www.unal.edu.co/icn/publicaciones/caldasia.htm CaldasiaWanntorp 28(2):221-225. & Klackenberg 2006 MORAE (), A NEW FROM COLOMBIA Gunnera morae (Gunneraceae), una nueva especie para Colombia

LIVIA WANNTORP Botaniska Institutionen, Stockholms universitet, SE-106 91 Stockholm, Sweden. [email protected].

JENS KLACKENBERG Naturhistoriska riksmuseet, Sektionen för fanerogambotanik, SE-104 05, Stockholm, Sweden. [email protected] ABSTRACT A Colombian species of Gunnera, G. morae L. Wanntorp & Klackenberg, previously mistakenly identified as Linden ex André, is presented and described. Molecular phylogenetic studies as well as morphological evidence have shown this to be different from the Brazilian true G. manicata. Key words. Flora of , flora of Colombia, Gunneraceae, Gunnera manicata, Gunnera morae.

RESUMEN Se describe una nueva especie de Gunnera de Colombia, G. morae L. Wanntorp & Klackenberg, que había sido previamente identificada de forma errónea como G. manicata. Nuevos estudios filogenéticos basados en caracteres moleculares y morfológicos demuestran que esta planta es diferente de la verdadera G. manicata de Brasil. Palabras clave. Flora de Brasil, flora de Colombia, Gunneraceae, Gunnera manicata, Gunnera morae. INTRODUCTION herteri Osten (subgenus Ostenigunnera Mattfeld). All species of Panke are similar in Gunnera L. is a subtropical genus of 30- morphology, being gigantic herbs with stout 40 species of which most occur in South and erect rhizomes surrounded by palmately and Central America. Five to six species lobed that sometimes reach up to inhabit New Zealand, while Southeast Asia, several meters in diameter. The inflorescences Tasmania and have a single endemic of Panke are compound spikes with many species each. Two species grow on Hawaii small, dimerous and wind pollinated flowers and one in Mexico (Bader 1961). The genus and the fruits are drupes, which are fleshy is currently divided into six monophyletic and brightly colored in some species while subgenera based on the geographical greenish and dry in others. distribution of the species as well as on morphology (Schindler 1905, Mattfeld Mora-Osejo (1984) presented a revision of 1933). Among these subgenera, Panke Gunneraceae in Flora of Colombia in which Schindler includes two Hawaiian species as 23 species were accepted. One of these was well as all South American species except for wrongly identified as Gunnera manicata G. magellanica Lam. and G. lobata Hooker Linden ex André, the most commonly f. (subgenus Misandra Schindler) and G. cultivated species of Gunnera. This species 221 Gunnera morae, a new species from Colombia was introduced into Europe from Southern Brazilian and the Colombian taxa belong to Brazil by the botanist and horticulturist two different clades, supporting the fact that Linden (1867), who, however, published the these actually constitute two different new introduction without a proper description species (Wanntorp et al. 2002a). In contrast, of the plant (Delchevalerie 1867). Some cultivated material and material from wild years later, André (1873) provided a valid Brazilian G. manicata had identical gene description for G. manicata based on the sequences, and therefore clustered together living material that had been introduced by in the tree. This clearly supports a Brazilian Linden and since propagated in La Muette origin for the cultivated G. manicata botanical garden in Paris. He wrongly stated, (Wanntorp et al. 2002b). A reexamination however, “Nouvelle Grenade” (present of the herbarium specimens that were used day Colombia) as place of origin for this for defining the Colombian ”Gunnera plant, although Linden (1867) wrote that manicata” (Mora-Osejo 1984) showed that his G. manicata was collected in Campos this taxon, although similar in habitus and in de Lages in Southern Brazil. Based on this macro-morphology to G. manicata, differs geographical mistake, in several later works, in some morphological characters. All e.g. in Flora of Colombia (Mora-Osejo 1984), species of Panke have conspicuous bracts Colombia has been wrongly reported as the (lepidophylls) that cover the rhizomes and place of origin for G. manicata. Mora-Osejo differ in shape, colour and size within the (1984) noted that the Colombian specimens genus. The bracts of G. manicata are up to examined by him did not really correspond in c. 12 cm long, whitish green, very thin and morphology to André’s description. This was except for the veins transparent when dry, explained by a theory that Linden’s original deeply laciniate with long lobes also from plant material might have been hybrids with near the base. These primary lobes (laciniae) G. brephogea Linden & André. Confident are often as long as 1/3 to 1/2 of the bract that his Colombian taxon was identical to G. and are in their turn laciniately divided into manicata, Mora-Osejo (1984) designated a secondary lobes with fimbriate margins. specimen of his Colombian taxon as neotype Only the adaxial side of the bract is hairy. for G. manicata Linden ex André. However, In contrast, the Colombian taxon has reddish due to both the geography and phylogenetic purple bracts, hairy on both sides, up to c. 25 studies (see below), this neotypification is cm long that are thicker and not transparent incorrect. Wanntorp et al. (2002b) designated when dry, at lower part without long primary a new type for G. manicata based on old lobes but with laciniae (up to 1/4 as long as cultivated material. See also Gandhi (2002). the bract, usually much less) at their margins Recently extensive phylogenetic studies, (Fig. 1F). Only larger bracts, 15 cm or more, mainly based on molecular characters, are sometimes divided into primary and were conducted on Gunnera (Wanntorp secondary laciniae. However, the margins are et al. 2002a, b). In the context of these not fimbriate or with few scattered fimbrillae studies, the ribosomal (DNA) ITS gene only. This gives the lepidophylls of the region was sequenced for the Colombian Colombian ”Gunnera manicata”, a much “G. manicata”, and for two accessions of coarser impression than the more plumy ones Brazilian G. manicata, one collected in the of G. manicata. In addition, the Colombian wild in Brazil and the other of cultivated taxon has perfect flowers (Fig. 1I), while in source (Wanntorp et al. 2002b). According the Brazilian G. manicata perfect flowers are to the retrieved phylogenetic trees, the mixed with unisexual ones.

