) Flower, three-quarter view. E) Flower, frontal view. F) Sepals dissected. G) Ovary, column, and lip, lateral view. H) Petal. I) Lip. J) Appendix of the lip. K) Anther cap and pollinia. L) Capsule. Plate by B. Téllez-Baños.

FIGURE 15. Lepanthes salvaterrae. Specimen from Xico, Veracruz, photograph by B. Téllez-Baños.

FIGURE 16. Distribution map of Lepanthes salvaterrae. The points correspond to the known localities of the species. Map by E. Licona.

FIGURE 17. A) Lepanthes brachystele. Specimen from Oaxaca, photograph by Glenn Pollard based on G. Pollard L-143. B) Lepanthes calopetala. Specimen from Xico, Veracruz, photograph by R. Carral-Domíngez.