Phytotaxa 601 (3): 263–284 ISSN 1179-3155 (print edition)

https://www.mapress.com/pt/
Article PHYTOTAXA
Copyright © 2023 Magnolia Press ISSN 1179-3163 (online edition)

https://doi.org/10.11646/phytotaxa.601.3.4

Over- and under-described: new species, new synonyms, and a new name in the
megadiverse genus Piper (Piperaceae) on the east Andean slopes
WILLIAM TRUJILLO1,5 *, M. ALEJANDRA JARAMILLO2,6*, DIEGO TORO3,7 & HENRIK BALSLEV4,8
1
Grupo Investigaciones territoriales para el uso y conservación de la biodiversidad. Fundación La Palmita, Centro de Investigación.
Cra 4 # 58-59 piso 2, Bogotá, Colombia.
2
Grupo DIVERSITAS, Facultad de Ciencias Básicas y Aplicadas, Universidad Militar Nueva Granada, km 2 Vía Cajicá–Zipaquirá,
Cajicá, Colombia.
3
Centro de Investigaciones Amazónicas CIMAZ-MACAGUAL, Laboratorio de Agrobiotecnología, Universidad de la Amazonia,
Florencia 180002, Colombia.
4
Department of Biology, Ecoinformatics and Biodiversity, Aarhus University, DK-8000 Aarhus C, Denmark.
5 �
williamtrujilloca@gmail.com; https://orcid.org/0000-0002-0170-016X
6 �
maria.jaramillo@unimilitar.edu.co; https://orcid.org/0000-0002-6539-4149
7 �
diegoalextoro@gmail.com; https://orcid.org/0009-0007-5970-8598
8 �
henrik.balslev@bio.au.dk; https://orcid.org/0000-0002-7101-7120
*
Corresponding authors: � williamtrujilloca@gmail.com, � maria.jaramillo@unimilitar.edu.co

Abstract

Three new species and a new name of Piper from the Amazonian slopes of the northern Andes are proposed and described.
Piper cajambrense var. caquetanum, described from Caqueta in Colombia, is elevated to species rank under the name
P. oteguanum and an emended description and an illustration are provided. Piper oteguanum is distinguished from P.
cajambrense var. cajambrense by its glabrous leaves and petioles without epidermal warty outgrowths. The new species
Piper rubrifolium has smooth internodes and petioles which easily distinguish it from the related species P. spoliatum and P.
cajambrense, which has internodes and petioles with epidermal warty outgrowths and fruiting spikes less than 15 cm long.
Piper caguanense differs from P. miguel-conejoanum by the smooth leaves that have evident idioblasts vs. leaves scabrous
and without evident idioblasts. Piper laperdizense is distinguished from related species (P. brachypodon, P .dichroostachyum,
P. dryadum, P. ottoniaefolium and P. cavendishoides) by its glabrous structures, leaves pinnatinerved from the lower third
and with no evident idioblasts. Furthermore, three names of taxa from the east Andean slopes in Colombia are synonymized:
Piper papillicaule under P. cuniculorum, Piper metanum var. villibracteum under P. mituense, and Piper calanyanum var.
sardinanum under P. morelianum.

Keywords: Piper caguanense, Piper oteguanum, Piper laperdizense, Piper rubrifolium, Piperaceae, north-western Amazon,
Colombia, Ecuador, Peru

Introduction

Piperaceae are pantropical and consist of approximately 4200 species in five genera (Wanke et al. 2007, Samain et
al. 2008). Piper with some 1500 species is one of the most species rich genera of flowering plants (Frodin 2004).
Approximately 412 Piper species have been recorded in Colombia (Bernal et al. 2019), where they are found particularly
in the Andes and the Pacific and Amazon lowlands. Piper species are abundant and diverse in the understory of tropical
forests around the world (Gentry 1990, Denslow et al. 2019, Draper et al. 2021). Piper fruits are an important food for
bats, which act as their main dispersers (Fleming 2004, Yohe et al. 2021) and their leaves and roots are widely used
as medicine by communities in the Amazon and around the world (Trujillo-C. & Gonzalez 2011, Mgbeahuruike et al.
2017, Vásquez-Ocmin et al. 2021). The economically most important species in the genus is Piper nigrum Linnaeus
(1753:28), which is widely known throughout the world as the source of black pepper (Parthasarathy et al. 2007).
Over the past ten years we have studied the species of Piper from the Amazon slope of the Andes in Colombia,
specifically from the department of Caquetá, in the field and in the herbarium. We have recorded about 90 species of Piper
in this area (Trujillo & Vargas 2022), of which 24 are endemic to Colombia and nine are restricted to the department of

Accepted by Marie-Stéphanie Samain: 28 Jun. 2023; published: 11 Jul. 2023 263

Caquetá. From the study area ten new species with mostly restricted distributions were recently discovered, described,
and published: Piper andakiense W.Trujillo-C. & Callejas (2015: 262), P. callejasii W.Trujillo-C. & M.A. Jaram.
(2021: 201), P. caranoense W.Trujillo-C. (2019: 60), P. resinaense W.Trujillo-C. (2021: 71), P. tarquiense W.Trujillo-
C. (2021: 73), P. pseudopeculiare W.Trujillo-C. (2021: 68), P. hoyoscardozii W.Trujillo-C. & M.A.Jaram. (2022: 29),
P. indiwasii W.Trujillo-C. & M.A.Jaram. (2022: 34), P. nokaidoyitau W.Trujillo-C. & M.A.Jaram. (2022: 39), and P.
velae W.Trujillo-C. & M.A.Jaram. (2022: 42).
Despite these recent discoveries, there are still many undescribed species on the megadiverse Amazon slope of the
Andes, but there are also many species which have been described several times. Our floristic studies on the Amazon
slope of the Andes have allowed us to clarify taxonomic problems contained in two classic works: the Piperaceae
– Flora of Peru (Trelease 1936) and The Piperaceae of northern South America (Trelease and Yuncker 1950). In the
Flora of Peru, Trelease described 284 new Piper species, most of them from the Amazon, an area that belongs to the
same biogeographic province as our study area in Caquetá. The Piperaceae of northern South America is a taxonomic
treatment of 454 Piper species, of which 246 were described as new to science, eight as new varieties, and 14 new
combinations and four new names were proposed (Trelease & Yuncker 1950). The new synonyms presented here
suggest that there is a need to continue reviewing and updating Piper’s taxonomy, especially in species-rich and poorly
studied areas such as the Andes in southern Colombia, where many species are known from only very few collections.
When there are only few collections, it may lead to errors in the interpretation of the species’ morphological variation,
especially when the collections lack material of both monopodial and sympodial axes or when the spikes are not fully
developed. The synonyms proposed here are examples of this difficulty. The case of Piper mituense Trel. & Yunck.
(1950: 382) and Piper metanum var. villibracteum Trel. & Yunck. (1950: 343) described from type specimens made by
the same collector, the same day and in the same locality, is an example of two separately published names that turn out
to be synonyms after a detailed review 84 years later. In another case described here, two Piper specimens that were
collected at the same locality, one day apart, and described as two different species, have now turned out to be the same
species.
The northeastern slopes of the Andes (towards the Amazon) are recognized as one of the main centers of vascular
plant diversity in the world (Barthlott et al., 2005). Scientists believe this is a result of the varied geography in this
region where the Andes, the Amazon, and the Llanos (the savannas extending between Colombia and Venezuela) meet
(Barthlott et al., 2005). The relief in this region is mountainous and varies from very broken to wavy, and the vegetation
corresponds to Tropical premontane wet forest (TPwf), and Tropical montane rain forest (TMrf) in Holdridge’s
classification (Holdridge 1978). The results presented here help fill the knowledge gap relating to taxonomic diversity,
specifically in the department of Caquetá, Colombia. We describe three species new to science, change the status of
one variety to species, and propose synonyms for three species.

