Species of Peperomia (Piperaceae) from the Saña River Valley, Peru

Guillermo Eloy Pino Infante; Marie-Stéphanie Samain; Joaquina Adelaida Albán Castillo; Luis Enrique Aarón Alomía Collazos

doi:10.3897/phytokeys.225.99277

Research Article

Species of Peperomia (Piperaceae) from the Saña River Valley, Peru

Guillermo Eloy Pino Infante^‡§, Marie-Stéphanie Samain^|, Joaquina Adelaida Albán Castillo^‡, Luis Enrique Aarón Alomía Collazos^§

‡ Universidad Nacional Mayor de San Marcos, Lima, Peru

§ Sociedad Peruana de Cactáceas y Suculentas, Lima, Peru

| Instituto de Ecología, A.C., Centro Regional del Bajío, Red de Diversidad Biológica del Occidente Mexicano, Pátzcuaro, Mexico

Corresponding author: Guillermo Eloy Pino Infante ( guillermo.pino@unmsm.edu.pe )

Academic editor: M. Alejandra Jaramillo

This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

Citation: Pino Infante GE, Samain M-S, Albán Castillo JA, Alomía Collazos LEA (2023) Species of Peperomia (Piperaceae) from the Saña River Valley, Peru. PhytoKeys 225: 1-40. https://doi.org/10.3897/phytokeys.225.99277

Abstract

The Saña River Valley in Northern Peru is unusual for the western slopes of the Peruvian Andes because of its nearly year-round regime of precipitation instead of marked seasonal dry winters. This results in unexpected plant diversity. We surveyed the species of Peperomia (Piperaceae), occurring in this valley from 300 to 3000 m elevation, based on the study of specimens from ten herbaria and field collections, resulting in a total of 81 accessions, of which 48 were made by the authors. We found 16 different taxa: Peperomia cacaophila, from Ecuador, is reported for the first time in Peru; P. cymbifolia, P. dolabriformis and P. emarginulata are reported for the first time for the Saña River Valley; other widely distributed species like P. fraseri, P. galioides, P. haematolepis, P. hispidula, P. inaequalifolia, P. microphylla, and P. rotundata were also found. Five species new to science are described: P. pilocarpa, P. riosaniensis, close to P. palmiformis from Amazonas; P. sagasteguii, related to P. trinervis, P. symmankii, close to P. ricardofernandezii from Piura, and P. vivipara, related to P. alata. A key to the species of Peperomia from the Saña River Valley, based on vegetative characters, is provided.

Resumen

El valle del Río Saña en el norte de Perú es inusual para las vertientes occidentales de los Andes peruanos por su régimen de precipitación ininterrumpido en lugar de inviernos secos estacionales marcados. Esto resulta en una diversidad de plantas inesperada. Revisamos las especies de Peperomia (Piperaceae), del valle del Río Saña entre los 300 a 3000 m de altitud, basándonos en el estudio de especímenes de diez herbarios y colecciones de campo, resultando en un total de 81 accesiones, de las cuales 48 fueron realizadas por los autores. Encontramos 16 taxones diferentes: Peperomia cacaophila, de Ecuador, se reporta por primera vez en Perú; P. cymbifolia, P. dolabriformis y P. emarginulata se reportan por primera vez para el valle del Río Saña; otras especies ampliamente distribuidas como P. fraseri, P. galioides, P. haematolepis, P. hispidula, P. inaequalifolia, P. microphylla y P. rotundata también se encontraron. Se describen cinco especies nuevas para la ciencia: P. pilocarpa, P. riosaniensis, cercana de P. palmiformis de Amazonas; P. sagasteguii, relacionada con P. trinervis, P. symmankii, cercana a P. ricardofernandezii de Piura, y P. vivipara, relacionada con P. alata. Se presenta una clave para las especies de Peperomia del Valle del Río Saña basada en caracteres vegetativos.

Keywords

Amotape-Huancabamba Zone, Andean cloud forest, Cajamarca, Fenestratae, Lambayeque, Panicularia, western slopes of the Andes

Palabras clave

Bosque montano andino, Cajamarca, Fenestratae, Lambayeque, Panicularia, Vertientes occidentales de los Andes, Zona Amotape-Huancabamba

Introduction

Peperomia (Ruiz & Pavón, 1794), was the name chosen to distinguish some species previously described by Linnaeus as Piper, and its first species were described in Peru (Ruiz and Pavón 1798). It is the second largest genus of the family Piperaceae after Piper, with a pantropical distribution of about 1,600 species (Samain et al. 2009; Mathieu et al. 2011; Samain et al. 2011). Most of them occur in the Neotropics, and approximately 400 species can be found in Peru (Trelease 1936). Due to its high species number, subgenera were proposed and used for classification, based only on morphology (Miquel 1843; Dahlstedt 1900) until this century, when their phylogeny was clarified using molecular studies (Wanke et al. 2006; Samain et al. 2007; Samain et al. 2009). Finally, Frenzke et al. (2015) published a new infrageneric classification as a reference for studies in the genus, based on fruit characters, macroscopic features, and molecular data, defining 14 monophyletic groups, and describing 5 new subgenera. Peru hosts representatives of all subgenera (Frenzke et al. 2015), and it also has the highest diversity of species in the world (Ulloa et al. 2017), many of which are still not described, whereas a considerable number of the currently accepted species are possible synonyms (unpublished results of the first author).

While some of the subgenera are highly diverse and widely distributed (e.g., Micropiper (Miq.) Miq., Leptorhynchum (Dahlst.) Trel. ex Samain, Pseudocupula Frenzke & Scheiris), others are small and restricted to the Neotropics, and even to some Andean countries (Colombia, Ecuador, Peru, and Bolivia) (e.g., Perlucida Scheiris & Frenzke, Phyllobryon (Miq.) Scheiris & Frenzke). Among these small groups, two of them were split from the former subgenus Panicularia sensu Dahlstedt, into two well-supported clades that are well characterized by morphological features. (Panicularia Miq. and Fenestratae Pino) with most of their species adapted to seasonal dry periods through succulence and circumscribed to Peru and Ecuador (Pino et al. 2020). Due to the lack of information, many species of these two groups need to be reviewed and new species are to be described. In the quest for these novelties, we explored the Saña River Valley, in the north of Peru, where we found many taxa together in a small area, that was not known to be diverse for Peperomia. Therefore, the objective of the present paper is to review the species of Peperomia present in the Saña River Valley, Peru, as a sample of the high diversity of this species existing in the country.

Materials and methods

Study area

The Saña (also spelled Zaña) River is a short course (ca. 125 km length) river on the western slopes of the northern Peruvian Andes, with two main tributaries: Udima River and Nanchoc River. Altogether they make up a total drainage area of 1754.7 m² (ANA 2010). The river rises in the district Calquis, province San Miguel, department Cajamarca, at approximately 3200 m elevation (ANA 1999), notoriously lower than similar rivers draining the Pacific Ocean basin.

After crossing the districts of Calquis and La Florida in the province of San Miguel and the district of Catache in the province of Santa Cruz (both provinces of the Cajamarca department), it enters the Oyotún district of the province of Chiclayo, Lambayeque department, after which it crosses the districts Nueva Arica, Cayaltí, Zaña, and Lagunas, as it flows to the sea across the coastal desert of the southernmost part of the Lambayeque department, close to the border with the department of La Libertad (Fig. 1).

Figure 1.

Study area A map of Peru and the localization of the departments Lambayeque and Cajamarca B close-up to show the Saña River and its basin (light green) shared by the provinces of Chiclayo (Lambayeque), San Miguel, and Santa Cruz (Cajamarca).

