Revista Mexicana de Biodiversidad Revista Mexicana de Biodiversidad 93 (2022): e934181 Taxonomy and systematics Lectotypification of Moussonia (Gesneria) triflora (Gesneriaceae) Lectotipificación de Moussonia (Gesneria) triflora (Gesneriaceae) Angélica Ramírez-Roa a, *, Laurence E. Skog b a Universidad Nacional Autónoma de México, Instituto de Biología, Departamento de Botánica, Herbario Nacional MEXU, Tercer Circuito, Ciudad Universitaria, Coyoacán, 04510 Ciudad de México, Mexico b Smithsonian Institution, National Museum of Natural History, Department of Botany, P.O. Box 37012, Washington D.C. 20013-7012, USA *Corresponding author: aramroa@ib.unam.mx (A. Ramírez-Roa) Received: 22 September 2021; accepted: 29 June 2022 Abstract Moussonia triflora was originally published as Gesneria triflora in 1842 by M. Martens and H. Galeotti without the designation of a nomenclatural type. The analysis of the nomenclatural history includes its being part of 3 genera and as 2 species. The species is one of the 24 taxa included in Moussonia and is known only from the states of Oaxaca and Veracruz in Mexico. The lectotypification of the species is proposed. In addition, an amended description is provided, along with a listing of the examined specimens cited, as well as a key to distinguish similar taxa. Keywords: H. G. Galeotti; J. J. Linden; Kohleria; Mexico; Oaxaca; Veracruz Resumen Moussonia triflora fue publicada como Gesneria triflora en 1842 por M. Martens y H. Galeotti sin designarse un tipo nomenclatural. Se analiza la historia nomenclatura del taxón y su pertenencia a 3 géneros y 2 nombres de especies. La especie es una de los 24 taxones incluidos en Moussonia y es conocida solamente de los estados de Oaxaca y Veracruz, México. Se propone la lectotipificación de la especie. Se presenta una descripción corregida junto con una lista de los ejemplares examinados y una clave de determinación de taxones relacionados. Palabras clave: H. G. Galeotti; J. J. Linden; Kohleria; México; Oaxaca; Veracruz ISSN versión electrónica: 2007-8706; Universidad Nacional Autónoma de México, Instituto de Biología. Open Access bajo la licencia CC BY-NC-ND (4.0) https://doi.org/10.22201/ib.20078706e.2022.93.4181 A. Ramírez-Roa, L.E. Skog / Revista Mexicana de Biodiversidad 93 (2022): e934181 2 https://doi.org/10.22201/ib.20078706e.2022.93.4181 Introduction Oaxaca, T. MacDougall 506 (US) (fig. 1), as well as C. Seler & E. G. Seler 5182 (B, not located), indicating this In one of their 19 articles entitled “Enumeratio last specimen was “cited by Fritsch”. synoptica plantarum phanerogamicarum, ab Henrico In 1975 Wiehler re-established Moussonia as a Galeotti in regionibus Mexicanis collectarum”, Martin genus distinct from Kohleria, and included M. triflora Martens and Henri Guillaume Galeotti in 1842 published (M.Martens & Galeotti) Hanst., indicating again Gesneria an enumeration of 14 species of Gesneriaceae mostly triflora M.Martens & Galeotti as the basionym and K. collected by Galeotti during his travels to Mexico from martensii as a synonym. No mention of a type or any other 1835 to 1840 (Crépin, 1883; Martens & Galeotti, 1842; specimen was given. McVaugh, 1978). Nine of those 14 species were new to More recently, Moussonia triflora was included in the science, and in the protologues of the new species the list of taxa in the reorganization of tribal and generic authors included: 1) an indication of the “nobis” (or boundaries of the Gloxinieae tribe by Roalson et al. “new”) status; 2) a description in Latin; 3) a discussion of (2005), as well as in the Index to the Gesneriaceae of the similar species and their differences from the new species; New World (Clark et al., 2020). Also, when one looks 4) information on the locality, altitude, and phenology; for Moussonia triflora (M.Martens & Galeotti) Hanst. in and 5) Galeotti’s collection number of the specimens that the International Plant Name Index (IPNI, 2021) (https:// would become the types of the new species. The entry for www.ipni.org/n/77221901-1), its basonym is shown to Gesneria triflora M.Martens & Galeotti, now Moussonia be Gesneria triflora M.Martens & Galeotti. Likewise, triflora (M.Martens & Galeotti) Hanst., includes all this when searching for this latter name (https://www.ipni. pertinent information except for an explicit citing of a org/n/381194-1), it is indicated that it is precisely a synonym Galeotti specimen that would at present be considered of