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Research Article
A new species of Piper (Piperaceae) with peltate leaves from Serranía de las Quinchas, Colombia
expand article infoM. Alejandra Jaramillo‡, Dayro Rodríguez-Duque§, Magda Escobar-Alba§
‡ Universidad Militar Nueva Granada, Cajicá, Colombia
§ Universidad Pedagógica y Tecnológica de Colombia-Universidad Nacional de Colombia, Tunja, Colombia

Corresponding author: M. Alejandra Jaramillo ( maria.jaramillo@unimilitar.edu.co )

Academic editor: Elton John de Lirio
© 2023 M. Alejandra Jaramillo, Dayro Rodríguez-Duque, Magda Escobar-Alba.
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.
Citation: Jaramillo MA, Rodríguez-Duque D, Escobar-Alba M (2023) A new species of Piper (Piperaceae) with peltate leaves from Serranía de las Quinchas, Colombia. PhytoKeys 227: 9-24. https://doi.org/10.3897/phytokeys.227.101405
Open Access

Abstract

Piper quinchasense is described and illustrated as a new species occurring in the understory of wet montane forest of the middle Magdalena Valley in Colombia, the easternmost portion of the Chocó Region. Its relationships are discussed with related taxa from the Macrostachys clade. An identification key for 35 Neotropical Piper species with peltate leaves is provided.

Resumen

Piper quinchasense se describe e ilustra como una nueva especie que ocurre en el sotobosque de bosques húmedos montanos del valle medio del Magdalena en Colombia, la porción más oriental del Chocó Biogeográfico. Se discuten sus relaciones con otras especies del clado Macrostachys. Se presenta una clave de identificación para 35 especies de Piper Neotropical con hojas peltadas.

Key words

Boyacá, Chocó Region, Macrostachys clade, Piperales, tropical montane forests

Palabras claves

bosque montano tropical, Boyacá, Chocó Biogeográfico, clado Macrostachys, Piperales

Introduction

Piper, with more than 2000 species (Callejas-Posada 2020), is one of the most species-rich genera among flowering plants (Frodin 2004). Piper is also a common element in the understory of Neotropical forests (Gentry 1990; Draper et al. 2021). Species of Piper are a key resource for bats of the genus Carollia (Fleming 2004; Bohlender et al. 2018; Yohe et al. 2021), and they are a critical element that supports diverse trophic networks that involve moths and parasitoid wasps (Wilson et al. 2012; Slinn et al. 2018). Piper’s enormous diversity of secondary metabolites is critical for their coexistence (Salazar et al. 2016a, 2016b) and diversification (Massad et al. 2022). And it is also of immense interest in the pharmaceutical industry (Perez Gutierrez et al. 2013; Salehi et al. 2019). This genus is, without doubt, an essential structural and trophic element of the understory and lower strata of Neotropical forests (Salazar et al. 2016b; Draper et al. 2021).

Molecular phylogenetic studies have been instrumental in reviving the infrageneric classification of the genus, but also identifying convergence in morphological traits (Jaramillo and Callejas-Posada 2004; Trujillo et al. 2022). Molecular phylogenetics (Jaramillo and Manos 2001; Jaramillo et al. 2008) have validated the monophyly of groupings proposed in the mid-1800s (Kunth 1839; Miquel 1843). The infrageneric classification was not used during the 1900s, probably as the expeditions in the early 1900s produced too many species to classify. Today we know that Piper clades and subclades are easy to recognize with a combination of key morphological characters (Jaramillo et al. 2008). A formal infrageneric classification of Piper based on phylogeny is under preparation (Callejas pers. comm.). Molecular phylogenetics have been instrumental in clarifying the relationships of Piper species with “axillary” inflorescences, a polyphyletic set of taxa from the Chocó Region that were wrongly merged in the genus Trianaeopiper Trel. (Trelease 1928). Molecular phylogenetics demonstrated that Trianaeopiper is not monophyletic, revealing that the diagnostic character axillary inflorescence – which are shortened sympodial branches, is convergent (Jaramillo and Callejas-Posada 2004). Similarly, molecular phylogenetics have further supported that Piper species with peltate leaves are part of at least four Neotropical Piper clades (Trujillo et al. 2022). Piper classification, evolution and ecological studies have been greatly enriched with molecular phylogenetics.