Unlike the Chancay-Reque River system located just north and the Jequetepeque River Valley South of it–both with typical coastal desertic valleys–the Saña River Valley is more humid on average, due to the influence of the Amotape-Huancabamba Zone (AHZ) (observation of the first author). This zone has been proposed as a biodiversity hotspot for its higher diversity levels more than adjacent zones (Weigend 2002). Its boundaries have been set from Río Jubones and Río Zamora systems in Ecuador to the Río Chicama system in Peru (La Libertad). Thus, its influence reaches the highlands of Lambayeque and particularly the Saña River Valley, where the humid forests on the western slopes of the Andes can only be explained by the relatively low elevation of the Andes in the AHZ, which allows the flow of moisture from the rainforests of the Amazon Basin on the east to the desert coasts of western Peru. Plants and animals typical of the Amazon Forest e.g. military macaws (Ara militaris L.) were reported here at the highest altitudes of the Saña River Valley (Taulis) before the anthropogenic deforestation of recent decades, (Koepcke 1961). Native forests of its middle elevations have been displaced by cultivated patches of the tropical bamboo Guadua angustifolia Kunth (Caña Guayaquil) which is extensively cultivated in northern Peru, but not in the adjacent valleys.

From the coast to the Andes, close to the riverbanks of the Saña River, from west to east, we find the localities of Lagunas, Mocupe, Zaña, Cayaltí, La Viña, Culpón, and Nueva Arica. After a detour of the road to the town of Nanchoc in the South, the river flows northwards to the town of Oyotún and Pan de Azúcar, turning later southeastward to El Espinal and La Florida, where a detour to the South moves the road away from the river to El Naranjo and Niepos. Beyond Nanchoc, there is a small relict forest called “La Oscurana” after the town of Bolívar, with four species of Peperomia reported, although only one was found in the Herbaria (Juárez et al. 2005). From La Florida, another small road continues east and then southwards along the Saña River leading to the former Taulis Estate, now a hamlet and no longer a cloud forest due to intense cultivation. From El Espinal on a short walk northwards, we find the Velo de Novia waterfall. Before reaching La Florida, after the bridge El Papayo over the Saña River, a detour to the northeast leads to the town of Monteseco, a former estate located south of a relict forest. From here, a trail leads to El Chorro Blanco waterfall and the former Udima Estate crossing the forest. River Udima, a northeastern affluent of Saña River, crosses the forest from east to west and flows into it at El Espinal.

The fauna and flora of the remaining forest North of Monteseco have been published by Cadle (1989), Cadle and McDiarmid (1990), and Sagástegui and Dillon (1991). The latter report 88 families, 200 genera, and over 326 species of ferns and flowering plants, among them nine species of Peperomia, in a cloud forest where native trees are covered with mosses, ferns, orchids and vines, and humidity is maintained by multiple streams and a waterfall. The forest has been partially destroyed by logging that occurred until the 1960s, and only patches of natural vegetation are left (Sagástegui and Dillon 1991). The Peruvian government has protected this area since 2010, declaring it as a reserved zone: “Zona Reservada Udima” (El Peruano 2010), and categorizing it as “Zona Reservada Udima en Refugio de Vida Silvestre, Bosques Nublados de Udima” (El Peruano 2011).

From 30 to 500 m elevations, the warm desertic climate with temperatures of up to 35 °C in summer (Clima.com) favors the abundance of cactus steppes intermingled with densely irrigated rice and sugar cane cultivation that almost dry up the river at this altitude. From 500 m upwards, the climate changes abruptly to temperate and humid with an annual average temperature of 18 °C. Most cacti grow below 500 m, but some species reach 1000 m of elevation. The unexpected diversity of species of cacti in such a coastal valley was described first by Mischler (1988) and more recently by Alomía (2020), who mentions 13 taxa: six are endemic to Peru but only the last one in the following list is endemic to the Saña River Valley; Armatocereus oligogonus Rauh & Backeb., Espostoa melanostele (Vaupel) Borg, Haageocereus icosagonoides Rauh & Backeb., Haageocereus pseudoversicolor Rauh & Backeb., Neoraimondia arequipensis (Backeb.) Buxb. and Rauhocereus riosaniensis Backeb. ssp. riosaniensis. Other plants adapted to this xerophytic environment are Bursera graveolens (Kunth) Triana & Planch, Mimosa pellita Kunth. ex Willd, Rauvolfia tetraphylla L., Vachellia aroma var. huarango (Ruiz ex J.F.Macbr.) Seigler & Ebinger and Vallesia glabra Link. (Sagástegui and Dillon 1991). From this altitude up to 2500 m, as precipitation increases, dense forests are seen, a combination of dry western Andean forests and eastern montane forests (observation by the first author). During the summer months (December to May), increased rain can cause downstream floods, that occasionally reach the lower basin, such as the one that wiped out the city of Zaña in 1720 (Angulo 2011).

Although the abovementioned studies have identified nine species of Peperomia in the relict humid forest north of the town of Monteseco and four species of Peperomia in the forest of La Oscurana, so far no research has been carried out for the rest of the valley, especially the drier areas where fewer species are supposed to occur. This study aims to assess the diversity of Peperomia species along the whole Saña River valley, adding new reports of already known species and describing new ones.

Plant material

This research was approved by the Bioethics Committee of the Facultad de Ciencias Biológicas de la Universidad Nacional Mayor de San Marcos, document 009-2021-CBE-FCB-UNMSM. Collections were enabled by the Servicio Nacional Forestal y de Fauna Silvestre (SERFOR) permit numbers 009-2009-AG-DGFFS-DGEFFS, 0124-2011-AG-DGFFS-DGEFFS and authorization n° AUT-IFL-2018-064 issued by RDG N° 491-2018-MINAGRI-SERFOR-DGGSPFFS extended by RD N° D000170-2021-MIDAGRI-SERFOR-DGGSPFFS-DGSPF. Fieldwork was carried out by the teams of the Ghent University Botanical Garden (Belgium) and the Technische Universität Dresden (Germany) in 2009, and later by the Museo de Historia Natural – Universidad Nacional Mayor de San Marcos in the years 2020–2022, along the Saña River Valley starting from Zaña city up to Niepos, including the departments of Lambayeque (prov. Chiclayo) and Cajamarca (prov. San Miguel, La Florida, and Santa Cruz). No collections were made in the protected zone of Udima.

Field observations were complemented with the study of herbarium specimens of ten herbaria: F, GB, HAO, HLL, HUA, HUT, NY, UC, US, and USM, checking the types online. (acronyms following Thiers 2022). A total of 81 collections were included, of which 48 were made by the authors.

Photographs with either Sony DSC-HX400V or Olympus TG-6 cameras were taken of plants in habitat and plants ex-situ (vegetative, reproductive, and young plants). Flowers and seeds were photographed either with macro lenses, OPPO Reno-7, or stereomicroscopes. Measurements were taken from fresh plants, photographs, and dry herbarium specimens. Whenever possible, three or more measurements per structure. Floral organ measurements were based on photographs using ImageJ 1.53t. Colors were based on photographs of living plants and notes on herbarium labels.

All localities were georeferenced using GPS coordinates (https://www.gps-coordinates.net/) and Google Maps. For herbarium specimens without coordinates, approximate coordinates were calculated using Google Earth. Maps of distribution were drawn using the same set of coordinates using QGIS v.3.26 Buenos Aires (Official website QGIS: https://www.qgis.org/en/site/).

Finally, the subgenus to which each species belongs is mentioned according to the infrageneric classification of the genus Peperomia by Frenzke et al. (2015), who assigned all species described until then to their respective subgenus based on morphological and/or molecular data. The subgeneric placement of the new species described in this paper was defined based on morphological characters.

