Systematic Botany (2014), 39(2): pp. 533–540 © Copyright 2014 by the American Society of Plant Taxonomists DOI 10.1600/036364414X680924 Date of publication 04/23/2014 Quipuanthus, a New Genus of Melastomataceae from the Foothills of the Andes in Ecuador and Peru Fabián A. Michelangeli,1,3 Carmen Ulloa Ulloa,2 and Karla Sosa1 1 The New York Botanical Garden, 2900 Southern Boulevard., Bronx, New York 10458 U. S. A. 2 Missouri Botanical Garden, P. O. Box 299, St. Louis, Missouri 63166-0299, U. S. A. 3 Author for correspondence (fabian@nybg.org) Communicating Editor: Jennifer A. Tate Abstract—The new genus Quipuanthus from the foothills of the eastern Andes of Ecuador and Peru is described. Quipuanthus seems to be related to Allomaieta, Alloneuron, and Wurdastom in the tribe Cyphostyleae, but the combination of characters as an herb with haplostemonous flowers, recurved style, inferior ovary and apically dehiscent capsular fruits is unique among the Melastomataceae. The new species Quipuanthus epipetricus is described and illustrated. Keywords—Cyphostyleae, haplostemony, Neotropics, new species, taxonomy. analyzed under parsimony and maximum likelihood. Parsimony analyses were performed in TNT (Goloboff et al. 2008) with 1,000 random addition sequences, holding five trees per replicate and the resulting most-parsimonious trees were then swapped to completion. Maximum likelihood analyses were performed with RAxML using default parameters (Stamatakis 2006; Stamatakis et al. 2008) and run through the CIPRES Science Gateway (http://www.phylo.org/; Miller et al. 2010). Bootstrap values were estimated on the ML tree also using RAxML based on 1000 searches run through the CIPRES Science Gateway. Voucher information and GenBank accession numbers for the two newly generated sequences are in Appendix 1. For all other voucher and GenBank accession numbers see Amorim et al. (2009), Michelangeli et al. (2011) and Goldenberg et al. (2012). Melastomataceae are among the five most diverse families of vascular plants both in Ecuador (Jørgensen and León-Yánez 1999) and Peru (Brako and Zarucchi 1993) with some ca. 570 and ca. 650 species, respectively, and novelties constantly emerging (Penneys and Jost 2009; Bussmann et al. 2010; Sagástegui Alva et al. 2010; Ulloa Ulloa et al. 2012; Michelangeli and Ulloa Ulloa 2013) with recent explorations of the amazing floras of these countries. During an intense curation of the Melastomataceae collection at the Missouri Botanical Garden in November 2012, we found specimens with peculiar inflorescences in a rosette-like plant, inferior ovaries and apically dehiscing fruits. Further studies and examination of additional specimens revealed that the flowers are haplostemonous, a character rather rare in the family, which usually have stamens double the number of petals (diplostemonous). We were not able to assign these specimens to any currently accepted genus in this diverse family. The combination of characters led us to believe it was necessary to place them in a new genus, Quipuanthus that we propose herein. Results The resulting data matrix and trees can be found under TreeBASE study number 14495. Parsimony (not shown) and likelihood analyses (Fig. 1) yielded highly congruent results differing only in the level of resolution within some higher nodes and in some short branches with low or no bootstrap support that collapsed under parsimony (i.e. the Triolena Naudin clade and the Cyphostyleae are not resolved as sister, but form a three-way polytomy with the Cambessedesia DC. and allies clade). However, both analyses unequivocally place Quipuanthus in a polytomy at the base of Allomaieta Gleason well within the tribe Cyphostyleae (Fig. 1). In fact, the partial