Material and methods

Specimens were collected during extensive botanical expeditions along the Amazon slope of the Andes in southern
Colombia, in the department of Caquetá from 2014 to 2021. Detailed observations in the field and collections allowed
the accurate description of growth habits and phenological stages. Botanical specimens were deposited in the herbaria
COAH, COL, HUA, and HUAZ (acronyms follow Thiers 2018). The new species were recognized by detailed
comparisons with morphologically similar species through literature review (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), examination of specimens (including type specimens) from COAH, COL, HUAZ, GH, NY, and US,
and digitized images of specimens available on the web (e.g., JSTOR Global Plants, https://plants.jstor.org/). The
measurements included in the descriptions below are based on herbarium specimens. To describe leaf architecture, we
used the 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 GeoCat (www.geocat.kew.org; Bachman
et al. 2011) with a 2 km2 cell size.
We collected silica gel-dried leaf tissue samples for DNA extractions from all the new species using a CTAB
method (Doyle and Doyle 1987). We amplified the nuclear ribosomal internal transcribed spacer (ITS) as described
by Jaramillo and Manos (2001) and aligned sequences using previous alignments as a guide (Jaramillo et al. 2008).
We conducted a preliminary analysis with >900 sequences available. Once it was clear where the new taxa are placed
in the global phylogeny, we selected a smaller number of sequences (representative taxa from each Neotropical clade)

264 • Phytotaxa 601 (3) © 2023 Magnolia Press TRUJILLO et al.

to describe the relationships of species discussed here. Maximum Likelihood (ML) phylogenetic and bootstrap (100
replicates) analyses were conducted using RAxML (Stamatakis 2014) that included species from the Asian tropics to
root the phylogeny.

Results

The phylogenetic analysis identified the relationships of four of the species discussed here (Table 1). The ML tree
placed Piper oteguanum and Piper rubrifolium within the Macrostachys clade, Piper laperdizense within Radula, and
Piper morelianum and Piper calanyanum within Peltobryon (Jaramillo et al. 2008) (Figure 1).

FIGURE 1. Phylogenetic relationships of the species Piper oteguanum, P. rubrifolium, P. laperdizense, P. calanyanum and P. morelianum
based on Maximum Likelihood analysis of nrITS sequences. Names with taxonomic novelties are shown in bold. Names of clades follow
Jaramillo et al. (2008), and Callejas-Posada (2020). Numbers above branches are ML bootstrap support values (>80).

Over- and under-described: piper Phytotaxa 601 (3) © 2023 Magnolia Press • 265
Table 1. GenBank accessions and voucher for ITS sequences for species discussed here (in bold) and other reference
taxa.
Species name Collection # Benbank Accession # Country Herbarium
Piper aduncum L. MAJ76 AF275157 Colombia DUKE
Piper amalago L. MAJ561 AF275186 México DUKE
Piper amplum (Kunth) Steud. MAJ804 EU581096 Brazil RB
Piper arboreum Aubl. MAJ839 OR133618  Brazil RB
Piper artanthe C. DC. MAJ83 EU581109 Colombia DUKE
Piper augustum Rudge MAJ122 AF275165 Colombia DUKE
Piper bartlingianum (Miq.) C.DC RE1267B AF275183 Guyana U
Piper biseriatum C. DC. EJT77 EF056228 Costa Rica MU
Piper brachypodon (Benth.) C. DC. MAJ728 OR133619  Colombia HUA
Piper cajambrense Trel. & Yunck MAJ768 AY326199 Colombia DUKE
Piper calanyanum Trel. & Yunck. WT 1979 OP508184 Colombia COAH
Piper caldense C. DC. MAJ805 EU581140 Brazil RB
Piper cavendishoides Trel. & Yunck MAJ70 AF275153 Colombia DUKE
Piper cernuum Vell. MAJ907 EU581150 Brazil RB
Piper cinereum C. DC. MAJ66 AF275190 Colombia DUKE
Piper cordulatum C. DC. SN EU581167 Panamá //
Piper darienense C. DC. MAJ103 AF275181 Colombia DUKE
Piper dichroostachyum Trel. & Yunck MAJ1317 OR133620  Colombia HUA
Piper filistilum C. DC. MAJ157 AF275155 Colombia DUKE
Piper flagellicuspe Trel. & Yunck MAJ65 AF275154 Colombia DUKE
Piper friedrichsthalii C. DC. MAJ584 AY326205 Colombia DUKE
Piper goesii Yunck. MAJ912 EU581220 Brazil RB
Piper hooglandii (I. Hutton & P.S. Green) M.A. Jaram. ROG8496 AF275192 New Zealand AK
Piper hoyoscardosii W. Trujillo-C & M. A. Jaram WT4127 OK235346 Colombia UMNG-H
Piper ladrillense Trel. AKM4193 OR133615  Costa Rica BM
Piper laperdizense W. Trujillo-C & M. A. Jaram ET 8073 OQ247984 Colombia UMNG-H
Piper lepturum Kunth. MAJ967 EU581277 Brazil RB
Piper longivillosum Trel. & Yunck MAJ64 EU581283 Colombia DUKE
Piper marsupiiferum Trel. FH040 OR133617  Colombia COAH
Piper morelianum Yunck. WT 4059 OP508185 Colombia UMNG-H
Piper munchanum C. DC. MAJ120 AF275164 Colombia DUKE
Piper nokaidoyitau W. Trujillo-C & M. A. Jaram WT4171 OK235339 Colombia COAH
Piper novogranatense C. DC. MAJ71 EU581317 Colombia DUKE
Piper obovatum Ruiz. & Pav. MAJ759 AY326213 Colombia DUKE
Piper obtusilimbum C. DC. EJT512 EU581328 Cultivated MU
Piper oteguanum W. Trujillo-C & M. A. Jaram WT 4039 OP508182 Colombia UMNG-H
Piper peltatum L. MAJ142 AF275169 Colombia DUKE
Piper peracuminatum C. DC. MAJ1086 OR133621  Panamá HUA
Piper pterocladum C. DC. TF4261 EU581171 Cultivated PTBG
Piper reticulatum L. MAJ62 AF275185 Colombia DUKE
......continued on the next page

266 • Phytotaxa 601 (3) © 2023 Magnolia Press TRUJILLO et al.

 TABLE 1. (Continued)
Species name Collection # Benbank Accession # Country Herbarium
Piper retrofractum Vahl MAJ395 AF275196 Vietnam DUKE
Piper rubrifolium W. Trujillo-C & M. A. Jaram WT 4029 OP508183 Colombia UMNG-H
Piper sabaletasanum Trel. & Yunck MAJ623 AY326217 Colombia DUKE
Piper sagittifolium C. DC. SN EU581376 Cultivated //
Piper sorsogonum C. DC. MAJ185 AY572320 Philipines DUKE
Piper spoliatum Trel. & Yunck MAJ60 AF275179 Colombia DUKE
Piper tricuspe C. DC. AG41 AY326224 Colombia CUCV
Piper trigonum C. DC. MAJ52 EU581416 Colombia DUKE
Piper truncatum Vell. MAJ941 OR133616  Brasil RB
Piper tuberculatum Jacq. MAJ710 AY326225 Ecuador DUKE
Piper umbellatum L. MAJ35 EU581433 Colombia DUKE
Piper umbricola C. DC. DSA305 OR133622  Costa Rica La Selva

Taxonomic treatment

Piper caguanense W. Trujillo-C. & M.A.Jaram. sp. nov., Figs. 2, 3, 4.

Type:—COLOMBIA: Caquetá, San Vicente del Caguán, road to Neiva, trail to Las Morras, 1031 m, 2°26’30”N, 74°47’2”W, 4 January
2011 [fr], W. Trujillo-C. & M. Calderón 1812 (holotype COAH!, isotype HUA!).
Diagnosis:—Piper caguanense differs from the morphologically similar P. miguel-conejoanum Trel. & Yunck. (1950: 287) by the scabrous
leaves with evident idioblasts vs. smooth leaves without evident idioblasts in P. miguel-conejoanum.

FIGURE 2. The localities of four new Piper species on the east Andes slopes of Colombia, and Ecuador.

Over- and under-described: piper Phytotaxa 601 (3) © 2023 Magnolia Press • 267
FIGURE 3. Piper caguanense. A. Sympodial branch. B. Detail of infructescence. C. Floral bract lateral view. D. Floral bract view from
above. E. Anthers. F. Fruit lateral view. G. Floral diagram. Illustration by Marcela Morales based on the type W. Trujillo-C. & M. Calderón
1812 (COAH).

268 • Phytotaxa 601 (3) © 2023 Magnolia Press TRUJILLO et al.

FIGURE 4. Piper caguanense. A. Sympodial branch; adaxial and abaxial leaf surfaces and a spike. B. Detail of the leaf base and prophyll.
C. Sympodial branch; adaxial and abaxial leaf surfaces and pubescent spikes and internodes. Photos by W. Trujillo from W. Trujillo 4133
(COAH).