Results

Taxonomy

The present study found 81 accessions of 16 different taxa of Peperomia in the Saña River Valley (Fig. 2). We here present an identification key for these taxa. Peperomia crystallina, P. hispiduliformis, and P. hartwegiana are included in the key due to the high probability of occurrence in the region. Characteristics of the species were taken from the present study as well as Pino (2004), Scheiris (2013), Steyermark (1984), and Trelease and Yuncker (1950).

Figure 2.

Distribution of the Peperomia species in the study area. Udima reserved zone in green shadow.

Key to the species of Peperomia from the Saña River Valley

1	Leaves papyraceous, stems friable, hispid, and transparent	2
–	Leaves succulent or semi-succulent, stems firm, opaque	4
2	Stems erect, trichotomously branched, spadices in zigzag-shape, seemingly sessile on leaves, fruits verrucose	Peperomia crystallina Ruiz & Pav.
–	Stems shortly erect, then subdecumbent, spadices straight, originating from upper leaf axils, hispid fruits	3
3	Branches inserted at less than 45°, leaves ovate-orbicular, apex and base obtuse, venation pinnate	Peperomia hispidula (Sw.) A. Dietr.
–	Branches inserted at 45–90°, leaves cordiform, apex round to obtuse, base cordate, venation palmate	Peperomia hispiduliformis Trel.
4	Lower leaves in basal rosette, cordate, only reproductive stem growing upwards	Peperomia fraseri C. DC.
–	Lower leaves not in a basal rosette, leaves growing along a long stem	5
5	Leaves alternate	6
–	Leaves opposite or whorled	14
6	Leaves vertically compressed, very succulent, adaxial size reduced, epidermis covering the internal hydrenchyma, and exposing the photosynthetic chlorenchymatous tissues to solar irradiance through the abaxial surface either directly or by reflection from the ground. (fenestra), inflorescence a panicle	7
–	Leaves not vertically compressed, adaxial size similar to abaxial side in size, succulent, semi-succulent or chartaceous	8
7	Leaves oval elliptic, papillated surface, gray-glaucous to reddish dull green	Peperomia cymbifolia Pino.
–	Leaves dolabriform, smooth surface, light green to glaucous	Peperomia dolabriformis Kunth.
8	Leaves succulent, spirally crowded at the distal end, stem erect, succulent, inflorescence a panicle	Peperomia riosaniensis Hutchison ex Pino, Samain & L.E. Alomía, sp. nov.
–	Leaves semi-succulent or chartaceous, all along the stem, inflorescence simple, geminated or loosely flowered	9
9	Stems many-branched distally and horizontally, plants densely pubescent	Peperomia sagasteguii Pino, Samain & L.E. Alomía, sp. nov.
–	Stems branching basally and at different angles, glabrous or puberulous	10
10	Leaves distichous, stem winged	Peperomia vivipara Pino, Samain & L.E. Alomía, sp. nov.
–	Leaves spirally attached	11
–	Leaves opposite or whorled	15
11	Leaf base cordate, subpeltate, stem succulent, tortuous, inflorescence a panicle of clusters of spadices	Peperomia symmankii Pino & Samain, sp. nov.
–	Leaf base cuneate, lamina elliptical to ovate or obovate, seeds sticky	12
12	Leaves with round, truncate, or emarginate apex	Peperomia haematolepis Trel.
–	Leaves with acute, obtuse to attenuate apex	13
13	Stems erect, short, quadrangular in section, leaves elliptic-obovate, spadices white, terminal in umbels of 3, whitish, 8–12 cm long	Peperomia emarginulata C. DC.
–	Stems trailing, long, terete, leaves elliptic ovate or obovate, spadix terminal simple, reddish or reddish green	14
14	Stems without wings, leaves ovate, acute to acuminate, succulent, canaliculated	Peperomia cacaophila Trel. & Yunck.
–	Stems with two small wings, leaves widely obovate,, apex obtuse, semisucculent, flat	Peperomia pilocarpa Pino, Samain & L.E. Alomía, sp. nov.
15	Leaves chartaceous, opposite or decussate, strongly pubescent leaves and stems	Peperomia rotundata Kunth
–	Leaves succulent, verticillate, glabrous to puberulous	16
16	Leaves orbicular to oval, inflorescence terminal simple, red	Peperomia hartwegiana Miq.
–	Leaves narrow elliptical to obovate, inflorescences yellow or green	17
17	Whorls of 4–5 leaves, small (usually less than 1 cm long), all homogeneous in size and thickness, inflorescence simple, terminal	Peperomia microphylla Kunth
–	Whorls of 6–8 leaves, juvenile and mature leaves of different size and thickness, inflorescences in terminal umbellas and axillary from distal nodes	18
18	Plants large (25–50 cm tall), stems 1.2–4.5 mm diam., basal leaves 6–12 mm long, terrestrial or lithophytic, above 2200 m	Peperomia galioides Kunth
–	Plants small (10–24 cm tall), stems 3.5–8 mm diam., basal leaves 4–7 mm long, epiphytical, below 2000 m	Peperomia inaequalifolia R. & P.

1. Peperomia cacaophila Trel. & Yunck., Piperac. N. South Amer. 2(610). 1950.

Fig. 3A–F

Type

Ecuador, prov. Los Ríos, cant. Quevedo, roadside at Quevedo, common at Cacao groves, Haught 2966 (holotype: US 1707621!, isotype: ILL).

Figure 3.

Peperomia cacaophila Trel. & Yunck A young plant hanging from a rock B long, trailing plant with distal inflorescence C plant ex-situ with several branches D detail of the leaves E spadix at the beginning of anthesis F mature spadix.

Distribution and habitat

Plants were known only from central Ecuador, prov. Los Ríos, Guayas, Esmeralda, and Pichincha, epiphyte on Theobroma or similar trees, in shaded warm forests from 200–1500 m. This is the first report for Peru.

Notes

The closest species we could compare to is P. topoensis Yunck., also from Ecuador, which is similar in its epiphytic, pendent, or assurgent habit with small, succulent, alternate ovate leaves, cordulate at the base and 7-plinerved, compared to the obtuse, subtruncate based and noticeable 3-nerved leaves of P. cacaophila. Peperomia nitida Dahlst. from Brazil, São Paulo (Campinas), a widely cultivated species, is similar but with larger, light green cordulate-based leaves, only stems slightly reddish. Peperomia portobellensis Beurl. from Panama is also similar in shape but with slightly larger leaves. No material other than the types of these species was available for further comparison.

This species belongs to Peperomia subg. Micropiper (Miq.) Miq. (Frenzke et al. 2015).

Specimens examined

Peru, dept. Cajamarca, prov. Santa Cruz, dist. Catache: Road to Hacienda Taulis, ca. 60 km east of Cayaltí on the Río Saña. 560 m, [06°50'41.5"S, 79°10'14.2"W], 28 Aug 1964, P. C. Hutchison 6323 (UC, US, F 1642092, USM 16451); Road from El Espinal to La Florida, 550 m, 06°51'23.0"S, 79°09'31.0"W, Jul 26 2020, G. Pino & L.E. Alomía 3234 (USM); Same road, 527 m, 06°51'16"S, 79°09'44"W, 28 Aug 2022, G. Pino et al. 3625 (USM 330878); idem, 536 m, 06°51'18.6"S, 79°09'41.6"W, 28 Aug 2022, G. Pino et al. 3630 (USM 330879).