Identifying Piper species continues to be difficult for the untrained eye, and many taxa are submerged in a few large, broadly distributed, but artificial taxa (Tebbs 1990, 1993). To the trained specialist, new Piper species are a common finding in the tropical forests, or even in the undetermined piles in the most important herbaria around the world. Additionally, molecular phylogenetics have confirmed the clade (subgeneric) affiliation of new species (Bornstein et al. 2014; Tepe et al. 2014; Trujillo et al. 2022). During our exploration to Serranía de las Quinchas we have identified a few undescribed species, one of the stands-out because it has peltate leaves, a characteristic rarely observed in the genus.

“Serranía de las Quinchas” is a small mountain spur west of the Cordillera Oriental in the middle Magdalena Valley, in Colombia. The region’s flora is particularly interesting as it combines its own floristic elements, mixed with taxa from Mesoamerica, the Chocó Region, and Amazonia (Balcázar-Vargas et al. 2000). Additionally, various endemic species and genera occur in the area (Rodríguez-Duque et al. 2021). The middle Magdalena Valley deserves close examination and more fieldwork to uncover the history of its biological richness. For centuries, the forest has been preserved because of its scarp ridges and high rainfall. However, the construction of a pipeline and road in the 1990s opened the area to colonization (Stiles and Bohórquez 2000). Today the region is threatened by the extension of agriculture and mining activities. Fortunately, Serranía de las Quinchas Regional Park was created in 2008 by Corpoboyacá (the regional environmental protection agency) to protect the remaining forest (Bohorquez-Osorio et al. 2020). Despite the deforestation in the region, new species are still being encountered. Here we describe a new species of Neotropical Piper with peltate leaves that we detected during our expeditions in this interesting location.

Materials and methods

Specimens were collected in Serranía de las Quinchas, located in the middle Magdalena Valley in Colombia, in the department of Boyacá. The sites visited range from 800–1200 m. a. s. l., the locality is dominated by humid montane forest (Fig. 1). Annual rainfall is 3333 mm on average; two dry seasons occur in January-February and June-August, and the average temperature is 26 °C ( Instituto de Hidrología, Meteorología y Estudios Ambientales, IDEAM). Detailed observations in the field, combined with examination of available herbarium collections, allowed the description of growth habits and phenological stages. Voucher specimens were deposited in the following herbaria HUA, H-UPTC, and UMNG-H (herbarium acronyms according to Index Herbariorum [Thiers continuously updated], and Instituto Humboldt – Red Nacional de Colecciones [http://rnc.humboldt.org.co/wp/]). In addition, taxonomic literature on Piper was examined (Trelease and Yuncker 1950, Steyermark 1984, Callejas-Posada 2020). The measurements included in the description below are based on herbarium specimens collected by the authors. Conservation status assessments employ the categories and criteria of the IUCN (2012, 2022). We calculated the extent of occurrence and area of occupancy using the R package ConR (Dauby et al. 2017).

Figure 1. 

Geographic distribution map of Piper quinchasense M.A.Jaram.

For preparing the key of Neotropical Piper species with peltate leaves we used the literature (Trelease and Yuncker 1950, Steyermark 1984, Callejas-Posada 2020, Carvalho-Silva et al. 2022) and type specimens were examined using digitized plant specimens available on-line from JSTOR Global Plants (https://plants.jstor.org/).

We extracted DNA from silica gel dried tissue, using the DNAeasy plant mini kit (Qiagen, Valencia, California, USA). The ITS region was amplified using one of two pairs of primers ITS5-ITS4, or LEU1-ITS4 (Baldwin 1992). Sequencing was contracted with GenCore (Universidad de los Andes, Bogotá, Colombia). The resulting sequences (Table 1) were manually aligned against previously obtained alignments (Jaramillo et al. 2008). We selected 43 ITS sequences from the large alignment to portray here. Three sequences from Piper species from Asia and the South Pacific were used as outgroups. Forty sequences of Neotropical Piper, comprising representatives of all clades, and 14 species with peltate leaves were selected. Maximum Likelihood (ML) phylogenetic and bootstrap (100 replicates) analyses were conducted using RAxML (Stamatakis 2014).