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This morphological evidence supports the distinct teeth of different sizes on young leaves, molecular results that the Colombian plant older leaves with smaller more rounded teeth. belongs to another species and should be Inflorescences sessile, 50-60 cm long after described as a new species, G. morae. The anthesis, pilose to sparsely pilose, dark red; description here presented is mainly based on rhachis 0.5-1.5 cm wide, deeply sulcate; lateral that of Mora-Osejo (1984). A description of branches 8-18 cm long, 1-1.5 cm wide, sulcate; the petals is added. The petals easily fall off bracts when present linear to thread-like, rarely and were not present on the material studied slightly broadened, 0.4-2 cm long, 0.1-1.5 mm by Mora-Osejo (1984), but can be studied on wide, with entire margins or a few teeth. the new material here chosen as type (Fig. perfect; sepals at anthesis triangular to oblong 1I-J). This new material is also the collection with irregular membranaceous often obtuse that was analyzed in the molecular study apex, 1.3-1.8 mm long, dark red, after anthesis (Wanntorp et al. 2002a, 2002b). The petals in persistent, triangular, usually acute, 0.6-0.8 G. morae are cucullate (Fig. 1I-J), similar to mm long; petals hooded, 2.0-2.3 x 1.3-1.6 mm, petals in other species of subgenus Panke. soon falling off; stamens 2; anthers elliptical to broadly elliptical, 1.3-1.8 mm long, 1.2-1.3 mm Gunnera morae L. Wanntorp & Klackenberg, wide, not or shortly apiculate; filaments very sp. nov. short. Ovaries (mature) cylindrical, 0.7-1.0 mm wide at base. Fruits subglobose, slightly TYPE: COLOMBIA. Tolima: Padua, km narrowed at top, 1.5-1.8 x 1.5-1.8 mm long, on 37 La Enea Vía Padua, 3370 m alt., 25 short pedicels. Mar 2001, G. Morales 1570 (holotype, S!; isotype, COL). Figure 1. Etymology. We have, with his agreement, named this plant after the late Colombian Diagnosis. Species haec Gunnerae manicatae botanist Mora-Osejo, who discovered this similis statura magna et rhizomatis foliis new species. 5-lobatis cordatisque, sed lepidophyllis purpureis usque 25 cm longis cum marginibus Distribution and ecology. Gunnera morae is laciniatis sed non-fimbriatis et floribus totis known from Colombia along the Cordillera perfectis differt. Central, where it grows at 2660-3525 m altitude. It has been collected in the provinces Description. Herbs of large size, with rhizomes of Antioquia, Quindío, Risaralda, Tolima and c. 4 cm wide. Bracts (lepidophylls) 10-25 Nariño. Specimens in flower or in young fruit cm long, c. 0.5-2 cm wide at base, laciniate, seen from March, July and October. larger bracts also lobed, finely hairy on both sides, shining, reddish purple, gradually Paratypes. COLOMBIA. Nariño: Túquerres narrowing into a filiform apex; margins with y Piedrancha, vereda de Chambú, Mora 4683 few fimbrillae or usually not fimbriate. (COL); ibidem, Mora 4684 (COL); Quindío: petioles of young plants 20-40 cm long, c. 1 La Línea, carretera Cajamarco, García-Barriga cm thick at base, sulcate, covered by scattered 21251 (COL); Risaralda: Cordillera Central, erect spines; blades orbicular, 5-lobate, cordate Municipio de Santa Rosa, entre la hacienda La at base; upper side areolate, scabrous, spotted Sierra y Termales de Santa Rosa, quebrada la dark red, lower side with dark red hairy veins; Sierra, Jaramillo et al. 5988 (COL); Tolima: lobes acute or truncate at apex, sometimes Carretera El Fresno a Manizales, arriba acuminate; largest sinuses rounded, cleft c. 1/3 de Padua, Uribe 3318 (COL); Antioquia: of the leaf radius, second largest sinuses cleft Municipio de Sonsón, Páramo de Sonsón, c. 1/4 of the leaf radius; margins dentate with 1977, Lozano et al. 2974 (COL). 223 Gunnera morae, a new species from Colombia

