INTRODUCTION
⌅Celastraceae R.Br. contains approximately 98 genera and 1264 species distributed worldwide. They are found in the tropics and subtropics, with some rare representatives in temperate regions (Loesener 1942LoesenerT.1942. Celastraceae. InEnglerH. & MattfeldJ. (eds.), Die Natürlichen Pflanzenfamilien Vol. 20b (A): 87–197. Preprint, Berlin, Duncker and Humblot.; Simmons & al. 2001aSimmonsM.P., ClevingerC.C., SavolainenV., ArcherR.H., MathewsS. & DoyleJ.J.2001a. Phylogeny of the Celastraceae inferred from 26S nuclear ribosomal DNA, phytochrome B, rbcL, atpB, and morphology. Molecular Phylogenetics and Evolution19: 353–366., 2001bSimmonsM.P., SavolainenV., ClevingerC.C., ArcherR.H. & DavisJ.I.2001b. Phylogeny of the Celastraceae inferred from phytochrome B gene sequence and morphology. American Journal of Botany88: 313–325.; Simmons 2004SimmonsM.P.2004. Celastraceae. InK.Kubitzki (eds.), The families and genera of vascular plants 6: 29–64. Springer Verlag, Berlin.). The delimitation of genus Maytenus Molina has been long debated particularly regarding the merger/segregation of species from Gymnosporia (Wight & Arn.) Hook.f. (Loesener 1942LoesenerT.1942. Celastraceae. InEnglerH. & MattfeldJ. (eds.), Die Natürlichen Pflanzenfamilien Vol. 20b (A): 87–197. Preprint, Berlin, Duncker and Humblot.; Brenan 1953BrenanJ.P.M.1953. Plants collected by the Vernay Nyasaland expedition of 1946. Memoirs of the New York Botanical Garden8: 191–256.; Exell 1953ExellA.W.1953. Tropical African plants: XXIII: Celastraceae. Kew Bulletin23: 103–104.; Hou 1955HouD.1955. A revision of the genus Celastrus. Annals of the Missouri Botanical Garden42: 215–302., 1962HouD.1962. Celastraceae I. Flora Malesiana, ser. 1, 6: 227–291.; Demissew 1985Demissew, S.. 1985. The genus Maytenus (Celastraceae) in NE tropical Africa and tropical Arabia. Acta Universitatis Upsaliensis. Symbolae Botanicae Upsaliensis25: 1–98.; Jordaan & van Wyk 1999JordaanM. & van WykA.E.1999. Systematic studies in the subfamily Celastroideae (Celastraceae) in southern Africa: Reinstatement of the genus Gymnosporia. South African Journal of Botany65: 177–181., 2003JordaanM. & van WykA.E.2003. Reinstatement of Gymnosporia (Celastraceae): Implications for the Flora Malesiana region. Telopea10: 155–167.; McKenna & al. 2011McKennaM.J., SimmonsM.P., BaconC.D. & LombardiJ.A.2011. Delimitation of the segregation genera of Maytenus. Systematic Botany36: 922–933.). Molecular data support that Gymnosporia (Wight & Arn.) Hook.f. is a natural group distinct from Maytenus (Simmons & al. 2001aSimmonsM.P., ClevingerC.C., SavolainenV., ArcherR.H., MathewsS. & DoyleJ.J.2001a. Phylogeny of the Celastraceae inferred from 26S nuclear ribosomal DNA, phytochrome B, rbcL, atpB, and morphology. Molecular Phylogenetics and Evolution19: 353–366., 2001bSimmonsM.P., SavolainenV., ClevingerC.C., ArcherR.H. & DavisJ.I.2001b. Phylogeny of the Celastraceae inferred from phytochrome B gene sequence and morphology. American Journal of Botany88: 313–325., 2008SimmonsM.P., CappaJ.J., ArcherR.H., FordA.J., EichstedtD. & ClevingerC.C.2008. Phylogeny of the Celastreae (Celastraceae) and the relationships of Catha edulis (qat) inferred from morphological characters and nuclear and plastid genes. Molecular Phylogenetics and Evolution48: 745–757.) and it has recently been reinstated to include all its thorny members, previously placed under Maytenus Molina. Gymnosporia is an Old World genus, comprising about 118 species and infraspecific taxa of dioecious trees, shrubs or dwarf shrubs, occurring in Africa, Madagascar and adjacent islands, southern Spain, the near Middle East, Afghanistan, Pakistan, India, Sri Lanka, Thailand, Vietnam, China, Taiwan, Ryukyu Islands (Japan), Malesia, Australia (Queensland), and the Polynesian Islands (Jordaan & van Wyk 1999JordaanM. & van WykA.E.1999. Systematic studies in the subfamily Celastroideae (Celastraceae) in southern Africa: Reinstatement of the genus Gymnosporia. South African Journal of Botany65: 177–181., 2003JordaanM. & van WykA.E.2003. Reinstatement of Gymnosporia (Celastraceae): Implications for the Flora Malesiana region. Telopea10: 155–167., 2006JordaanM. & van WykA.E.2006. Sectional classification of Gymnosporia (Celastraceae), with notes on the nomenclatural and taxonomic history of the genus. Taxon55: 515–525.; Simmons 2004SimmonsM.P.2004. Celastraceae. InK.Kubitzki (eds.), The families and genera of vascular plants 6: 29–64. Springer Verlag, Berlin.).