Table 1.
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Genbank accessions for the new species of Piper.

Taxon Voucher Genbank Accession #
Piper quinchasense M. A. Jaram. MAJ1807 OQ354973
MAJ1939 OQ354974

Taxonomic treatment

Piper quinchasense M.A.Jaram., sp. nov.

urn:lsid:ipni.org:names:77320207-1

Type

Colombia. Boyacá: Otanche, vereda Las Quinchas, Sector La Y, Finca Lote Terreno, 5°48'17"N, 75°15′24"W, 1210 m, 17 Mayo 2022 [fl], M. A. Jaramillo et al. 1807 (holotype: HUA; isotypes: UPTC, UMNG-H). Figs 1–3.

Description

Piper quinchasense is similar to P. parianum, it differs from the latter in having all leaf blades peltate (vs. leaves deeply lobed to peltate), and inflorescence peduncle 4–5 cm long, (vs. peduncle 1–2.7 cm long).

Shrub , 3 m tall, branched in the upper portion only, exhibiting stilt roots (Fig. 2C). Internodes 5–10 × 2.5–4.6 cm, green when young and becoming brown when maturing, tomentose, idioblasts not evident. Prophylls not seen. Petioles 4.5–8.5 cm long, vaginate the entire length (Figs 2D, 3J), tomentose, idioblasts evident. Leaf blades (28) 35–49 × 7.5–19 cm, oblong-lanceolate, base obliquely-peltate, asymmetric to truncate, peltate, petiole inserting 2.5–10.5 cm from the margin, on the adaxial surface leaf blade is depressed and umbonate above petiole insertion (Fig. 2E), blade medially asymmetric, apex long attenuate, green on the adaxial surface and green-silver on the abaxial surface when alive (Fig. 2E, F), coriaceous, chartaceous when dry, drying dark maroon on the adaxial surface and ochre on the abaxial surface, glabrous with visible idioblasts on the adaxial surface (Fig. 3K), sparsely tomentose on the blade and veins densely tomentose on the abaxial surface (Fig. 3I), eciliate, margin folds towards the abaxial surface appearing to form irregular spaced teeth; pinnately nerved to the distal third, 4–6 pairs of secondary veins, curved and ascending, diverging in angles that decrease towards the apex (from 80–30 degrees) and spacing slightly decreasing towards the base, tertiary veins forming areoles 1.2–1.8 × 0.5–1 cm, rectangular and not uniform in size, perpendicular to secondary nerves, nerves not impressed on the adaxial surface, elevated on the abaxial surface. Inflorescences a simple spike, terminal, pendulous; peduncle 4.5–5 cm long, tomentose, green, idioblasts not evident; rachis length in flower 12–19 cm × 3–4 mm, rachis length in fruit 25–27cm. Floral bracts cucullate, sagitate from above (Fig. 3F), 1.8–2.3 × 0.9–1.0 mm, pellucid dotted, pedicel fimbriate on the distal portion (Fig. 3E), forming bands around the spike (Fig. 3B). Flowers sessile with 4 stamens (Fig. 3C), filaments 0.25–0.40 mm long, anthers 0.3–0.6 × 0.16–0.26 long, with connective glabrate, longitudinally dehiscent, dithecous (Fig. 3D), ovary four carpellate, four stigmas sessile, not persistent in fruit, 0.4–0.5 mm long (Figs 2B, 3G). Fruits obpyriform, 2.1–2.4 × 1.0–1.6 mm, glabrous, green when alive, brown when dry (Fig. 3G). Seeds smooth, pellucid dotted, obpyriform, 1.9–2.1 × 1.0–1.3 mm, glabrous, brown when dry (Fig. 3H).

Figure 2. 

Piper quinchasense M. A. Jaram. A habit B magnified view of inflorescence C stilt roots D sheathing petiole E adaxial surface of leaves F abaxial surface of leaves and inflorescence. Photographs by D. Rodríguez-Duque.

Figure 3. 

