PHYTOTAXA ISSN 1179-3163 (online edition)

Phytotaxa 217 (1): 087 091 www.mapress.com/phytotaxa/ ISSN 1179-3155 (print edition) Copyright 2015 Magnolia Press Article PHYTOTAXA ISSN 1179-3163 (online edition) http://dx.doi.org/10.11646/phytotaxa.217.1.8 A new species of Ternstroemia (Pentaphylacaceae) from La Amistad Binational Park and World Heritage Property, Costa Rica and Panama D. Santamaría Aguilar 1, Q. Jiménez-Madrigal 3 2, 4, * & A. K. Monro 1 Current address: Herbaria, Harvard University, 22 Divinity Avenue, Cambridge, Massachusetts 02138-2020 U.S.A 2 Department of Life Sciences, The Natural History Museum, London, SW7 5BD, UK 3 Unidad de Gestión Ambiental, Empresa de Servicios Públicos de Heredia, Heredia, Costa Rica 4 Department of Identification & Naming, The Royal Botanic Gardens, Kew, TW9 3AB, UK * Corresponding author: a.monro@kew.org Abstract Ternstroemia amistadensis Q. Jiménez & D. Santam., a new tree species from La Amistad International Park in Costa Rica and Panama, is described and illustrated. Ternstroemia amistadensis is assessed as Near Threatened, It may be may be distinguished from other species in the genus based on its obovate, coriaceous and very shiny leaf laminae with obtuse-rounded apices and bearing many small black punctations below. Resumen Se describe y se ilustra Ternstroemia amistadensis Q. Jiménez & D. Santam., un arbolito procedente del Parque Internacional La Amistad, Costa Rica-Panamá. La especie se distingue por sus láminas foliares obovadas, obtuso-redondeadas (a veces emarginadas) en el ápice, muy coriáceas y brillantes en el haz y con numerosos y notorios puntos negros en el envés. Key words: Theaceae, Ericales, Mesoamerica, Central America, taxonomy. Introduction Ternstroemia Mutis ex L. f. (1782: 264) is the most species-rich genus of the angiosperm family Pentaphylacaceae, comprising ca. 100 species distributed throughout tropical and subtropical America, Africa, SE and East Asia (Weitzman et al. 2004, Berry & Weitzman 2005, Mabberley 2008). South America, with ca 60 species, is the centre of species diversity for the genus in the Neotropics (Every 2009). Seven species are recorded from Mesoamerica (Boom 1989, Weitzman et al. 2004). Ternstroemia can be distinguished from other Costa Rican and Panamanian Pentaphylacaceae genera by the alternate or subopposite leaves which are arranged spirally and clustered towards the branch tips, the axillary, usually solitary, perfect flowers with a campanulate corolla, many stamens (>40) and a superior ovary and the fruit irregularly dehiscent containing large seeds with a sarcotesta. Materials & Methods Types and specimens at A, BM, CR, F, GH, INB, LPB, MO, MOL, NY, PMA, SCZ and USM were examined together with images from the Global Plants Initiative (GPI). All herbarium acronyms are those of Thiers (2015). Flowers were rehydrated by placing in ammonia hydroxide for one day and then in water until they were sufficiently soft and pliable to be examined under a dissecting microscope. Rehydrated material was then returned to the herbarium specimen. Material was examined under a Leica StereoZoom 5 binocular microscope at X 40. Accepted by Tim Utteridge: 19 Jan. 2015; published: 22 Jun. 2015 87

Ternstroemia amistadensis Q. Jiménez & D. Santam., sp. nov. (Fig. 1, 2) Ternstroemia amistadensis is most similar to the Peruvian T. pachytrocha Kobuski (1942: 320) with which it shares black punctations on the lower leaf surface, glandular-denticulate bracteoles, bilocular ovaries and undivided styles. Ternstroemia amistadensis may be distinguished from T. pachytrocha by the former s larger and obovate leaf laminae with more conspicuous nervation on both surfaces, ovoid fruit with thinner pericarp and smaller seeds that are fewer per fruit. Figure 1. Ternstroemia amistadensis Q. Jiménez & D. Santam.: A, fertile branch with fruit, with enlarged view of lower leaf surface with black punctations. B, flower bud. C, immature fruit, with an insertion highlighting the unequal bracteoles borne opposite or subopposite the base of the calyx. D, mature fruit, with section cleared to reveal seeds. E, seeds. Illustration based on images of living material and herbarium specimens of Santamaría 6706 (INB). 88 Phytotaxa 217 (1) 2015 Magnolia Press Aguilar ET AL.