rbcL sequence of Quipuanthus is identical to two known sequences of Allomaieta (A. hirsuta (Gleason) Lozano and A. pancurana Lozano), and thus its placement with these taxa is to be expected. The accession of Wurdastom hexamera is resolved as sister to Wurdastom cuatrecasii (Wurdack) B. Walln. Materials and Methods We revised the collections deposited at the Missouri Botanical Garden (MO), The New York Botanical Garden (NY), the Pontificia Universidad Católica del Ecuador (QCA), Herbario Nacional del Ecuador (QCNE), and at Universidad del Azuay (HA), as well as electronic images made available to us from the Museo de Historia Natural de la Universidad Mayor de San Marcos (USM) and the Universidad Nacional de Trujillo (HUT). Morphological examinations and measurements were done from dried herbarium specimens and flowers were rehydrated. Habit details were observed on color photographs made available to us. Seeds and leaves from herbarium specimens were observed under Scanning Electron Microscopy, mounted on aluminum stubs, coated with gold-palladium for three min with a HUMMER 6.2 Sputter Coater (Aratech LTD). Specimens were observed and photographed on a JEOL, JSM 5410LV. In order to ascertain the phylogenetic position of Quipuanthus within the family, genomic DNA was extracted from herbarium material (see Appendix 1) using the DNeasy plant mini kit from QIAGEN (Valencia, California) following the manufacturer’s instructions with the following exceptions: 5 ml of ascorbic acid (0.1 mg/ml) was added to the lysis buffer and samples were incubated at 45
for 24 h; DNA was eluted with 50 ml of elution buffer. Several attempts were made to amplify different loci but only the 50 end of rbcL was successfully amplified and sequenced following the protocols of Fay et al. (1997) and Clausing and Renner (2001). This rbcL sequence was incorporated into the plastid matrix of Goldenberg et al. (2012) with the addition of an rbcL sequence for Wurdastom hexamera (Wurdack) B.Walln., and the resulting matrix Taxonomic Treatment Quipuanthus Michelangeli & C. Ulloa, gen. nov.—TYPE: Quipuanthus epipetricus Michelangeli & C. Ulloa. Herbs. Inflorescence a unilateral pedunculate cyme. Flowers 5-merous, the calyx open in bud and persistent in fruit with external teeth well developed; haplostemonous, only the antisepalous stamens present, these truncate with a wide apical pore. Style recurved. Ovary inferior, 3-locular. Fruit capsular, apically dehiscent by bivalved openings. Herbs with short stems growing on rocks, rosette-like; stems short, up to 10 cm with condensed internodes up to 533 534 SYSTEMATIC BOTANY [Volume 39 Fig. 1. Maximum likelihood phylogeny of the Melastomataceae based on plastid sequence data (rbcL, ndhF and rpl16 gene) showing the position of Quipuanthus within the tribe Cyphostyleae. Numbers above branches represent bootstrap values. 2014] MICHELANGELI ET AL: QUIPUANTHUS, A NEW GENUS OF MELASTOMATACEAE + + + + + + + 5 mm, fleshy, pink inside (cut), completely concealed by a mix of densely packed coarse stramineous simple trichomes, 9–12 ca. 0.5 mm and sparse, minute (< 0.1 mm) sessile glands. Leaves with petioles 5–26 cm long, pinkish, the entire length covered by simple trichomes 4.5–18 mm long and sparse, minute (< 0.1 mm) sessile glands; leaf blades round to ovate (4.5–)9–26.5 (4.2–)7.5–30 cm, the base deeply cordate sometimes with the lobes slightly overlapping, the sinuses 1.8–4.2 cm, the apex broadly acute to shortly apiculate (then the tip up to 15 12 mm), the margin entire to obscurely crenulate, ciliate, the trichomes simple 4.5–12 mm long; venation acrodromous, with 4–5 pairs of secondaries, the outermost pair often only evident in the basal sinuses, then becoming marginal