2. Peperomia cymbifolia Pino, Pino, Peperomias de Cajamarca: 16–17, 50–51 f. 3 A–N. 2004.

Fig. 4A–F

Type

PERU, dept. Cajamarca, prov. San Pablo, dist. San Pablo, the road to Chilete, 2080 m, 14 May 1964, P.C. Hutchison & J. K. Wright 5076 (holotype: UC 301913!, isotype: USM 16457!).

Figure 4.

Peperomia cymbifolia Pino var. occidentalis Pino & L.E. Alomía var. nov. A plant in habitat B plant ex situ C comparison of the leaves of P. cymbifolia var. cymbifolia (left) and var. occidentalis (right) D young plants showing development E cultivated plant showing inflorescence F seeds of P. cymbifolia var. cymbifolia (right) and var. occidentalis (left).

Distribution and habitat

Montane scrubs of the department of Cajamarca (Prov. San Pablo, Contumazá, Cajamarca, Chota, and San Marcos) between 1800 to 2800 m.

Notes

The samples collected were determined as P. cymbifolia (Pino 2004) However, comparing the plants of the Saña River Valley and the plants from the type locality we found some differences summarized in Table 1. These differences do not support the description of a new species but are sufficient to describe a new variety of this species. Moreover, while previous collections grow from 1800 to 2800 m in colder and moister places, this variety seems to be adapted to low altitudes of 500 to 600 m with higher temperatures and drier periods. It is easy to cultivate outdoors in the city of Lima, quite different from var. cymbifolia, which requires a cool greenhouse. This species belongs to Peperomia subg. Fenestratae Pino (Frenzke et al. 2015).

Table 1.

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Comparison of the main differences between P. cymbifolia var. cymbifolia and var. occidentalis (Data from var. cymbifolia taken from Pino (2004), and from live plants).

Features	P. cymbifolia var. cymbifolia	P. cymbifolia var. occidentalis
Height (vegetative) in cm	10–20	5–15
Leaf (shape)	Adaxial side slightly canaliculate, Abaxial side convex.	Adaxial side strongly canaliculate, Abaxial side straight to slightly convex.
Leaf (shape)	Lateral faces flat to slightly convex or concave.	Lateral faces more constantly convex.
Leaf (length in cm)	(1.6–) 2.2–3.4	(1.4–) 1.9–4
Leaf (vertical height of lateral sides in mm)	9–13, flat to concave, nerves conspicuous	7–10, flat to convex, nerves obscure
Rachis of the spadix (length in cm)	(2–)3.8–6	7–15
Fruit (shape/color)	Globose, yellowish-green to brown	Ovoid to ellipsoid, light olive green
Fruit (length in mm)	0.7–0.8	0.6–0.8
Fruit (diam. in mm)	0.6–0.7	0.45–0.55
Fruit (stylopodium)	Broad conical, light brown,	Broad cylindrical, light brown – orangish,
	0.1–0.2 mm long,	0.2–0.25 mm long,
	0.2–0.3 mm diam.	0.2–0.35 mm diam.
Altitude range (m)	1800–2800	500–600

Peperomia cymbifolia Pino var. occidentalis Pino & L.E.Alomía, var. nov.

urn:lsid:ipni.org:names:77317651-1

Diagnosis

Peperomia cymbifolia var. occidentalis differs from the type variety mainly in size, plants are smaller and more compact; Leaves are slightly longer (4 cm maximum compared to 3.6 cm long), their lateral faces are not as high (7–10 mm compared to 9–13 mm), and they are more constantly convex than concave, its abaxial side is less convex, sometimes even straight; Its fruits are smaller and narrower, (0.6–0.8 × 0.45–0.55 mm compared to the 0.7–0.8 × 0.6–0.7 mm), light olive green instead of yellowish green, and with a relatively larger stylopodium. Populations of this variety are not only separated geographically from the type variety but mostly by altitude and climate: they grow 1200 m below the lowest occurrence of the type variety, at least 10 °Celsius warmer. The epithet reminds the localization of this variety, the most occidental of all three varieties of P. cymbifolia. (P. cymbifolia var. cymbifolia ,var. goodspeedii , and var. occidentalis).

Type

Peru, dept. Cajamarca, prov. Santa Cruz, dist. Catache: road from Espinal to La Florida, 521 m, 06°51'14.2"S, 79°09'45"W, 28 Aug 2022, G. Pino & L.E. Alomía 3634 (holotype: USM 330881!).

Distribution and habitat

Plants grow from 450 to 600 m, on exposed rock crevices facing southwards between the towns of El Espinal and La Florida in Department Cajamarca, province of Santa Cruz. A similar collection, very likely the same taxon, was found in Chota at 750 m, exactly 20 km to the North of the Saña River Valley in a straight line.

Phenology

Inflorescences appear from June to February; fruits ripen from October to March.

Additional specimens examined

Peru, dept. Cajamarca, prov. Santa Cruz, dist. Catache: Road from El Espinal to La Florida, 582 m, 06°51'13.7"S, 79°09'16.5"W, 26 Jul 2020, G. Pino & L.E. Alomía 3232 (USM 330872); Same road, 591 m, 06°51'13.4"S, 79°09'18.4"W, 28 Aug 2022, G. Pino et al. 3638 (USM 330883). Prov. Chota, dist. Chimbán, Road from Cumbil to Llama, [between Potrerillo and Maychil], border of the road, on rocks, 750 m, [06°31'56"S, 79°11'51"W], 21 May 1965, A. López & A. Sagástegui s/n (HUT 5520). Dept. Lambayeque, prov. Ferreñafe, dist. Incahuasi: Before Laquipampa, 721 m, 06°20'31.5"S, 79°26'41"W, 08 Mar 2023, G. Pino & J.E. Romero 3840 (USM).

3. Peperomia dolabriformis Kunth, Nov. Gen & Sp. [Humboldt] 1: 50, tab. 4. 1815.

Fig. 5A

Piper dolabriforme Poir., Lam. Encycl. Meth. Suppl. 4: 464. 1816. Type: Based on Peperomia dolabriformis Kunth.

Type

Peru, dept. Piura, prov. Huancabamba: “in Peruviae calidis, ad ripas flumini Guancabamba” [in the warmth of Peru, on the banks of the Huancabamba River], Humboldt s/n (holotype: B [lost?], isotype P!).

Figure 5.

Peperomia species that can be found in the Saña River Valley A P. dolabriformis B P. fraseri C P. galioides D P. haematolepis E P. hispidula F P. inaequalifolia G P. microphylla H P. rotundata.

Distribution and habitat

Warm valleys of the Western slopes of the Andes of departments Lambayeque, La Libertad, and Cajamarca. Huancabamba, Marañón and Crisnejas Valleys in departments Piura, Cajamarca, Amazonas, La Libertad and Ancash (Sihuas) in Peru. In Ecuador, it occurs in the provinces of Loja and Zamora-Chinchipe.

Note

This species belongs to Peperomia subg. Fenestratae Pino (Frenzke et al. 2015).

Specimens examined

Peru, dept. Lambayeque, prov. Chiclayo, dist. Oyotún: Canyon above Chiclayo near Mocupa [Mocupe, 391 m, 06°48'58.7"S, 79°12'01.6"W], Johnson Cactus Garden, prepared at the BG of the University of California at Berkeley, accession 52.498, 24 Sep 1958, Larrabie s/n (F1567768, NY, UC088167, US2301205) Dept. Cajamarca, prov. Santa Cruz, dist. Catache: road from Oyotún to El Espinal, 397 m, 06°49'46.7"S, 79°11'11.7"W, 28 Aug 2022, G. Pino & L.E. Alomía 3613 (USM 330874); same road, 440 m, 06°50'39.9"S, 79°10'23.1"W, 28 Aug 2022, G. Pino & L.E. Alomía 3621 (USM 330875).