Moussonia triflora (M.Martens & Galeotti) Hanst., the holotype. without mentioning other associated names. However, if Gesneria triflora was accepted and included in one searches for Kohleria martensii Fritsch (https://www. monographs by workers on Gesneriaceae. For example, ipni.org/n/133786-2), then that name is nomenclaturally Hanstein (1865) transferred the species from Gesneria to the linked only to G. triflora M.Martens & Galeotti without genus Moussonia, including as information on its origin a mention of M. triflora. In addition, if one attempts “Patria: Mexico: Galeotti”, and citing G. triflora (hence the to search for Gesneria triflora Hook., no results are basionym). Fritsch (1913) later reduced the genus Moussonia to found. Nevertheless, searching in the platform of one of the sections of Kohleria. However, as there already Tropicos.org (2022), for the various species’ names existed the combination K. triflora (Hook.) Regel, based on mentioned above one finds a lack of clarity. When one G. triflora Hook. (Hooker, 1847), which currently searches for M. triflora (https://www.tropicos.org/ corresponds to K. hondensis (Kunth) Hanst. (Kvist & name/50081504), one finds that Hanstein is indicated as Skog, 1992), Fritsch then established K. martensii Fritsch as the author of the species’ name but not noting that M. replacement name for G. triflora M.Martens & Galeotti, Martens & Galeotti were the authors of the basionym nor even though the latter was the earlier homonym. In the same whether Moussonia triflora Hanst. is legitimate or work, Fritsch stated that he had not seen the type specimen illegitimate. In addition, on the page for M. triflora, 2 for G. triflora M.Martens & Galeotti and suggested that names appear, the “Replaced synonym” Gesneria triflora Galeotti may not have collected the species. Under his M.Martens & Galeotti, considered there as an illegitimate proposed replacement name K. martensii indicated a single name and, G. triflora Hook., indicated as a legitimate representative specimen from Veracruz: “Im feuchten Walde, “blocking name”. When one searches for G. triflora Cerro de Chocaman hinter Cordoba”, M.Martens & Galeotti (https://www.tropicos. org/ C. Seler & E.G. Seler 5182 (B, not located). name/50081505), 2 “New names” are shown, Kohleria Conrad Morton, in 1967, published a synopsis of martensii indicated as an illegitimate name and M. triflora the genus Kohleria for Mexico, mostly following with no indication of legitimacy or not. Furthermore, Fritsch’s (1913) concept of the genus, and in the when one searches on the website for Kohleria martensii publication K. martensii was considered to be rare and (https://www.tropicos.org/name/50081503), the name is local species in Veracruz and Oaxaca. Morton included recognized as nom. illeg. superfl., replacing the “Replaced Gesneria triflora M.Martens & Galeotti (which was synonym” G. triflora M.Martens & Galeotti, and the the replaced name), indicating “Mirador, Veracruz, “Blocking name” G. triflora Hook, with no mention Mexico” as the type locality, and “Galeotti” as the of the latter name as being legitimate or illegitimate. collector. He included 3 additional representative In addition, this page includes M. triflora Hanst. as the specimens: Mount Orizaba, Veracruz, E. W. Nelson “Nomenclaturally correct name”. It should be noted that 294 (US), Comaltepec, above Vista Hermosa, Distr. Ixtlán, K. martensii was not an illegitimate name when it was A. Ramírez-Roa, L.E. Skog / Revista Mexicana de Biodiversidad 93 (2022): e934181 3 https://doi.org/10.22201/ib.20078706e.2022.93.4181 Figure 1. Moussonia triflora (M. Martens & Galeotii) Hanst. MacDougall 506 (US). published, according to Art. 52.1 of the International Code to locate neither the type specimen supposedly collected of Nomenclature (Turland et al., 2018), as it was not a by Galeotti nor the specimen Seler & Seler 5182 reported superfluous name, but a replacement name for G. triflora by Fritsch (1913) and Morton (1967) under Kohleria M.Martens and Galeotti, as the blocking name Kohleria martensii. However, new information about their Galeotti’s triflora (Hook.) Regel (= Kohleria hondensis) had been and Linden’s travels in México, and their location of published earlier based on Gesneria triflora Hook. their collections now make it possible to propose the Finally, in the revision of the genus Moussonia, lectotypification of M. triflora in this work. An amended Ramírez-Roa (2017) included M. triflora as 1 of the 23 description is provided below, as well as additional recognized species, mentioning that the author was unable specimens and images not included in the 2017 work. A. Ramírez-Roa, L.E. Skog / Revista Mexicana de Biodiversidad 93 (2022): e934181 4 https://doi.org/10.22201/ib.20078706e.2022.93.4181 Materials and methods hirsute-tomentose to hirsutulous with acroscopic trichomes (fig. 5E-F); calyx lobes deltoid (fig. 5E) to Pertinent specimens were reviewed from the triangular-lanceolate, 2.5-5 mm long, 1.5-3 mm wide, collections at A, BH, BM, C, CAS, CHAPA, ENCB, F, hirsute to hirsutulous with acroscopic trichomes, reddish to FCMEX, GH, HAL, IEB, INEGI, K, LL, M, MEXU, greenish-brown; corollas tubular, 2.4-3.5 cm long, almost MICH, MO, NY, OXF, P, SEL, TEX, UAMIZ, erect in the calyx, tube 2-2.8 cm long, 4-5 mm wide at the UC, US, and XAL (herbarium acronyms base, ampliate towards the limb, more or less ventricose, according to Thiers, 2020), and Galeotti collections throat slightly constricted, 0.8-1.2 cm wide at the throat, from Veracruz and information were requested from red-orange, not translucent, densely hirsute to hirsutulous B, BR, GOET, and GZU. outside, generally with reddish trichomes, throat yellow, with red elongated maculae; corolla lobes 3-5.5 mm high, Results obovate, slightly elongate, reddish-orange with reddish lines, margin erose; stamens 2.4-3 cm long, exserted, Moussonia triflora (M.Martens & Galeotti) Hanst., filaments and anthers yellowish; nectary annular, 5-lobed; Linnaea 34: 286. 1865. Gesneria triflora M.Martens & ovary sericeous with whitish trichomes, style 2.4-2.7 cm Galeotii, Bull. Acad. Brussels 9 (2): 33. 1842; Type: long; stigma stomatomorphic. Fruit an ellipsoid-rostrate Mexico. [Veracruz:] Jalapa, [Jean J.] Linden 442 (lectotype capsule (fig. 5G), 1-1.5 cm long, 3-6 mm wide, ribbed, hic designatus: BR! [https://www.botanicalcollections.be/ dry, brown, investiture hirsute-tomentose, with acroscopic specimen/BR0000013476335] (fig. 2); isolectotypes: BR! hyaline to reddish trichomes, or glabrescent; seeds elliptic, [https://www.botanicalcollections.be/specimen/BR00000 0.4-0.6 mm long, 0.1-0.2 mm wide, striate, reddish-brown. 13476342] (fig. 3); MICH!) (fig. 4), not Gesneria triflora Hook., Bot. Mag., pl. 4342. 1847 [= K. hondensis (Kunth) Taxonomic summary Hanst.)]. Distribution and ecology. Moussonia triflora has been Kohleria martensii Fritsch, Bot. Jahrb. Engler 50: 428. found in cloud forest, Pinus forests, and Pinus-Quercus 1913. forests, in humid and shady places in Oaxaca and Veracruz, Subshrubs 0.2-2 m in height, with pendulous Mexico at an elevation of 694 to 2,100 m. The abundance flexible stems, generally reddish, investiture pubescent of plants of this species at the collection sites can be scarce to pubescence of lax acroscopic trichomes, but also to abundant and plants are apparently endemic. appearing to be almost glabrous, the few lenticels ellipsoid, Phenology. Flowering and fruiting specimens have flat, almost imperceptibly reddish. Leaves opposite, been collected from December to May. elliptic-lanceolate to elliptic, sometimes elliptic-obovate, Additional specimens examined: Mexico. Oaxaca, moderately anisophyllous, asymmetric, larger blades in a Mun. Santiago Comaltepec, La Esperanza, G. Juárez pair 7.5-15 cm long, 2.7-5.8 cm wide, smaller blades et al. 3151 (MEXU); distrito Ixtlán, about km 103 on 4.5-5.5 cm long, 2-2.7 cm broad, herbaceous when fresh, Mex. 175, T. MacDougall 506 (MEXU, US); carr. 175 apex acuminate, base generally attenuated, oblique, de Valle Nacional a Esperanza, km 83-84, adelante de margin shortly serrate towards the apex, ciliate, Esperanza, junto y antes del mirador, A. Ramírez-Roa et upper surface dark green to greenish-brown, investiture al. 145 (MEXU, MO); 38 km al S de Valle Nacional, strigillose, with lax cream-colored trichomes (fig. 5A), sobre la carretera a Oaxaca, J. Rzedowski 34094 (ENCB); underside sparsely to densely strigillose to strigose on 3 km al S de la Esperanza, rumbo a Oaxaca, O. Téllez veins, greyish-green or yellowish-brown to reddish- & J.L. Villaseñor 6745 (MEXU); Dto. Ixtlán, 3.6 km al brown (fig. 5B), veins cream-colored, secondary SO de la Esperanza, carr. Tuxtepec-Oaxaca., R. Torres veins 7-9; petioles 0.7-3.5 cm long, canaliculate, & L. Cortes 11686 (MO); along half-way between Valle appressed-pilose. Inflorescences of umbellate cymes