PhytoKeys 20 | ; 99-| 22 (2022) A peer-reviewed open-access journal & HC a $¢PhytoKeys https:/ / Pp hyto keys -pen soft.net Launched to accelerate biodiversity research Resurrection of Passiflora acuminata DC. and synonymization of P. tolimana Harms, P. gleasonii Killip, P. metae M. Bonilla, C. Aguirre & Caetano (Passifloraceae) following a study of their morphology and ecogeography Maxime Rome', Geo Coppens d’Eeckenbrugge”’, John Ocampo Pérez*, Mathew Rees>® | Jardin du Lautaret, CNRS, Université Grenoble Alpes, Grenoble, France 2 CIRAD, UMR AGAP Institut, Avenue Agropolis, F-34398 Montpellier, France 3 UMR AGAP Institut, Univ. Montpellier, CIRAD, INRAE, Institut Agro, F-34398 Montpellier, France 4 Universidad Nacional de Colombia, Palmira, Valle del Cauca, Colombia 5 University of Edinburgh, Kings Building, Edinburgh EH9 3E UK 6 Royal Botanic Garden Edinburgh,20a, Inverleith Row, Edinburgh, EH3 5LR, UK Corresponding author: Geo Coppens d’Eeckenbrugge (geo.coppens@cirad.fr) Academic editor: Hanno Schaefer | Received 8 March 2022 | Accepted 27 May 2022 | Published 23 June 2022 Citation: Rome M, Coppens d'Eeckenbrugge G, Ocampo Pérez J, Rees M (2022) Resurrection of Passiflora acuminata DC. and synonymization of P tolimana Harms, P gleasonii Killip, P metae M. Bonilla, C. Aguirre & Caetano (Passifloraceae) following a study of their morphology and ecogeography. PhytoKeys 201: 99-122. https://doi. org/10.3897/phytokeys.201.83316 Abstract Within the very uniform series Laurifoliae, Passiflora acuminata (treated as a synonym of P /aurifolia in the Flora of China), P. tolimana, P gleasonii and P metae appear particularly similar. A review of their descrip- tions and the associated specimens confirms their lack of morphological differentiation and leads us to formally resurrect P acuminata and place the three other taxa under its synonymy. This taxonomic move is also supported by a revision of 72 additional geolocalized specimens (for a grand total of 78) and an analysis of their distribution and habitats. In fact, the bioclimatic space corresponding to the specimens previously assigned to P acuminata encompasses that of all specimens previously assigned to the three other taxa under study. The species range covers a wide region, comprising the lower Amazon and the north of its basin, mostly below 200 m, and, to the west, in the upper Amazon, the Orinoco basin, and along the Andean foothills and valleys, from Venezuela to Peru, at elevations between 100 and 2200 m. In the lowlands, the species appears associated with white sand savannas and water courses. A more complete description is proposed for the species, including its unusual fusiform and slightly ribbed fruit. Another rare trait in the series Laurifoliae is that the outer corona filaments tend to be longer than the corolla. Copyright Maxime Rome et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 100 Maxime Rome etal. / PhytoKeys 201: 99-122 (2022) Keywords Laurifoliae, subgenus Passiflora, taxonomy Introduction Passiflora acuminata DC., P tolimana Harms, P gleasonii Killip, and P metae M. Bonil- la, C. Aguirre & C.M. Caetano are very similar taxa that belong to series Laurifoliae Killip ex Cervi, 1997 (subgenus Passiflora, supersection Laurifoliae (Killip ex Cervi) Feuillet & J.M. MacDougal, 2003), which counted 29 species in the classification of MacDougal and Feuillet (2004). The whole series shows a remarkable morphological and ecological uniformity, rendering its taxonomy extremely difficult (Killip 1938). Revising the criteria that delimit it, Rome and Coppens d’Eeckenbrugge (2017) ex- cluded five species from the series Laurifoliae. The remaining 24 species are glabrous to pubescent, have stems terete to angular, wingless, sometimes corky on old parts with stipules linear to setaceous, early deciduous. Their leaves are unlobed, oblong- lanceolate, entire to glandular-serrulate, not peltate, and have petioles with 0-2 discoid to oblong sessile glands. Flowers are pendent with two outer series of long campanulate filaments, an involucre of three glandular bracts, free