Abstract
We describe four new species of Piper from the Amazonian slopes of the northern Andes. Piperhoyoscardozii is distinguished from similar climbing species, P.dryadum and P.flagellicuspe, by its longer peduncles. The Amazonian species Piperindiwasii is distinguished from P.scutilimbum from Panama and northern Colombia by the narrowly spatulate leaf base extension. Pipernokaidoyitau is characterised by the presence of larger leaves and longer spikes than similar species, P.anonifolium and P.hostmannianum. Finally, P.velae is characterised by cordulate leaf bases in all nodes, petioles 0.8–1.5 cm long and pubescent fruits, which easily distinguish it from the related species, P.holdridgeanum.
Keywords: Colombia, Ecuador, north-western Amazon, Peru, Piperhoyoscardozii , Piperindiwasii , Pipernokaidoyitau , Pipervelae , Piperaceae
Introduction
Piper L. is the most diverse and representative genus in Piperaceae, encompassing ca. 2600 species (Callejas-Posada 2020). Piper represents an extraordinary diversification amongst early diverging angiosperms. It is a Pantropical group (Jaramillo and Manos 2001) and its highest diversity lies in the Neotropics (ca. 1800 species; fideUlloa-Ulloa et al. 2018 onwards). The growth forms of the genus are very variable and are most commonly perennial shrubs and suffrutices, growing mainly in the understorey of rainforests. Additionally, other Piper species are lianas, rarely caulescent herbs, hemi-epiphytes or trees not taller than 15 m (Callejas-Posada 2020). Piper is characterised by simple, alternate leaves and jointed stems with enlarged nodes; many species produce pearl bodies on the leaves or stems (Tepe et al. 2004), but the most distinctive morphological feature of the genus is the production of solitary spikes, upright or pendant, which contain dozens to thousands of apetalous flowers. Many Piper species are rich in essential oils, which can be found in many tissues and organs, including the fruits, seeds, leaves, branches, roots and stems (Salehi et al. 2019). Although very few species have significant global economic value, many are locally important for their use by native populations for medicinal and culinary purposes (Colvard et al. 2006; Cáceres and Kato 2014).
Piper has an extensive history of taxonomic and nomenclatural instability. The leading causes for that uncertainty are the challenges of interpreting morphological characteristics and the limited geopolitical circumscription of taxonomists’ work (Ramírez-Amezcua 2016; Callejas-Posada 2020). Piper flowers are small and morphologically homogeneous and the number of stamens and their position, characters of diagnostic importance, are challenging to see or interpret. Not all stamens develop at anthesis (Tucker 1982a, b) and the anthers, articulated at the filaments, fall off quickly after anthesis (Valentin-Silva et al. 2018). Many Piper species present foliar dimorphism and many specimens do not include leaves of both monopodial and sympodial axes. Description of new species, based on incomplete specimens, has led to the publication of superfluous names that later need to be submerged in synonymy (Ramírez-Amezcua 2016). Furthermore, because floristic treatments are often limited to specific countries, many species are given several names across borders (Ramírez-Amezcua 2016). Extensive fieldwork and comparisons with collections (including digitised collections) are essential to improve the taxonomy of this species-rich genera.
Observing species in the field is imperative to ensure detailed descriptions and complete specimens that support an excellent Piper taxonomy. Numerous essential characters cannot be included or are lost in dry specimens. Fortunately, in the last decade, W. Trujillo has collected and studied more than one thousand (1000) specimens from the Amazonian slopes of the Colombian Andes. Some of them have become type specimens of recently-described taxa (Trujillo and Callejas 2015; Trujillo and Jaramillo 2019, 2021). Continued work by the authors will bring to light more diversity in the future.
The present contribution describes four new Piper species from the Amazonian slopes of the Andes in Colombia, Ecuador and Peru. We use the nuclear ribosomal internal transcribed spacer (ITS) to determine their phylogenetic relationships within Piper. We increase to 419 the number of Piper species known from Colombia (Bernal et al. 2015).
Materials and methods
Fieldwork was conducted along the Amazonian slopes of the Andes in southern Colombia, in the Department of Caquetá, during 2010–2020. We collected silica gel-dried leaf tissue samples for DNA extractions. Detailed observations in the field and examination of available herbarium collections were used to describe growth habits and phenological stages accurately. We deposited botanical specimens in COAH, COL, HUAZ and HUA (acronyms according to Thiers 2019). Detailed comparisons with morphologically similar species allowed us to recognise the four new species. Besides reviewing the literature (Trelease 1936; Trelease and Yuncker 1950; Brako and Zarucchi 1993; Jørgensen and León-Yánez 1999; Jørgensen et al. 2014; Jardim Botânico do Rio de Janeiro 2020), we examined specimens through visits to Herbaria COAH, COL, HUA, HUAZ, GH, NY, PMA, RSA and US and digitised plant specimens available on the web (e.g. JSTOR Global Plants, https://plants.jstor.org/). Measurements included here were taken from specimens collected in Colombia. To describe leaf architecture, we used terminology proposed in the Manual of Leaf Architecture (Ellis et al. 2009). To assess conservation status, we calculated area of occupancy (AOO) and extent of occurrence (EOO) using R and the package ConR (Dauby et al. 2017).
We extracted DNA from all the new species (Table 1) using a CTAB method (Doyle and Doyle 1987). We amplified the nuclear ribosomal internal transcribed spacer (ITS) according to Jaramillo and Manos (2001) and aligned sequences using previous alignments as a guide (Jaramillo et al. 2008). First, we included the new sequences in our 900+ PiperITS alignment, to determine the relationships of the new Piper species. This preliminary analysis (not shown) served to select representatives of each major clade of Piper (Jaramillo et al. 2008) for the analysis presented here. Forty-two species of Piper, 35 from the Neotropics and seven from Asia were selected to provide comparisons with the new taxa. Maximum Likelihood (ML) phylogenetic and bootstrap (100 replicates) analyses were conducted using RAxML (Stamatakis 2014) using species from the Asian tropics to root the phylogeny.
Table 1.
GenBank accessions for new species and reference taxa. Other GenBank accessions are available in original manuscripts (Jaramillo et al. 2008). New species in bold.