Gymnosporia cassinoides (L’Hér.) Masf. (based on Celastrus cassinoides L’Hér.), popularly known as “peralillo”, “peralito”, “acebuche” or “árbol negro” (the leaves resemble those of a pear tree, genus Pyrus, “peral” in Spanish) (Kunkel 1971KunkelG.1971. Nombres vernáculos de la flora Gran Canaria (incluyendo especies asilvestradas). Cuadernos de Botánica Canaria, Supl.2: 1–64., 1986KunkelG.1986. Diccionario Botánico Canario. Manual etimológico. Edirca S.L.Las Palmas de Gran Canaria.; Pérez de Paz & Hernández Padrón 1999Pérez de PazP.L. & Hernández PadrónC.E.1999. Plantas medicinales o útiles en la flora canaria. Aplicaciones populares. Francisco Lemus, Editor, La Laguna.), is a hermaphrodite and small tree, with leathery leaves, alternate, oval, irregularly toothed edges (serrated), dichasiale inflorescences, well-developed petals and obconic fruits, trigonous and dehiscent (capsules), pale greenish to brown, and seeds having a white aril at the base. The specific epithet “cassinoides” alludes to the resemblance to a species of Ilex (I. cassinoides Link) from India, whose vernacular name is “cassine” (Kunkel 1986KunkelG.1986. Diccionario Botánico Canario. Manual etimológico. Edirca S.L.Las Palmas de Gran Canaria.).
Gymnosporia cassinoides is a Canarian endemic plant, recorded from all the islands of the archipelago except Lanzarote (i.e., El Hierro, La Palma, La Gomera, Gran Canaria, Tenerife, and Fuerteventura) (Masferrer 1881MasferrerH.1881. Recuerdos botánicos de Tenerife, o sea, datos para el estudio de la flora canaria. Anales de la Real Sociedad Española de Historia Natural10: 139–230.; Pitard & Proust 1908PitardJ. & ProustL.1908. Les Iles Canaries. Flore de l’archipel. P. Klincksieck, Paris.; Ceballos & Ortuño 1951CeballosL. & OrtuñoF.1951. Estudio sobre la vegetación y la flora forestal de las Canarias occidentales. Madrid.; Lems 1960LemsK.1960. Floristic botany of the Canary Islands. Sarracenia5: 1–94.; Sunding 1971SundingP.1971. A new name to the Canarian Peralillo tree. Cuadernos de Botánica Canaria13: 8–10.; Reyes-Betancort & Guerra 2010Reyes-BetancortJ.A. & GuerraA.S.2010.Gymnosporia cryptopetala Reyes-Bet. & A. Santos (Celastraceae), a new species from the Canary Islands. Candollea65: 189–196.). It is a medicinal plant (Pérez de Paz & Hernández Padrón 1999Pérez de PazP.L. & Hernández PadrónC.E.1999. Plantas medicinales o útiles en la flora canaria. Aplicaciones populares. Francisco Lemus, Editor, La Laguna.) due to its antibacterial effect, among others (e.g., dermatics, cystostatic, emollient, antirheumatic) (González & al. 1989GonzálezA.C., JiménezI.A., RaveloA.G.Luis, J.G. & Bazzocchi, I.L.1989. β-Agarofurane sesquiterpene esters from Maytenus canariensis. Phtochemistry. 