and faint; tertiary veins percurrent and perpendicular to the primary and secondary veins, with finely reticulate veins of higher order; abaxial surface flat, greenish-white, the primary and secondary veins slightly raised and pinkish lilac, glabrous on the surface, densely to more rarely sparsely pubescent, with a mix of simple trichomes up to 8.5 mm long (up to 12 mm long and denser at the base of the nerves) and sparse, minute (< 0.1 mm) sessile glands, these also present in all the higher order veins; adaxial surface flat, bright green, appearing glabrous but with sparse, minute (< 0.1 mm) sessile glands, and also occasionally with very sparse appressed simple trichomes up to 12 mm long. Inflorescences unilateral pedunculate cymes 8–26 cm long (including the peduncle), the peduncle 6–20 cm, ca. 1 mm wide and pinkish while in flower, becoming 2.2–2.5 mm wide and stramineous when the fruits are mature, clustered in the upper nodes of the short stem, occasionally bifurcated ca. 2/3 the way up the first flower, glabrous or with sparse simple trichomes up to 6 mm long. Bracteoles absent or early caducous. Flowers 5-merous; hypanthia 5.2–8.1 mm long, ca. 1.3 mm wide, mostly cylindrical, light green, slightly 10-costate, the upper portion flaring outwards, up to 4.5 mm at the apex of the calyx. Calyx ca. 0.7 mm long, the sepals broadly deltoid, fused 2/3 of the way, the teeth ca. 1.3 mm long, subulate, and projecting above the sepals; petals 14–17 22–25 mm, broadly obovate to round, entire, glabrous, pink; stamens 5, only the antisepalous set present, the filaments 2.1–2.5 mm long, the anthers 1.8–2 ca. 0.6 mm, obovate and truncate, with a broad apical pore, the connective not prolonged but slightly expanded basally. The style 1.9–2.4 mm long, distally dilated to ca. 0.45 mm wide, and completely recurved so the stigma is oriented downwards. Ovary inferior, 3-locular. Fruits capsular, 10–18 3.5–5 mm, conspicuously 10-costate, apically dehiscent through three distinct bivalved openings, the valves ca. 1.5 mm, blunt, the calyx and teeth persistent, the ribs fibrous, stramineous and persistent. Seeds numerous, narrowly ovoid to slightly pyramidal, 0.6–1.2 0.28– 0.35 mm, the testa smooth; under light microscope the hilum end dark brown, the chalazal end translucent, appearing hollow. Figures 2–4. Quipuanthus epipetricus Michelangeli & C. Ulloa, sp. nov. —TYPE: PERU. Amazonas: Bagua, Imaza, Comunidad Aguaruna de Wanás (km 92 carretera Bagua-Imacita): Cerros Chinim, 650–750 m, 26 Aug 1996 (fl, fr), C. Dı́az, A. Peña, L. Tsamajain & M. Roca 8017 (holotype USM!; isotypes: HUT!, K!, MO!, NY!). The diagnosis and description of the new species are the same as that of the genus. 535 Etymology—The generic name is a combination of the Quechua quipu = cords used as record-keeping devices, and anthos = flower (Greek). The quipu (or kipu) were long, knotted strings used by pre-colonial Andean societies to encode information. The overall appearance of the longpedunculate inflorescences and infructescences, emerging from the short stems, with flowers and fruits like knots on strings, and the puzzling combination of characters in this genus, reminded us of this enigmatic system used by the Andean societies, long before the arrival of the Spanish writing system. The specific epithet of this new species alludes to the fact that all specimens known have been collected growing on top of rocks. Ecology and Distribution—Quipuanthus epipetricus is known from two populations on the foothills of the Eastern Andes mountains: one in the Cordillera de Galeras, Napo province (border with Orellana), Ecuador, and another in the Imaza district, province of Bagua, department of Amazonas, Peru (Fig. 5). In both