| Taxon | Voucher | GenBank Accession ITS | Collection * | Publication |
|---|---|---|---|---|
| Piperabalienatum Trel. | MAJ 552 | EU581075 | DUKE | Jaramillo et al. (2008) |
| Piperabbreviatum Opiz | MAJ 203 | EU581076 | DUKE | Jaramillo et al. (2008) |
| Piperaduncum L. | MAJ 76 | AF275157 | DUKE | Jaramillo & Manos (2001) |
| Piperalatabaccum Trel. & Yunck. | AL 1177 | KJ930372 | INPA | Molina-Henao et al. (2016) |
| Piperalbispicum C. DC. | MAJ 388 | AY572317 | DUKE | Jaramillo & Callejas (2004) |
| Piperalbozonatum C. DC. | MAJ 697 | AY326195 | DUKE | Jaramillo & Callejas (2004) |
| Piperamalago L. | MAJ 561 | AF215186 | DUKE | Jaramillo & Manos (2001) |
| Piperamplum Kunth | MAJ 804 | EU581096 | RB | Jaramillo et al. (2008) |
| Piperanonifolium Kunth | AL 1242 | EU581084 | MG | Jaramillo et al. (2008) |
| EJT 527 | EU581101 | MU | Jaramillo et al. (2008) | |
| Piperarboretum Aubl. | MN 1565 | EU581106 | RB | Jaramillo et al. (2008) |
| Piperbiolleyi C DC. | CD 10896 | EU581128 | SRP | Jaramillo et al. (2008) |
| Piperbrachypodon (Benth.) C. DC. | MAJ 757 | AY326198 | DUKE | Jaramillo & Callejas (2004) |
| Piperbreviamentum C.DC. | MAJ 221 | EU581134 | DUKE | Jaramillo et al. (2008) |
| Pipercallosum Ruiz & Pav. | MJK 161 | EU581142 | SPF | Jaramillo et al. (2008) |
| Pipercandollei Sodiro | EJT 1449 | EF056237 | MU | Jaramillo et al. (2008) |
| Pipercapense L. | CD 11004 | EU581143 | SRP | Jaramillo et al. (2008) |
| Pipercavendishoides Trel. & Yunck. | MAJ 70 | AF275153 | DUKE | Jaramillo & Manos (2001) |
| Pipercinereum C. DC. | MAJ 66 | AF275190 | DUKE | Jaramillo & Manos (2001) |
| Pipercocornarum Trel. & Yunck. | RC 12493 | AY326203 | HUA | Jaramillo & Callejas (2004) |
| Pipercordulatum C. DC. | BCI-EL | EU581167 | Jaramillo et al. (2008) | |
| Piperdarienense C. DC. | MAJ 103 | AF275181 | DUKE | Jaramillo & Manos (2001) |
| Piperdilatatum Rich. | MAJ 858 | EU581179 | RB | Jaramillo et al. (2008) |
| Piperdryadum C. DC. | EJT 1047 | EU581186 | MU | Jaramillo et al. (2008) |
| Piperexcelsum G. Forst. | N. V. | EF635476 | Jaramillo et al. (2008) | |
| Piperfilistilum C. DC. | MAJ 157 | AF275155 | DUKE | Jaramillo & Manos (2001) |
| Piperflagellicuspe Trel. & Yunck. | MAJ 65 | AF275154 | DUKE | Jaramillo & Manos (2001) |
| Piperhartwegianum (Benth.) C. DC. | MAJ 781 | AY326207 | DUKE | Jaramillo & Callejas (2004) |
| Piperhispidum Sw. | MCS 304 | EU581241 | RB | Jaramillo et al. (2008) |
| Piperholdridgeanum W. C. Burger | CD 10865 | EU581247 | SRP | Jaramillo et al. (2008) |
| JF 9128 | EU581248 | MO | Jaramillo et al. (2008) | |
| Piperhostmanianum C. DC. | SM 25228 | EU581251 | NY | Jaramillo et al. (2008) |
| EJT 573 | EU581249 | MU | Jaramillo et al. (2008) | |
| Piperhoyoscardozii | WT 4127 | OK235346 | UMNG | This study |
| WT 4004 | OK235347 | UMNG | ||
| WT 4099 | OK235348 | UMNG | ||
| Piperimberbe Trel. & Yunck. | AB 983 | EU581255 | SEMO | Jaramillo et al. (2008) |
| Piperindiwasii | ET 7254 | OK235342 | JBB | This study |
| EJT 1438 | EU581390 | MU | Jaramillo et al. (2008) | |
| Pipermaxonii C. DC. | EJT 370 | EF056270 | MU | Jaramillo et al. (2008) |
| Pipermultiplinervium C. DC. | MAJ 139 | AF275168 | DUKE | Jaramillo & Manos (2001) |
| Pipernokaidoyitau | WT 4171 | OK235339 | COAH | This study |
| WT 4258 | OK235340 | COAH | ||
| ET 7249 | OK235341 | COAH | ||
| Pipernovogranatense C. DC. | MAJ 71 | EU581317 | DUKE | Jaramillo et al. (2008) |
| Piperpeltatum L. | MAJ 564 | EU581335 | DUKE | Jaramillo et al. (2008) |
| Piperretrofractum Vahl. | MAJ 395 | AF275196 | DUKE | Jaramillo & Manos (2001) |
| Pipersanctum (Miq.) Schltdl. ex C. DC. | AB 744 | EU581382 | SEMO | Jaramillo et al. (2008) |
| Piperscutilimbum C. DC. | WT 4151 | OK235343 | UMNG | This study |
| Piperscutifolium Yunck. | MAJK 281 | EU581389 | SPF | Jaramillo et al. (2008) |
| Pipersorgosonum C. DC. | MAJ185 | AY572320 | DUKE | Jaramillo & Manos (2001) |
| Piperspoliatum Trel. & Yunck. | MAJ 60 | AF275179 | DUKE | Jaramillo & Manos (2001) |
| Pipersubscutatum C. DC. | EJT 1604 | EU581406 | MU | Jaramillo et al. (2008) |
| Pipertrianae C. DC. | MAJ 662 | EU581413 | DUKE | Jaramillo et al. (2008) |
| Pipertruncatum Vell. | MAJ 937 | EF056291 | RB | Jaramillo et al. (2008) |
| Pipertuberculatum Jacq. | MAJ 710 | AY326225 | DUKE | Jaramillo & Callejas (2004) |
| Piperumbellatum L. | MAJ 35 | EU581433 | DUKE | Jaramillo et al. (2008) |
| Piperumbricola C. DC. | EJT 1014 | EU581435 | MU | Jaramillo et al. (2008) |
| Pipervelae | WT 4058 | OK235344 | UMNG | This study |
| WT 3995 | OK235345 | UMNG | ||
| Piper viçosanum Yunck. | MAJ 809 | EU581440 | RB | Jaramillo et al. (2008) |
| Piperyanaconasense Trel. & Yunck. | MAJ 774 | AY326229 | DUKE | Jaramillo & Callejas (2004) |
*Collections: COAHH, Herbario Amazónico Colombiano; DUKE, Duke University Herbarium, UMNG: Herbario Universidad Militar Nueva Granada; HUA, Herbario de la Unviersidad de Antioquia; INPA, Instituto Nacional de Pesquisas da Amazônia; MG, Museu Paraense Emílio Goeldi; MU, Miami University Herbarium; NY, New York Botanical Garden Herbarium; RB, Herbario Jardim Botânico do Rio de Janeiro; SEMO, Southeast Missouri State University Herbarium; SPF, Herbario da Universidade de São Paulo; SRP, Snake River Plains Herbarium (University of Idaho).
Results
Phylogenetic analyses identified the relationships of the new species described here (GenBank accession numbers are provided in Table 1). The ML tree placed Piperhoyoscardozii and Pipernokaidoyitau within the Radula clade, Piperindiwasii species within Oxodium (= Schilleria, Callejas-Posada 2020) and Pipervelae forms a clade with Piperholdridgeanum W. C. Burger (1971: 144–145), sister to the Macrostachys clade (Fig. 1).
Figure 1.
Phylogenetic relationships of species described in this manuscript, based on Maximum Likelihood analysis of nrITS sequences. New species are shown in bold. Species with peltate leaves are underlined. Numbers above branches are ML bootstrap support values (> 80).