Type: COSTA RICA. Limón, Z. P. Río Banano. Cuenca del Estrella. Valle de la Estrella, Fila Matama. Cerca de 11 km SW del pueblo de Aguas Zarcas. Punto 10 C. Bosque con predominio de árboles de Drimys, Oreopanax; sotobosque con Piper, Geonoma, Miconia, 09º47 49 N, 83º09 43 W, 1400 1500 m, 29 Oct 2007 (fruit), D. Santamaría, D. Solano, M. Moraga, C. Godínez & A. Rodríguez 6706 (holotype: INB; isotypes: BM, CR, F, GH, K, MO, NY, PMA). Small tree, 5 8 m, the slash reddish; twigs cylindrical, subopposite or verticillate, glabrous. Leaves spirally arranged and clustered toward the branch tips, petiolate; petioles 0.3 1.0 cm, adaxially flattened, glabrous; laminae 5.5 11 2.4 7.1 cm, obovate, coriaceous, glabrous on both surfaces; midrib weakly sunken on the upper surface, prominent and rounded on the lower surface; lateral nerves 4 6 pairs, borne at 45 60 to the midrib, weakly sunken on both surfaces; lower surface with black punctuations visible to the naked eye, ca. 0.3 mm, ± crateriform; upper surface prominently papilose-tuberculate, ca. 0.1 mm; base cuneate, occasionally obtuse; margin entire, somewhat revolute; apex obtuse to occasionally rounded (sometimes emarginate). Flowers axillary, bracteate, solitary, bisexual; pedicel 10 35 mm long, thick 2 3 mm at apex, straight to weakly curved; bracteoles 2, borne opposite or subopposite the base of the calyx, unequal, 2 3 3 5 mm, D -shaped, both inner and outer surface glabrous; outer surface rugose, the margins sparsely glandular-denticulate; apices apiculate or rounded; sepals 5, lobed; lobes imbricate in 2 whorls; external whorl of lobes 7 10 8 10 mm in flowers (in fruits 8 11 10 12 mm), broadly ovate broadly ovate to very broadly ovate; internal whorl of lobes 7 10 7 9.5 ( 12) mm in flowers (in fruit 11 14 9 10 mm), broadly ovate; inner and outer surfaces glabrous; outer surface granular-rugose; margins entire, membranous, brown, rarely inconspicuously glandulardenticulate toward the base (external whorl), the apices rounded; petals 5, 6 9 4 mm, reported as white, fused for the basal ca. 4 mm, ovate; margin entire; apex rounded or acute; stamens ca 43 55; filaments 0.5 2 mm, ± biseriate, free from the petals and separate from each other; anthers 0.5 1.1 mm, the connective extending 1 1.3 mm long; ovary 4 3.5 mm, round and complanate with 2 locules, each bearing 8 11 ovules. Immature(?) fruit 1.3 2.3 1 1.5 cm, spherical to ovoid, brownish-green; pericarp 0.2 0.3 cm thick; style persistent, 0.1 0.3 cm, straight or weakly curved; seeds 3 or 4 by fruit, ca 15 7 9 mm, the sarcotesta purple in fresh material, brown in herbarium material. Etymology: Ternstroemia amistadensis is named after the La Amistad Binational Park and World Heritage Property, the only known localities. Phenology: Flowering material of Ternstroemia amistadensis has been collected in March, April and August and fruiting material between August and October. The area is relatively unseasonal but has variable short dry seasons of 2 4 weeks in February-March and a shorter dry period in August. Distribution and habitat: Ternstroemia amistadensis is known only from the Caribbean slopes of La Amistad Binational Park, Costa Rica and Panama. Here it has been recorded in semi-open areas and edges of wet evergreen forest and cloud forest at elevations of 1050 1700 m with an Extent of Occurrence of ca 14,000 km 2 and associated with Drimys granadensis L. f. (Winteraceae), Oreopanax sp. (Araliaceae) and forest understory characterised by Piper sp. (Piperaceae), Geonoma sp. (Arecaceae) and Miconia sp. (Melastomataceae). Conservation status: Using IUCN criteria (IUCN 2001), Ternstroemia amistadensis is considered Near Threatened (NT). Ternstroemia amistadensis has an Extent of Occurrence of 14,000 km2 (IUCN criteria B1 <20,000 km 2 ) and is known from only six localities (IUCN criteria Ba, number of locations 10). At these localities, the populations of this species comprise a few