28: 173–175., 1992GonzálezA.G., JiménezI.A. & RaveloA.G.1992. Triterpenes from Maytenus canariensis and synthesis of a derivative from Betulin. Phtochemistry31: 2069–2072., 1996GonzálezA.G., AlvarengaN.L., RaveloA.G., JimenezI.A., BazzocchiI.L., CanelaN.J. & MoujirL.M.1996. Antibiotic phenol nortriterpenes from Maytenus canariensis. Phtochemistry43: 129–132.; Huang & al. 2021HuangY.Y., ChenL., MaG.X., XuX.D., JiaX.G.DengF.S.,LiX.J. & Yuan, J.Q.2021. A review on phytochemicals of the genus Maytenus and their bioactive studies. Molecules26: 4563.). In the Eastern most islands (Lanzarote and Fuerteventura) (Kunkel 1973KunkelG.1973. Nuevas adiciones para la flora de las Islas Orientales. Cuadernos de Botánica Canaria18–19: 25–31., 1974KunkelG.1974. Resultados de dos viajes a Fuerteventura y Lanzarote. Cuadernos de Botánica Canaria20: 17–23., 1975KunkelG.1975. Inventario de los Recursos Naturales Renovables de la Provincia de Las Palmas (Islas Canarias, España). Cabildo Insular de Gran Canaria y Excma. Mancomunidad Interinsular de las Palmas., 1976KunkelG.1976. Enumeración de las plantas vasculares del Parque Nacional de Timanfaya (Lanzarote) con notas adicionales. Cuadernos de Botánica Canaria26–27: 41–58., 1977KunkelG.1977. Endemismos Canarios. Inventario de las Plantas Vasculares Endémicas en la Provincia de Las Palmas. Ministerio de Agricultura.; Santos & Fernández 1984SantosA. & FernándezM.1984. Notas florísticas de las islas de Lanzarote y Fuerteventura (I. Canarias). Anales del Jardín Botánico de Madrid41: 167–174.) a second species of the genus has been described, i.e., G. cryptopetala Reyes-Bet. & A.Santos belonging to sect. Tenuispinae, which shows a close relationship with geographically distant species such as G. tenuispina (Sond.) Szyszyl. and G. emarginata (Willd.) Thwaites (Reyes-Betancort & Guerra 2010Reyes-BetancortJ.A. & GuerraA.S.2010.Gymnosporia cryptopetala Reyes-Bet. & A. Santos (Celastraceae), a new species from the Canary Islands. Candollea65: 189–196.).
Gymnosporia cassinoides was traditionally known as Maytenus canariensis (Loes.) G.Kunkel & Sunding (Sunding 1971SundingP.1971. A new name to the Canarian Peralillo tree. Cuadernos de Botánica Canaria13: 8–10.), a new combination at new rank for M. dryandri var. canariensis Loes.; the name “Maytenus cassinoides” (based on the L’Héritier’s name Celastrus cassinoides) could not be used because to the older homonym M. cassinoides (Poir.) Urb. used to name a plant of the West Indies. The name M. dryandri var. canariensis Loes. is a replacemente name for Celastrus cassinoides; Loesener (1942LoesenerT.1942. Celastraceae. InEnglerH. & MattfeldJ. (eds.), Die Natürlichen Pflanzenfamilien Vol. 20b (A): 87–197. Preprint, Berlin, Duncker and Humblot.) published “M. dryandri (Lowe) Loes. (Catha dryandri Lowe, Fl. Madeira [1868] 109), typische Form auf Madeira, var. canariensis Loes. auf den Canaren (Celastrus cassinoides L’Hér. 1786; Catha cassinoides Webb et Berth. (Webb & Berthelot 1842WebbP.B. & BerthelotS.1842. Histoire Naturelle des Îles Canaries. Ouvrage publié sous les auspices De M. le Ministre de lInstruction Publique. Tome troisième. Deuxième partie. Phytographia canariensis. Sectio II. Béthune, éditeur, rue de Vauirard.); G. cassinoides Masf.; non M. cassinoides Urban 1904)”, and therefore M. dryandri var. canariensis can be treated as a new name (nomen novum) published as an explicit substitute (avowed substitute) for a legitimate previously published name (Celastrus cassinoides), which is its replaced synonym.