Taxonomic treatment
. Piper hoyoscardozii
W. Trujillo-C & M. A. Jaram. sp. nov.
92B8727A-6424-595F-B81F-BE14E19F1C47
urn:lsid:ipni.org:names:77303983-1
Figure 2.
The locality of four new Piper species from the Andes eastern slopes of Colombia, Ecuador and Peru.
Figure 3.
PiperhoyoscardoziiA sympodial branch B magnified view of the infrutescence C floral bract view from above D floral bract side view E anthers and distal portion of the filament F fruit side view G seed H floral diagram. Illustration by Marcela Morales based on F. Hoyos 049 COAH.
Figure 4.
PiperhoyoscardoziiA growth form, scandent shrub B upper leaf surfaces and spike C infructescence D leaf base and prophyll. Photos from F. Hoyos 049 by F. Hoyos
Type.
Colombia. Caquetá: Florencia, vereda Sucre, 1076 m elev., 1°47'50"N, 75°38'50"W, 18 Oct 2020 [fr], F. Hoyos 049 (Holotype COL, Isotype COAH, HUA, UMNG)
Diagnosis.
Piperhoyoscardozii W. Trujillo & M. A. Jaram. is similar to P.dryadum C. DC. (1891:221) and P.flagellicuspe Trel. & Yunck. (1950:59) from which it is easily distinguished by peduncles 2–3 cm long, spikes long-apiculate and fruit with stigmas sessile vs. peduncle 0.5–1 cm long, spikes not-apiculate and fruit with stigmas on a short style in P.flagellicuspe and P.dryadum.
Description.
Shrub with sarmentose branches. Internodes (1)3–7 cm long, smooth, green, pubescent, trichomes pluricellular, uniseriate, 1–2.3 mm long, idioblasts not visible. Prophylls caducous, 1.2–1.5 cm long, green-whitish, pubescent, trichomes pluricellular, uniseriate, 0.2–1.0 mm long, dispersed on the abaxial surface, idioblasts not visible. Petioles uniform in size along all axes, 0.5–0.8 cm long, vaginate on the basal half, smooth and pubescent. Leaf-blades membranaceous, drying black, uniform in shape and size on all nodes, (5)6–7 × (13)15–17 cm, elliptic, symmetric, base cordate to rounded, apex acuminate; leaf blade smooth, pubescent on both surfaces, trichomes pluricellular, uniseriate, 0.5–2.3 mm long, dispersed on the adaxial surface, along first and second order nerves and dispersed on the areolas and third order nerves of the abaxial surface, eciliate; pinnately nerved from the lower 1/3, 2–3 nerves on each side, with spacing uniform or decreasing and angle increasing gradually towards the base, eucamptodromous, tertiary veins percurrent. Inflorescence and infructescence a simple spike, erect; peduncle 2–3 cm long, pubescent, green; rachis in flower 4–7 cm long, rachis in fruit 7–9 cm long, rachis with a 10–15 mm long, sterile green apical extension, fruits densely grouped along the rachis. Floral bracts cucullate, reddish in flower, triangular from above, 0.4–0.6 × 0.7–0.8 mm, glabrous on the adaxial surface, margin fimbriate, bracts forming bands around the spike. Flowers with four stamens, filaments 0.6–0.8 mm long, anthers 0.4–0.6 × 0.5–0.7 mm, longitudinally dehiscent, dithecous, with connective not protruding, glabrate, idioblasts not evident, black when dried. Sessile stigmas. Fruits rectangular, laterally compressed, green when alive and black when dry, 0.9–1.2 × 1.5–1.9 mm, pubescent, partially immersed in the rachis, with persistent sessile stigmas, 0.05–0.1 mm. Seeds oblong, laterally compressed, brown, smooth, 0.8–1 × 0.8–1.1 mm.
Distribution and habitat.
Piperhoyoscardozii is a shade-loving sarmentose shrub that grows on trees and rocks. It is known from the Amazonian slope of the Andes in southern Colombia and Ecuador, between 1000–1500 m in elevation (Fig. 2).
Phenology.
Flowering specimens were collected in July. Fruiting specimens were collected in August.
Etymology.
This species name is dedicated to Fernando Hoyos Cardozo, a great companion during our floristic explorations of the Amazonian foothills and who collected the type specimen of this species in Caquetá.
Conservation status.
This species is known from six specimen collections representing two subpopulations. The locations where it occurs are threatened by deforestation and expansion of the agriculture frontier, especially extensive cattle ranching. The extent of occurrence (EOO) of 876 km2 and area of occupancy (AOO) of 16 km2 are small, which, together with the continuing decline in quality of habitat, suggests it is Endangered [EN B1a+B2a].
Phylogenetic relationships.
Piperhoyoscardozii belongs to the large clade Radula. A group of medium-size shrubs, mostly self-supporting, but some species are herbs or lianescent shrubs; leaves are plinerved or pinnately nerved. Flowers are densely arranged in spikes forming banding patterns and inflorescences can be erect or distally curved. Furthermore, this species is closely related to the clade of sarmentose shrubs occurring in wet tropical forests in Central America and the western slopes of the west Cordillera in Colombia (the latter corresponds largely to the Chocó Region) that includes P.brachypodon C. DC. (1869:327), P.cavendishioides Trel. & Yunck. (Trelease and Yuncker 1950: 85), P.dryadum, P.flagellicuspe, P.ottoniifolium C. DC. (1866:213), P.oxystachyum C. DC. (1898:255) and P.novogranatense C. DC. (1869:313) (Jaramillo et al. 2008).
Discussion.
Piperhoyoscardozii is a sarmentose shrub, a habit not commonly observed amongst Piper species in the study region (eastern slope of the Andes). The phylogeny (Fig. 1) places P.hoyoscardozii sister to other climbing Piper species occurring on the western slope of the Andes and wetter parts of Mesoamerica. Here we provide a comparative table for the climbing Piper species included in the phylogeny (Table 2). P.hoyoscardozii is easily differentiated because its spikes (in flower and fruit) have a long peduncle and a long apiculate apex.
Table 2.
Comparison of Piperhoyoscardozii with related species of scandent habit.
| Species | Internodes indument | Secondary nerves branch | Peduncle length | Rachis length | Spike a pex | Distribution |
|---|---|---|---|---|---|---|
| P.brachypodon (Benth.) C. DC. | glabrous | from the lower half | 1 cm | 5–7 cm | obtuse | Chocó Region |
| P.cavendishioides Trel. & Yunck. | tomentulose | from the lower one-fourth or one-third | 1 cm | 7–8 cm | obtuse | Chocó Region |
| P.dryadum C. DC. | pubescent | from the lower third | 0.5–1 cm | 5–6 cm | obtuse | Mesoamerica and Chocó Region |
| P.flagellicuspe Trel. & Yunck. | velvety | from the lower half | 0.5 cm | 4 cm | obtuse | Chocó Region |
| P.novogranatense C. DC. | glabrous | from the lower third | 0.5 cm | 4 cm | obtuse | Chocó Region |
| P.ottoniaefolium C. DC. | glabrous | near the base | 0.5–1 cm | 6–8 cm | obtuse | Chocó Region |
| P.xanthostachyum C. DC. | glabrous | from the lower third | 0.5–1 cm | 5–6.5 | obtuse | Mesoamerica and western slope of the Andes |
| P.hoyoscardozii W. Trujillo & M. A. Jaram. | pubescent | from the lower third | 2–3 cm | 7–9 cm | long apiculate | Eastern slope of the northern Andes |
Specimens examined.