places, the short-stemmed plants grow appressed to rocks, sometimes on vertical walls, with the abaxial side always facing the substrate, in low- to midelevation rain forest, between 1,000–1,400 m in Ecuador and between 400–730 m in Peru. Even though the two populations are over 500 km apart, based on vegetative and fruit morphology, there is no doubt that they belong to the same species. The Cordillera de Galeras is part of the SumacoNapo Galeras National Park in northeastern Ecuador and is an outcrop mostly formed of Cretaceous limestone (Neill 1999). The two Peruvian populations were collected only some 30 km apart along the 5N road in the Imaza district close to the village of Chiriaco, in forested ravines. Conservation Status—The two populations of Quipuanthus epipetricus so far known have an area of occupancy (AOO) of 63 km2 (36 km2 in Ecuador and 27 km2 in Peru) in evergreen moist forests. The extent of occurrence (EOO) is 6,284 km2. Only the Ecuadorian population is protected under Ecuador’s National System of Protected Areas in a pristine forested area with limited access; the Peruvian population was collected in ravines immediately surrounded by crop fields and although extensive areas of forest were observed on larger satellite imagery, it is unknown if suitable habitat is available. In terms of our current knowledge, and taking into consideration the overall distribution and the specialized habitat of the species it is assigned a provisional IUCN conservation status of Endangered (IUCN 2011). Additional Specimens Examined—ECUADOR. Napo [border with Orellana]: Archidona. Parque Nacional Sumaco Napo-Galeras, Cordillera de Galeras, bloque 19, lı́nea 30, Compañı́a Triton, 0
530 S 77
330 W, 1,400 m, 24 Abr 1996 (fr), E. Freire & J. Cerda 499 (MO, QCNE n.v.); Archidona. Parque Nacional Sumaco Napo-Galeras, Cordillera de Galeras, bloque 19, lı́nea 30, Compañı́a Triton, 0
530 S 77
330 W, 1,600 m, 24 Abr 1996 (fr), E. Freire & J. Cerda 518 (MO, QCNE [2 sheets]); Parque Nacional Sumaco Napo-Galeras, Cordillera de Galeras, 0
50.30 S 77
33.60 W, 1,080 m, 24 Oct 2006 (fr) J. Homeier et al. 2477 (GOET n.v., MO, NY, QCA n.v., QCNE), 15 Abr 2006 (fr), J. Homeier & M. Unger 2330 (GOET n.v., MO, NY, QCA, QCNE n.v.); Cordillera de Galeras, Parque Nacional Sumaco Napo-Galeras, western base of Galeras massif, near park boundary, in Mushullacta community forest, 00
490 5100 S 077
380 2700 W, 1,200 m, 2 Mar 2003 (fr), D. Neill et al. 14201 (MO n.v., QCNE); Cordillera de Galeras, Parque Nacional Sumaco Napo-Galeras, steep east slope of Galeras massif, below military base on trail to Arapino, 00
490 5700 S 077
310 2600 W, 1,600 m, 10 Mar 2003 (fr), D. Neill et al. 14207 (AAU n.v., CAS n.v., MO, QCNE). PERU. Amazonas: Bagua: 43 km (by road), northeast of Chiriaco. Along roadside from Chiriaco to Puente Venezuela, 320–730 m, 5 Nov 1978 (fl, fr), J. Barbour 4451 (MO); Bagua, Imaza, Quebra El Almendro, 5
180 S 78
200 W, 400 m, 9 Feb 1999 (fl), R. Vázquez et al. 26112 (MO, NY, US); 536 SYSTEMATIC BOTANY [Volume 39 Fig. 2. Quipuanthus epipetricus: A. Flowering and fruiting plant. B. Inflorescence detail. C. Flower before anthesis, petals removed, and detail of the calyx. D. Anthers before anthesis. E. Flower at anthesis, petals removed and hypanthium in longitudinal section. F. Anther, side view. G. Style. H. Fruiting plant. I. Detail of infructescence. J. Mature capsule with opened valves. (A, I, from Dı́az et al. 8017 (NY), B–D, J from Vázquez 27573 (MO), E–G from Vázquez et al. 26112 (NY), H from van der Werff et al. 14552 (MO)). 2014] MICHELANGELI ET AL: QUIPUANTHUS, A NEW GENUS OF MELASTOMATACEAE 537 Fig. 3. Quipuanthus epipetricus, habitat and fruiting plant (Homeier 2477). Photograph courtesy of J. Homeier. Bagua, Imaza, Quebra El Almendro, 5