Nacional to Oaxaca, north of Cerro Pelón, H. Wiehler et of 3 flowers on most of the plant (rarely fewer) (fig. al. 9120 (MEXU, MO, SEL, US). Veracruz, Mun. Acajete, 5C); peduncles (3.5-) 5-11.4 cm long, slender, generally Rincón de Sedeño, C. Gallardo & P. Rodriguez 3399, reddish, pubescent-velutinous to shortly velutinous with 3510 (MEXU); por el camino de Plan de Sedeña hacia acroscopic trichomes; pedicels 3.3-6.5 cm, slender, Mesa de La Hierba, D.J. Sevilla et al. DJ953 (MEXU); generally reddish, shortly velutinous with acroscopic Plan de Sedeño, entrada a Plan de Sedeño, orillas de rio trichomes, trichomes generally with reddish walls; en cañada, P. Zamora & J. López 3864 (MEXU, XAL). bracts subulate to ensiform (fig. 5D) 3-10 mm long, 0.5-1 Mun. Coatepec, La Crotadura, falda E del Cofre de Perote, mm wide. Flowers with the floral tube obconic, 2.5-6 G. Castillo 22457 (MEXU, XAL), 21408, 22666, 22742 mm long, 2.5-3 mm wide, reddish to greenish-brown, (XAL); en el punto conocido como Cruz de Duela, C. A. Ramírez-Roa, L.E. Skog / Revista Mexicana de Biodiversidad 93 (2022): e934181 5 https://doi.org/10.22201/ib.20078706e.2022.93.4181 Figure 2. Gesneria triflora M.Martens & Galeotti, Lectotype. Linden 442 (BR) [https://www.botanicalcollections.be/specimen/ BR0000013476335] Gallardo et al. 3048 (MEXU); Loma Alta, V.E. Luna Mun. Comapa, road Fortín-Cardel, between Huatusco and 937 (MEXU, XAL); Loma Alta, 4 km al E de 5 Palos, Mirador, along roadside, H. Wiehler et al. 9103 (MEXU, V.E. Luna 1422 (XAL). Mun. Chimaltengo, road from US). Mun. Coscomatepec, Cerro La Magdalena, 3 km al S Jalapa to Misantla, at a small bridge crossing a stream de Tozongo, J.L. Martínez & A. Hernández 1297 (XAL). 0.5 mi NE of Chimaltengo, J.V. La Frankie 1064 (GH). Mun. Ixhuacán de los Reyes 2 km (by road) W of Ixhuacán A. Ramírez-Roa, L.E. Skog / Revista Mexicana de Biodiversidad 93 (2022): e934181 6 https://doi.org/10.22201/ib.20078706e.2022.93.4181 Figure 3. Gesneria triflora M.Martens & Galeotti, Isolectotype. Linden 442 (BR) [https://www.botanicalcollections.be/specimen/ BR0000013476342]. de los Reyes and 2.2 km (by road) E of Ayahualulco, M. Coscomatepec, El Nacimiento, F. Ventura 3467 (ENCB, Nee & C. Taylor 25968 (F, NY, XAL). Mun. Nogales, F, MICH, NY). Mun. Tlalnehuayocan, 1 km above and Nogales, E. Matuda S-110 (MICH). Mun. Orizaba, NW of San Andrés Tlalnehuayocan, M. Nee et al. 26159 Orizaba, M. Botteri 282 (GH). Mt. [monte] Orizaba, (F, NY, US, XAL). Mun. Xalapa, Paredes al N de Jalapa, East slope, E.W. Nelson 294 (GH, US). Mun. San Juan carretera a Mizantla, P. Tenorio et al. 3652 (MEXU, MO). A. Ramírez-Roa, L.E. Skog / Revista Mexicana de Biodiversidad 93 (2022): e934181 7 https://doi.org/10.22201/ib.20078706e.2022.93.4181 Figure 4. Gesneria triflora M.Martens & Galeotti, Isolectotype. Linden 442 (MICH). Mpio. Xico, gorge at Puente Acabaloya, ca. 1 km SE of and thin peduncles and pedicels, deltoid to triangular- Xico Viejo and 5 km NW of Xico, along trail between lanceolate calyx lobes, tubular corollas red-orange, the 2, remnant of ones on steep slopes of gorge, M. Nee yellow inside, ampliate toward the limb, and ellipsoid- & Taylor 26259 (F, NY). rostrate fruits. Although the name of Moussonia triflora indicates the Remarks presence of an inflorescence with 3 flowers in each cyme, Moussonia triflora is distinguished by its flexible and it is not the only species in Moussonia that might have appearing almost glabrous stems, but generally with short 3-flowered cymes, for example the recently published, pubescence, umbellate cymes of 3 flowers, evidently long M. pedunculata (Brandegee) Ram.-Roa (Ramirez-Roa & A. Ramírez-Roa, L.E. Skog / Revista Mexicana de Biodiversidad 93 (2022): e934181 8 https://doi.org/10.22201/ib.20078706e.2022.93.4181 Figure 5. Moussonia triflora (M. Martens & Galeotii) Hanst. A, Detail of the upper surface of the leaf showing the strigillose investiture; B, detail of the underside of the leaf showing the trichomes and stomates; C, umbellate cymes of tree flowers showing the slender peduncle and pedicels; D, detail of the stem showing the bracts of the inflorescence; E, detail of the calyx and the base of the flower; F, detail of the hirsutulous investiture of the floral tube; G, immature ellipsoid-rostrate capsule. MacDougall 506 (US). Skog, 2021) has 3 flowers. It is important to note that fourth flower may not develop (Ramírez-Roa, 2017). The one may see 3 flowers in any cyme of any branch of following key may help to distinguish M. triflora from species which normally have cymes of 4 flowers, as the other species with 3 and 4 flowers per cyme. A. Ramírez-Roa, L.E. Skog / Revista Mexicana de Biodiversidad 93 (2022): e934181 9 https://doi.org/10.22201/ib.20078706e.2022.93.4181 Figure 6. A, Moussonia costaricensis Klotzch ex Oerst., umbellate cymes of 4 flowers; B, Moussonia papillosa Oerst. ex Hanst., detail of the flower and calyx; C, Moussonia pedunculata (Brandegee) Ram.