Colombia: Caquetá, Florencia, vereda Tarqui, monumento Divino Niño, 1570 m elev., 1°50'0.3"N, 75°39'52.8"W, 30 Aug 2020 [fl], W. Trujillo & F. Hoyos 4120 (COAH, UMNG).; vereda Tarqui, quebrada Tarqui, 1530 m elev., 1°50'28"N, 75°39'42"W, 20 Aug 2020 [42], W. Trujillo 4099 (COAH); corregimiento El Caraño, vereda Sucre. 1076 m elev., 1°47'50.8"N, 75°38'50.5"W, 8 Jul 2014 [fl], W. Trujillo 3130 (COAH); vereda Sucre, Finca campamento Sucre. 1°46'52"N, 75°39'5.1"W. 1050 m elev,. 5 Jul 2012 [fl]. W. Trujillo & C. Malambo 2400; vereda Sucre, vía antigua Florencia-Huila, 1°47'50.8"N, 75°38'50.3"W, 1000 m elev., 24 Sep 2020 [fl], F. Hoyos 042 (COAH). Ecuador. Napo, Parque Nacional Sumaco-Galeras, 0°50'S, 77°34'W, 1090 m elev., 27 Oct 2005 [fr], J. Homeier & M.A. Chinchero 2000 (MO).
. Piper indiwasii
W. Trujillo-C & M. A. Jaram. sp. nov.
138BB73E-0D57-5BDD-893D-64DB4B7025B4
urn:lsid:ipni.org:names:77303984-1
Figure 5.
PiperindiwasiiA sympodial branch B lower leaf surfaces and details of the petiole C magnified view of the infructescence D floral bract view from above E floral bract side view F fruit side view G seed side view. Illustration by Marcela Morales based on Angulo 1550, COAH.
Figure 6.
PiperindiwasiiA sympodial branch upper leaf surfaces B sympodial branch lower leaf surfaces C erect spikes. Photos from E. Trujillo 7249 by G. Delgado.
Type.
Colombia. Caquetá, municipio de San José del Fragua, ronda de bosque cerca al balneario Villa Collazos sobre el rio Fragua, 1°20'04"N, 75°59'28"W, 395 m elev., 14 May 2020, M. Angulo 1550 (Holotype COL, Isoptype COAH, HUA, HUAZ, UMNG).
Diagnosis.
Piperindiwasii W. Trujillo & M. A. Jaram. can be distinguished from P.scutilimbum C. DC. (1920a:242) by many attributes. Piperindiwasii has 1–1.7 cm long petioles, 7–8 pairs of secondary veins and a narrowly spatulate leaf base extension, 0.4–0.9(1.5) cm wide, vs. P.scutilimbum, which has a 4–6 cm long petioles, 10–12 secondary veins and an obtuse and rounded leaf-base extension, 2.5–4 cm wide. Piperindiwasii occurs in the Amazon watershed, on the eastern foothills of the Andes, while P.scutilimbum occurs west of the Andes in Panama and extends to Sierra Nevada de Santa Marta in northern Colombia.
Description.
Shrub, up to 2 m tall. Internodes 2–4(5) cm, canaliculate, green, glabrous. Prophylls not seen. Petioles are uniform in size along all axes, 1–1.7 cm long, vaginate along their entire length, canaliculate, glabrous. Leaf-blades coriaceous, drying grey to brown, uniform in size along all axes, (4.5)6–8.5 × (11)14–18 cm, elliptic, symmetric, leaf-base peltate, symmetric, with a narrowly spatulate extension, 0.4–0.9 × (0.5)1–2.3(2.5) cm, the leaf-base extension covering the petiole on sympodial nodes and orientated towards the axis on monopodial nodes, apex acuminate; leaf-blade glabrous on both surfaces, eciliate; pinnately nerved throughout, the nerves 7–8 on each side, brochidodromous, with spacing decreasing towards the base and angle uniform throughout, tertiary veins random reticulate. Inflorescences and infructescences a solitary spike, erect; peduncle 0.9–1.5 cm long, glabrous, green; rachis length in fruit (5)6–20 cm, fruits loosely grouped along the rachis. Floral bracts cucullate, triangular from above, 0.3–0.5 × 0.5–0.8 mm, glabrous on the adaxial surface, margin eciliate, not forming bands around the spike. Flowers with three stamens, filaments 0.5–0.8 mm long, anthers 0.3–0.5 × 0.2–0.3 mm long, longitudinally dehiscent, dithecous, with connective not protruding, glabrate, idioblasts not evident; stigmas 3, 0.05–1.5 mm long, sessile. Fruits obpyriform, green when alive and brown when dry, 0.6–0.8 × 0.9–1.2 mm, glabrous, partially immersed in the rachis, with stigmas persistent, 0.05–1.5 mm long, sessile. Seeds obpyriform, black.
Distribution and habitat.
Piperindiwasii is known from the Amazonian slopes of the Andes in Colombia (Departments of Putumayo, Caquetá and Guaviare), Ecuador (Provinces of Guayas, Napo, Orellana, Pastaza and Sucumbíos) and Peru (Provinces of Amazonas), from 200 to 1,608 m elevation (Fig. 2). It occurs in lowland (sometimes along riverbanks) and lower montane forests. It is a shade-loving species that grows in the understorey and the edges of trails of preserved forests.
Phenology.
Fruiting specimens were collected from December to June and August to October. Flowering samples were collected in March, April, May and December.
Etymology.
Piperindiwasii is named after the Inga word meaning “House of the Sun”. Ingas or Inganos are an indigenous group belonging to the Quechua linguistic family. The Ingas of the Amazon foothills are made up of migratory groups from the Peruvian and Ecuadorian Amazon, the Mocoas and some survivors of the Andaquíes. The clans are united by their location, cosmovision and the culture of “yajé” (Banisteriopsiscaapi) . Furthermore, the type specimen of this species was collected in the Alto Fragua Indi-Wasi National Park, located in San José del Fragua, Caquetá-Colombia.
Conservation status:
Piperindiwasii is not endangered. It is known from 11 subpopulations and 12 localities; it has an EOO of 1,478,359 km2 and an AOO of 52 km2. According to IUCN guidelines, it is of Least Concern (LC) as the region where it occurs is threatened by deforestation and its conservation status should be monitored.
Phylogenetic relationships.