Shrub, 1.5 m tall. Internodes 5–9 cm, smooth, reddish, pubescent, with evident idioblasts throughout the plant except
on the connective of the anthers. Prophylls 1.8–2.2 cm, green, pubescent, caducous. Petioles uniform in shape and size
along axes, 0.5–1 cm long, vaginate at the base, smooth, pubescent. Leaf blades coriaceous, dark green above, fresh
green below, brown when dry, uniform in shape and size along axes, 12–17 cm × 5.5–8.8 cm, ovate, blade medially
symmetric, base rounded, equilateral, insertion asymmetric; leaf blade pubescent on both surfaces with trichomes
(1–)1.5–2 mm, scabrous above, ciliate; pinnately nerved from the lower half of the midvein; secondary veins 4–5 pairs,

Over- and under-described: piper Phytotaxa 601 (3) © 2023 Magnolia Press • 269
eucamptodromous, with spacing decreasing and angle increasing towards the base; tertiary veins percurrent; apex
acuminate. Inflorescences and infructescences simple spikes, terminal, erect; peduncles 4–4.5 cm long, pubescent,
green; fruiting rachis 7–9 cm × 0.2 cm, fruits densely grouped along the rachis. Floral bracts cucullate, triangular
when seen from above, 0.3–0.5 × 0.3–0.3 mm, glabrous on the adaxial surface, margin fimbriated, forming bands
around the spike. Flowers with three stamens, filaments 0.3–0.5 mm long, anthers 0.15–0.35 × 0.1–0.3 mm long,
longitudinally dehiscent, with two thecae, with connective not protruding, glabrate, idioblasts not evident; stigmas 3,
0.05–0.2 mm long, sessile. Fruits rectangular in lateral view, laterally compressed, green when fresh and brown when
dry, 0.8–1.2 × 1.2–1.5 mm, glabrous, partially immersed in the rachis, with stigmas persistent 0.05–0.2 mm long,
sessile. Seeds rectangular, laterally compressed, yellow.
Distribution:—Piper caguanense is endemic to the eastern slopes of the Andes towards the Amazonas basin in
Colombia at 600–1200 m elevation in Tropical premontane wet forests (TPwf).
Habitat and ecology:—Piper caguanense is a shade-loving species growing in the understory of well-preserved
and secondary forests.
Phenology:—Flowering samples were collected in March. Fruiting specimens were collected in October, January,
and February.
Etymology:—The species is named for the type locality at San Vicente del Caguán, in Caquetá where the Caguán
river is born.
Conservation status:—This species is known from four specimens representing two subpopulations. The locations
where it occurs are threatened by deforestation and expansion of the agricultural frontier. The area of occupancy
(AOO) is 8 km2 which is small and together with the continuing decline in quality of habitat, suggests that this new
species is Critically Endangered [CR B2a].
Comments:—Piper caguanense exhibits the traits of the Radula clade which has leaves that are pinnately nerved
from the lower half of the midvein and spikes with bracts forming bands around the rachis (Jaramillo et al. 2008). After
several intents we were not able to obtain ITS sequences for this species, thus, molecular evidence is lacking to confirm
its phylogenetic position. Piper caguanense and P. miguel-conejoanum are both villous shrubs with long peduncles
but Piper caguanense has scabrous leaves with evident idioblasts whereas P. miguel-conejoanum has smooth leaves
without evident idioblasts.
Additional specimens examined:—COLOMBIA: Caquetá, Florencia, trail to Sucre, 1076 m, 1°47’50”N,
75°38’50”W, 18 October 2020, W. Trujillo & F. Hoyos 4123 (COAH!); same locality, 12 December 2020, W. Trujillo
et al. 4133 (COAH!); same locality, 24 September 2020, F. Hoyos & W. Trujillo 039 (COAH!).

Piper laperdizense W. Trujillo-C. & M.A.Jaram. sp. nov. Figs. 2, 5, 6.

Type:—COLOMBIA: Caqueta Florencia, trail to Tarqui, Gabinete, 1°52’50.5”N, 74°40’56”W, 2400 m, 4 January 2022 [fr], W. Trujillo &
F. Hoyos 4449 [holotype COL!, isotypes COAH!, HUA!].
Diagnosis:—Piper laperdizense can be separated from related lianescent species P. novogranatense C. DC. (1869: 313) and P. brachypodon
C. DC. (1869: 327) by its pinnatinerved leaves from the lower third vs. leaves pinnatinerved from the lower half or up to ¾ of the
blade. It can be differentiated from P. dryadum C. DC. (1891: 221) by its glabrous leaves (vs. leaves pubescent). Additionally, P.
laperdizense can be easily differentiated from P. ottoniifolium C. DC. (1866: 213) by having pinnatinerved leaves, secondary nerves
> 2 pairs, branched from the lower third (vs. leaves plinerved, secondary nerves up to 2 pairs, branched near the base). Finally, P.
laperdizense differs from P. cavendishioides Trel. & Yunck. (1950: 85) by its leaves with no evident idioblasts, base acute, bracts
triangular-cucullate vs. leaves with red-idioblasts evident, base rounded, abruptly acute, bracts rounded-subpeltate.

Lianescent shrub, reaching 3m high. Internodes 2.5–6 cm long, smooth, green, glabrous, idioblasts not evident in any
part of the plant. Prophylls 2.8–3.5 cm long, green to white, glabrous, caducous. Petioles (0.5–)0.8–1.2 cm long, vaginate
at the base, smooth, glabrous. Leaf blades coriaceous, green when alive above, almost glaucous below, olivaceous
when dry, uniform in shape and size along all axes, 3.3–5.5 × (10–)11–14 cm, blade elliptic, medially symmetric, base
acute, smooth, glabrous on both surfaces, eciliate, pinnately nerved from the lower third of the midvein, three pairs
of secondary veins, eucamptodromous, with spacing decreasing and angle increasing towards the base, tertiary veins
randomly reticulate, apex acuminate. Inflorescences and infructescences simple spikes, terminal, erect; peduncles
0.7–1 cm long, glabrous, green; flowering rachis 4–6 cm, fruiting rachis 5–8 cm long. Floral bracts cucullate at
anthesis, triangular when seen from above, 0.2 × 0.1 mm, not forming bands around the rachis, fimbriate. Flowers
with 3 stamens, filaments 0.4–0.7 mm long, anthers 0.3 × 0.3 mm, with two thecae, with longitudinal dehiscence and
a glabrous connective; stigmas three, sessile. Fruit obpyriform, 1.4–1.8 × 0.4–0.6 mm, immersed in rachis, glabrous,
stigmas 3, to 0.1 mm long, sessile, light green in fresh material and brown when dry.

270 • Phytotaxa 601 (3) © 2023 Magnolia Press TRUJILLO et al.

FIGURE 5. Piper laperdizense. A. Sympodial branch. B. Adventitious roots at nodes. C. Third order nerves in leaves. D. Detail of
infructescence. E. Anthers and connective. F. Floral bract seen from above. G. Fruit lateral view. H. Floral diagram. Illustration by Marcela
Morales based on the type specimen W. Trujillo & F. Hoyos 4449 (COL).

Distribution:—Piper laperdizense is endemic to the eastern cordillera of the Andes in Colombia along the
Amazon slope at 2000–2400 m elevation, in the Tropical montane rain forest (TMrf).
Habitat and ecology:—It is a shade-loving species growing in the understory of montane rain forest.
Phenology:—Flowering specimens were collected in December and fruiting specimens in January.
Etymology:—The type specimen was collected in the municipality of Florencia in Caquetá. The epithet
laperdizense refers to the name “La Perdiz”, which is an old name of the settlement that would later become the city
of Florencia.
Conservation status:—This species is known from four specimens that were all collected in the same population.
The location where it occurs is threatened by deforestation. The extent of occurrence (EOO) is 74 km2 and area of

Over- and under-described: piper Phytotaxa 601 (3) © 2023 Magnolia Press • 271
occupancy (AOO) is 12 km2, which is small, which together with the continuing decline in quality of its habitat,
suggests that it is Critically Endangered [CR B1a+B2a].
Comments:—Piper laperdizense has glabrous, elliptic leaves that are 10–15 cm long and have acute bases and
spikes that are 5–10 cm long. It can be separated from the other morphologically similar species because it is lianescent
with smooth and green internodes and petioles.

FIGURE 6. Piper laperdizense. A. Lianescent habit. B. Sympodial branch, abaxial leaf surfaces. C. Adventitious root at nodes. D. Leaf
base and prophyll. Photos of the type specimen by William Trujillo based on W. Trujillo & F. Hoyos 4449 (COL).

272 • Phytotaxa 601 (3) © 2023 Magnolia Press TRUJILLO et al.