4. Peperomia emarginulata C.DC. DC. Prodr. 185. 16(1): 433. 1869.

Fig. 6A–F

Type

Peru, dept. [Huánuco], Casapí, In Peruvia, sylvis Andium occident. [In Perú, western Andes forests], Mathews 1687 (lectotype: K, designated by Mathieu and Callejas (2006: 351; isolectotype: E, K!)).

Figure 6.

Peperomia emarginulata C.DC A colony growing on fallen Guadua angustifolia leaves B exposed plant growing on a rock C plant ex-situ showing 4-angled stems and inflorescences D detail of the leaves E terminal umbella with spadices F spadices at the beginning of anthesis G mature spadix with seeds.

Distribution and habitat

Plants are reported from Ecuador and Peru, in montane forests from 300 to 1500 m.

Notes

We compared the three species with features closer to our findings (Table 2) The plants we found are in the range of all three, but according to this table, it matches P. emarginulata. Peperomia angularis and Peperomia caucana from Colombia should be reviewed in the future to prove if they are conspecific with P. emarginulata. Peperomia garcia-barrigana Trel. & Yunck. and P. gabinetensis Trel. & Yunck. also show similarities that deserve further analysis, being mostly different in size. Some specimens in herbaria were also determined as P. glabella (Sw.) A. Dietr., a widely distributed species, very close to this group, that is relatively small.

Table 2.

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Comparison of some features of the descriptions of P. angularis, P. emarginulata, and P. caucana with the specimens of P. emarginulata studied. All data according to Trelease and Yuncker (1950), complemented with Steyermark (1984) for P. angularis and Trelease (1936) for P. emarginulata.

Features	P. emarginulata of Saña River Valley	P. angularis	P. emarginulata	P. caucana
Height in cm	15–45	25 or more	30 or more	15 or more
Stem (diam. in mm)	3–5	1–3	4–5	2
Internodes in cm	1.5–4	1–3	1–3	1–3
Petiole (length in mm)	5–15	5–10 (–22)	5–15	5–10
Leaf (shape)	Elliptic -obovate, lanceolate, subrhomboidal.	Elliptic, elliptic- oblanceolate or obovate.	Elliptic-lanceolate or obovate, somewhat rhomboidal.	Elliptic-obovate.
Leaf (apex)	Acute to shortly acuminate, emarginulate.	Protracted, blunt or acutish.	Shortly acute to attenuately acuminate, blunt and emarginulate.	Shortly attenuate, blunt or acutish.
Leaf (length in cm)	3.5–6	2–6 (–9)	3–6 (–11)	4–6.5
Leaf (width in cm)	2.5–4	1–2.5	2.5–4.5	2–3.5
Peduncle (length in cm)	1–2	1.4–2 (–4.5)	0.5–1	2–3
Spadix (length in cm)	8–12	5–7 (–10.5)	–12	5–7

This species belongs to Peperomia subg. Micropiper (Miq.) Miq. (Frenzke et al. 2015).

Specimens examined

Peru. Dept. Cajamarca, prov. Santa Cruz, dist. Catache: El Papayo, El Espinal, Monteseco, 600 m, [06°51'31.7"S, 79°06'46.8"W], 18 Jul 1984, A. Sagástegui, E. García & S. Leyva 12354 (HUT, F 1960604, MO 3226489); Monteseco, 3 km NE, 1800 m, [06°50'42.3"S, 79°06'17.1"W], 5 May 1987, J. Santisteban C. & J. Guevara B. 013 (HUT, F 1994832). Same place: 13 May 1987, J. Santisteban C. & J. Guevara B. 055 (HUT 24750, F 1995336). Path in Monteseco Forest, 1350 m. [06°51'17.8"S, 79°06'33.3"W], 10 Oct 1993, S. Leiva et al. 911 (HAO, F 2142594); Road from Cayaltí to La Florida, 549 m, 06°51'23.2"S, 79°09'31.1"W, 1 Mar 2009, G.Mathieu & L. Symmank 161 (USM 255771); Road from El Espinal to La Florida, 519 m, 06°51'13.5"S, 79°09'45"W, 28 Aug 2022, G. Pino et al. 3623 (USM 330876); Same road, 527 m, 06°51'16"S, 79°09'44"W, 28 Aug 2022, G. Pino et al. 3627 (USM 330877); Idem, 536 m, 06°51'18.6"S, 79°09'41.6"W, 28 Aug 2022, G. Pino et al. 3632 (USM 330880); Prov. San Miguel, dist. Niepos: Road from Naranjo to La Florida, 1316 m, 06°53'08.6"S, 79°06'45.8"W, Jul 26 2020, G. Pino & L.E. Alomía 3218 (USM 330868); Same road, 1291, 06°53'08.7"S, 79°06'49.1"W, Jul 26 2020, G. Pino & L.E. Alomía 3219; Same road, 1265 m, 06°53'04.6"S, 79°06'51.2"W, Jul 26 2020, G. Pino & L.E. Alomía 3220; Idem, 1213 m, 06°52'54.2"S, 79°06'51.9"W, Jul 26 2020, G. Pino & L.E. Alomía 3221; Idem, 1210 m, 06°52'53.6"S, 79°06'51.0"W, Jul 26 2020, G. Pino & L.E. Alomía 3222; Idem, 1176 m, 06°52'49.2"S, 79°06'52.5"W, Jul 26 2020, G. Pino & L.E. Alomía 3223; Idem, 1130 m, 06°52'44.1"S, 79°07'00.0"W, Jul 26 2020, G. Pino & L.E. Alomía 3224; Idem, 1118 m, 06°52'42.0"S, 79°07'01.1"W, Jul 26 2020, G. Pino & L.E. Alomía 3225; Idem, 1067 m, 06°52'32.9"S, 79°07'10.4"W, Jul 26 2020, G. Pino & L.E. Alomía 3226; Road from La Florida to Niepos, 1297 m, 06°53'6.9"S, 79°06'47.6"W, 28 Aug 2022, G. Pino et al. 3642 (USM 330885).

5. Peperomia fraseri C.DC., J. Bot. 4: 134. 1866.

Fig. 5B

Peperomia rugosa Tafalla. Flora Huayaquilensis 1:12. 1989, nom. ined.

Peperomia resediflora Linden & André, Ill. Hort. 17: 135–137. fig. 26. 1870. [As resedaeflora] Type: (iconotype): Illustr. Hortic. 17: fig. 26.

Peperomia fraseri var. resedaeflora C.DC., Bull. Herb. Boissier 5: 703. 1897.Type: Based on Peperomia resediflora Linden & André.

Peperomia fraseri var. peltata Yunck., Piperac. N. South Amer. 2: 740. 1950. Type: Ecuador, Loja, Horta-Naque, Espinosa 859 (holotype: ILL, isotype: NY).

Peperomia treleasei Standl. & Steyerm., Fieldiana, Bot. 24(3): 273. 1952. Type: Guatemala (cultivated), Steyermark 46298 (holotype: F).

Peperomia fraseri var. guayasana Trel., nom. herb. [Mathieu (2007)]. Type: Ecuador, Guayas, Haught 3022 (holotype: US).

Type

Ecuador, sine loco: Fraser s/n, (lectotype, designated by Mathieu and Callejas (2006: 351–352: G-DC!, isotypes: BM, G-DC)).