Piperindiwasii belongs to the Neotropical clade Oxodium. Species in this group are shrubs, sometimes sarmentose. They have plinerved or pinnately nerved leaves, leaf bases are often acute or cordate and they have lax inflorescences with loosely arranged flowers (Jaramillo et al. 2008). Piperindiwasii is closely related to P.scutilimbum. Nucleotide difference between the two PiperindiwasiiITS accession is 5 bp out of 641 (0.8%), while these sequences have a 27 bp (4%) dissimilarity with P.scutilimbum (collected in the type locality). Sequence data and leaf material were available for five species of Neotropical Piper with peltate leaves. P.subscutatum C. DC. (1869:321) and P.imberbe Trel. & Standl. (Standley and Steyermark 1952:303–304) are in the Oxodium clade as P.indiwasii and P.scutilimbum; P.scutifolium Yunck. (1966:123–124) is part of the Ottonia clade; and P.maxonii C. DC. (1920b:16), P.hartwegianum (Benth.) C. DC. (1869: 369) and P.candolleiSodiro (1905:202) are members of the Macrostachys clade. Neotropical Piper species with peltate leaves do not form a monophyletic group and are part of at least four clades: Oxodium, Pothomorphe, Macrostachys and Ottonia (Fig. 1). Further studies are needed to shed light on the convergence of this trait.
Uses by communities.
Various common names are used for P.indiwasii amongst indigenous communities in Ecuador: a) “ñahui tapa panga” (closed eye leaf) (D. Irvine & L. Cejua 1125, F, QCA); when a patient is sick and the eyes are closed even when awake, leaves are wrapped around tobacco and the smoke blown over the eyes of the patient; b) “uchi-ampar” a Shuar name, plant used against parasites, leaves and roots are used (Guerrero 171; Herrera 288, MO, QCNE); c) “palu sera aula” or the grandmother of palu sera, which is used for relieving toothache (D. Irvine 959, F, QCA).
Discussion.
After reviewing the specimens identified as P.scutilimbum from Panama and northern Colombia vs. specimens from the eastern (Amazonian) Andes slope in Colombia, Ecuador and Perú (Fig. 2), we found a consistent difference in the leaf base shape between collections from both regions. The obtuse and rounded leaf base extension of the type specimen from the Sierra Nevada de Santa Marta coincides with that of specimens collected in Panama in contrast to the narrowly spatulate base extension seen in P.indiwasi. The leaf base shape, combined with other morphological characters and geographical distribution, clearly allows Piperindiwasii to be proposed as a new species. We provide a comparative table of morphological characters for species of Piper with peltate leaves that belong to the Oxodium clade (see Table 3).
Table 3.
Comparison of Piperindiwasii with species of Piper with peltate leaves belonging to the Oxodium clade.
| Species | Leaf dimorphism* | Leaf base extension width | Leaf base extension shape | Leaf width | Geographic distribution |
|---|---|---|---|---|---|
| P.imberbe Trel. & Standley | all leaves peltate | 1.2–1.5 cm | rounded | 4.5–8 cm | Mesoamerica |
| P.scutilimbum C.DC. | all leaves peltate | (2)2.5–4 cm | obtuse and rounded | 8–14 cm | Mesoamerica and northern Andes |
| P.subscutatum (Miq.) C. DC | present | 2–4 cm | obtuse and rounded | 25–28 cm | Eastern slope of the northern Andes and Amazonia |
| P.indiwasii W. Trujillo & M. A. Jaram. | all leaves peltate | 0.4–0.9 cm | narrowly spatulate | 4–18 cm | Eastern slope of the northern Andes |
* Leaves can be peltate or not.
Specimens examined.
Colombia. Caquetá, San José del Fragua, ronda de bosque cerca al balneario Villa Collazos sobre el rio Fragua, 1°20'04"N, 75°59'28"W, 400 m elev., 29 Jun 2011 [fr], W. Trujillo et al. 1999 (COAH); Belén de los Andaquies, vereda las verdes, cerro Monserrate, entrada por dos quebradas, 1°36'38"N, 75°53'23"W, 700 m elev., 24 Jun 2011 [fr], W. Trujillo et al. 1990 (COAH); Belén de los Andaquies, Parque Natural Municipal Andaqui, cabeceras del rio pescado, 1°41'52"N, 75°54'15"W, 1608 m elev., 25 Jan 2017 [fr], N. Castaño et al. 8734 (COAH, HUA); Belén de los Andaquies, Parque Natural Municipal Andaqui, sector entre filo seco y la bocana de la quebrada las verdes, 1°37'13"N, 75°53'46"W, 600–800 m elev., 7 Feb 2017 [fr], N. Castaño et al. 9659 (COAH, HUA); Florencia, vereda Damas Arriba, finca el Mirador, 1°37'56"N, 75°41'49"W, 750 m elev., 14 Feb 2002 [fr], M. Correa et al. 2853 (COAH, UDBC); Guaviare, El Retorno, cerca del Retorno, granja de la Corporación Araracuara, zona ligeramente disectada, bosque intervenido, 1 Mar 1994 [fl], P. Stevenson 1168 (COAH); Putumayo, Villagarzón, corregimiento la Castellana, vereda la Pradera, finca el Cairo, bosque intervenido a borde de quebrada, 0°52'23"N, 76°45'27"W, 600 m elev., 10 Dec 1999 [fr], C. Marín & D. Cárdenas 1997 (COAH); Villagarzón, vereda la Kofaina, 1°01'00"N, 77°17'00"W, 550–700 m elev., 2 Sep 1993 [fr], A. Cogollo et al. 6830 (COAH, JAUM, MO); Ecuador. Napo, Estación Biológica Jatun Sacha, 1°04'00"S, 77°37'00"W, 450 m elev., 10 Oct 2007 [fr], J. Homeier et al. 2834 (MO, QCA, QCNE, GOET); Estación Biológica Jatun Sacha, rio Napo, 8 km debajo de Misahualli, 1°04'00"S, 77°36'00"W, 450 m elev., 17 Jan – 6 Feb 1987 [fr], C. Cerón 638 (MO, HUA); Estación Biológica Jatun Sacha, rio Napo, 8 km debajo de Misahualli, 1°04'00"S, 77°36'00"W, 450 m elev., 19–28 Mar 1987 [fr], C. Cerón 1063 (HUA, MO); 9 km rio debajo de puerto Misahualli y 2 km al sur de la cuenca del rio Chinguipino, 1°05'00"S, 77°36'00"W, 430 m elev., 10 Mar 1985 [fr], D. Neill et al. 6054 (ECUAMZ, HUA, MO, NY, QCNE, US); Tena, Estación Biológica Jatun Sacha, rio Napo, 8 km E of puerto Misahualli, 1°04'00"S, 77°36'00"W, 400 m elev., 18 May 1985 [fr], W. Palacios 421 (AAU, HUA, MO, NY, QCNE, US); Tena, Estación Biológica Jatun Sacha, along S bank of rio Napo, 8 km E of puerto Misahualli, 1°04'00"S, 77°36'00"W, 450 m elev., 1 Apr 1992, T. Croat 73366 (HUA, MO) [fr]; along road between Tena and Puyo, 61.5 km N of Puyo, 1°11'36"S, 77°52'34"W, 500 m elev., 22 Dec 1979 [fl], T. Croat 49657 (MO); Archidona, Parque Nacional Galeras a 1.5 km de la comunidad Santa Rosa de Arapino, 00°51'00"S, 77°31'00"W, 1230 m elev., 3 Apr 1996 [fr], H. Vargas & P. Grefa 951 (HUA, MO, QCNE); San José de Payamino 40 km W of Coca, 00°30'S, 77°20'W, 26 Apr 1984 [fl], D. Irvine & H. Jipa 959 (F, QCA); 1 May 1984, D. Irvine & L. Cejua 1125 (F, MO, QCA); Sucumbíos, Dureno, comunidad Cofan al sur del rio Aguarico, 20 km al este de Lago Agrio, 00°05'00"N, 76°40'00"W, 350 m elev., 27 Dec 1988 [fr], C. Cerón et al. 5824 (MO, HUA); Dureno, comunidad Cofan al sur del rio Aguarico, 20 km al este de Lago Agrio, 00°05'00"N, 76°40'00"W, 350 m elev., 27 Dec 1988 [fr], C. Cerón et al. 5827 (MO, HUA); Francisco de Orellana, Orellana, comunidad Shuar Tiguano al sur del Coca por la vía al Pindo, 00°44'58"S, 76°46'55"W, 300 m elev., 6–12 May 2004 [fl], W. Guerrero & A. Herrera 171 (MO, QCNE); Orellana, comunidad Shuar Tiguano al sur del Coca por la vía al Pindo, 00°44'58"S, 76°46'55"W, 300 m elev., 11 May 2004 [fr], A. Herrera & W. Guerrero 288 (MO, QCNE); Parque Nacional Yasuni, Rio Tiputini al noroeste de la confluencia con el Rio Tivacuno, este de la carretera Repsol-YPF, km 32 hacia NPF, Sendero Botánico Guiyero, 00°38'S, 76°30'W, 200—300 m elev., 26 Feb 2002 [fr], G. Villa 1350 (F, QCA, US); Peru. Amazonas, Condorcanqui, Santiago, Cerros Kampankis, Serranía entre los Rios Santiago y Morona, desde Río Marañón hasta frontera con Ecuador, Campamento 1: Pongo Shenin, 03°07'01.52"S, 77°46'55.14"W, 520 m elev., 3 Aug 2011 [fr], I. Huamantupa 15217 (USM, F).