Piper quinchasense M. A. Jaram. A sympodial branch, showing both the abaxial and adaxial surface of the leaves B magnified view of inflorescence C floral diagram D anther E floral bract, abaxial view F floral bract view from above G fruit H seed I magnified view of leaf abaxial surface J sheathing petiole K magnified view of leaf adaxial surface. Illustration by Ariadna Valenzuela, based on M. Escobar-Alba 764, and photographs by D. Rodríguez-Duque.

Phylogenetic relationships

Piper quinchasense belongs to the Macrostachys clade (Fig. 4). A group of shrubs and treelets reaching 7–8 (15) m tall, petioles sheathing above the middle or throughout their length, pinnately nerved leaves, mostly larger than 30cm long, inflorescences mostly pendulous (erect in some species), and flowers forming bands around the spike (Jaramillo et al. 2008). Species with peltate leaves have evolved independently in clades Macrostachys, Pothomorphe, Oxodium (=Schilleria, Callejas 2020) and Ottonia. Peltate leaves are known to occur in the two large genera of Piperaceae: Piper and Peperomia. They are more common in plants that grow in the shade of humid tropical forests (Wunnenberg et al. 2021). There is still much to learn about the functional morphology and anatomy of Piper species with peltate leaves.

Figure 4. 

Phylogenetic relationships of Piper quinchasense M. A. Jaram. and 14 species of Neotropical Piper with peltate leaves (in bold). Topology based on Maximum likelihood analysis of nrITS sequences. Numbers on branches are maximum likelihood bootstrap support values (>90%).

Distribution and habitat

The species is only known from the type locality Serranía de las Quinchas (Fig. 1). Piper quinchasense is a tall shrub in the understory of very humid forests; it often grows near streams. The occurrence of stilt roots (Fig. 2C), a character not commonly seen in Piper, suggests it is adapted for flooded areas near watercourses. It also is a very resilient plant that will produce adventitious roots if tumbled and it will continue growing, or even resprout, from a fallen leaf. Resprouting is common among shade-tolerant Piper (Lasso et al. 2009).

Phenology

Flowering specimens were collected in March, and May. Fruiting specimens were collected in October.

Etymology

The epithet quinchasense, refers to Serranía de las Quinchas, the type locality for this species. According to locals, Quinchas derives from the indigenous groups “Quinchos” that inhabited the region.

Conservation status

This species is known only from one population in the type locality (Fig. 1). The extent of occurrence (EOO) of 8 km2 and area of occupancy (AOO) of 8 km2 are very small. The locality is under threat of disappearing for the extension of agricultural and mining activities (Rodríguez-Duque et al. 2021), which suggests it is Endangered [EN B1a]. Serranía de las Quinchas was declared a Regional Park in 2008; since its creation, the park extension has been reduced by 20% as the buffer zones were eliminated. Furthermore, the constant threat of coal mining makes the locality highly exposed to deforestation.

Additional specimens examined

Colombia. —Boyacá: Otanche, vereda Las Quinchas, Sector La Y, Finca Lote Terreno, 5°48'17"N, 75°15′24"W, 26 October 2022, [fr] M. A. Jaramillo et al 1939 (HUA, UMNG-H); Boyacá: Otanche, Parque Regional Natural, Serranía de las Quinchas, 5°48′45.5"N, 75°15′22.2"W, [st] 14 June 2021, Magda Escobar-Alba et al. 489 (UPTC); Parque Regional Natural Serranía de las Quinchas, vereda las Quinchas, Finca Chorro Negro, 5°49′7.2"N, 75°14′57.3"W, [fl] 3 March 2022, Magda Escobar-Alba et al. 762 (UPTC).

Notes

Piper quinchasense is a handsome species that differs from related Macrostachys taxa in having long lanceolate-oblong leaves. It is morphologically similar to P. parianum from which it differs in having mononomorphic leaves (all of them peltate) vs. leaves dimorphic, some peltate and others are deeply lobed. P. parianum is only known in the isolated cloud forests of “Peninsula de Paria”, a region located in the eastern portion of Coastal Cordillera in the extreme northeast of Venezuela. The flora of Paria Peninsula is characterized by the high occurrence of endemic species (Steyermark and Agostini 1966; Meier 2011) and several types of vegetation communities (Beard 1946). The flora of this region has many affinities to plants of Trinidad and Tobago (Selaginella hartii Hieron.; – Selaginellaceae; 5 Turrill – Acanthaceae: sensu Baksh-comeau et al. 2016), and the forests of the Northeastern sector of Guayana Shield in Venezuela and the Guianas (e.g., Besleria insolita C.V. Morton – Gesneriaceae; Meier 2011). Serranía de las Quinchas, on the other hand, has the most floristic affinities with the Chocó region (e. g. Dimerocostus cryptocalyx N. R. Salinas & Betancur, Salinas and Betancur 2004). The regions have similar habitats driven by high precipitations despite their divergent biogeographical affinities.