-Roa, detail of the flower and calyx; D, Moussonia pendula (C.V.Morton) Ram.-Roa, flower and calyx; E, Moussonia skutchii (C.V.Morton & D.N.Gibson) Wiehler, detail of the flowers; F, Mousonia triflora (M.Martens & Galeotti) Hanst., detail of the flower and calyx. A. Cortesy of Gerardo Salazar Chávez from Guerrero, Mexico. B. Liebman 9302 (US). C. Purpus 6664 (US). D. Courtesy of Claudia Rodríguez Flores, Las Joyas, Jalisco, Mexico. E. Skutch 906 (BM). F. MacDougal 506 (US). A. Ramírez-Roa, L.E. Skog / Revista Mexicana de Biodiversidad 93 (2022): e934181 10 https://doi.org/10.22201/ib.20078706e.2022.93.4181 Key to Moussonia species with normally 3 or 4 flowers in the umbellate cymose inflorescences. 1a. Umbellate cymes of 3 flowers. 2a. Pedicels 2.5-3 cm long; bracts elliptic; calyx lobes lanceolate (fig. 6C); Chiapas and Guatemala .............................................................................................................. M. pedunculata (Brandegee) Ram.-Roa 2b. Pedicels 3.3-6.5 cm long; bracts subulate to ensiform; calyx lobes deltoid to triangular-lanceolate (fig. 6F); Oaxaca and Veracruz ......................................................................................................M. triflora (M.Martens & Galeotti) Hanst. 1b. Umbellate cymes of 4 flowers. 3a. Leaf upper surface with obvious rugose surface or only towards the margin. 4a. Floral tube densely tomentose with yellowish trichomes; calyx lobes deltoid to triangular (fig. 6A), with acuminate apex; Guerrero, Chiapas, and Guatemala ...................................................................... M. costaricensis Klotzch ex Oerst. 4b. Floral tube villous with whitish trichomes; calyx lobes lanceolate-elliptic (fig. 6B), with acuminate apex; Veracruz and Puebla. ..............................................................................................................................M. papillosa Oerst. ex Hanst. 3b. Leaf upper surface with no obvious rugose surface. 5a. Calyx lobes lanceolate (fig. 6D), red outside; capitate trichomes; Jalisco ..........M. pendula (C.V.Morton) Ram.-Roa 5b. Calyx lobes shortly ovate (fig. 6E), yellowish; trichomes not capitate; Chiapas and Guatemala ..............................................................................................M. skutchii (C.V.Morton & D.N.Gibson) Wiehler Discussion Our not finding any Galeotti collections of Gesneria triflora in all the places where his collections are known, Although M. Martens and H. Galeotti (1842) published it seems appropriate to accept Fritsch’s (1913) suggestion Gesneria triflora without designating the holotype, the that Galeotti may not have collected any material of species is validly published considering the Articles the species. However, it difficult to find a reason for 7.9 and 38.1 in the International Code of Nomenclature his not finding any of this species. It is known that (Turland et al., 2018), because it has a protologue which Galeotti explored Mexico from 1835 to 1840, gathering included a Latin description, locality, phenology, as well between 7000 and 8000 specimens, many of them new to as comments on its relationship with other taxa of the science (Crépin, 1883; Diagre, 2004), including several genus. However, the nomenclatural type designation is from Oaxaca and Veracruz, states from which Gesneria pending (Article 7.2), which is presented below. triflora is now known. In addition, about 200 Galeotti To justify the selection of the Linden 442 (BR) to numbers are known to be from El Mirador, in Veracruz be the lectotype for Gesneria triflora M.Martens & (McVaugh, 1978), the locality given by Martens and Galeotti (Article 9.3), we consider the historical evidence Galeotti for Gesneria triflora: “Cette espèce se trouve avec and analyze the protologue. As it was mentioned, the la précédente dans les environs de la colonie allemande publication by Martens and Galeotti (1842), was about de Mirador, à 3,000 