. Piper nokaidoyitau
W. Trujillo-C & M. A. Jaram. sp. nov.
6726D4AE-2BBF-5514-9CAE-A0CC4C3EAF5D
urn:lsid:ipni.org:names:77303985-1
Figure 7.
PipernokaidoyitauA sympodial branch B details of the inflorescence C floral bract viewed from above D floral bract side view E flower side view F stamen with glabrous connective G fruit side view H floral diagram. Illustration by Marcela Morales based on F. Hoyos 046 (COL).
Figure 8.
PipernokaidoyitauA sympodial branch, lower leaf surfaces B sympodial branch, upper leaf surfaces and spikes C single infructescence. Photos from F Hoyos & W Trujillo 046 by F Hoyos
Type.
Colombia, Caqueta; Florencia, corregimiento el Caraño, vereda Sucre, 01°47'50.8"N, 75°38'50.5"W, 1020 m elev., 25 Oct 2020 [fr], F. Hoyos & W. Trujillo 046 (Holotype COL, Isotype COAH, UMNG, HUAZ, HUA)
Diagnosis.
Pipernokaidoyitau W. Trujillo & M. A. Jaram., can be separated from the similar species P.hostmannianum (Miq.) C. DC. (1869:287), by its prophylls up to 2.4 cm long, leaves 12–20 cm long vs. prophylls 2.8–3.5 cm long, leaves 21–26 cm long in P.hostmannianum.
Description.
Shrub up to 3 m tall; internodes (2.5–)3–4.5 cm long, canaliculate superficially, green, glabrous. Prophylls 2.8–3.2 cm long, green, glabrous, caducous, swollen in the basal portion (observable in live plants). Petioles uniform in size along all nodes, (0.7–)1–1.2(–1.5) cm long, sheathing at the base, smooth, glabrous. Leaf-blades coriaceous, drying black, uniform in shape and size along all axes, (4.5–)5–8 × (18) 21–26 cm, ovate, asymmetric, base rounded, glabrous on both surfaces, eciliate; pinnately nerved throughout, 4–5 ascending nerves on each side, eucamptodromous, with spacing decreasing and angle increasing towards the base, tertiary veins random, reticulate; apex acuminate. Inflorescences simple spikes, erect; peduncle 1–1.5 cm long, glabrous, green; rachis (7.5)10–12 × 0.3 cm in flower, 11–13 × 0.4–0.5 cm in fruit, flowers densely grouped along the rachis, forming bands around the spike. Floral bracts cucullate, heart-shaped from above, 0.2–0.35 × 0.4–0.7 mm, glabrous centrally on the abaxial surface, margin densely fimbriate. Flowers with three stamens, filaments 0.3–0.5 mm long, anthers 0.1–0.25 × 0.2–0.3 mm, transversally dehiscent, dithecous, with connective not protruding, glabrate, idioblasts not evident, colour black when dried; stigmas 3, 0.1–0.25 mm long, sessile. Fruits obpyriform in side view and triangular from above, green when alive, black when dry, 0.5–0.7 × 0.8–0.9 mm, glabrous, partially immersed in the rachis, with stigmas sessile and persistent.
Distribution and habitat.
Pipernokaidoyitau is known from the lower montane forests in the eastern slopes of the Andes in Colombia, ca. 1,100 m elevation (Fig. 2), the Department of Caquetá. It is a shade-loving species that grows in the understorey of preserved forest.
Phenology.
Flowering specimens were collected in September and October. Fruiting specimens in October.
Etymology.
Pipernokaidoyitau is named after the Huitoto name for Piper plants, “Nokaido yitau”. It means “the powers of the toucan” because these are sacred and medicinal plants used against fever, body and headaches and as anti-inflammatories.
Phylogenetic relationships.
Pipernokaidoyitau belongs to the Radula clade of Neotropical Piper (Jaramillo et al. 2008). Specifically, P.nokaidoyitau is sister to the Isophyllon subclade, within Radula. Isophyllon species are mostly self-standing shrubs with coriaceous leaves, pinnately nerved, acute or obtuse bases and flowers densely arranged in erect inflorescences. Isophyllon species occur in the Atlantic Forest, Central America and the Amazon Region.
Conservation status.
This species is known from four specimen collections representing two subpopulations. The locations where it occurs are threatened by deforestation and expansion of the agriculture frontier. The area of occupancy (AOO) of 8 km2 is small, which, together with the continuing decline in habitat quality, suggests it is Endangered [EN B1a+B2a].
Comments.
Leaves of Pipernokaidoyitau are pinnately nerved throughout with 4–5 ascending secondary veins on each side. This characteristic is shared by other species belonging to the Radula clade, specifically subclade Isophyllon (Jaramillo et al. 2008). We compared P.nokaidoyitau with similar species that occur on the eastern slope of the northern Andes (Table 4).
Table 4.
Comparison of Pipernokaidoyitau with related species that have leaves pinnately nerved throughout and occur in the eastern slope of the northern Andes.
| Species | Prophylls length | Leaf shape | Leaf base shape | Leaf length | Infructescence length | Geographic distribution |
|---|---|---|---|---|---|---|
| P.anonifolium Kunth | 0.5–0.8 cm | elliptic | cuneate | (10)14–17 cm | 3–3.5 cm | Amazonia |
| P.hostmannianum C. DC. | 1–2 cm | elliptic | rounded | 12–20 cm | 10–11 cm | Amazonia |
| P.nokaidoyitau W. Trujillo & M. A. Jaram. | 3–3.5 cm | ovate | rounded | 21–26 cm | 11–13 cm | Eastern slope of the northern Andes |
| P.mastersianum C. DC. | 0.4–0.8 cm | ovate | rounded | 9–12 cm | 4–9 cm | Amazonia |
Specimens examined.