A key to species of Neotropical Piper with peltate leaves is presented below.

New status

While preparing the key for peltate Neotropical Piper, we realized some species deserve new status. New status and new names are proposed for two species.

Piper neovenezuelense M.A.Jaram., stat. nov et, nom. nov.

urn:lsid:ipni.org:names:77320208-1

Basionym

Piper veraguense C. DC. var. venezuelense Steyerm. Fl. Venez. 2(2): 590 (1984). Type: Venezuela, Edo. Trujillo, 16 km de Boconó a lo largo de la carretera a Biscucuy, 1850 m, 11 aug 1964, F. Breteler 4082 (Holotype; VEN; Isotype: MER). Non Piper venezuelense C.DC., J. Bot. 4: 216 (1866).

Note

The epithet P. neovenezuelense is proposed here to replace Piper veraguense C. DC. var. venezuelense Steyerm. Because Piper venezuelense C. DC. (De Candolle 1866: 216) is already in use.

Etymology

The new epithet neovenezuelense honors the intention of J. Steyermark to highlight the occurrence of this species in Venezuela.

Steyermark (1984) synonymized under P. veraguense C. DC. various distinct species: P. albert-smithii Trel. & Yunck., and P. mutisii Trel. & Yunck., and proposed that P. veraguense C. DC. has three varieties: P. veraguense var. veraguense, P. veraguense var. mutisii (Trel. & Yunck.) Steyermark, and he added P. veraguense var. venezuelense Steyermark (Steyermark 1984). These species are similar in their overall morphology and potentially form a species complex. Unfortunately, we do not have sequence data to test the latter hypothesis. Here we consider them separate species. P. veraguense has glabrous leaves with nerves puberulent on the abaxial surface, not pellucid dotted (Callejas-Posada 2020); P. albert-smithii is distinct because it has fleshy warts below the nodes, and leaves are glabrous (Trelease and Yuncker 1950); and P. mutisii has fleshy warts below the nodes, nerves are dense velvety puberulent on abaxial surface; P. neovenezuelense has glabrous internodes without warty outgrowths, and nerves pilose on abaxial surface.

Piper andersii M.A.Jaram., stat. nov. et, nom. nov.

urn:lsid:ipni.org:names:77320209-1

Piper mikanianum Steud. var. peltatum Yunck., Bol. Inst. Bot. (São Paulo) no. 3: 54 (1966). Type: Brazil, Minas Gerais, Caldas, A. F. Regnell II 256*, 9 Jul 1864. Non Piper peltatum L.

Note

The epithet P. andersii is proposed here to replace Piper mikanianum Steud. var. peltatum Yunck. because Piper peltatum L., (Sp. Pl. 1: 30 1753) is already in use.

Etymology

The new epithet andersii honors Anders Fredrik Regnell (1807—1884), Swedish physician and botanist who established himself in Minas Gerais (Brazil) and collected the type specimen for this species.

T. Yuncker provided a key to four varieties of Piper mikanianum (Kunth) Steud: P. mikanianum (Kunth) Steud var. mikanianum; P. mikanianum f. clausum characterized by the closed sinus and overlapping lobes; P. mikanianum var. pilosius C. DC. with leaves and stems strongly pilose with hairs up to 1mm long; and P. mikanianum var. peltatum Yunck with peltate leaves and nerves minutely hirtellous on the abaxial surface (Yuncker 1972). Yuncker had mentioned that this latter variety might deserve species rank, we propose to make the change suggested and propose the new name P. andersii. Furthermore. P. mikanianum occurs in the states of Minas Gerais, Paraná and Rio Grande do Sul in Brazil and in Argentina, P. andersii, with peltate leaves has only been registered from the state of Minas Gerais.