at the base and measuring more than 1 cm long. Since this revision, one more species corresponding to these criteria has been described in Colombia as P gustaviana Ocampo & Molinari, 2017. Laurifoliae species also share habitat preferences, in humid neotropical forests, at low to medium elevations in Central and South America. Furthermore, several species, like P nitida Kunth, P riparia Mart. ex Mast., P ambigua Hemsl., show very extensive distributions (e.g. across the Amazonian basin and/or along the tropical Andes), which favors geographical differentiation and may lead to redundant descriptions in different countries. Rome and Coppens d’Eeckenbrugge (2019) demonstrated such a situation for P crenata Feuillet & Cremers, P emiliae Sacco, P fernandezii L.K. Escobar and P pergrandis Holm-Niels. & Lawesson, and included them in the synonymy of P riparia. Here, we present a similar analysis on 2 acuminata, P. tolimana, P gleasonii, and P metae. Among these four taxa, the most ancient one is P acuminata, described from Bra- zil by de Candolle (1828). His very short Latin diagnosis translates as follows: “leaves glabrous, ovate-lanceolate, acuminate, entire; petiole with two glands at the apex; bracts oblong, obtuse, and entire.” De Candolle mentioned a specimen of this species conserved at the National Museum of Natural History of Paris. The latter, without number, had been collected by an anonymous botanist. In Flora Brasiliensis, Masters (1872) placed P acuminata as a synonym of P /aurifolia L. It was not until 1938 that the species was revived by Killip in Zhe American species of Passifloraceae, with a de- tailed description and citations of several herbarium specimens from Brazil, on which the floral structure can be easily examined. As did Killip, Tillett (2003) distinguished P. acuminata from P laurifolia by its radii in two equal series (vs. two unequal series). However, in 2007, this species was again treated as a synonym of P /aurifolia in the Flora of China (Flora of China Editorial Committee 2007). This rearrangement was Passiflora acuminata DC. and its synonyms 101 not followed, and many authors (e.g. Vanderplank and Zappi 2011, Silva et al. 2013, 2017, Bonilla Morales et al. 2016) still mention it as a distinct species. In January 1886, in the municipality of Dolores (Tolima, Colombia), at elevations of 1400-1800 m, Lehmann collected a passionflower with greenish white flowers and a corona of purple filaments. He was careful to note that its glossy dark green leaves had a tough texture and, often, brown nervation. The fruit seemed to remain dark green at maturity with a pleasant and sweet taste. Lehmann drew it on the label as a relatively fusiform and ribbed fruit (Plate 1F), not resembling the globular yellow fruits generally observed in series Laurifoliae. Finally, he mentioned that the species is quite common but seems to produce fruit only rarely. A few years later, Harms (1894) described meticulously Lehmann’s specimen as Passiflora tolimana. For the corona, he mentioned that the outermost filaments are slightly longer than the petals, while the innermost filaments are sometimes fused, forming a membrane. In 1924, Killip described Passiflora gleasonii from a herbarium specimen collected by J.S. De La Cruz in 1923, on the banks of the Pomeroon River in British Guyana. In the original description, he noted that the species had only three series of filaments, with two outer subequal series, capillary, 4-5 cm long, and an inner series, narrowly linear, 1 mm long. Finally, he mentioned that this species is close to Passiflora nitida Kunth, but differs from the latter by the outer filaments being narrower and bracts narrowed at both ends, resembling those of Passiflora vitifolia Kunth. In 2016, Bonilla Morales et al. described P metae from two very close sites (within 1 km) at Villavicencio in the Meta department of Colombia. Their description and diagnostic key to other Colombian species underline the glabrous ovary (vs. pubescent in P tolimana and P gleasonii); however, most characters remain the same. In the