Key to Piper laperdizense and related species

1 Leaves pinnatinerved up to ¾ of the blade.............................................................................................................P. novo-granatense

– Leaves pinnatinerved from the lower third or from the lower half.....................................................................................................2
2 Leaves pinnatinerved from the lower third.........................................................................................................................................3
– Leaves pinnatinerved from the lower half....................................................................................................................P. brachypodon
3 Rachis of the infructescence < 2.5 cm................................................................................................................... P. dichroostachyum
– Rachis of the infructescence > 2.5 cm.................................................................................................................................................4
4 Internodes and leaves pubescent.........................................................................................................................................P. dryadum
– Internodes and leaves glabrous or puberulent.....................................................................................................................................5
5 Leaves plinerved, secondary nerves 2 pairs, branched near the base . ........................................................................ P. ottoniifolium
– Leaves pinnatinerved, secondary nerves > 2 pairs, branched from the lower third............................................................................6
6 Leaves with red-idioblasts evident, base rounded, abruptly acute, bracts rounded-subpeltate............................... P. cavendishioides
– Leaves with no evident idioblasts, base acute, bracts triangular-cucullate...................................................................P. laperdizense

Additional specimens examined:—COLOMBIA: Caquetá, Florencia, Gabinete, near Caqueta Huila border, km
39 along old road Florencia-Guadalupe, 2400 m, 1°52’50.5”N, 75°40’56”W, 29 August 2020 [fl], W. Trujillo & F.
Hoyos. 4103 (UMNG!); same locality, 13 December 2020 [sterile], W. Trujillo & F. Hoyos. 4131 (COAH!, UMNG!).
Old road Florencia–Neiva, 1°52’7”N, 75°40’14”W, 2070 m, 13 December 2020 [Fl], W. Trujillo et al. 4142 (COAH!,
HUA!).

Piper oteguanum W. Trujillo-C. & M.A.Jaram., stat. nov., nom. nov., Figs. 1, 2, 7, 8.

Type:—COLOMBIA: Caquetá, Río Orteguaza, 2 August 1926, G. Woronow & S. Juzepczuk 6364 (holotype ILL!).
Basionym:—Piper cajambrense var. caquetanum Trel. & Yunk., The Piperaceae of northern South America 1: 27. 1950.

Shrubs 3 m tall, loosely branched. Internodes 8–10 cm long on monopodial axes, 5–7 cm long on sympodial axes,
smooth, light green in fresh specimens and yellow when dry, glabrous, idioblasts not evident in any part of the plant.
Prophylls not developed. Petioles (8–)12–15(–17.5) cm long, margins vaginate for ½ to 4/5 of the length of the petiole,
glabrous, slightly canaliculate. Leaf blades uniform in shape and size along all axes, (32–)35–48(–53) × (16–)20–(33–)37
cm long including the basal extension, elliptic, base cordate, inequilateral on both monopodial and sympodial axes,
sinus closed, with one lobe (5–)7–9 cm long and exceeding the other, the longest lobe rotundate and fully overlapping
the petiole, the shorter one rotundate and shorter than the petiole, base insertion symmetric, blade basally and medially
symmetric, glabrous on both sides, coriaceous, pinnately nerved from the lower half of the midvein with one or two
minor veins along the distal third, 6–7 pairs of secondary nerves, festooned brochidodromous, the three or four proximal
pairs diverging between the base and the lower third, the fifth and sixth pair diverging from the distal third; higher
order veins reticulate, apex acuminate, leaf blade fresh dark green above, light green below in living material, light
yellow to olive green when dry. Inflorescences and infructescences simple spikes, terminal, pendulous; peduncles
6–7 × 2–3 mm, glabrous, light green; rachis (27–)33–43 cm long; fruits sessile, densely arranged. Floral bracts
cucullate at anthesis, triangular to rectangular when seen from above, 1.8–2.1 × 0.7–0.9 mm, forming bands around the
rachis, fimbriate. Flowers with four stamens, filament 4.1–4.6 mm long, anthers 0.5 × 0.2 mm, with two thecae, with
longitudinal dehiscence and a pellucidly dotted connective; stigmas three, on a 0.4–0.7 mm long style. Fruits drupes,
pyramidal, apically obtuse, 2.4–2.6 × 1–1.3 mm, immersed in rachis, papillate to rugulose, with aristate, 0.6–0.9 mm
long style, 3–4 stigmas to 0.6 mm long, glabrous, black when dry and light green in fresh material.
Taxonomic history:—Piper cajambrense var. caquetanum Trel. & Yunk. (1950: 27) was published by Trelease
and Yuncker (1950) based on a specimen from the municipality La Montañita on Río Orteguaza, department of Caquetá
in Colombia (G. Woronow & S. Juzepczuk 6364, ILL). Based on examination of the original herbarium material and
new field collections made in the vicinity of the type locality, we provide morphological and molecular evidence that
suggest P. cajambrense var. caquetanum should be treated as a separate species. It can be distinguished from the typical
variety, P. cajambrense var. cajambrense, in having glabrous leaves and petioles without epidermal warty outgrowths.
The geographic distributions of the two entities are distinct; P. cajambrense var. cajambrense occur in the rainforests
of the Pacific region of Colombia below 50 m elevation, whereas P. cajambrense var. caquetanum, here elevated to
species as P. oteguanum, occurs in the premontane forest and foothills on the Amazonian slope of the Andes. Apart from
our own collections from Colombia, it is also known from two locations in Ecuador. Since the epithet “caquetanum”
is already in use as Piper caquetanum Yunck. (1957: 530), a new name is presented. The new epithet, oteguanum,
honors the Otegua indigenous people who inhabited the headwaters of the Orteguaza River in Caquetá, and from which

Over- and under-described: piper Phytotaxa 601 (3) © 2023 Magnolia Press • 273
the river derives its name. The Otegua lived in the region during the 16th century and the Spanish conquest. The type
specimen of this species was collected in a locality on the Orteguaza River.

FIGURE 7. Piper oteguanum. A. Sympodial branch. B. Detail of papillae on leaf abaxial surface. C. Petiole on sympodial branch. D.
Petiole on monopodial branch. E. Detail of infructescence. F. Floral bract adaxial view. G. Stamen. H. Detail of the anthers and connective.
I. Fruit lateral view. J. Floral diagram. Illustration by Marcela Morales based on W. Trujillo & C. Malambo 2406 (COL).

274 • Phytotaxa 601 (3) © 2023 Magnolia Press TRUJILLO et al.

FIGURE 8. Piper oteguanum A. monopodial and sympodial branches and inflorescences. B. Detail of flowering spike showing long
filaments and stigmas. Photos by William Trujillo based on W. Trujillo 2075 (COAH).

Distribution:—Piper oteguanum is distributed along the Amazonian slope of the Andes of Colombia and Ecuador,
at 600–1500 m elevation in the Tropical premontane wet forest (TPwf).
Habitat and ecology:—It is a shade-loving species in the understory of well-preserved and secondary forests.
Phenology:—Flowering specimens were collected in February, August, and October and fruiting specimens were
collected in May, July, and October and.
Conservation status:—This species is known from eight collections representing three subpopulations. The
locations are threatened by deforestation and expansion of the agricultural frontier. The extent of occurrence (EOO) is

Over- and under-described: piper Phytotaxa 601 (3) © 2023 Magnolia Press • 275
2695 km2 and area of occupancy (AOO) is 20 km2, which, together with the continuing decline in quality of its habitat,
suggests that it is Endangered [EN B1a+B2a].
Phylogenetic relationships:—Piper oteguanum belongs to the Macrostachys clade, which is a group of shrubs or
treelets with sheathing petioles 2/3 or entire length on monopodial and sympodial branches, pinnately nerved leaves,
and mostly long inflorescences with densely arranged flowers forming bands around the rachis, flowers with four long
stamens (Jaramillo et al. 2008). Piper oteguanum together with P. obtusilimbum C. DC. (1905: 105), P. marsupiiferum
Trel. (1936: 189), and P. cochleatum Sodiro (1905: 203) form a monophyletic clade with strong support (Figure 1,
BS=99). Species in this subclade are distributed in northwestern Amazonia in Colombia, Ecuador, and Peru. These taxa
have a strongly asymmetrical leaf base, with one side of the leaf forming a pronounced lobe that overlaps the petiole.
Piper oteguanum is distinct from the others in having glabrate leaves. Furthermore, our phylogenetic reconstructions
shows that P. cajambrense, from the Pacific coast of Colombia, is not closely related to these Amazonian taxa (Figure
1). On the contrary, P. cajambrense shares a recent common ancestor with other species distributed in the Pacific
lowlands of Colombia such as P. spoliatum Trel. & Yunck. (1950: 118) (Figure 1, BS=85).
Comments:—The four species in the Macrostachys clade to which P. oteguanum is related have stylose fruits and
long stigmas. Piper oteguanum differs from P. obtusilimbum and P. cochleatum in having glabrous leaves and it differs
from the similar P. marsupiiferum in having smooth leaves vs. bullate leaves.