Colombia. Caquetá, municipio de Florencia: corregimiento El Caraño, vereda El Caraño, finca Las Brisas, 01°44'14.7"N, 75°40'35.3"W, 1116 m elev., 18 Oct 2013 [fl], W. Trujillo et al. 3005 (COL); Corregimiento El Caraño, finca Las Brisas, 01°44'14.7"N, 75°40'35.3"W, 1116 m elev., 18 Oct 2013 [fl], W. Trujillo et al. 3022 (COL); Municipio de Belén de los Andaquíes: río Pescado, Parque Natural Andaqui, sector sur, 01°36'31"N, 75°55'16"W, 950 m elev., 25 Jun 2013, W. Trujillo et al. 2791 (HUAZ). Cauca. Municipio de Piamonte, corregimiento de Miraflor, vereda La Florida, camino a la reserva La Cristalina, 01°04'59.6"N, 76°28'08.2"W, 1146 m elev., 06 Jan 2021 [fr], E. Trujillo et al. 7249 (CUVC, JBB).
. Piper velae
W. Trujillo-C & M. A. Jaram. sp. nov.
4BAD96EB-8763-5066-959F-55F6618FD24D
urn:lsid:ipni.org:names:77303989-1
Figure 9.
PipervelaeA sympodial branch B magnified view of the infructescence C floral bract view from above D floral bract side view E fruit side view F stamen side view G Seed side view H floral diagram. Illustration by Marcela Morales based on W Trujillo 905
Figure 10.
PipervelaeA sympodial branch upper leaf surfaces and spike B magnified view of the leaf base and prophyll C magnified view of the spike D magnified view of the prophyll. Photos from W. Trujillo 2039 by William Trujillo.
Type.
Colombia, Caquetá, Belen de los Andaquies, corredor resguardo La Cerinda, PNN Alto Fragua Indiguazi, etnia Embera Katio, 1°36'08.6"N, 75°51'49.1"W, 470 m elev., 03 Oct 2007, W. Trujillo et al. 905 (Holotype COAH, Isotype HUAZ).
Diagnosis.
Pipervelae W. Trujillo & M. A. Jaram. can be distinguished from the related species P.holdridgeanum W.C. Burger by its elliptic leaves with cordulate leaf bases at all nodes, petioles 0.8–1.5 cm long, fruits cylindrical and pubescent vs. leaves cordate to elliptical with leaf bases that are rounded at fertile nodes and cordate at sterile nodes, petioles that are variable in size from 1–5 cm long, fruits rounded and glabrous in P.holdridgeanum. It can be separated from similar species P.cornifoliumKunth (1815 [1816]:52) because it has leaves pinnately nerved in the lower half of the blade and pubescent fruits vs. leaves pinnately nerved in the lower third of the blade and glabrous fruits in P.cornifolium.
Description.
Shrub up to 1.5 m tall. Internodes 2–8.5 cm long, smooth, green, tomentulose, idioblasts not evident. Prophylls 1.2–2 cm long whitish, tomentulose, caducous. Petioles variable along all axes; on monopodial axes 1–1.5 cm long, vaginate to 3/4 of the length, smooth, tomentulose; on sympodial axes 0.8–1.2 cm long, vaginate at the base, smooth, tomentulose. Leaf-blades coriaceous, drying black, uniform in shape and size along all axes, 6–7(11) × 12–15(19) cm, elliptic, symmetric, base cordulate, basal extension asymmetrical; leaf blade smooth, tomentulose on the abaxial surface and glabrous adaxially, eciliate; pinnately nerved from the lower half, 4–5 ascending nerves on each side, festooned brochidodromous, with spacing decreasing and angle increasing towards the base, tertiary veins percurrent; apex acuminate. Inflorescences and infructescence a solitary spike, terminal, erect; peduncle 0.8–1.5 cm long, tomentulose, green; rachis in flower not seen, rachis in fruit 6–8.5 cm long, fruits densely grouped along the rachis. Floral bracts cucullate, triangular from above, 0.15–0.25 × 0.3–0.4 mm, glabrous on the adaxial surface, margin fimbriate, not forming bands around the spike. Flowers with four stamens, filaments 0.2–0.4 mm long, anthers 0.2–0.3 × 0.15–0.25 mm, longitudinally dehiscent, dithecous, shorter than filament, with connective not protruding, glabrate, idioblasts not evident, colour not seen. Stigmas 3, on a short style. Fruits cylindrical, laterally compressed, green when alive and black to brown when dry, 0.8–1.2 × 1–1.3 mm, pubescent on the tip, partially immersed in the rachis, with stigmas persistent, 0.07–0.12 mm long, on a short style, 0.1–0.3 mm long. Seeds 0.4–0.6 × 0.9–1.1 mm, rectangular, laterally compressed, obtuse, black.
Distribution and habitat.
Pipervelae occurs in the eastern slopes of the Andes, from 250–1,500 m in elevation, spreading from wet lowland to wet premontane forests. It occurs in the Colombian Departments of Caquetá, Meta, Cauca and Putumayo. In lowland forests, it occurs in dense terra firme forests. In premontane forests, it grows mostly on moderate slopes, sometimes occurring on steep slopes and rocky substrates. It is a shade-loving species, growing in the understorey and it is also found in forest gaps.
Phenology.
Flowering specimens were collected in February, April, May, June, July and October. Fruiting specimens in January, April, June, July, September, October and December.
Etymology.
Pipervelae is named in honour of Huber Fernando Vela, M.D., a social and environmental leader of Caquetá who was murdered in 2021. Dr Vela and sponsored Piper collections by WT during 2020. Huber Fernando was the leader of the Nature Reserve Romi Kumu, where 30 ha of forest were restored in 2020. The type specimen of P.velae occurs in the region that Dr Vela loved and helped conserve and restore.
Conservation status.
This species is known from 41 specimen collections representing 12 subpopulations. It occurs in 18 locations threatened by deforestation. The extent of occurrence (EOO = 40,810 km2, below the EOO to be considered Vulnerable, VU) and area of occupancy (AOO = 96 km2), suggest it is of Near Threatened [NT B1a+B2a].
Phylogenetic relationships.
P.velae is sister to P.holdridgeanum and these form a clade sister to Macrostachys (Jaramillo et al. 2008). P.velae and P.holdridgeanum have sheathing petioles to ¾ of their length and tightly-arranged flowers. The marked foliar dimorphism between leaves on sterile (monopodial) and fertile (sympodial) nodes distinctive of P.holdridgeanum have obscured its relationships (Callejas-Posada 2020). Here, we present phylogenetic evidence for its placement sister to Macrostachys. Both species, P.velae and P.holdridgeanum require further study to understand their morphological affinities.
Discussion.
Pipervelae can be confused with P.cornifolium, because of its cordulate leaf base; however, these taxa are distinguished, based on the leaf venation pattern and fruit pubescence (see Table 5). We also compare P.velae to closely-related P.holdridgeanum; further studies will help us corroborate this relationship and find morphological similarities.