present article, we compare the descriptions of P acuminata with those of P tolimana, PR metae and P gleasonii and study the type specimens and other herbarium specimens cited in their descriptions. Field observations of flowers and fruits corresponding to P gleasonii from Venezuela, P acuminata from Para (Brazil), and P tolimana from Ecuador, are presented. We thus show that they cannot be distinguished on their morphology. Based on the examination of 78 geolocalized field observations and herbarium specimens, we analyze their distribution and habitat, using Ecological Niche Modeling tools and available label information. Finally, we resurrect formally P acuminata, provide a complete and precise description of it, and place the names of P tolimana, P metae and P. gleasonii in its synonymy. Materials and methods Morphological analyses We first compare the descriptions of P acuminata by Killip (1938) and Tillett (2003), P. tolimana by Harms (1894) and Killip (1938), the original description of P metae by Bonilla Morales et al. (2016) and those of P gleasonii by Killip (1924, 1938) and Tillett (2003). This comparison is focused on traits relative to stems, stipules, petiole, leaves, 102 Maxime Rome etal. / PhytoKeys 201: 99-122 (2022) bracts, calyx, corolla, corona (outer and innermost series of filaments), operculum, li- men, ovary, and fruit. A second comparison bears on 15 of the 17 vouchers associated with the above-men- tioned descriptions, focusing on the following traits: petiolar gland position, plant pu- bescence, bract size, and perianth parts, as observable or mentioned on the voucher label. The specimen Ducke 10528, cited in Killip’s description of P acuminata, could not be obtained for examination. The specimen Bonilla 197 (paratype of P metae) had not been deposited at CUVC. No voucher was associated to Tillett’s description of P gleasonii. We further include our own observations on living material from Ecuador (Sucum- bios), Colombia (Meta, Santander), Venezuela (Mérida, Tachira, Zulia), and Para (Bra- zil), other vouchers (newly determined), as well as good and precisely geolocalized photographs from several botanists and/or passionflowers enthusiasts. Examined herbarium specimens The following list consists of specimens that were previously assigned either to one of the four taxa under study (P acuminata, P tolimana, P gleasonii, and P metae), or to other Passiflora species, and now re-assigned to or kept as P acuminata by the authors. Previously assigned to Passiflora acuminata Brazil: Amazonas: unknown collector s.n. (holotype, P). Lago de Tefé, Manua, 13 Jul 1973, Lleras 16644 (NY). Manaus, along road to Caracarai, 16 Aug 1986, Croat 62222 (MO). Para: Para, Ilha de Marajé, Salvaterra, 2 Mar 2012, Costa 534 (MG). 27 Oct-7 Nov 1929, Killip 30272 (US). Obidos, 19-20 Jul 1934, Swallen 5095 (US). Ilha do Mosqueiro, 3-9 Nov 1929, Killip 30572 (US). Santarém, Alter do Chao, between 1998 and 2005, Knowles 1555 (MG). Belém, near Instituto Agronémico do Norte, 28 Oct 1959, Kuhlmann 386 (SPF). Rodovia Belém-Brasilia, Km 93, 14 Sept 1959, Kuhlmann 361 (SPF). Santarém, 16 Sept 1999, Cordeiro 4089 (MG). Belém, Parque Estadual do Utinga, 19 Sept 2010, Silva 10 (MG). Belém, Reserva do Mocambo, 23 Nov 2009, Kerpel 8 (MG). Municipio Almeirim, Mount Dourado, 23 Dec 1986, Pires 1560 (UPCB). South of Instituto Agronémico do Norte, Belém, 25 Nov 1942, Archer 7864 (K). Road from Belém to Viguié, 6 Apr 1961, Aubréville 198 (P). Municipio de Oriximina, Rio Trombetas, 22 Apr 2007, Salomao 902 (MG). Belém, Parque Estadual do Utinga, 19 Nov 210, Silva 06 (MG). Belém, on lands of IAN, 20 Jun 1944, Silva 243 (IAN). Santarém, shore of Amazon, 15 Mar 1857, Spruce 760 (K). Belém, Fazenda Murutuct, 22-23 May 1924, Zerny s.n. (W). Porto Trombetas, 26 Apr 1987, Knowles s.n. (INPA). South of Belém near mouth of Rio Guama, 14 Aug 1986, Croat 62142 (MO). Cachoeira Porteira, 17 Jan 1991, Knowles 1696 (INPA). Oriximina, 23 Jan 2013, Koch 528 (RB). Roraima: Amazonas (now Roraima), Caracarai. estrada Manaus, 10 Mar 1978, Silva 4557 (NY). Colombia: Antioquia: Municipio San Carlos, Corregimiento Norte del Samana, 12 Nov 1989, Escobar 8835 (HUA). Meta: Municipio de Restrepo, Sector Mirador, 23 Jan 2018, Passiflora acuminata DC. and its synonyms 103 Ocampo 48 (CUVC). French Guiana: Monts Tumuc Humac, 30 Aug 1972, De- eranville 1430 (CAY). Guyana: Demarara-Mahaica region: Along Linden-Soesdyke Highway, 7 Dec 1986, Pipoly 9142 (US). Berbice savanna near Takama Army Base, 14 Aug 1993, Henkel 2520 (NY). Cuyuni-Mazaruni; Kako River, 12 May 12 2009, Wurdack 4954 (NY). Upper Demerara-Berbice region: Linden-Soesdyke Highway, between Dora and Maibia Creek, 21 Jan 1987, Pipoly 9707 (US). Previously assigned to Passiflora tolimana Colombia: El Valle: La Cumbre, 2000 meters, 14-19 May 1922, Pennell 5754 (N), Killip 5594 (N), 1 Oct 1922, Killip 11679 (N). Guaviare: 1844, Goudot 10 (P). Santander: Municipio de Zapatoca, vereda La Cacica, Reserva La Montafia Magica “El Poleo”, parte alta, 28 Jul 2014, Diaz 829 (MEDEL). Tolima: Dolores, 23 Jan 1886, Lehmann 6060 (K). Ecuador: Sucumbios: Shushufundi, 4 Dec 2014, Rome 538 (LYJB). Previously assigned to Passiflora gleasonii Brazil: Acre: Municipality of Rio Branco, road to Xapuri, 24 Oct 1980, Cid 3035 (NY). Colombia: Guaviare: Municipality of San José del Guaviare, Inspeccién de Puerto Arturo, 25 Aug 1995, Cardenas 6533 (COL). Guainia: Maimachi, Serrania del Naquén, 11 Apr 1993, Madrifan 1014 (MO). Guyana: Pomeroon district, Pomeroon River, 14 Jan 1923, De La Cruz 2963 (GH). Essequibo River: Moraballi Creek, 14 Sept 1929, Sandwith 254 (K). Peru: Pasco: Oxapampa, 7 Jul 2003, Werff 18100 (MO). Venezuela: Amazonas: Department of Rio Negro, 9 Mar 1984, Liesner 16501 (MO). Tachira: East of Ayari, 7 Nov 1979, Steyermark 119472 (MO), Cerro de Cuite, along Quebrada La Colorada, 8 Nov 1979, Steyermark 119657 (MO), Cerro of Cuchilla, La Pabellana, 6 Nov 1979, Steyermark 119423 (MO). Zulia: 6 km east north-east of Rio de Oro, 28 Mar 1982, Liesner 13280 (MO). Previously assigned to Passiflora metae Colombia: Meta: Municipality of Villavicencio: road Cao Pendejo, 541 m, 23 Dec 2013, (fl), M. Bonilla, J. Mosquera, K. Pulido & A. Cajar 167 (Holotype CUVC)). Previously assigned to Passiflora ambigua Colombia: Santander: entre Duitama y Virolin, 6 Nov 1979, Escobar 3056 (HUA). Previously assigned to Passiflora laurifolia Brazil: Amazonas: Manaus; reserva campina - BR174 km 45, Benson 8282 (NY). Ig- arapé Ipiranga, 4 Jul 1993, Ribeiro 997 (INPA). Estrada para Igarapé do Tinga, 7 Aug 104 Maxime Rome et al. / PhytoKeys 201: 99-122 (2022) 1996, Sothers 883 (INPA). Para: Cachoeira Porteira, 16 Jan 1991, Knowles 1687 (INPA). Mineragao Rio do Norte, Porto Trombetas, Mina Saraca, 6 Jun 1999, Mi- randa 397 (NY). Colombia: Caqueta: Orillas del Rio Caguan, 12 Apr 1953, Romero- Castaneda 3963 (COL). Cundinamarca: Ubala B. Inspeccidn de Policia San Pedro de Jagua, Vereda Soya, 5 Jul 1998, Fernandez 16513 (RB). Meta: Cordillera de la Macarena, 30 Dec 1950, Idrobo 868 (COL). Previously assigned to Passiflora nitida Brazil: Para: Municipio de Oriximina, Rio Trombetas, 11 Jul 1980, Cid 1461 (INPA). Colombia: Antioquia: San Carlos, corregimiento El Jordan, 29 Sept 1989, Velasquez 241 (HUA). San Carlos, Vereda Patio Bonito, Alto El Cerrén, 17 Feb 1998, Correa 85 (HUA). Caqueta: Municipio de Doncello, vereda de Buena Vista, 21 May 2003, Castano 1649 (COL). Santander: Barbosa, bosque en BellaVista, 24 Nov 1940, Pérez Arbeldez 8124 (COL). Previously assigned to Passiflora riparia Ecuador: Morona Santiago: ca. 32.5 km S of Gualaquiza on road to Zamora, 4 Feb 1984, Knapp 6242 (QCNE). Napo: Archidona, 27 Jun 1968, Holm-Nielsen 1040 (AAU). Previously assigned to Passiflora aff. guazumifolia Colombia: Boyaca: Municipio Pajarito, corregimiento de Corinto, 16 Oct 1967, Lozano 935 (COL). Previously assigned to Passiflora sp. Peru: Madre de Dios: Manu, Aguas Calientes, 13 May 1984, Knapp 6441 (K). Geolocalized data collected by botanists and/or on iNaturalist Passiflora acuminata Brazil: Acre: Porto Acre (D. S. Menezes, https://www.inaturalist.org/observa- tions/41238406). Amazonas: BR174, 1.5 km N of Presidente Figueiredo (A. Adair). AM-240, 1 km S of Vila de Balbina (A. Adair). 