Distribution and habitat

Ecuador (120–2100 m, prov. Loja, El Oro, Guayas, Chimborazo and Manabí). Peru, 500–1600 m in departments Tumbes, Piura, Lambayeque, and eastern Cajamarca along the Saña River valley, on rocky soil, shaded by trees and shrubs. This is the southernmost location of this species.

Specimens examined

Peru, dept. Lambayeque, prov. Chiclayo, dist. Oyotún: road from Oyotún to El Espinal, 515 m, 06°48'30.3"S, 79°11'08"W, 26 Jul 2020, G. Pino & L.E. Alomía 3236 (USM 325348); Dept. Cajamarca, prov. Santa Cruz, dist. Catache: road from Cayaltí to La Florida, 8.4 km from La Florida, 497 m, 06°50'51.6"S, 79°09'53.7"W, 1 Mar 2009, G.Mathieu & L. Symmank 160 (USM 255770); road from Espinal to La Florida, 521 m, 06°51'14.2"S, 79°09'45"W, 28 Aug 2022, G. Pino et al. 3636; Same road, 527 m, 06°51'16"S, 79°09'44"W, 28 Aug 2022, G. Pino et al. 3626. Same road, 570 m, 06°51'14.1"S, 79°09'23.1"W, 26 Jul 2020, G. Pino & L.E. Alomía 3231 (USM 325349).

Note

This species belongs to Peperomia subg. Panicularia Miq. (Frenzke et al. 2015).

6. Peperomia galioides Kunth, Nov. Gen & Sp. [Humboldt] 1: 58, tab. 17. 1815.

Fig. 5C

Piper galioides Poir., Lam. Encycl. Meth. 4: Suppl. 4: 470. 1816. Type: Based on Peperomia galioides Kunth.

Peperomia suaveolens Hamilton, Prodr. Pl. Ind. Occ.:2. 1825. Type: Cuba, (holotype: P- Herbarium Desvaux?).

Piper mollugo Willd. [Miquel] 1843. Type: Based on Peperomia galioides Kunth.

Peperomia jamesoni Regel, Bull. Soc. Nat. Moscou 31: 544. 1858. Type: Ecuador, Jameson 814 (holotype: G [not designated], isotypes: K, TCD).

Peperomia agapatensis C.DC., Prodromus 16: 455. 1869. Type: Perú, Junín, Agapata, Lechler 1934 (holotype: G-DC; isotypes: G, GOET, K, MO, P, UPS, W).

Peperomia galioides Kunth var. longifolia C.DC., Prodromus 16 (1): 464. 1869. Type: Bolivia, Mandon 1120 (syntypes: G-DC, B).

Peperomia galioides Kunth var. menkeana (Miq) C.DC., Prodromus 16 (1): 464. 1869. Type: Brazil, Menke 125 (holotype: BR).

Peperomia galioides Kunth var. nigro -punctulata C.DC., Prodromus 16 (1): 464. 1869. Type: Brazil, Wawra 441, Ecuador, Jameson s.n. (syntype: K).

Peperomia galioides Kunth var. aprica Henschen. Nov. Act. Soc. Sci. Upsal. ser.3,8(2): 37. 1873. Type: Brazil, Minas Gerais, Caldas (Holotype: not designated).

Peperomia galioides Kunth var. umbrosa Henschen, Nov. Act. Soc. Sci. Upsal. ser.3,8(2):37. 1873. Type: Brazil, Minas Gerais, Caldas (Holotype: not designated).

Peperomia subcorymbosa Sodiro, Contrib. al Conoc. Fl. Ecuador. Monogr. 1 ed. 2:181. 1901. Type: Ecuador, Quito, El Alacatzo, Sodiro s.n. (holotype: Q, isotypes: G-DC, QPLS).

Peperomia galioides Kunth var. saxicola C.DC., Bull. Her. Boissier ser. 2,1: 360. 1901. Type: Brazil, Schwacke 10279 (holotype: G).

Peperomia anisophylla C.DC., Bot. Jahrb. 40: 266. 1908. Type: Perú, Lima, Matucana, Weberbauer 72 (holotype: B, isotype: G).

Peperomia galioides var. aromatica C.DC., Bot. Jahrb. Syst. 40: 266. 1908. Type: Peru. Ancash, Caraz. Weberbauer 3227 (holotype: B [lost?]; isotype: G-DC).

Peperomia galioides Kunth var. minutifolia C.DC. Ann. Cons. Jard. Bot. Gen. 21: 262. 1920. Type: Ecuador, Quito, Nanegal, Sodiro 2/52 (holotype: G-DC).

Peperomia granata Trel., Fedde Rep. Sp. Nov. 23: 24. 1926. Type: Cuba, Pico Turquino, Ekman 14590 (lectotype: ILL [designated by Saralegui (2004)]; isolectotype: S).

Peperomia galiifolia Trel., Mem.N.Y. Bot.Gard. 7: 227. 1927. Type: Bolivia, La Paz, Pongo de Quime, White 166 (holotype: NY).

Peperomia okarana Trel., Bull. Bot. Club. 55: 170. 1928. Type: Bolivia, La Paz, Larecaja, Okara, Tate 970 (holotype: NY).

Peperomia artatiflora Trel., Flora of Peru [Macbride] 13(2): 23. 1936.Type: Perú, Palca, Stevens 47 (holotype: ILL).

Peperomia brachyiula Trel., Flora of Peru [Macbride] 13(2): 26. 1936. Type: Perú, Lima, Matucana, Macbride 129 (holotype: F, isotypes: G, ILL).

Peperomia ceapanana Trel., Flora of Peru [Macbride] 13(2): 28. 1936. Type: Perú, Cusco, Paucartambo, Herrera 1468 (holotype:US).

Peperomia chillonensis Trel., Flora of Peru [Macbride]. 13(2): 30. 1936. Type: Perú, Lima, Canta, Obrajillo. Pennell 14413 (holotype: F).

Peperomia dendroides Trel., Flora of Peru [Macbride]. 13(2): 38. 1936. Type: Perú, Junín, Chaglla, Macbride 3640 (holotype: F, isotypes: G, ILL, US).

Peperomia distractiflora Trel., Flora of Peru [Macbride] 13(2): 40. 1936. Type: Perú, Junín, Challhuapuquio, Stevens 214 (holotype: ILL).

Peperomia longispica Trel., Flora of Peru [Macbride] 13(2): 59. 1936. Type: Perú, Huánuco, Muña, Macbride 3928 (isotype: BM).

Peperomia sanbuenaventurana Trel., Flora of Peru [Macbride] 13(2): 91. 1936. Type: Perú, Lima, Canta, San Buenaventura. Pennell 14566 (holotype: PH).

Peperomia trullifolia Trel., Flora of Peru [Macbride] 13(2): 102. 1936. Type: Perú, Cusco, Ollantaytambo, Cook 401 (holotype: US).

Peperomia inaequalifolia Ruiz & Pav. var. galioides (Kunth) Pino, Avonia 28(2): pages 45–49 (f.16–25). 2010. Type: Based on Peperomia galioides Kunth.

Type

Colombia, dept. Cundinamarca, prov. Tequendama: “Crescit in montanis Regni Novogranatensis, juxta cataractam Tequendamae, alt. 1300 hexap. Floret Augusto” [Grows in mountains of Colombia, close to the Tequendama Waterfall, 2400 m, Flowers in August], Humboldt s/n, (isolectotype, designated by Saralegui (2004: 341: B-W [00762])).

Distribution and habitat

Montane forests of Mexico, Central America, Venezuela, Colombia, Ecuador, Peru, Bolivia, and Brazil, on rocks or terrestrial, always above 2200 m.

Note

This species belongs to Peperomia subg. Micropiper (Miq.) Miq. (Frenzke et al. 2015).