Key to Neotropical Piper species with peltate leaves

1 Leaves subpeltate, petiole inserted slightly inside the leaf margin 2
– Leaves peltate, petiole inserted 0.5–11 cm from the leaf margin 12
2 Leaves ovate-lanceolate or elliptic, up to 18 cm long, petioles terete 3
– Leaves ovate, ovate –oblong or elliptic, more than 20 cm long, petioles sheathing 5
3 Inflorescence a raceme with glabrous to sparsely pubescent rachis 4
– Inflorescence a spike, with pubescent rachis P. klotzschianum (Kunth) C. DC.
4 Leaves elliptic and glabrous without hirtellous intramarginal nerve P. brumadinense M. Carv.-Silva & E. Guim.
– Leaves ovate, intramarginal nerve hirtellous on abaxial surface P. ovatum Vahl
5 Petioles sheathing only at the base, nerves puberulent on the abaxial surface 6
– Petioles sheathing over half their length, leaves and nerves glabrous or pubescent 7
6 Leaves 35–20 cm long, base lobate P. omega Trel.
– Leaves 10–20 cm long, base barely cordate P. marginecontinuum Callejas
7 Leaf base rounded or cordate, inflorescences erect 8
– Leaf base lobate, inflorescences pendulous 9
8 Leaf ovate, leaf base rounded, peduncle up to 0.5 cm long P. palenquense Callejas
– Leaf broadly ovate, leaf base deeply cordate, sinus open, peduncle 0.7–1 cm long P. vallicola C. DC.
9 Leaves more than 35 cm long, inflorescences 50–60 long 10
– Leaves up to 26 cm long, inflorescences up to 20 cm long 11
10 Leaves up to 50cm long, sinus closed, longer lobe overlapping the petiole, peduncle 1.5–2.7cm long P. caracasanum Bredem. ex Link
– Leaves up to 35cm long, sinus open, peduncle 7 cm long P. gualeanum C. DC.
11 Petioles shortly pubescent, inflorescences 10–20 cm long P. calcariforme Tebbs
– Petioles with trichomes forming lines, inflorescences 6–9 cm long P. hebetifolium W. C. Burger
12 Plants herbaceous, inflorescences spikes, arranged in umbels P. peltatum L.
– Shrubs, suffrutex, or climbers, inflorescences solitary racemes or spikes 13
13 Leaves 10–30 cm long, inflorescences erect 14
– Leaves 35 cm long or longer, inflorescences erect or pendulous 31
14 Lianescent vines 15
– Shrubs or suffrutex 17
15 Internodes, petioles and leaves glabrous, leaf blade ovate or oblong-elliptic, smooth 16
– Internodes, petioles and abaxial leaf surface pilose, leaf blade obovate, slightly bullate P. parmatum Dressler
16 Leaf blades with evident idioblasts on abaxial surface, spikes 1–2 cm long, with obtuse apices P. foreroi Gentry
– Leaf blades without visible idioblasts, spikes 5–6 cm long with mucronate apices P. peltifolium Callejas
17 Leaf base lobed or cordate 18
– Leaf base rounded, obtuse or scutellate 24
18 Leaves rounded –ovate, leaf base deeply cordate P. andersii M.A.Jaram., stat nov. nom nov.
– Leaves ovate, leaf base cordate or lobed 19
19 Shrubs densely crisp –villous P. copeyanum (C.DC) Trel.
– Shrubs glabrous 20
20 Idioblasts visible on both surfaces P. subscutatum (Miq.) C. DC.
– Idioblasts not visible 21
21 Petioles vaginate at the base, flowers not forming bands on the spikes 22
– Petioles vaginate half the length or more, flowers forming bands on the spikes P. maxonii C. DC.
22 Leaves 5–10 cm wide, petiole inserted ca. 1 cm from the margin P. jacaleapaense Callejas
– Leaves 15 –20 cm wide, petiole inserted 5–6 cm from the margin 23
23 Leaves broadly ovate, glabrous on both surfaces P. veraguense C. DC.
– Leaves elliptic-oblong to ovate, nerves on the abaxial surface pilose P. neovenezuelense M.A.Jaram., stat. nov. nom nov.
24 Leaves glabrous 25
– Leaves pubescent at least abaxially on veins 27
25 Leaves oblong-lanceolate, leaf base rounded P. imberbe Trel.
– Leaves ovate, leaf base obtuse or spatulate 26
26 Leaf base narrowly spatulate P. indiwasii W. Trujillo & M.A.Jaram.
– Leaf base broadly obtuse P. scutilimbum C. DC.
27 Apex short acuminate, inflorescence obtuse 28
– Apex long attenuate, inflorescences mucronate P. tuerckheimii C. DC.
28 Flowers forming bands around the spike P. hammelii Callejas
– Flowers laxly arranged in a spike or raceme 29
29 Plant glabrous, inflorescences a spike P. scutifolium Yunck.
– Villous shrub, inflorescence a raceme 30
30 Petiole inserted ca. 1 cm from margin, with visible idioblasts on leaf lamina P. cariacicaense M. Carv.-Silva & E.F.Guim.
– Petiole inserted 0.5 cm from margin, without visible idioblasts on leaf lamina P. carautensei E.F.Guim. & M. Carv.-Silva
31 Leaves broadly ovate, petiole vaginate to the middle 32
– Leaves ovate, lanceolate or oblong, petiole vaginate above the middle 33
32 Leaf base rounded, inflorescence short-apiculate P. mutisii Trel. & Yunck.
– Leaf base cordate, inflorescence obtuse P. grandilimbum C. DC.
33 Internodes warty above nodes, leaves glabrous P. albert-smithii Trel. & Yunck.
– Internodes smooth, leaves pubescent or tomentose, at least on the abaxial surface 34
34 Leaf strongly bullate P. hartwegianum (Benth.) C. DC.
– Leaves not bullate, or occasionally softly bullate 35
35 Leaves elliptic-ovate, apex acute or short acuminate 36
– Leaves oblong-lanceolate, apex gradually acuminate or long attenuate 37
36 Heteromorphic trichomes of intermixed on the abaxial surface, inflorescence up to 43 cm long P. peltilimbum Yunck.
– Trichomes of uniform length on the abaxial surface, inflorescences 50–60 cm long P. candollei Sodiro
37 Leaves 14–26 cm wide, leaf base deeply cordate sub-peltate to obliquely peltate P. parianum Yunck.
– Leaves 7.5–19 cm wide, leaf base obliquely peltate P. quinchasense M.A.Jaram., sp. nov.