5 km N of Oriximina (A. Adair). 1 km NE of Obidos along road (A. Adair). Rodovia Eng. Fernando Guilhon, Santarém (A. Adair). Para: Santa Barbara do Para (B. Ferreira, https://www.inaturalist.org/observa- tions/34108563 / L. FE Matos (https://www.inaturalist.org/observations/805 18762). Colombia: Antioquia: Municipio de Remedios, corregimiento La Fragua (J. Res- trepo, — https://www.flickr.com/photos/220 12266@N02/9906490006/in/photostre- Passiflora acuminata DC. and its synonyms 105 am/). Meta: Villavicencio, Reserva Forestal Vanguardia (J. Ocampo P.). Putumayo: Puerto Leguizamo (M. Molinari & M. Wettges). Ecuador: Zamora-Chinchipe: Yantzatza (A. M. Hualpa Erazo, https://www.inaturalist.org/observations/62721105). Venezuela: Bolivar: La Escalera (M. L. Watson, https://www.inaturalist.org/obser- vations/41457638). Mérida: La Blanca (M. Molinari). Guayabones (M. Molinari). Tachira: Santo Domingo airport (M. Molinari). Colén-San Felix road (M. Molinari). Passiflora tolimana Colombia: Cundinamarca: (N. B. Uribe, https://www.inaturalist.org/observa- tions/65638815). Passiflora metae Colombia: Cundinamarca: El Colegio (geolocalization from the photographer, H. Svoboda) https://uk.inaturalist.org/photos/). Meta: Buenavista, E of Villavicencio (G. S. Castro, https://www.inaturalist.org/observations/7356149). Analyses of distribution and habitat After the analyses of descriptions and reference materials showed that the four spe- cies under study could not be distinguished on a morphological basis, a geographical database was constituted, including the localizations of 78 herbarium specimens, pho- tographs and observations listed above. ‘These sites were mapped and a global distribu- tion model was developed, using the MAXENT 3.4.1 software and 19 bioclimatic var- iable layers from the Worldclim 2.1 database at a 2’30” grid resolution (corresponding roughly to 4.4 x 4.6 km at Equator; https://www.worldclim.org/data/worldclim21. html; Fick and Hijmans 2017). MAXENT identifies potential distribution areas based on their similarity in climatic conditions compared to those at the sites where the spe- cies has already been observed, hence modeling where conditions are suitable for their development. It infers the probability distribution of maximum entropy (i.e., closest to uniform) subject to the constraint that the expected value of each environmental vari- able (or its transform and/or interactions) under this estimated distribution matches its empirical average (Phillips et al. 2006). A logistic threshold value equivalent to the 10" percentile training presence was retained to separate climatically favorable areas from marginally fit areas. Thresholds of 33 and 67% training presence were used to discriminate “very good” and “excellent” climates. Furthermore, those bioclimatic co- variates that most contributed to the model were extracted for the collection sites, and submitted to a Principal Component Analysis to characterize the bioclimatic spaces corresponding respectively to the previous assignations of specimens. Three environmental parameters were extracted from specimen labels and/or observed from Google Earth (when highly precise geographical coordinates, from direct GPS meas- ures, were available): proximity of water courses, white sands, and degree of perturbation. 106 Maxime Rome etal. / PhytoKeys 201: 99-122 (2022) Results and discussion Comparative analysis of descriptions The comparison between the descriptions of P acuminata and the other three taxa is presented in Table 1. The color and pubescence of bracts, and the pubescence of sepals are not given in the table because the descriptions are incomplete regarding these cri- teria. These traits will be treated in the analysis of herbarium specimens. ‘The series of filaments appear banded with white and purple in all descriptions. As P tolimana and P. gleasonii descriptions were based on herbarium materials, they lack information on perianth color. In the case of P tolimana, the