From the nomanenclatural point of view, the name Gymnosporia cassinoides has not yet been typified. For this reason, the purpose of this paper is to propose its typification to ensure nomenclatural stability. This contribution is a further step in our work on the nomenclature of the genus Gymnosporia (e.g., Ferrer-Gallego & Laguna 2020Ferrer-GallegoP.P. & LagunaE.2020. Typification of Celastrus senegalensis and C. europaeus (Celastraceae, Celastroideae). Acta Botanica Malacitana45: 197–202.).
MATERIALS AND METHODS
⌅The protologue of Celastrus cassinoides was analyzed to identify original material pertinent to the typification. The taxonomic identity of the proposed type was verified against the traditional concept and the current usage of the name. In the following account herbarium acronyms follow Thiers (2024Thiers, B.2024[continuously updated]. Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium, New York. Available from: http://sweetgum.nybg.org/science/ih/ [accessed: 15 Sep 2024] [continuously updated]). The typification of the name is in accordance with the rules and recommendations of the Shenzhen Code (Turland & al. 2018TurlandN.J., WiersemaJ.H., BarrieF.R., GreuterW., HawksworthD.L., HerendeenP.S., KnappS., KusberW.-H., LiD.-Z., MarholdK., MayT.W., McNeillJ., MonroA.M., PradoJ., PriceM.J. & SmithG.F. (eds.) 2018. International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum Vegetabile 159. Glashütten, Koeltz Botanical Books.).
RESULTS AND DISCUSSION
⌅L’Héritier’s protologue (1789: 4) of Celastrus cassinoides consisted of a short diagnosis: “C. [Celastrus] inermis, foliis ovatis utrinque acutis laxe dentatis perennantibus, floribus axillaribus”, followed by the provenance “Habitat in Maderâ, Nivariâ. Masson.” In the protologue was cited an illustration, as “2. CELASTRUS cassinoides. Tab. 10.” (see L’Héritier, 1790 [May]: Tab. 10) that can be considered original material for typification purposes.
The illustration of this species mentioned by the author in the protologue was published by L’Héritier in May 1790 (Tab. 10). It is an excellent drawing of a stem with leaves, flowers and fruits, with several details of the flowers, fruits, and seeds (Fig. 1). L’Héritier published his work Sertum anglicum in various fascicles in different years. According to Stafleu & Cowan (1981StafleuF.A. & CowanR.S.1981Taxonomic literature. Ed. 2, 3. Bohn, Scheltema & Holkema, Utrecht.: 3, no. 4492) in 4 fascicles between 1789 and 1792 (Fasc. 1, p. [i–iv], 1–36, tab. 1–2 (princ. I.1789L’HéritierC.-L.1789–1792. Sertum anglicum seu plantae rariores quae in hortis juxta Londinum, imprimis in horto regio Kewensi excoluntur, ab anno 1786 ad annum 1787 observatae. P.F. Didot, Paris, 36 pp. + 34 tab. [4 fasc., between 1789 y 1792 [cf. F.A. Stafleu & R.S. Cowan, “Taxonomic literature” ed. 2, n.º 4.492]; the date on the cover is 1788: Fasc. 1, pp. [i–iv], 1–36, tab. 1–2 (princ. I.1789) Fasc. 2, tab. 3–12 (V.1790) Fasc. 3, tab. 13–24, 15bis (IV.1792) Fasc. 4, tab. 25–34 (fin. 1792)].), Fasc. 2, tab. 3–12 (V.1790) Fasc. 3, tab. 13–24, 15bis (IV.1792) Fasc. 4, tab. 25–34 (fin. 1792)). According to ICN Art. 9.4 (Shenzhen Code, Turland & al. 2018TurlandN.J., WiersemaJ.H., BarrieF.R., GreuterW., HawksworthD.L., HerendeenP.S., KnappS., KusberW.-H., LiD.-Z., MarholdK., MayT.W., McNeillJ., MonroA.M., PradoJ., PriceM.J. & SmithG.F. (eds.) 2018. International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum Vegetabile 159. Glashütten, Koeltz Botanical Books.), original material comprises: (a) those specimens and illustrations (both unpublished and published prior to publication of the protologue) that the author associated with the taxon, and that were available to the author prior to, or at the time of, preparation of the description, diagnosis, or illustration with analysis validating the name; (b) any illustration published as part of the protologue.