pieds de hauteur. Fl. Rouges. the gesneriads collected by Galeotti in Mexico and it is Mai-sept.” important to emphasize that each species, whether new or It is also known that Galeotti had been at the Mirador not, has included a Galeotti collection, except for Gesneria locality from at least September 1838 to March 1839 triflora. It might be suggested that the lack of indication of (McVaugh, 1978). This place was a colony of German the type is due to an omission of the authors. For this reason, immigrants, mainly from 1834 to 1837 (Méndez-Maín, collections by Galeotti were sought from the herbarium at 2011), in which Galeotti and other explorers had their the Meise Botanic Garden, Belgium (BR), from which the “center of operations” during their explorations in most complete set of Galeotti is known to exist (McVaugh, Mexico, thanks to the hospitality of Carl C. Sartorius 1978). But no specimens of the species have been found who hosted all of them in his property “El Mirador” there, and according to curators at B, G, NY, OXF, and W (McVaugh, 1978; Ossenbach, 2009; Schätti et al. 2018; a search was also unsuccessful (Ramírez-Roa, 2017), and Schuermans-Ceulemans et al., 2006). Interestingly, one more recently at E, K, MO, P, and US, where there are of these other explorers was Jean Jules Linden who was some duplicates of Galeotti material (Stafleu & Cowan, traveling and collecting in Mexico in the company of 1976). Recently, we asked Dr. Sofie De Smedt, Curator August Ghiesbreght and Nicolas Funck, and had likely and Project Coordinator from Meise Botanic Garden (BR) crossed paths with Galeotti in at least 2 places, i.e., at to look for any specimen of Galeotti from Veracruz that Citlaltépetl or Pico de Orizaba in August 1838 (McVaugh, was among determined or undetermined Gesneriaceae, but 1978; Diagre, 2004; Schätti et al., 2018) and/or at again no specimens of the species were found. El Mirador. A. Ramírez-Roa, L.E. Skog / Revista Mexicana de Biodiversidad 93 (2022): e934181 11 https://doi.org/10.22201/ib.20078706e.2022.93.4181 Because of Linden’s interests were directed more to can be seen in “Enumeratio synoptica ...” of 1842, the plants’ horticultural value and their possible sale, he sought authors only designated Galeotti’s material for the family, mainly orchids, but also other plants with horticultural but Galeotti probably did not want to leave out the one potential (Schuermans-Ceulemans et al., 2006; Stafleu species of Gesneria that he did not collect. But it is likely & Cowan, 1981a), Linden could have given the plant that the specimens of Linden 442 in BR were the only to Galeotti before the latter returned to Europe in 1840 specimens that Martens and Galeotti had at the time to (Crépin, 1883; McVaugh, 1978, Schuermans-Ceulemans prepare the description and to name Gesneria triflora. et al., 2006). It is interesting to note that Linden did The specimen selected as the lectotype needs to be provide specimens to Galeotti on at least 2 other compared to the information in the original publication occasions, namely: 1, Ipomoea lindenii M.Martens & (Martens & Galeotti, 1842). Recommendation 9A.1 and Galeotti [Convolvulaceae] (Bull. Acad. Roy Sci. Bruxelles particularly the 9A.2. of the International Code (Turland 12(2): 264. 1845), the type deposited at K, on which et al., 2018), indicate that the chosen the lectotype should both Galeotti and Linden labels can be seen; and 2, the adhere as closely as possible to the protologue. Some acknowledgements of Linden, Funck, and Ghiesbreght inconsistencies can be seen. in Richard and Galeotti’s (1845) monograph on orchids First, the locality on the specimen Linden 442 (BR of Mexico, in which at the beginning it says: “D’après 0000013476335) is “Jalapa”, while in the protologue the les échantillons, notes et dessins de MM. Galeotti, name of the German colony Mirador is given as locality. Linden, Funck, Ghiesbreght”. Reading further, however, it is written that Gesneria Reviewing material from the Martens’ herbarium triflora M.Martens & Galeotti is found in the same area deposited at the Meise Botanic Garden in Belgium (BR), as “G. elongata H. et B.”, the species on the page before the only institution in which an herbarium of Martens is in the same work and in which the locality “la Ville de known (Stafleu & Cowan, 1981b), we find the hoped- Xalapa du bourg de Huatusco et de la colonie allemande for specimen, a collection by Linden, J.J. Linden 