description underes- timates the perianth size and lacks the size of the bracts and the operculum. As shown in Table 1, the comparison of the different descriptions allows no clear distinctions that could justify the consideration of different species. Stems are rounded to angulate, which is the commonest situation in passion flowers. Stipules appear to be short, as compared to the common size in Laurifoliae, which is likely related to their ob- servation on dry materials, and slightly longer for P metae, but the information on stipule shape and width or with the scale given in the drawing is very inconsistent for this taxon, hampering any particular conclusion. Petioles are short too, under 2 cm, and their glands are placed close to apex, with the relative exception of Killip’s description of P tolimana. For leaves and bracts, their variation in size, shape and distribution of marginal glands is negligible, falling well within the range of variation observed in other widespread species of series Laurifoliae (e.g. P riparia; Rome and Coppens d’Eeckenbrugge 2019). Similarly, taking into account the effect of desiccation on herbarium descriptions, no taxa show a distinct range of variation for sepals, petals, and corona filaments. Kil- lip’s description of P gleasonii is unique in mentioning no awn on sepals, which would be exceptional for the whole subgenus Passiflora; and it is contradicted by Tillet’s de- scription. In fact, minute awns can be seen on sepals of the isotypes conserved at the Gray herbarium and the Missouri Botanical Garden (897955). The respective lengths given for the corona elements clearly indicate that the two outer series of filaments are longer than sepals and petals, an observation that was reported only in Harm’s descrip- tion of P tolimana. The description of the innermost series of filaments, intermediate in size, is not explicit in the descriptions of Harms and Killip, however these authors mention their possible fusion at base, which implies that they are not as short as, and denser than, those intermediate series with few filaments 1-2 mm long. Indeed, the analysis of the P gleasonii holotype shows that the operculum described by Killip is a fourth series of striped filaments, more or less fused. Apart from this confusion in Killip’s description, no significant variation has been reported for the operculum and limen. The mention of a 6 mm-long limen by Bonilla Morales et al. (2016) is not sup- ported by the longitudinal flower section drawn and photographed in their paper. In fact, the limen is reduced to a ring widening the trochlea, as in the description of Tillett (2003). Finally, the only potentially significant difference is the report of a glabrous ovary for P metae in the last row of Table 1. Passiflora acuminata DC. and its synonyms 107 Table |. Comparison of published descriptions of P acuminata, P gleasonii, P metae, and P. tolimana. Killip’s description stems terete or subangular stipules | length (mm) 4-6 4-6 8 P. acuminata | P. acuminata Tillet’s description P. tolimana Harms’ original description rounded to subangular P. tolimana Killip’s description stem purplish, 5-angled P. gleasonii Killip’s original description P. gleasonii Tillett’s description P. metae original description of Bonilla et al. terete, striate, glabrous 13 2: linear-falcate | narrow-linear,| linear, upper linear, setaceous linear, falcate, falcate part slightly glandular- glandular broader and __| serrate toward irregularly apex glandular-serrate petiole 1-1.5 1.1-1.3 1-1.5 1.7-2 stipe of - sessile sessile or sessile glands sligthly stipitate gland at apex at apex at apex in upper half | below apex 2-5 mm at apex position below apex leaves size (cm) 7-14 x 2-5 7-16 x 2-8 10-12 x 3-4 10-20 x 3-10 up to 16x9 9-16 x 4.4-9 | 6-19 x 2.2-5.5 apex tapering acute or abruptly abruptly obtuse, cuspidate gradually to an abruptly acuminate, acuminate abruptly acute apex acuminate, mucronate acuminate mucronate base rounded or acute, narrowed at | truncate at base] truncate to cuneate acutish rounded, or base shallowly subcordate