mature individuals. The majority of the Extent of Occurrence for the species occurs within the La Amistad Binational Park. Whilst this is part of a UNESCO World Heritage site, the forest has been impacted by encroachment from cattle ranching, illegal mining exploration and the construction of a number of hydroelectric stations around the Park s periphery. We suggest that this meets IUCN criteria Bb(iii) of continuing decline in area, extent and/or quality of habitat (IUCN 2001). Using the IUCN methodology, our Global Conservation Assessment for Ternstroemia amistadensis is classed as Vulnerable (VU) based on criteria B1 Ba and Bb(iii). Given that much of the Extent of Occurrence has been poorly explored, we have revised this classification to Near Threatened (NT). Discussion: Ternstroemia amistadensis can be distinguished from other Mesoamerican Ternstroemia species by its coriaceous obovate leaf laminae with obtuse to rounded and occasionally emarginate apices, shiny upper surfaces, black-punctate lower surfaces, conspicuous lateral nerves in dried material and large granular-rugose sepals. T. amistadensis is most similar to T. pachytrocha from Peru and T. macrocarpa Triana & Planchon (1862: 269) from Colombia and Ecuador, which both share with T. amistadensis the characters of a 2-locular ovary, entire style and relatively large fruit. Ternstroemia amistadensis may be distinguished from T. pachytrocha and T. macrocarpa on the basis of leaf, petiole, pedicel and fruit morphology as summarised in Table 1. In the Theaceae treatment (Jiménez 2015) for the Manual de Plantas de Costa Rica T. amistadensis was treated as Ternstroemia. sp. A. A new species of Ternstroemia Phytotaxa 217 (1) 2015 Magnolia Press 89

Figure 2. Living material of Ternstroemia amistadensis Q. Jiménez & D. Santam.: A, leafy branch, showing lower surface of leaves with dark punctations. B, decomposed fruit and sepals. C, mature fruit. Images are of living material (prior to pressing) of Solano et al. 5721 (MO). Photos by Daniel Solano Peralta. Additional specimens examined: Costa Rica. Limón: Cantón de Limón. Zona Protectora Río Banano, cuenca del Banano, Valle de la Estrella, Fila Matama, ca. 11 km SW del pueblo de Aguas Zarcas, 09º48 06 N, 83º09 57 W, 1400 1500 m, 29 Oct 2007 (but fl., fr.), D. Solano et al. 4799 (BM, CR, GH, INB, MO); Cantón de Limón. Matama, El Progreso. Cabeceras del Río Cariei, Fila Matama, 09º47 20 N, 83º08 18 W, 1400 m, 27 Apr 1989 (fl.), G. Herrera & A. Chacón 2817 (INB, BM, CR, GH, MO) Cantón de Talamanca. Bratsi, Amubri, Alto Lari, Kivut, afluente innominado del Río Lari, margen izquierda, 09º22 50 N, 83º05 10 W, 1500 m, 21 Mar 1992 (fl.), G. Herrera 5414 (BM, CR, GH, INB, MO, US); Cantón de Talamanca. Bratsi. Talamanca, Bratsi, Fila de exploración minera entre Río Sukut y Río Carbri. Cerro Schtomat, 09º22 12 N, 82º56 24 W, 1050 m, 13 Jul 1989 (fl.), G. Herrera 3266 (INB, CR). PanamA. Bocas del Toro: Changuinola, Parque Internacional La Amistad, Falso Fábrega, 09º09 59.0 N, 82º39 21.3 W, 1700 m, 6 Aug 2008 (fr.), D. Solano et al. 5721 (INB, MO). Acknowledgements The authors would like to thank Darwin Initiative grant 15-027 (DEFRA, UK Government) for providing field access; botanical artist Claudia Aragón, for the illustration; Frank González for logistical support in the field; Daniel Solano, for images of this species, company and assistance in the field; and the curators of A, BM, CR, F, GH, INB, LPB, MO, MOL, NY, PMA, SCZ and USM, for providing access to their collections, and the two reviewers, Michael Grayum (MO) and Stefan Dressler (FR) for their comments, corrections and suggestions. In addition, the senior author would like to thank the Missouri Botanical Garden and an Elizabeth E. Bascom Grant for supporting a visit to MO, and the curators and staff in the Harvard University Herbaria for their support and hospitality. 90 Phytotaxa 217 (1) 2015 Magnolia Press Aguilar ET AL.