Specimens examined

Peru, dept. Cajamarca, prov. Santa Cruz, dist. Catache: Udima, herbácea, creciendo sobre piedra, [herb growing on rocks] 2500 m, [06°49'38.3"S, 79°06'01.1"W], 20 May 1987, J.Santisteban C. & J. Guevara B. 087 (HUT, F1995293). Prov. San Miguel, dist. Niepos: Road from La Florida to Niepos, 2240 m, 06°54'49"S, 79°08'17.7"W, 28 Aug 2022, G. Pino & L.E. Alomía 3647 (USM 330888); Dist. Calquis: Taulis Playa (Agua Blanca) 2650 m, [06°56'06"S, 78°59'23"W], 3 Jul 1986, J. Mostacero et al. 1143 (HUT 22520, MO).

7. Peperomia haematolepis Trel., Field. Mus. Pub. Bot. [Macbride] 13(2):52. 1936.

Fig. 5D

Type

Peru, dept. Junín, prov. Chanchamayo, dist. San Ramón: Hacienda Chalhuapuquio, Stevens 212 (holotype: ILL).

Distribution and habitat

Plants are reported from Brazil, the Guyanas, Venezuela, Colombia, Ecuador, and Peru from 1000 to 2000 m, epiphytic in montane forests shaded by the canopy. Most of the collections in Peru are from the Amazon Basin; this is the first report for a Pacific Ocean draining Andean valley.

Notes

This species belongs to Peperomia subg. Oxyrhynchum (Dahlst.) Samain (Frenzke et al. 2015).

Specimen examined

Peru, dept. Cajamarca, prov. Santa Cruz, dist. Catache: Upper Río Zaña Valley, ca. 5 km above Monteseco, on the path below the campsite, lower reaches of evergreen tropical montane forest, 1450 m, [06°51'10.8"S, 79°06'09.4"W], 19 Mar 1986, M.O. Dillon & A. Sagástegui 4425 (US 3338748, F, GB 0167869, HUA 49003).

8. Peperomia hispidula (Sw.) A. Dietr., Sp. Pl. 1: 165. 1831.

Fig. 5E

Piper hispidulum Sw., Prodr. 15. 1788. Type: Jamaica, Swartz s.n. (holotype not designated: S) [Basionym for Peperomia hispidula (Sw.) A.Dietr.] .

Acrocarpidium hispidulum Miq., Syst. Piperac. 54. 1843 Type: Based on Piper hispidulum Sw.

Acrocarpidium sellowianum Miq., Syst. Piperac. 55. 1843 [as ‘sellovianum’] Type: Brazil, Brasilia, Sellow s.n. (holotype: B [lost?], lectotype: U [designated Zanotti et al. (2012: 133)], isolectotypes: G, K, W).

Peperomia tenera Miq., Fl. Bras. [Martius] 4(1): 19. 1852. Type: Based on Acrocarpidium sellowianum Miq.

Peperomia muscophila C.DC., Journ. Bot. 4: 133. 1866. [as ‘muscophylla’] Type: Mexico, Tafalla s.n (holotype: G; isotypes: G-DC).

Peperomia hispidula C.DC., Prodr.16:1:397. 1869. Based on Piper hispidulum Sw.

Peperomia hispidula var. sellowiana (Miq.) Dahlst. Kongl. Svenska Vetenskapsakad. Handl. 33(2): 14 . 1900. Type: Based on Acrocarpidium sellowianum Miq.

Peperomia hispidula var. swartziana Dahlst., Kongl. Svenska Vetenskapsakad. Handl. 33(2): 13.1900. Type: Based on Peperomia hispidula C.DC.

Peperomia hispidula var. swartziana f. barbensis Dahlst., Kongl. Svenska Vetenskapsakad. Handl. 33(2): 14 .1900. Type: Costa Rica, Hoffmann 54 (holotype: B [lost?]).

Peperomia perhispidula C.DC., Bot. Jahrb. Syst. 40: 257. 1908). Type: Peru, Huacapistana, Weberbauer 2014 (holotype: B, F G-DC).

Peperomia hispidula var. muscophila (C.DC.) C.DC., Candollea 1: 335 .1923. Type: based on Peperomia muscophila C.DC.

Peperomia hispidula var. perhispidula Trelease & Yuncker, Candollea 1: 335. 1923. Type: Based on Peperomia perhispidula C.DC.

Peperomia hispidula var. ellipticifolia Trelease & Yuncker, Piper. North. S. Amer. 2: 705. Pl. 626. 1950. Type: Colombia, Suratá, Killip & Smith16580 (holotype: GH; isotypes: ILL, MA, NY, US).

Type

Jamaica, In nemorosis humidis montium altissimorum coeruleorum Jamaicae. [In shaded moist elevated forest on Blue Mountains Peak of Jamaica, 2200 m, 18°6'N, 76°40'W], Swartz s.n. (holotype not designated: S!)

Distribution and habitat

Terrestrial on humus near water courses, shaded, 1500–3000 m, in evergreen montane forests of West Indies, Mexico, Central America, and South America (except Chile). All the other collections in Peru are from the eastern slopes of the Andes; this is the only report for the western slopes.

Note

This species belongs to Peperomia subg. Hispidula Frenzke & Scheiris (Frenzke et al. 2015).

Specimen examined

Peru, dept. Cajamarca, prov. Santa Cruz, dist. Catache: Al norte del Chorro Blanco (Monteseco), borde de acequia [North of Chorro Blanco (Monteseco), border of stream] 1500 m, [06°50'58.6"S, 79°05'53.7"W], 20 Jan 1989, S. Leyva G. 008 (F 2016256).

9. Peperomia inaequalifolia Ruiz & Pav., Flora Peruviana & Chilensis 1: 30, plat. 46 fig.a. 1798.

Fig. 5F

Piper inaequalifolium Vahl., Enum. Pl. 1: 355. 1804. Type: Based on Peperomia inaequalifolia Ruiz & Pav.

Piper aromaticum Willd., Enum. Pl. Hort. Berol. Suppl.: 3. 1813. Type: Not mentioned.

Peperomia chrysotricha Miq., Syst. Pip.:163. 1843. Type: Dombey 933 (holotype: P; isotype: U).

Peperomia parvula Sodiro, Piperac. Ecuator. 1: 139. 1900. Type: Sodiro s/n (Holotype not designated: G, P, Q).

Peperomia fasciculata Sodiro, Contrib. al Conoc. Fl. Ecuador Monog.1 ed. 3: 181. 1901. Type: Based on Peperomia parvula Sodiro.

Peperomia galioides Kunth var. minutifolia C.DC., Ann. Cons. Jard. Bot. Genève 21: 262. 1920. Type: Ecuador, Valle Nonegal. A. Sodiro 2/52 (holotype: G-DC).

Peperomia atocongona Trel., Publ. Field Mus. Nat. Hist., Bot. Ser. [Macbride J.F.] 13(2): 24. 1936. Type: Peru. Lima: Atocongo. F.W. Pennelll 14752 (holotype: PH).

Peperomia limaensis Trel., Publ. Field Mus. Nat. Hist., Bot. Ser. [Macbride J.F.] 13(2): 58. 1936. Type: Peru. Lima: San Gerónimo. J.F.Macbride 5920 (holotype: F).

Peperomia pseudogalapagensis Trel., Publ. Field Mus. Nat. Hist., Bot. Ser. [Macbride J.F.] 13(2): 79. 1936. Type: Peru. Lima: San Gerónimo. Wilkes Expl. Exped. 5920 (holotype: GH; isotype: K).