442 de Mirador” is indicated. Thus, it is clear that the locality (https://www.botanicalcollections.be/specimen/BR00 of the lectotype is included in the distribution that the 00013476335), which corresponds to Gesneria triflora protologue indicates. M.Martens & Galeotti and is chosen here as a lectotype Second, the shape of the leaves in the protologue is for the following reasons: 1) the specimen has 2 labels given as “folia oppositis ovato-lanceolatis acuminatis”, (fig. 2), 1 printed and placed in the specimen’s lower right which corresponds to the shape of the largest leaves of the corner, on which it is indicated that the material is part of lectotype specimen, but the smaller leaves and in general the Martens herbarium; and 2) another label, handwritten, those observed on the isolectotypes at BR and MICH are placed in the lower left corner, where one can read the elliptic-lanceolate to elliptic, following the concepts of name of the species “Gesneria triflora Nobis”, indicating Hickey and King (2000) and Lawrence (1951). Martens that it is a new species. Also, handwritten are the locality and Galeotti (1842) probably took examined the largest “fl mexic (Jalapa)”, the collector, and collection number leaves or more mature leaves which are ovate-lanceolate, “Linden No. 442”. and not the shape of the smaller or younger ones, although Regarding the calligraphy of this handwritten label, it should be mentioned that the great majority of the Dr. Sofie De Smedt (pers. comm.) told us that it could specimens included in the additional material, also have be either Martens or Galeotti, but not Linden who wrote elliptic or elliptic-lanceolate blades. the label. One can see similar handwriting in the work of Third, the shape of the calyx lobes was described in Dorr (1986) on the collections of Jean-Baptiste Duerinck the protologue as “lobus linearibus acuminatis”, but in (1809-1857). His (Dorr’s) figure 3 shows handwritten the Linden specimens the lobes are deltoid to triangular labels, one of which is from the Martens herbarium, a with acute apices. To this point in which the protologue collection of plants that Duerinck sent Martens as a gift differs significantly from what we would now describe the for determination, and is in Martens herbarium in BR. The shape when examining the lectotype, we must consider calligraphy of the label in Dorr’s figure 3 is very similar that terminology has evolved for describing the shapes of to the label of Linden 442 (BR 0000013476335), mainly various structures in botany over almost 2 centuries. For in the letters “l”, “r”, and “s”. example, usages in describing compound leaves, some From the information seen on the Linden 442 specimen inflorescences, the peduncle, the pedicel, the pericarp, types and its residing in the Martens herbarium at BR, it is likely of pubescence and, for example, for the term “lanceolate”, that Martens and Galeotti used that Linden 442 to describe Beentje (2016) mentions the change in concept between their Gesneria triflora. It is likely that the authors of the Linnaeus and Lindley. For the term “linear”, Dr. Beentje species did not mention the Linden specimen because as (pers. comm.) commented that Lindley (1799-1865) when A. Ramírez-Roa, L.E. Skog / Revista Mexicana de Biodiversidad 93 (2022): e934181 12 https://doi.org/10.22201/ib.20078706e.2022.93.4181 describing orchids, had in mind structures that we would search for the specimen in the Berlin herbarium, B now define as narrowly elliptic, whereas Linnaeus when (E. Lücking, pers. comm.), GOET (Marc using the terms “linearis “ and” linearibus” in Species Appelhans, pers. comm.), and in GZU (A. Scharfetter, Plantarum (1753, p. 618), to describe the leaves of pers. comm.) failed. Had a specimen actually been at Antirrhinum canadense L. [Plantaginaceae], one must B, it was probably destroyed during World War II think of something a little wider than what is currently Allied bombing raids (Hiepko, 1978, 1987), as all of the considered as “linear”. Gesneriaceae specimens were presumed lost. Among the What Martens and Galeotti (1842) defined as “lobus many specimens examined during the research on linearibus acuminatis”, following Beentje (2016), means a Moussonia, no specimens have been found of the Selers’ narrow structure much longer than it is wide with parallel collection or from municipality of Chocamán. However, margins and a rounded apex. They apparently considered material has