cordate margin - entire or entire or remotely and | subentire to glandular very finely remotely and shallowly shallowly denticulate- obscurely glandular- glandular- serrate denticulate serrulate, or serrulate- subentire dentate shape lanceolate lanceolate- oblong- oblong- oblong- elliptic or oblong- ovate or lanceolate lanceolate lanceolate to lanceolate oblong- lanceolate- lanceolate ovate peduncle] length (cm) 3-4 1-5 2.5-3 up to 3 up to 5 - 3-3.5 bracts size(mm) | 25-40 x 10-15 | 15-40 x 5-15 s 25-35x 20 | about 20x 4-5 | 20-35x 4-14] 15-20 x 4.5-5 obtuse at apex }| narrowly spatulate- oblong-elliptic, oblong- ensiform elliptic to obovate to cuspidate, elliptic, base oblong oblong with an acuminate narrowed, acute apex apex rounded, abruptly cuspidate- acuminate margin entire or | crenate-serrate glandular | glandular-serrate | basal half with glandular glandular | with 2-3 glands serrate at apex large marginal glands, finely serrate sepals 20-30 x 6-10 about 20 15-20 x 7 30-35 x 10 30-35 x 10 30-32 x 9-10 narrow oblong lanceolate, lanceolate, lanceolate, lanceolate obtuse obtuse round at apex whitish green greenish white - - adaxially green, with purple without, light abaxially white spots violet within sub-apical yes yes yes awn petals size (mm) about 15 mm |} about 15 mm| similar to sepals | similar to 20x5 20x5 30-32 x 7-8 long long or narrower __| sepals, sligthly narrower color - white or - light violet - - white lavender 108 Maxime Rome et al. / PhytoKeys 201: 99-122 (2022) P. acuminata | Pacuminata| P.tolimana | P.tolimana | P.gleasonii | P. gleasonii P. metae Killip’s Tillets | Harms’ original] _Killip’s Killip’s original| —_Tillett’s original description | description | description description description description | description of Bonilla et al. series of | number of 5 5 or 6 - 4or5 3 3 5 filaments series outer series 30-40 25-40 slightly longer subequal; 38-40 length (mm) than petals slighlty exceeding petals inner series 2 series with | 3 or 4 series, shorter than inner one series linear, | linear, 1-2 3d series 2-3 few filaments, the 2 or 3 outer series, filaments 1mm mm mm, 4th series setaceous, outer few, the innest much shorter, 0.8-1 mm, barely 1 mm_ | setaceous, less | sometimes fused | united at base innermost series long, innermost] than 1-2 mm, 7.2-7.5 mm, filaments innermost inclined towards subulate, subulate, androgynophore 5-6 mm 5-7 mm oper- length (mm) 4-5 4-5 3 culum membranous, incurved, membranous, | membranous, | membranous, | cleft to base in] membranous, slithtly fimbriate | inflexed at base, | inflexed at cleft to the segments to 3 strongly incurved, margin erect, | base, margin | base into linear | mm wide | incurved, margin minutely short-toothed | erect, short- | segments nearly fimbriate, white fimbriate toothed 3 mm wide at base, red at apex limen tubular, closely thick base with annular, borne close 6 mm long surrounding two rings, the closely to base of base of first above base | surrounding | gynophore, gynophore of receptacle, | the base of the} barely 2mm the second 1.5 | gynophore, | high, denticulate mm from the | about 12 mm first long, bearing a thickened, annular process about 1.5 mm above its base ovary ovoid, minutely ovoid, tomentose, ellipsoidal, ovoid, finely ovoid, finely | ovary 5-6 x 3 puberulent minutely | elliptic, about 5 short- ferruginous- | ferruginous- | mm, ellipsoid, puberulent mm long tomentose tomentullous velutinous glabrous, yellowish green fruit globose to unknown ovoid, 7 x 3.5 cm Analysis of specimens associated to the descriptions Table 2 presents the 15 reference specimens cited in the descriptions of P acuminata, P. gleasonii, P tolimana and P metae that could be examined. Six of them were sterile or in buds. The type of P acuminata shows a very degraded flower. Thus, while we perceive the presence of two equal outer series of filiform filaments, as well as pubescent bracts with glands, it is impossible to observe the pubescence of the ovary and the presence of the inner series of filaments. 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