140 4000 S 78
210 3400 W, 360 m, 11 Feb 2002 (fl, fr), R. Vázquez et al. 27573 (MO); Bagua, Imaza, Quebrada El Almendro, 5
140 4000 S 78
210 2400 W, 430 m, 9 Mar 1998 (fr), H. van der Werff & R. Rojas 14552 (MO, NY); Bagua, Imaza, Quebrada El Almendro, 5
140 4000 S 78
210 3400 W, 400 m, 17 Mar 2001 (fr), H. van der Werff et al. 16151 (COL, M, MO, NY, USM). Discussion Previous to this description, most specimens of Quipuanthus had been determined as Salpinga Mart. ex DC., Triolena Naudin, or Monolena Triana ex Benth. & Hook. f. Quipuanthus resembles all of these genera in the herbaceous habit, long pedunculate inflorescences with unilateral cymes, and capsular fruits (Wurdack 1964; Almeda and Dodero 1991; Wurdack et al. 1993; Warner 2002). However, Salpinga, Monolena, and Triolena all have diplostemonous flowers (with 10 anthers) with developed anther appendages and completely superior ovaries. Additionally, the capsules of Monolena and Triolena are proportionally much shorter and obviously three-sided (Almeda and Dodero 1991; Warner 2002). Salpinga, Monolena, and Triolena have traditionally been placed in the tribe Bertolonieae (Cogniaux 1891; Renner 1993), but recent analyses based on DNA sequence data have shown that Salpinga is probably better placed in the Merianieae (Goldenberg et al. 2012), indicating that the herbaceous rosette habit with long unilateral cymes may have evolved more than once in the family. Haplostemonous flowers and capsular fruits derived from inferior ovaries are seldom encountered in Neotropical Melastomataceae (Goldenberg and Amorim 2006; Amorim et al. 2009; Michelangeli et al. 2011), and together they are only known to occur in three genera: Allomaieta Gleason, Alloneuron Pilger, and Wurdastom Wallnöfer. These three genera were considered part of the tribe Miconieae, but have been recently grouped under the tribe Cyphostyleae (Michelangeli et al. 2011). All 20 species in Cyphostyleae grow at mid- to low-elevation in inter-Andean valleys from Colombia to Peru (Wallnöfer 1996; Lozano and Becerra-deLozano 1999; Wallnöfer 1999). Given the character combination and geographic distribution Quipuanthus seems well positioned within the Cyphostyleae. However, the character combination of Quipuanthus does not match any of the three genera as currently defined. Quipuanthus differs from all other genera in the Cyphostyleae in that the calyx is opened in bud and that the fruit opens by apical valves and does not break apart when the seeds are released (Michelangeli et al. 2011). The seeds of Quipuanthus differ remarkably from any known Cyphosteleae, or any other Melastomataceae: they have a very smooth testa and appear to be hollow, with the embryo positioned only on the hilum 538 SYSTEMATIC BOTANY [Volume 39 Fig. 4. Quipuanthus epipetricus, SEM of seeds and abaxial leaf surface. A. Seed. B. Seed with the chalazal end ruptured showing embryo on the hilum end. C. Abaxial leaf surface showing trichomes of tertiary veins (all from van der Werff 14552 (NY)). 2014] MICHELANGELI ET AL: QUIPUANTHUS, A NEW GENUS OF MELASTOMATACEAE 539 Because we only have a partial sequence for Quipuanthus for a fairly conserved locus, the fact that there are no differences between it and two species of Allomaieta should not be taken as a justification to describe this new species within the latter genus. As Quipuanthus is not embedded within Allomaieta (just not resolved) and its inclusion in Allomaieta would considerably change the current boundaries of the latter, we feel that the creation of this new genus is warranted. Acknowledgments. This research has been supported in part by the National Science Foundation (DEB-0818399 