Key to Piper oteguanum and related species in the Macrostachys clade

1. Leaves bullate............................................................................................................................................................ P. marsupiiferum

– Leaves smooth.....................................................................................................................................................................................2
2. Abaxial leaf surfaces, petioles, and internodes glabrous................................................................................................ P. oteguanum
– Abaxial leaf surfaces, petioles, and internodes pubescent..................................................................................................................3
3. Peduncles 4–5 cm long, rachis > 8 cm long................................................................................................................ P. obtusilimbum
– Peduncles 2.5 cm long, rachis < 5 cm long.................................................................................................................... P. cochleatum

Additional specimens examined:—COLOMBIA: Caquetá Belén de los Andakíes: Parque Natural Municipal
Andakí, La Mina, Las Verdes canyon, 1°37’50”N, 75°54’23”W, 730 m, 1 February 2017 [fl], N. Castaño et al. 9322
(COAH!). Florencia: Trail to Sucre, Finca campamento Sucre, 1°46’52”N, 75°39’5.1”W, 1050 m, 5 July 2012 [fr], W.
Trujillo & C. Malambo 2406 (COL!); El Caraño, Las Brisas farm, 1°44’14.7”N, 75°40’35.3”W, 1116 m, 18 October
2013 [fl], W. Trujillo et al. 3007 (COL!); El Caraño, Las Brisas farm, 1°44’14.5”N, 75°40’35.4”W, 1100 m, 10 May
2012 [fr], W. Trujillo & O. Perdomo 2075 (COAH!, COL!); El Caraño, trail to Quindío, 550 m, 16 July 2014 [fr], W.
Trujillo et al. 3183 (COL!); El Caraño, trail to Sucre, 1076 m, 1°47’5.8”N, 75°38’50.5”W, 8 July 2014, W. Trujillo
et al. 3189 (COL!). ECUADOR: Napo-Pastaza, between Tena and Archidona, 9 October 1939 [fr], E. Asplund 9176
(NY!); Napo, Misahualli, along Misahualli river, 1°02’S, 77°40’W, 12 August 1990 [fl], B. Bennett 4489 (NY!).

Piper rubrifolium W. Trujillo-C. & M.A.Jaram. sp. nov., Figs. 1, 2, 9, 10.

Type:—COLOMBIA: Caquetá, Florencia, trail to Tarqui, La Ruidosa ravine, 1°52’8”N, 75°39’40”W, 2027 m, 16 July 2014 [fr], W.
Trujillo et al. 3139 (holotype COAH!].
Diagnosis:— Piper rubrifolium is closely related P. sabaletasanum Trel. & Yunck. (1950: 104), P. spoliatum (1950: 118) and P. cajambrense.
It differs from P. cajambrense and P. spoliatum in having smooth internodes and petioles (vs. internodes and petioles with epidermal
warty outgrowths in the latter). It can also be easily differentiated from P. sabaletasanum by its smooth leaves vs. bullate leaves in
the latter. Piper rubrifolium has young leaves that are red on the underside, another trait that characterizes this species.

Shrub to 4 m tall. Internodes smooth, green to red, glabrous to puberulent, idioblasts not evident in any part of the
plant. Prophylls lacking. Petioles uniform in shape and size along all axes, 4–6 cm long, vaginate along their entire
length, smooth, glabrous to puberulent, green to burgundy colored. Leaf blades chartaceous, brown when dry and
red when fresh on the underside in young leaves, 12.5–16 × 29–37 cm, variable in shape along axes; on monopodial
axes, blade ovate, medially symmetric, base lobate, with lobes 1–1.5 cm long, symmetric, and not overlapping the
petiole, sinus open, equilateral, basal insertion symmetric; on sympodial axes, blade elliptic, medially symmetric, base
auriculate, lobes asymmetric, the shorter lobe 0.7–3 cm long, the other lobe 2–2.2 cm longer, the longer one partially
overlapping the petiole, sinus closed, inequilateral, insertion symmetric; leaf blade smooth, glabrous to puberulent on
the abaxial surface, eciliate; pinnately nerved from the lower half to 3/4 of the midvein; secondary veins 4–5 pairs,
festooned brochidodromous, with spacing decreasing and angle increasing towards the base; tertiary veins randomly

276 • Phytotaxa 601 (3) © 2023 Magnolia Press TRUJILLO et al.

reticulate to percurrent; apex acuminate. Inflorescences and infructescences simple spikes, terminal, erect; peduncles
3–4 cm long, glabrous, green; flowering rachis 9–14 cm and fruiting rachis 25–28 cm long, fruits densely grouped
along the rachis. Floral bracts cucullate, triangular when seen from above, 0.6–0.8 × 0.3–0.5 mm, glabrous on the
adaxial surface and margin, not forming bands around the spike. Flowers with four stamens, filaments 0.8–1.2 mm
long, anthers 0.10–0.16 × 0.08–0.12 mm, longitudinal dehiscent, with two thecae, with connective not protruding,
glabrate; stigmas 3, 0.03–0.07 mm long, sessile. Fruits one-seeded berries, globose, 2–4 × 1.8–4 mm, glabrous,
partially immersed in the rachis, with stigmas persistent, 0.03–0.07 mm long, sessile, green when alive and black when
dry. Seeds ovoid, dark brown.

FIGURE 9. Piper rubrifolium. A. Sympodial branch with a spike near the apex. B. Fruit lateral view. C. Detail of the infructescence. D.
Floral bract from above. E. Floral diagram. Illustration by Marcela Morales based on the type specimen, W. Trujillo et al. 3139 (COAH).

Over- and under-described: piper Phytotaxa 601 (3) © 2023 Magnolia Press • 277
FIGURE 10. Piper rubrifolium. A. Sympodial branch showing abaxial and adaxial leaf surfaces. B. Abaxial and adaxial leaf surfaces,
petioles and a young spike. C. Fruiting spike. Photos by William Trujillo from the type, W. Trujillo 3139 (COAH).

Distribution:—Piper rubrifolium is endemic to the eastern cordillera of the Andes along the slopes towards the
Amazon basin at 1500–2000 m elevation, in the Tropical montane rain forest (TMrf) .
Habitat and ecology:—It is a shade-loving species growing in the understory of rainforests.
Phenology:—Flowering and fruiting specimen were collected in July.
Etymology:—The name Piper rubrifolium is derived from the Latin rubrum meaning red and folia meaning
leaves, referring to burgundy color underneath on young leaves.
Conservation status:—This species is known from three specimens representing two populations. Its habitat is
threatened by deforestation and expansion of the agricultural frontier. The extent of occurrence (EOO) is 398 km2 and
its area of occupancy (AOO) is 12 km2, which, together with the continuing decline in quality of the habitat, suggests
that it is Endangered [EN B1a+B2a].

278 • Phytotaxa 601 (3) © 2023 Magnolia Press TRUJILLO et al.

 Phylogenetic relationships:—Piper rubrifolium belongs to the Macrostachys clade (Jaramillo et al. 2008). In our
phylogeny this species forms a monophyletic group with taxa known to occur in the Chocó region, i.e., P. spoliatum,
P. sabaletasanum, and P. cajambrense (Figure 1).
Comments:—Piper rubrifolium is red on the abaxial surface of immature leaves and mature petioles, the petioles
and internodes are smooth without epidermal warty outgrowths, and the leaf base has short lobes which are the most
important characteristics that separate Piper rubrifolium from similar species.

Key to Piper rubrifolium and related species in the Macrostachys clade

1. Leaves bullate............................................................................................................................................................P. sabaletasanum

– Leaves smooth.....................................................................................................................................................................................2
2. Petioles with epidermal warty outgrowths..........................................................................................................................................3
– Petioles smooth (without epidermal warty outgrowths)................................................................................................. P. rubrifolium
3. Leaves oblanceolate, narrowed to the base....................................................................................................................... P. spoliatum
– Leaves broad–elliptic, wide at the base........................................................................................................................ P. cajambrense

Additional specimens examined:—COLOMBIA: Caquetá, Florencia, trail to Campo Hermoso Alto, 3rd tunnel,
1°44’56”N, 75°44’36”W, 2037 m, 19 March 2020 [sterile], W. Trujillo & L. Sandoval 4029 (UMNG-H!). Putumayo,
Mocoa, road Sibundoy-Mocoa at El Mirador, 1°04’11”N, 76°44’41”W, 2000 m, 9 July 1998 [fl], H. Mendoza & F.
Quevedo 6035 (HUAZ!).