The description of this plant was published in 1789 (fascicle 1) and the illustration “Tab. 10” of Celastrus cassinoides in 1790 (fascicle 2). However, in the protologue was mentioned “2. CELASTRUS cassinoides. Tab. 10.” and therefore L’Héritier associated this illustration (unpublished at the time that was published the protologue) with the taxon and it was therefore available to the author at the time of preparation of the description of the species. Therefore, this illustration is original material of C. cassinoides and elegible for lectotypification. However, according to Lowe (1862LoweR.T.1862. A Manual Flora of Madeira. Part II, Subclass. II. Calyciflorae: 107–109. John van Voorst, London.), this illustration represents heterogeneous material.
On the other hand, L’Héritier (1789) mentioned in the protologue at least two concrete gatherings collected by Francis Masson, from Madeira, and also from “Nivariâ”. Nivaria or Ninguaria were the names by which the Romans knew the island of Tenerife, in reference to the snow perched on the Teide Volcano.
Francis Masson (1741–1805) was a Scottish botanist and gardener, in the 1760s he went to work at Kew Gardens. As an under-gardener he was the first official plant collector from the Royal Botanic Gardens of Kew, appointed by Sir Joseph Banks. Between 1776 and 1779 he collected extensively on the Macaronesian archipelagos of Madeira, the Azores and the Canaries (Francisco-Ortega & al. 2008Francisco-OrtegaJ., Santos-GuerraA., CarineM.A. & JarcisC.E.2008. Plant hunting in Macaronesia by Francis Masson: the plants sent to Linnaeus and Linnaeus filius. Botanical Journal of the Linnean Society157: 393–428.). Whereas Masson’s living plants were destined to Kew, the herbarium specimens were mainly sent to Sir Joseph Banks and are now preserved at BM. Duplicates of Masson’s collections are in several herbaria, e.g., AWH, BR, CGE, DBN, HAL, LD, LINN, MO, OXF, P, PH, UPS (incl. Thunberg herbarium) (e.g., Stafleu & Cowan 1981StafleuF.A. & CowanR.S.1981Taxonomic literature. Ed. 2, 3. Bohn, Scheltema & Holkema, Utrecht.; Rodrigues de Moraes 2013Rodrigues de MoraesP.L.2013. The collection of Lauraceae in the Herbarium of Henri Van Heurck (AWH). Plant Ecology and Evolution146: 360–383.). The Royal Botanic Gardens, Kew, had no herbarium in the 18th century and as noted by Stafleu & Cowan (1976StafleuF.A. & CowanR.S.1976. Taxonomic literature. Ed. 2, 1. Bohn, Scheltema & Holkema, Utrecht.), I have not been able to locate any specimen of Celastrus cassinoides from the Canary Island or Madeira collected by Masson in the K herbarium (David Goyder, pers. comm.).
According to Francisco-Ortega & al. (2008Francisco-OrtegaJ., Santos-GuerraA., CarineM.A. & JarcisC.E.2008. Plant hunting in Macaronesia by Francis Masson: the plants sent to Linnaeus and Linnaeus filius. Botanical Journal of the Linnean Society157: 393–428.), there is only record of Masson’s stay in Tenerife in the first half of 1778 (from February to August), with shipments of plant material (with accompanying letter) on 19 March 1778 and 4 May of the same year, in both cases addressed to Banks. And according to the list consulted by Francisco-Ortega & al. (2008Francisco-OrtegaJ., Santos-GuerraA., CarineM.A. & JarcisC.E.2008. Plant hunting in Macaronesia by Francis Masson: the plants sent to Linnaeus and Linnaeus filius. Botanical Journal of the Linnean Society157: 393–428.), he sends Canarian material (from Tenerife) of “Celastrus rotundifolia” (which they assimilate to Maytenus canariensis, the only species of this genus growing in Tenerife) on 23 November 1778 (but sent from Madeira). The Spanish material (BM000829286 and BM000829287) was collected at “Barranca Honda” [Barranco Hondo], a gorge located in the ‘Acentejo’ region, in northern Tenerife, where the species is locally common (Francisco-Ortega & al. 2008Francisco-OrtegaJ., Santos-GuerraA., CarineM.A. & JarcisC.E.2008. Plant hunting in Macaronesia by Francis Masson: the plants sent to Linnaeus and Linnaeus filius. Botanical Journal of the Linnean Society157: 393–428.).