and DEB-1140409). The Davidson family supported Ulloa’s visit to the Ecuadorian herbaria. We also wish to thank Daniela Cevallos (QCA), Diana Fernández (QCNE), Eric Rodrı́guez (HUT) and Asunción Cano and Diego Paredes (USM) for locating material at their respective herbaria and providing digital images, and Jim Solomon (MO) and the staff at NY for facilitating loans. Jürgen Homeier provided plant material and photographs of these collections in the field. Burgund Bassüner prepared the base map and provided the Conservation Status. Bobbi Angell prepared the beautiful line drawing. Frank Almeda provided tissue material from Wurdastom hexamera which was collected as part of the PBI-Miconieae project; the PBI project in Colombia has the required permits for collection as well as for molecular research (Contrato de Acceso a Recurso Genético para Investigación Cientı́fica sin Interés Comercial) for members of the Melastomataceae family, granted by Ministerio de Ambiente y Desarrollo Sostenible and Autoridad Nacional de Licencias Ambientales (ANLA). Two anonymous reviewers provided very helpful comments that greatly improved an earlier version of this manuscript. Literature Cited Fig. 5. Distribution map of Quipuanthus epipetricus. end of the seed (Fig. 4A, B). Hollow appendages are present in some species of Miconieae, but these are formed by empty spaces in between the cells, and the seed themselves appear to be hard and the nature of the appendage is not obvious unless the seed is sectioned (Martin et al. 2008; Martin and Michelangeli 2009). In Quipuanthus the appendage is soft, translucent (features not obvious under SEM) and the chalazal end resembles a sac (see Fig. 4B for a photograph of the ruptured sac). The seeds of the remaining members of the Cyphostyleae are not smooth, and pyramidal, pyramidal with the testa bearing tubercules on the edges of the chalazal (the tubercules small in Wurdastom). The seeds of some species of Wurdastom have a very long appendage on the chalazal end, but this appendage is not hollow (see seed SEM images for all other genera of Cyphostyleae at Michelangeli et al. 2009; http://sweetgum .nybg.org/melastomataceeae). Quipuanthus has the recurved style of Allomaieta, but lacks the calyptrate calyx and differs in inflorescence morphology (unilateral cymes vs. dichasial cymes) and anther morphology (truncate with a broad terminal pore vs. apiculate with a subterminal small pore) (Lozano and Becerrade-Lozano 1999; Michelangeli et al. 2011). Quipuanthus shares with Wurdastom leaves with round to cordate bases and truncate anthers, but lacks barbellate hairs and differs in inflorescence and style morphology (Wallnöfer 1996). Lastly, Quipuanthus and Alloneuron both have unilateral cymes and some species of Alloneuron have external calyx teeth, but the two differ in leaf venation and style morphology (Wallnöfer 1996). Almeda, F. and G. U. Dodero. 1991. 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Alloneuron Pilg. (Melastomataceae): some additions. Annalen des Naturhistorischen Museums in Wien Serie B Botanik und Zoologie 101B: 593–598. Warner, R. H. 2002. Systematics of the genus Monolena (Melastomataceae) in Central America. Proceedings of the California Academy of Sciences 53: 95–116. Wurdack, J. J. 1964. Certamen Melastomataceis VIII. Phytologia 9: 409–426. Wurdack, J. J., S. S. Renner, and T. Morley. 1993. Melastomataceae. Flora of the Guianas. Koenigstein, Germany: Koeltz Scientific Books. Appendix 1. Taxon, Collector, collection number (herbarium) and GenBank accession number for rbcL sequences newly generated for this study. Quipuanthus epipetricus. van der Werff et al. 14552 (NY), KF407947; Wurdastom hexamera. Almeda 10279 (CAS), KF407948.