Table 5.
Comparison of Pipervelae with closely-related species P.holdridgeanum and morphological similar species P.cornifolium.
| Species | Petiole length | Leaf shape | Leaf base | Secondary nerves branched | Fruit shape | Fruit pubescence | Geographic distribution |
|---|---|---|---|---|---|---|---|
| P.cornifolium Kunth | 0.8–1.2 cm | obovate-elliptic | cordulate | from the lower third | rounded | pubescent | Northern Andes |
| P.holdridgeanum W.C. Burger | 1–5 cm | cordate to elliptic | rounded to cordate | from the lower half | rounded | glabrous | Mesoamerica |
| P.velae W. Trujillo & M. A. Jaram. | 0.8–1.5 cm | elliptic | cordulate | from the lower half | cylindrical | pubescent | Eastern slope of the northern Andes |
Specimen examined.
Colombia: Caquetá: Belén de los Andaquíes: Parque Bosque Microcuenca La Resaca, sendero Alto Sarabando, 800 m elev., 1°27'29"N, 75°53'1.8"W, 26 Oct 2010, D. Cárdenas et al. 40791 (COAH); vereda Las Verdes, río Pescado, margen izq. Parque Natural Municipal Andakí, 700 m elev., 1°36'7.2"N, 75°54'10"W, 28 Oct 2010, D. Cárdenas et al. 40896 (COAH); sector Paramillo, camino entre Acevedo - Belén de Andaquíes, 1400 m elev., 1°40'58"N, 75°54'21"W, 23 Jul 2011, D. Cárdenas et al. 41848 (COAH); cerca del río Pescado, vereda Los Angeles, 1225 m elev., 1°34'49.7"N, 75°54'17"W, 9 Jul 2011, L. Martinez 24 (COAH); río Pescado, Parque Natural Andakí, sector Sur, 950 m elev., 1°36'31"N, 75°55'16"W, 25 Jun 2013, W. Trujillo et al. 2792 (COAH). Cartagena del Chairá: vereda Laguna del Chairá, 240 m elev., 1°15'23.9"N, 74°48'49.8"W, 28 Sep 2007, W. Trujillo et al. 853 (COAH). Florencia: corregimiento El Caraño, vereda Alto Brasil, camino hacia la quebrada Yumal, 321 m elev., 1°39.543'N, 75°36.128'W, 22 Oct. 2012, A. Jimenez et al. 4 (HUAZ); río Hacha, vereda El Caraño, finca Marsella, 500 m elev., 1°41'47"N, 75°37'16"W, 26 Jun 2010, D. Cárdenas et al. 24884 (COAH, HUAZ); vereda El Paraíso, antigua vía Florencia-Neiva, zona de cordillera baja, 756 m elev., 1°45'7.4"N, 75°39'56.9"W, 20 Oct 2010, D. Cárdenas et al. 40568 (COAH); vereda El Paraíso Bajo, vegetación de cordillera baja, 800 m elev., 1°45'0.2"N, 75°37'8.2"W, 22 Oct 2010, D. Cárdenas et al. 40595 (COAH); Centro de investigaciones Macagual, 258 m elev., 1°29'59.8"N, 75°39'22.6"W, 13 Dec 2008, W. Trujillo et al. 1214 (COAH); vereda El Caraño, 1116 m elev., 1°44'14.7"N, 75°40'35.3"W, 18 Oct 2013, W. Trujillo et al. 3002 (COAH). La Montañita: vereda Los Morros, reserva Las Dalias, 300 m elev., 1°29'21.5"N, 75°24'17.6"W, A. Meneses 15 (HUAZ); vereda Itarca, Reserva Natural Itarca, 340 m elev., 1°32'53"N, 75°28'20"W, 30 Oct 2010, D. Cárdenas et al. 40956 (COAH); vereda Itarca, Reserva Natural Itarca, 330 m elev., 1°32'34.5"N, 75°28'19"W, 26 Apr 2011, N. Castaño et al. 3142 (COAH); Reserva Las Dalias, 382 m elev., 1°29'21.6"N, 75°24'17.6"W, 41050, W. Trujillo et al. 2086 (COAH). San Vicente del Caguán: inspección Guacamaya, vereda La Música, margen izquierda del río Caguán, 600 m elev., 2°20'46"N, 74°54'12.4"W, 12 Jul 2015, D. Cárdenas et al. 44401 (COAH). Cauca: Piamonte: La Libertad, 700 m elev., 1°6'41.78"N, 76°28'46.5"W, 22 Feb 2010, W. Trujillo et al. 1271 (COAH). Santa Rosa: Inspección de Santa Marta, vereda Diamante Alto, 1150 m elev., 1°14'N, 76°36'W, 22 Jun 2002, B. Ramírez 16061 (COAH). Meta: San Juan de Arama: sector del chorro Santo Domingo, Parque Nacional Natural Sierra de La Macarena, 520 m elev., 3°15'23"N, 73°57'50"W, 27 Oct 2019, D. Cárdenas et al. 52002 (COAH); vereda Monserrate Bajo, finca El Paraíso, cerca al caño Las Ninfas, 590 m elev., 3°20'33.75"N, 73°57'3.06"W, 4 Apr 2004, L. Carvajal et al. 173 (COAH). Putumayo: Mocoa: vereda Medio Afán, camino Serranía El Churumbelo, sector Nororiental, 900 m elev., 1°10'39"N, 76°38'47"W, 4 Oct 2000, D. Cárdenas et al. 12221 (COAH); vereda San José del Pepino, 1°5'40.17"N, 76°37'49.05"W, 21 Jun 1997, R. López et al. 2514 (COAH); vereda San José del Pepino, 1°5'40.17"N, 76°37'49.05"W, 21 Jun 1997, R. López et al. 2610 (COAH). Orito: Santuario de Flora y Plantas Medicinales Ingui-Ande, vereda Líbano, sector de don Reinaldo, 900 m elev., 0°41.62'N, 77°3.789'W, 1 Oct 2015, D. Cárdenas et al. 45416 (COAH); Santuario de Flora y Plantas Medicinales Ingui-Ande, 1004 m elev., 0°41.627'N, 77°3.801'W, 27 Sep 2015, D. Cárdenas et al. 45454 (COAH); vereda El Líbano, 832 m elev., 0°38'15"N, 77°4'17.9"W, D. Cárdenas et al. 51291 (COAH).
Supplementary Material
Acknowledgements
We thank Marcela Morales for botanical illustrations and Viviana Vargas for preparing Figures 3, 6, 8 and 10. ET and WT thank Vicerrectoría de Investigaciones and Facultad de Ciencias Básicas of the Universidad de la Amazonia for support in conducting floristic research in Caquetá. We thank Norida Marín from Herbario Amazónico Colombiano (COAH) and Hector García from Herbario Universidad de Magdalena (UTMC) for providing photographs of herbarium specimens.
Citation
Trujillo W, Trujillo ET, Ortiz-Morea FA, Toro DA, Jaramillo MA (2022) New Piper species from the eastern slopes of the Andes in northern South America. PhytoKeys 206: 25–48. https://doi.org/10.3897/phytokeys.206.75971
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