been seen from the municipality of San Juan an intermediate form between the concepts of Lindley and Coscomatepec, region of Veracruz contiguous to the Linnaeus, now coinciding with the term “deltoid” (Hickey municipality of Chocamán, F. Ventura A. 3467 & King, 2000) or “triangular” (Beentje, pers. comm., (ENCB, F, MICH, NY), which is included in the list Moreno, 1984; Radford et al., 1974) and what we use here. of additional specimens examined. On the other hand, with respect to the Linden 442 material, 2 very important aspects must be pointed out. Acknowledgements One of them is the error that appears in JStor where it is indicated that the herbarium in which the Linden 442 We want to express our appreciation to several curators material named Gesneria triflora from Mexico is deposited. who searched their collections or helped to locate the type GENT is indicated instead of BR (error confirmed by material for Gesneria triflora; Kanchi. N. Gandhi (Harvard Sofie De Smedt, pers. comm.). The other point of possible University) for his illuminating nomenclatural observations confusion is the existence of at least one other set of that guided the decision in making the typification; Sofie specimens collected by Linden with number 442. There De Smedt (BR) for her valuable information provided are 7 specimens that correspond to the type material of the about H. Galeotti and J. Linden, as well as for the species Alchemilla verticillata Field & Gard. [Rosaceae] comments about the label on the Linden specimens and collected in April 1842. in Caracas, Venezuela (holotype: the material in JStor. De Smedt also put us in contact K, isotypes: P, BM, U, GH, G, JE). Sofie De Smedt (BR) with Denis Diagre-Vanderpelen (Meise Botanical Garden, informed us “it is very common to find duplicate numbers Belgium), who is thanked for information about the lives for the Linden specimens. He made 3 trips to Central and of Galeotti and Linden; Henk Beentje (K) for his insightful South America and every time he started from collection comments about the shape of the calyx lobes on the type number 1. The first, December 1835 to March 1837, a trip material; Robert. Lücking (B), Marc Appelhans (GOET), to Brazil (from Rio de Janeiro to the forest of the Mato and Astrid Scharfetter (GZU) for searching for specimens Grosso and Minas Gerais) with Nicolas Funck collected by Seler & Seler; María del Rosario García (1816-1896) and Auguste Ghiesbreght (1810-1893). The Peña (MEXU) for facilitating the loans; Gerardo Arévalo second September 1837 to December 1840, a trip to Cuba Guzmán (IB Library) for his advice and assistance with and Mexico (from Veracruz to the tip of Yucatan) also bibliography and citations; Mauricio Mora and Maribel with Nicolas Funck and Auguste Ghiesbreght, and along Paniagua (IB) for the management and maintenance of with Henri Galeotti (1814-1858), where they were the first the Gesneriaceae database; Gerardo and Elsa Varela for to climb Mt. Orizaba. The third trip was October 1841 to their technical support during visits to foreign herbaria; to December 1844 a trip to Venezuela and Colombia with Julio César Montero (IB) for the graphic material; to the his step-brother Louis-Joseph Schlim (1819-1863). After a staff of national and foreign herbaria for their support in brief passage in Jamaica in March 1844, they returned to consulting and loaning specimens; and to the anonymous Brussels in December’’. Hence with the knowledge of reviewers for their valuable comments that greatly enriched Linden’s numbering system, and citation of names, this work. collections localities, and the herbarium location of the types, there should be no confusion about the specimens References with the same collection numbers. Finally, regarding the specimen C. Seler & E.G. Seler Beentje, H. (2016). The Kew plant glossary. Kew, UK: Royal 5182, cited by Fritsch (1913) and again by Morton (1967) for Botanic Gardens. Kohleria martensii, Ramírez-Roa (2017) indicated that the Clark J. L., Skog, L. E., Boggan, J. K., & Ginzbarg, S. (2020). specimen was not located in the follow herbaria (BM, CAS, F, G, GH, JE, K, MEXU, MO, NY, P, U, or US), and a recent Index to names of New World members of the Gesneriaceae A. Ramírez-Roa, L.E. Skog / Revista Mexicana de Biodiversidad 93 (2022): e934181 13 https://doi.org/10.22201/ib.20078706e.2022.93.4181 (subfamilies Sanangoideae and Gesnerioideae). 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