New synonyms

Piper cuniculorum Trel. & Yunck., The Piperaceae of northern South America 1: 234. 1950.

Type:—COLOMBIA: Putumayo, Colombia–Ecuador border, near the confluence of the San Miguel and Conejo rivers, 300 m, J.
Cuatrecasas 10906 (holotype US!, isotype ILL!).
= Piper papillicaule Trel. & Yunck., The Piperaceae of northern South America 1: 224. 1950. Type:—COLOMBIA: Putumayo, Colombia–
Ecuador border at the intersection of rivers San Miguel and Conejo, 300 m, J. Cuatrecasas 10917 (holotype US!, isotype COL!)
syn. nov.

Comments:—In the The Piperaceae of northern South America (Trelease & Yuncker 1950) both P. cuniculorum Trel.
& Yunck. (1950:234) and P. papillicaule Trel. & Yunck. (1950:224) are described as having leaves that are less than
20 cm long, pinnately nerved from the lower half of the midvein, and glabrous on both sides. Both species were
separated from other species in the diagnosis and the key by the combination of having 4–5 secondary nerves on
each side, bracts peltate and short-fringed, and 10 cm long spikes. In addition, other characters are shared by the two
species, for example, both have glabrous leaves, discrete papillae on the internodes as seen on type specimens, even
if not mentioned in the description, and they are pinnately nerved from the lower half of the midvein, their spikes are
2–4 mm thick × 10 cm long and their fruits are obpyriform with sessile stigmas. Detailed examination of the two type
collections revealed their similarities except for the length of the internodes (2–6 cm long in P. papillicaule vs. 1.5–3.5
cm long in P. cuniculorum), which is a character that varies with age of the branch or the nodes. Additionally, the type
specimens of both P. cuniculorum and P. papillicaule were collected at the Colombia–Ecuador border at the junction of
the rivers San Miguel and Conejo. Our conclusion is that there are no consistent morphological characters that justify
a separation of two species, and thus P. papillicaule is synonymized under P. cuniculorum. Since both names were
published in the same publication, we selected P. cuniculorum as the accepted name because it has the most complete
type specimen.
Distribution and habitat:—This species is distributed along the slopes of the Andes towards the Amazon in
Colombia and Ecuador, at 300–1100 m elevation in Tropical premontane wet forests (TPwf). It is a heliophilous
species that grows along roadsides.
Additional specimens examined: —COLOMBIA: Caquetá, Florencia, near Universidad de la Amazonia,
1°36’23.45”N, 75°39’59”W, 270 m, 14 September 2009, J. Alzate 8 (HUAZ!); First bridge in urban area, middle basin
of Hacha river, 1°38’707”N, 75°36’978”W, 318 m, 28 March 2005, C. Blanco et al. 267 (HUAZ!). Puerto Rico, trail to
San Rafael, Buena Vista farm, 1340 m, 1°54’21.7”N, 75°15’40.4”W, 23 March 2002, J. Díaz et al. 370 (COAH!); Trail
to La Estrella, Pensilvania farm, 550 m, 1°56’14.5”N, 75°12’32.3”W, 22 March 2002, M. Correa et al. 3040 (COAH!).

Over- and under-described: piper Phytotaxa 601 (3) © 2023 Magnolia Press • 279
San Vicente del Caguan: Road from Neiva to trail to Campanas, 725 m, 2°33’50”N, 74°45’26”W, 04 January 2011, W.
Trujillo et al. 1801 (COAH!).

Piper mituense Trel. & Yunck. The Piperaceae of northern South America 1: 382. 1950.

Type:—COLOMBIA: Vaupés, Mitú, 200 m, 14 September 1939, E. Pérez Arbeláez & J. Cuatrecasas 6810A (holotype US!).
= Piper metanum var. villibracteum Trel. & Yunck. The Piperaceae of northern South America 1: 343. 1950. Type:—COLOMBIA: Meta,
Mitú, at Vaupés boundary, 200 m, 14 September 1939, E. Pérez-Arbeláez & J. Cuatrecasas 6810 (holotype US!). syn. nov.

Comments:—Piper mituense Trel. & Yunck. (1950: 382) was described from the collection E. Pérez Arbelaez and J.
Cuatrecasas 6810 and P. metanum var. villibracteum Trel. & Yunck. (1950: 343) was described from E. Pérez-Arbelaez
and J. Cuatrecasas 6810A. The two collections were made at the same locality and on the same day. The protologue of
P. mituense highlighted: “the villous twigs, pinnately nerved, oblong-elliptic leaves, and short spikes with substylose
ovaries characterize this species”, while for P. metanum var. villibracteum the authors note: “Differs from the species
in having somewhat less pubescent twigs, slightly thicker leaves, and with the floral bracts rounded and densely
long-fringed”. In addition, P. metanum is remarkably different from the variety because it has spikes 4–6 cm long and
erect (vs. spikes 1–1.8 cm long and pendulous). The protologues do not recognize the similarity between the two taxa
discussed here. After a detailed examination, we conclude that there are no notable morphological differences that
allow considering the two specimens as belonging to different species, for which reason we propose to synonymize the
variety P. metanum var. villibracteum under the species P. mituense, giving priority to the species name.
Distribution and habitat:—Piper mituense is distributed in the Amazon of Colombia and Ecuador, at elevations
below 500 m. It is a shade-loving species growing in the understory.
Additional specimens examined:—COLOMBIA: Caquetá, Albania, trail to Florida 1, El Porvenir farm, 13 March
2010, W. Trujillo et al. 1358 (HUAZ!). Florencia, trail to Sebastopol, 5 September 2010, L. Zúñiga 02 (HUAZ!); Center
Macagual, Monitoreo research station, 30 June 2008, M. Correa et al. 6648 (HUAZ!); San Martin, trail to El Venado,
Macagual Research Center of Universidad de la Amazonia, 246 m, 01°29’59.3”N, 75°39’21.9”W, 30 September 2011,
M. Correa 7120 (HUAZ!). Milán: Bombay, La Juansoco canyon, 230 m, 1°13’54.9”N, 75°29’30.3”W, 18 September
2007, W. Trujillo et al. 713 (HUAZ!, COAH!). ECUADOR: Napo, Añangu, south bank of Río Napo 95 km downstream
from Coca, 246 m, 0°32’S, 76°23’W, 4 July 1985, H. Balslev et al. 60620 (AAU!).

Piper morelianum Yunck., Svensk Botanisk Tidskrift 51: 532. 1957. Fig. 11.

Type:—COLOMBIA: Caquetá, Morelia, 300 m. K. Sneidern, 1195 (holotype S!).

= Piper calanyanum var. sardinanum Trel. & Yunck., The Piperaceae of northern South America 1: 67. 1950. Type:—COLOMBIA:
Caquetá, Florencia, Cerro de La Sardina, 500 m, J. Cuatrecasas 8900 (US!). syn. nov.

Comments:—In The Piperaceae of northern South America, Trelease and Yuncker published P. calanyanum var.
sardinanum Trel. & Yunck. (1950: 67) based on a specimen collected at Florencia in the Department of Caquetá,
Colombia, giving the following diagnostic characters: “subshrub, 1–2 m tall, upper leaves 4–5 cm wide × 15–18 cm
long, lower leaves to 7.5 cm wide × 20 cm long, upper petioles short and winged by the decurrent blade”. Detailed
examination of the type specimens of P. calanyanum Trel. & Yunck. (1950: 66) and P. calanyanum var. sardinanum, the
latter species has long apiculate spikes, a character not shared by the typical variety P. calanyanum var. calanyanum.
Yuncker described P. morelianum Yunck. (1957: 532) based on a specimen collected at Morelia in the same Department
of Caquetá, Colombia, approximately 20 km to the southwest from the type locality of P. calanyanum var. sardinanum.
In the diagnosis, Yuncker recognized in P. morelianum the morphological similarity with P. parapeltobryon Trel. ex
Gleason (1931: 356), and he pointed to the following differences: “but differs in its leaf shape and acute to obtusish
rather than wedge-shaped base and with the petiole vaginate to the blade.” These differences are notable, but Yuncker
missed the similarities of P morelianum with P. calanyanum var. sardinanum. When examining the type specimens for
both taxa they share the persistent petiolar margin, a decurrent leaf base on the petiole, secondary nerves irregularly
spaced, prophylls 2–2.5 cm long, and long apiculate pendular spikes. Specimens of P. morelianum, that we collected
in the Department of Caquetá over the past few years, were all found in the understory and at forest edges, and they
all had red peduncles and adaxially lustrous leaves. Based on this collective evidence we conclude, that P. calanyanum
var. sardinanum should be treated as a synonym P. morelianum.