The British Museum (BM) holds three relevant specimens for the name Celastrus cassinoides collected by Masson (BM000829286, BM000829287, and BM000838914). The sheet BM000829286 bears a specimen with leaves and four fruits, and a handwritten label: “Teneriffe. Fr. Masson” (Fig. 2). The sheet BM000829287 bears a specimen with leaves, fruits, and seeds, and is annotated (on the back of the sheet) “Teneriffe. Fr. Masson” (Fig. 3). The geographical locality “Teneriffe” and the author “Fr. Masson” agree with the locality and author given in the protologue “Nivariâ” and “Masson”.
On the other hand, the sheet BM000838914 bears a specimen with leaves and fruits, and a handwritten label: “Madera. Fr. Masson” (Fig. 4). This specimen is part of the original material, but is not suitable for designation as the lectotype, as it currently belongs to another species, and therefore a lectotypification using this specimen would be nomenclaturally disruptive This specimen was selected as the lectotype of Maytenus umbellata (R.Br.) Mabb. (Mabberley 1981MabberleyD.J.1981. The name of the madeiran Maytenus (Celastraceae). Taxon30: 485–486.: 486).
These three specimens can be treated as syntypes mentioned by L’Héritier in the protologue, because the information match between the original annotations on the labels of the herbarium sheets and the protologue (“Habitat in Maderâ, Nivariâ. Masson”), (i.e., “Nivariâ” [Tenerife]. Fr. Masson” and the specimens BM000829286 and BM000829287, and “Madera. Fr. Masson” and the specimen BM000838914). I have not been able to locate other Celastrus cassinoides material from Masson’s collection in other herbaria (e.g., AWH, BR, CGE, DBN, HAL, LD, LINN, MO, OXF, P, PH, UPS).
In conclusion, among the original elements, the illustration cited in the protologue and the specimens at BM, I designate as the lectotype of the name Celastrus cassinoides the specimen BM000829287 (Fig. 3). This specimen shows the diagnostic characters of Gymnosporia cassinoides: with leathery leaves, alternate, oval, irregularly toothed edges (serrated), bark grayish brown, young twigs not grooved; unarmed (the plant may rarely have axillary thorns); inflorescence equal to or shorter than leaves (3–7-flowered); capsule obconical-trigona.
Gymnosporia cassinoides (L’Hér.) Masf., Anales Soc. Esp. Hist. Nat. 10: 176. 1881. ≡ Celastrus cassinoides L’Hér. in Sert. Angl.: 4. 1789 [basionym] ≡ Catha cassinoides (L’Hér.) Webb & Berthel., Hist. Nat. Iles Canaries 3(pt. 2, sect. 2, livr. 67): 142. 1842. ≡ Maytenus dryandri var. canariensis Loes. in Engler, Nat. Pflanzenfar., ed. 2, Aufl. 20b: 140. 1942 [nom. nov.] ≡ Maytenus dryandri var. canariensis (Loes.) G. Kunkel & Sunding in Kunkel, Cuad. Bot. Canar. 13: 9. 1971; in Kunkel, Cuad. Bot. Canar., Supl. 2: 62. 1971. Type (lectotype designated here):—[Spain] Canary Islands “Nivariâ” [Tenerife], [Barranco Hondo, 1778, see Francisco-Ortega & al. 2008], F. Masson s.n. (BM000829287 [photo!]; image of the lectotype, see Fig. 3). Isolectotype: BM000829286 (Fig. 2).