Acknowledgements

We thank Juan E. Carvajal Cogollo for his assistance, Ariadna Valenzuela Zúñiga for preparing the illustration, A. F. Majin for field assistance, and Jenny Peña Varon for preparing the distribution map. We thank G. Aymard-Corredor, M. Carvalho-Silva, E. Tepe, E. J. Lirio and one anonymous reviewer for comments on previous versions of the manuscript. We thank K. Gandhi for his expert advice on nomenclatural issues regarding the new names proposed here. Special thanks to the Galvis Family: Don Lucindo, Doña Edilsa and Leidy for hosting our field expeditions in Serranía de las Quinchas. Don Lucindo’s expert guidance in the forest was key to finding the plants. This work was supported by project “BPIN No. 2020000100003 – Investigación de la biodiversidad de Boyacá: complementación y síntesis a través de gradientes altitudinales e implicaciones de su incorporación en proyectos de apropiación social de conocimiento y de efectos de cambio climático.” Funds for field expeditions were also provided by Universidad Militar Nueva Granada as part of the Plant Taxonomy and Systematics course. Gencore (Sequencing Center) at Los Andes University partially subsidized the sequencing analysis.

Additional information

Conflict of interest

No conflict of interest was declared.

Ethical statement

No ethical statement was reported.

Funding

No funding was reported.

Author contributions

Conceptualization: MAJ. Investigation: DRD, MEA. Writing – original draft: MAJ.

Author ORCIDs

M. Alejandra Jaramillo https://orcid.org/0000-0002-6539-4149

Dayro Rodríguez-Duque https://orcid.org/0000-0002-3829-3377

Magda Escobar-Alba https://orcid.org/0000-0002-3756-9132

Data availability

All of the data that support the findings of this study are available in the main text or Supplementary Information.

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