280 • Phytotaxa 601 (3) © 2023 Magnolia Press TRUJILLO et al.

FIGURE 11. Piper morelianum. A. Sympodial branch; showing adaxial and abaxial leaf surfaces and spikes. B. Infructescence with red-
wine colored peduncle. Photos by W. Trujillo from W. Trujillo 1086 (COAH).

Piper calanyanum and P. morelianum are part of the Peltobryon clade (Fig. 1, Jaramillo et al 2008), a group of
shrubs with pinnately nerved leaves along the entire midvein length, spikes erect or pendular, flowers forming banded
patterns around the spikes. However, these species are not closely related.
Distribution and Habitat:—Piper morelianum is distributed along the eastern slopes of the Andes towards the
Amazon in Colombia. It is a local endemic, known only from the Department of Caqueta at elevations of 500–1100 m.
It is a shade-loving species growing in the understory.
Additional specimens examined: —COLOMBIA: Caquetá, Albania, trail to Florida 1, El Porvenir farm, 13
March 2010, W. Trujillo et al. 1359 (HUAZ!). Belén de los Andaquies, Parque Bosque Microcuenca La Resaca,

Over- and under-described: piper Phytotaxa 601 (3) © 2023 Magnolia Press • 281
1°26’23.5”N, 75°53’24.5”W, 482 m, 11 October 2007, W. Trujillo 1086 (COAH!, HUAZ!); Road from los Ángeles to
trail to las Verdes, 450–650m, 1°35’21”N, 75°52’7.3”W, 22 June 2011, W. Trujillo et al. 1978 (COAH!). Curillo, trail
to Los Conquistadores, 240 m, 1°1’16.8”N, 75°54’50”W, 29 April 2010, W. Trujillo et al. 1651 (HUAZ!). Florencia,
Research center Macagual, 1°37’N, 75°31’W, 260 m, 1 June 2012, M. Gordillo 029 (HUAZ!); San Antonio de Atenas,
Embera Chami indigenous camp, 1°44’58”N, 75°34’50”W, 600 m, 19 March 2012, D. Castro et al. 47 (HUAZ!);
Trail to La Estrella, 01°36.377’N, 75°34.824’W, 390 m, 18 October 2003, M. Correa et al. 3729 (HUAZ!); Las Brisas
farm, trail to Santander, road to Morelia, 450–550 m, 1°31’59”N, 75°42’23”W, 30 June 2010, W. Trujillo et al. 1672
(COAH!). La Montañita, trail to Itarca, 330 m, 1°32’34.5”N, 75°28’19”W, 26 April 2011, N. Castaño et al. 3140
(COAH!).

Discussion

Continued systematic sampling of Piper is needed in eastern slopes of the northern Andes, in Colombia, Ecuador
and Peru. On the one hand, it will unveil the diversity along the northern slopes towards the Llanos, a region largely
unexplored because of the armed conflict in Colombia. On the other hand, to the south in Ecuador and Peru, to
integrate the knowledge across countries.

Acknowledgements

We thank Marcela Morales for the botanical illustrations and Viviana Vargas for preparing Figures 2, 4, 6, 8, 9 and
10 and Arturo Salas and Yasodara Salas for help with the etymology of the new Piper names. WT’s work in the
AAU herbarium was supported by the project Amazonian Biogeography which is funded by the Danish Council for
Independent Research - Natural Sciences, grant #9040-00136B to HB. Sequences were partly funded by the Fundación
para la Promoción de la Investigación y la Tecnología project # 4674. Gencore (Sequencing Center) at Los Andes
University partially subsided the sequencing analysis. Laboratorio de Agrobiotecnología of the Universidad de la
Amazonia generated part of the sequences used in this study.

References

Bachman, S., Moat, J., Hill, A.W., de Torre, J. & Scott, B. (2011) Supporting Red List threat assessments with GeoCAT: Geospatial
conservation assessment tool. ZooKeys 150: 117–126.
https://doi.org/10.3897/zookeys.150.2109
Barthlott, W., Mutke, J., Rafiqpor, D., Kier, G. & Kret, H. (2005) Global Centers of Vascular Plant Diversity. Nova Acta Leopoldina 92:
61–83.
Bernal, R., Gradstein, S.R. & Celis, M. (eds.) (2019) Catálogo de plantas y líquenes de Colombia. Instituto de Ciencias Naturales,
Universidad Nacional de Colombia, Bogotá. Available from: http://catalogoplantasdecolombia.unal.edu.co (accessed 8 May 2022)
Brako, L. & Zarucchi, J.L. (1993) Catalogue of the flowering plants and gymnosperms of Peru. Monogr. Monographs in Systematic
Botany from the Missouri Botanical Garden 45: 1–1286.
Callejas-Posada, R. (2020) Piperaceae. In: Davidse, G., Ulloa-Ulloa, C., Hernández-Macías, H.M. & Knapp, S. (eds.) Flora Mesoamericana.
Missouri Botanical Garden Press, Saint Louis.
De Candolle, C. (1866) Piperaceae novae. Journal of Botany. British and Foreign 4: 210–219.
De Candolle, C. (1869) Piperaceae. Prodromus Systematis Naturalis Regni Vegetabilis 16: 235–471. [Paris]
De Candolle, C. (1891) Primitiae Florae Costarricensis. Premier Fascicule, Piperaceae. Bulletin de la Société Royale de Botanique de
Belgique 30: 196–235.
De Candolle, C. (1905) Piperaceae. Piper L. Verhandlungen des botanischen Vereins der Provinz Brandenburg 74: 105–106.
Denslow, J.S., Chaverri, S. & Vargas, R.O. (2019) Patterns in a species‐rich tropical understory plant community. Biotropica 51: 664–
673.
https://doi.org/10.1111/btp.12704
Doyle, J.J. & Doyle, J.L. (1987) A rapid DNA isolation procedure for small quantities of fresh leaf tissue. Phytochemical Bulletin 19:
11–15.

282 • Phytotaxa 601 (3) © 2023 Magnolia Press TRUJILLO et al.