Figure 1. Gunnera morae. A, habit; B, leaf; C-D, part of leaf (C, abaxial side), (D, adaxial side); E, leaf ; F, bract (lepidophyll); G, bract of the inflorescence; H, part of inflorescence with ca 4 flowers removed; I, flower; J; petal. (A-J Morales 1570, A-E from living plants in greenhouse). Drawn by Emma Hultén, Stockholm. 224 Wanntorp & Klackenberg

ACKNOWLEDGMENTS MATTFELD, J. 1933. Weiteres zur Kenntnis der Gunnera Herteri Osten. Ostenia: 102-118. The first author of the paper would like to MORA-OSEJO, L. E. 1984. . thank H-E. Wanntorp and M. Källersjö for Pages 1-173 in: Pinto P. & P. M. Ruiz encouragement and advice. Thanks to Emma (eds.). Flora de Colombia, 3. Instituto de Hultén for the illustrations, to E. Forero, Ciencias Naturales, Bogotá, Colombia. F. González and the National Colombian SCHINDLER, A. K. 1905. Halorrhagaceae. Herbarium (COL) for loan of herbarium Pages 1-133 in: Engler, A. (ed.). Das material and support. Pflanzenreich IV. 225. Verlag von Wilhelm Engelmann, Leipzig, Germany. LITERATURE CITED WANNTORP, L., WANNTORP, H-E. & M, KÄLLERSJÖ. 2002 (a). Phylogenetic ANDRÉ, E. 1873. Quelques plantes rares. L’ Relationships of Gunnera based on Nuclear Illustration Horticole 20: 156-158. Ribosomal DNA ITS Region, rbcL and rps16 BADER, F. W. J. 1961. Das Areal der Gattung intron Sequences. Systematic Botany 27 (3): Gunnera L. Botanische Jahrbücher für 512-521. Systematik, Pflanzengeschichte und WANNTORP, L., WANNTORP, H-E. & M., Pflanzengeographie 80 (3): 281-293. KÄLLERSJÖ. 2002 (b). The identity of DELCHEVALERIE 1867. Gunnera manicata. Gunnera manicata Linden ex André – Revue Horticole 219. Resolving a Brazilian-Colombian enigma. GANDHI, K. 2002. Nomenclatural corrections Taxon 51: 493-497. and comments in Taxon (3). Taxon 51: 845. LINDEN, J. J. 1867. Plantes nouvelles introduites, par M. J. Linden. Belgique Recibido: 15/04/2006 Horticole 17: 102-105. Aceptado: 09/10/2006

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