Table 1. Diagnostic comparison of Ternstroemia amistadensis, T. pachytocha and T. macrocarpa. T. amistadensis T. pachytrocha T. macrocarpa Petiole length 0.3 1.0 cm 0.9 1.1 cm up 2 cm* Lamina outline obovate elliptic oblong-obovate, elliptic-obovate Lamina length 5.5 11 cm 6 8 cm 9 16 cm Nervation prominence Prominent on both surfaces Weakly raised on upper Weakly raised on both surface surface only surface Pedicel length 10 35 mm 17 25 mm 25 45 mm* Pedicel diameter at their thickest 2 3 mm 3 3.5 mm 4 5 mm point Bracteole arrangement and shape D-shaped, opposite or Ovate, opposite or medium prominent on upper Triangular, subopposite or clearly as, viewed from above subopposite alternate Petal length 6 9 mm Unknown 13 15 mm* Outer fruiting calyx lobe 8 11 10 12 mm 13 17 14 15 9 10 11 Fruit shape Ovoid Globose Conical, ovoid to depressed globose Fruit length 1.3 2.3 cm 1.7 2.5 cm* 2.5 cm* Fruit diameter 1 1.5 cm 2.2 2.5 cm* up 2.5 cm* Pericarp thickness 2 mm 5 6 mm* 1 mm Seed no. by fruit 3 or 4 5 9 16 18* Seed size (long) 15 mm 6 8 mm* 7 8 mm* * Information from Kobuski 1942. References Berry, P.E. & Weitzman, A.L. (2005) Ternstroemiaceae. In: Steyermark, J.A., Berry, P.E. & Holst, B.K. (Eds.) Flora of the Venezuelan Guayana. Vol. 9. Missouri Botanical Garden Press, St. Louis, pp. 300 308. Boom, B.M. (1989) New species of Ternstroemia (Theaceae) from the Guayana Highland. Brittonia 41 (2): 136 142. http://dx.doi.org/10.2307/2807518 Every, J.L.R. (2009) Pentaphylacaceae. In: Milliken, W., Klitgård, B. & Baracat, A. (2009 onwards), Neotropikey - Interactive key and information resources for flowering plants of the Neotropics. Available from: www.kew.org/neotropikey (accessed 1 January 2014) IUCN [International Union for Conservation of Nature and Natural Resources Species Survival Commission] (2001) IUCN Red List Categories, Version 3.1. International Union for Conservation of Nature and Natural Resources, Gland and Cambridge, 70 pp. Jiménez, Q. (2015) Theaceae. In: Hammel, B.E., Grayum, M.H., Herrera, C. & Zamora, N. (Eds.) Manual de Plantas de Costa Rica. Vol. 8. Dicotiledóneas (Sabiaceae Zygophyllaceae). Monographs in Systematic Botany from the Missouri Botanical Garden, pp 393 406. Kobuski, E.C. (1942) Studies in the Theaceae XII Notes on the South American Species of Ternstroemia. Journal of the Arnold Arboretum 23: 298 343. Linnaeus, C. (1782 [1781 publ. Apr. 1782]) Supplementum Plantarum Systematis Vegetabilium Editionis Decimae Tertiae, Generum Plantarum Editiones Sextae, et Specierum Plantarum Editionis Secundae. Vol. 39. Editum a Carolo a Linné, Brunsvigae [Braunschweig], 264 pp. Mabberley, D.J. (2008) Mabberley s Plant-book: A Portable Dictionary of Plants, their Classifications, and Uses, Third edition. Cambridge University Press, Cambridge, 1021 pp. Thiers, B. (2015) Index Herbariorum: a global directory of public herbaria and associated staff. New York Botanical Gardens Virtual Herbarium. Available from: http://sweetgum.nybg.org/ih (accessed January 2015) Triana, J.J. & Planchon, J.E. (1862) Annales des Sciences Naturelles; Botanique, Série 4 18: 259. Weitzman, A.L., Dressler, S. & Stevens, P.F. (2004) Ternstroemiaceae. In: Kubitzki, K. (Ed.) The Families and Genera of Vascular Plants. Flowering Plants. Dicotyledons. Vol. 6. Springer-Verlag, Berlin, pp. 450 460. http://dx.doi.org/10.1007/978-3-662-07257-8_47 A new species of Ternstroemia Phytotaxa 217 (1) 2015 Magnolia Press 91