Draper, F. C., Costa, F. R. C., Arellano, G., Phillips, O. L., Duque, A., Macía, M. J., ter Steege, H., Asner, G. P., Berenguer, E., Schietti, J.,
Socolar, J. B., Souza, F. C. D., Dexter, K. G., Jørgensen, P. M., Tello, J. S., Magnusson, W. E., Castilho, C. V., Monteagudo-Mendoza,
A., Fine, P. V. A., Ruokolainen, K., Honorio Coronado, E. N., Aymard, G., Dávila, N., Sánchez Sáenz, M., Rios Paredes, M. A., Engel,
J., Fortunel, C., Paine, C. E. T., Goret, J.-Y., Dourdain, A., Petronelli, P., Allie, E., Guevara Andino, J. E., Brienen, R. J. W., Pérez,
L. C., Manzatto, Â. G., Paniagua Zambrana, N. Y., Molino, J.-F., Sabatier, D., Chave, J., Fauset, S., Garcia Villacorta, R., Réjou-
Méchain, M., Berry, P. E., Melgaço, K., Feldpausch, T. R., Sandoval, E. V., Vasquez Martinez, R., Mesones, I., Junqueira, A. B.,
Roucoux, K. H., de Toledo, J. J., Andrade, A. C., Camargo, J. L., del Aguila Pasquel, J., Santana, F. D., Laurance, W. F., Laurance, S.
G., Lovejoy, T. E., Comiskey, J. A., Galbraith, D. R., Kalamandeen, M., Navarro Aguilar, G. E., Vega Arenas, J., Amasifuen Guerra,
C. A., Flores, M., Flores Llampazo, G., Torres Montenegro, L. A., Zarate Gomez, R., Pansonato, M. P., Moscoso, V. C., Vleminckx,
J., Valverde Barrantes, O. J., Duivenvoorden, J. F., de Sousa, S. A., Arroyo, L., Perdiz, R. O., Cravo, J. S., Marimon, B. S., Marimon
Junior, B. H., Carvalho, F. A., Damasco, G., Disney, M., Vital, M. S., Stevenson Diaz, P. R., Vicentini, A., Nascimento, H., Higuchi,
N., Van Andel, T., Malhi, Y., Ribeiro, S. C., Terborgh, J. W., Thomas, R. S., Dallmeier, F., Prieto, A., Hilário, R. R., Salomão, R. P.,
da Costa Silva, R., Casas, L. F., Guimarães Vieira, I. C., Araujo-Murakami, A., Ramirez Arevalo, F., Ramírez-Angulo, H., Vilanova
Torre, E., Peñuela, M. C., Killeen, T. J., Pardo, G., Jimenez-Rojas, E., Castro, W., Galiano Cabrera, D., Pipoly, J., de Sousa, T. R.,
Silvera, M., Vos, V., Neill, D., Núñez Vargas, P., Vela, D. M., Aragão, L. E. O. C., Umetsu, R. K., Sierra, R., Wang, O., Young, K. R.,
Prestes, N. C. C. S., Massi, K. G., Huaymacari, J. R., Parada Gutierrez, G. A., Aldana, A. M., Alexiades, M. N., Baccaro, F., Céron,
C., Muelbert, A. E., Grandez Rios, J. M., Lima, A. S., Lloyd, J. L., Pitman, N. C. A., Valenzuela Gamarra, L., Cordova Oroche, C.
J., Fuentes, A. F., Palacios, W., Patiño, S., Torres-Lezama, A. & Baraloto, C. (2021) Amazon tree dominance across forest strata.
Nature Ecology & Evolution 5: 757–767.
https://doi.org/10.1038/s41559-021-01418-y
Ellis, B., Daly, D., Hickey, L.J., Mitchell, J., Wilf, P. & Wing, S. (2009) Manual of leaf architecture. Cornell, New York, 190 pp.
https://doi.org/10.1079/9781845935849.0000
Fleming, T.H. (2004) Dispersal ecology of Neotropical Piper shrubs and treelets. In: Dyer, L.A. & Palmer, A.D.N. (eds.) Piper: a model
genus for studies of phytochemistry, ecology, and evolution. Kluwer Academic-Plenum Publishers, New York, pp. 58–75.
https://doi.org/10.1007/978-0-387-30599-8_4
Frodin, D.G. (2004) History and Concepts of Big Plant Genera. Taxon 53: 753–776.
https://doi.org/10.2307/4135449
Gentry, A.H. (1990) Floristic Similarities and differences between southern Central America and upper and central Amazonia. In: Gentry,
A.H. (ed.) Four Neotropical Rainforests. Yale University Press, New Haven.
Gleason, H.E. (1931) Botanical Results of the Tyler-Duida Expedition. Bulletin of the Torrey Botanical Club 58: 356.
Holdridge, L.R. (1978) Ecología basada en zonas de vida. IICA. Serie de libros y materiales educativos. San José-Costa Rica, 231 pp.
Jaramillo, M.A. & Manos, P.S. (2001) Phylogeny and the patterns of floral diversity in the genus Piper (Piperaceae). American Journal
of Botany 88: 706–716.
https://doi.org/10.2307/2657072
Jaramillo, M.A, Callejas-Posada, R., Davidson, C., Smith, J.F., Stevens, A.C, Tepe, E.J, Editor, C. & Plunkett, G.M. (2008) A phylogeny
of the tropical Genus Piper using ITS and the chloroplast intron psbJ – petA. Systematic Botany 33: 647–660.
https://doi.org/10.1600/036364408786500244
Jardim Botânico do Rio de Janeiro (2020) Flora do Brasil. Available from: http://floradobrasil.jbrj.gov.br/ (accessed 12 January 2021)
Jørgensen, P.M. & León-Yánez, S. (1999) Catalogue of the vascular pants of Ecuador. Monograph in Systematic Botany from the Missouri
Botanical Garden 75: 1–1181.
Jørgensen, P.M., Nee, M.H. & Beck, S.G. (2014) Catálogo de las plantas vasculares de Bolivia. Missouri Botanical Garden Press, St.
Louis Missouri, pp. 1744.
Mgbeahuruike, E.E., Yrjönen, T., Vuorela, H. & Holm, Y. (2017) Bioactive compounds from medicinal plants: Focus on Piper species.
South African Journal of Botany 112: 54–69.
https://doi.org/10.1016/j.sajb.2017.05.007
Parthasarathy, V.A., Sasikumar, B., Nair, R.R. & George, K.J. (2007) Black pepper: botany and horticulture. Horticultural reviews-
westport then New York 33: 173.
https://doi.org/10.1002/9780470168011.ch3
Samain, M., Mathieu, G.,Wanke, S., Neinhuis, C. & Goetghebeur, P. (2008) Verhuellia revisited—unravelling its intricate taxonomic
history and a new subfamilial classification of Piperaceae. Taxon 57: 583–587.
https://doi.org/10.2307/25066024
Sodiro, L. (1905) Contribuciones al conocimiento de la Flora Ecuatoriana. Anales de la Universidad Central del Ecuador 19: 192–204.
https://doi.org/10.5962/bhl.title.45473
Stamatakis, A. (2014) RAxML version 8: A tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics 30:

Over- and under-described: piper Phytotaxa 601 (3) © 2023 Magnolia Press • 283
 1312–1313.
https://doi.org/10.1093/bioinformatics/btu033
Thiers, B. (2019) Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual
Herbarium. Available from: http://sweetgum.nybg.org/science/ih/ (accessed 8 May 2022)
Trelease, W. & Yuncker, T.G. (1950) The Piperaceae of northern South America. University of Illinois Press, Indiana, 434 pp
Trelease, W. (1936) Piperaceae. In: Macbride, J.F. (Ed.) Flora of Peru. Publications of Field Museum of Natural History. Botanical Serie,
Chicago, 253 pp.
Trujillo, W. & González, V.H. (2011) Plantas medicinales utilizadas por tres comunidades indígenas en el noroccidente de la Amazonía
colombiana. Mundo Amazónico 2: 283–305
Trujillo, W. & Callejas, R. (2015) Piper andakiensis (Piperaceae) una especie nueva de la vertiente amazónica de la cordillera oriental de
Colombia. Caldasia 37: 261–269.
https://doi.org/10.15446/caldasia.v37n2.54379
Trujillo, W. & Jaramillo, M.A (2019) An emended description of Piper cochleatum and a new species of Piper (Piperaceae) from the
Amazonian slopes of the Colombian Andes. Phytotaxa 408: 59–68.
https://doi.org/10.11646/phytotaxa.408.1.4
Trujillo, W. & Jaramillo, M.A. (2021) Taxonomic novelties of Piper (Piperaceae) from the Amazonian slopes of the Northern Andes.
Brittonia 73: 67–76.
https://doi.org/10.1007/s12228-020-09638-5
Trujillo, W., Trujillo, E. & Jaramillo, M.A. (2021) Piper callejasii (Piperaceae), a new species from the eastern slopes of the Andes,
Northern South America. Novon: A Journal for Botanical Nomenclature 29 (1): 200–205.
https://doi.org/10.3417/2021659
Trujillo, W. & Vargas, V. (2022a) Las especies de Piper en la vertiente Amazónica de los Andes, Caquetá. Guía de campo. Editorial
Unviersidad de la Amazonia, Florencia Colombia, 268 pp.
Trujillo, W., Trujillo, E, Ortiz-Morea, F., Toro, D. & Jaramillo, M.A. (2022b) 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
Yuncker, T. (1957) New taxa of South American Piperaceae. Svensk Botanisk Tidskrift 51: 529–543.
Yuncker, T. (1959) Four new species of Piperaceae from Ecuador. Svensk Botanisk Tidskrift 53: 45–48.
Vásquez-Ocmín, P.G., Gadea, A., Cojean, S., Marti, G., Pomel, S., Van Baelen, A.C., Ruiz-Vásquez, L., Ruiz Mesia, W., Figadère, B.,
Ruiz Mesia, L. & Maciuk, A. (2021) Metabolomic approach of the antiprotozoal activity of medicinal Piper species used in Peruvian
Amazon. Journal of Ethnopharmacology 264: 113262.
https://doi.org/10.1016/j.jep.2020.113262
Wanke, S., Jaramillo, M.A., Borsch, T., Samain, M., Quandt, D. & Neinhuis, C. (2007) Evolution of Piperales--matK gene and trnK intron
sequence data reveal lineage specific resolution contrast. Molecular phylogenetics and evolution 42: 477–497.
https://doi.org/10.1016/j.ympev.2006.07.007
Yohe, L.R., Leiser-Miller, L.B., Kaliszewska, Z.A., Donat, P., Santana, S.E. & Dávalos, L.M. (2021) Diversity in olfactory receptor
repertoires is associated with dietary specialization in a genus of frugivorous bat. G3 Genes|Genomes|Genetics 11: jkab260.
https://doi.org/10.1093/g3journal/jkab260

284 • Phytotaxa 601 (3) © 2023 Magnolia Press TRUJILLO et al.