Type

Peru, dept. Lima, prov. Lima, dist. Rimac: “Habitat Limae in collibus altis in Amancaes copiosa, dicitur Congona Zimarrona” [Grows in Lima in high hills, abundant in Loma de Amancaes, it is called Congona Cimarrona], Ruiz & Pavón s/n: MA [29573]

Distribution and habitat

Coastal valleys of Peru and Galápagos (Ecuador) between 200 and 2000 m, in dry forests, almost always epiphytic on trunks, occasionally on rocks.

Note

This species belongs to Peperomia subg. Micropiper (Miq.) Miq. (Frenzke et al. 2015).

Specimens examined

Peru, dept. Cajamarca, prov. Santa Cruz, dist. Catache: Monteseco, 3 km NE, epífito, sobre tronco de árbol [epiphytic, on tree trunk] 1850 m, [06°50'41.4"S, 79°06'14.4"W], 5 May 1987, J. Santisteban C. & J. Guevara B. 004 (HUT 24701, F1995288); Arriba de Monteseco, sobre árbol de Erythrina [Above Monteseco, on Erythrina tree] 1350 m, [06°51'17.8"S, 79°06'33.3"W], 10 Oct 1993, S. Leyva et al. 904 (HAO, F 2165067) Prov. San Miguel, dist. Niepos: Road from La Florida to Monte Seco (Naranjo), 15.6 km from La Florida, 1700 m, 06°54'08"S, 79°08'15"W, 1 Mar 2009, G. Mathieu & L. Symmank 166. (USM 255775). Road from Naranjo to La Florida, 1316 m, 06°53'08.6"S, 79°06'45.8"W, 26 Jul 2020, G. Pino & L.E. Alomía 3227 (observed). Same road, 1291 m, 06°53'08.7"S, 79°06'49.1"W, 26 Jul 2020, G. Pino & L.E. Alomía 3228 (observed). Idem. 1265 m, 06°53'04.6"S, 79°06'51.2"W, 26 Jul 2020, G. Pino & L.E. Alomía 3229 (observed). Road from La Florida to Niepos, 1297 m, 06°53'06.9"S, 79°06'47.6"W, 28 Aug 2022, G. Pino & L.E. Alomía 3640. (USM 330884).

10. Peperomia microphylla Kunth, Nov. Gen & Sp. [Humboldt] 1: 57, tab. 15, fig. 2. 1815.

Fig. 5G

Piper microphyllum (Kunth) Poir., Lam. Encycl. Meth. Suppl. 4: 469. 1816. Type: Based on Peperomia microphylla Kunth.

Peperomia aphylla C.DC., Prodr. 16(1): 456. 1869. Type: Colombia. Quindío, Jameson 690 (holotype: G; isotypes: BM, K).

Peperomia gilbertii Trel., Publ. Field Mus. Nat. Hist., Bot. Ser. [Macbride J.F.] 13(2): 50.1936. Type: Peru. Cusco: Ollantaytambo, Cook & Gilbert 740 (holotype: US).

Peperomia chachopoana Trel., Cat. Fl. Venez. 1:244. 1945, Nom. Nud. Type: Venezuela. Mérida. (2800 m) Pittier 13165 (holotype: NY; isotypes: F, M, NY, US).

Type

Colombia, dept. Quindío: “Crescit in lapidosis frigidis Andium Quinduensis, juxta El Boquerón del Páramo, alt. 1650 hexap. Floret Octobri” [Grows in cold rocky places of the Andes of Quindiu, close to El Boquerón del Páramo, 3000 m, Flowers in October], Humboldt s/n, (lectotype, designated by Mathieu and Callejas (2006: 341: B!; isolectotype: P)).

Distribution and habitat

Montane forests of the Andes of Venezuela, Colombia, Ecuador, and Peru, on rocks or more frequently epiphyte.

Note

This species belongs to Peperomia subg. Micropiper (Miq.) Miq. (Frenzke et al. 2015).

Specimen examined

Peru, dept. Cajamarca, prov. Santa Cruz, dist. Catache: ca. 4.2 Km (por aire) [in a straight line] NE Monteseco, 2500 m, [06°49'47.1"S, 79°06'22.4"W], 20 May 1987, J. Santisteban C. & J. Guevara B 084 (HUT 24800, F1995295).

11. Peperomia pilocarpa Pino, Samain & L.E. Alomía, sp. nov.

urn:lsid:ipni.org:names:77317652-1

Fig. 7A–G

Type

Peru, Dept. Cajamarca, prov. San Miguel, dist. Niepos: road from La Florida to Niepos, 1613 m, 06°54'03.3"S, 79°07'33.5"W, 26 Jul 2020, G. Pino & L.E. Alomía 3217 (USM 333265!); Same collection, 28 Aug 2022, G. Pino et al. 3646 (USM 333266!).

Figure 7.

Peperomia pilocarpa A plant in habitat with assurgent stems B decumbent plant C young plants showing terminal inflorescence D plant ex-situ in anthesis showing normal inflorescence and inflorescence with galls E detail of leaves F spadix at the beginning of anthesis G mature spadix with galls and seeds. Details: (above, normal seed; below, gall with trichomes).

Diagnosis

Perennial, semi-succulent, prostate caespitose herb similar in habit to P. cacaophila, but differs mainly in leaf shape, which is widely obovate to elliptic or rotundate and sometimes slightly acuminate, compared to the ovate lamina and constantly acuminate apex of this species, leaves are flatter and less succulent, slightly puberulous compared to the canaliculate, glossy leaves of P. cacaophila. Stems are not terete as in this species but with two low prominent wings. Seeds are dimorphic, normal seeds are very similar in shape to the seeds of P. sagasteguii, modified seeds (probably galls) are conspicuous, large up to 2.5 mm long, bright green to brownish, densely covered with white trichomes.

Description

Perennial semi succulent terrestrial or semiepiphytic herb, living in the shade and on abundant decayed vegetal matter, 10–15 cm tall, up to 25 cm when flowering. Roots basal and from basal nodes, very fibrous, light gray, 0.2–0.3 mm diam., 1–3 cm long. Stem mainly prostate, straight, procumbent when vegetative and decumbent sometimes at tips, up to 45 cm long or more, subterete, 2.5–3.5 mm diam. at the base, gradually tapering to 1.5–2 mm at the apex, dull olive green, reddish where exposed, with two not very prominent longitudinal wings decurrent with leaf petioles, internodes 1–5 cm, rarely with alternate branches from the base or every 10–15 cm. Leaves alternate, spirally attached one per internode, glabrous to slightly puberulous, present all along the stem; petiole circular in section, 0.8–1.5 cm long, 1–1.5 mm diam., with two lateral not prominent wings decurrent with stem, straight or slightly recurvate, lamina flat, widely obovate to oblong, (2–) 3–6 cm long, 1–3.5 mm thick, (1–)1.5–2.5 cm wide at distal third, (1.5–)2.8–3.8 cm wide at the middle, (1–)2–3 cm wide at proximal third, apex acute, obtuse in some leaves and slightly acuminate in larger leaves, subemarginate at tip, base rounded; adaxially dull green, 3-palmatinerved, nerves slightly depressed; margin completely entire; abaxially light green, obscurely 3-palmatinerved, central nerve and proximal thirds of lateral nerves somewhat darker in color. Inflorescence a single terminal spadix appearing from November to December; peduncle terete or slightly funnel-shaped distally, sometimes reddish and obscurely furrowed, 7–12 mm long, 2–2.5 mm diam., with a small oval bract at the base, similar to leaves, rachis 6–10 cm long, 1.5–2.5 mm diam., light green. Floral bracts narrowly oval to subpentagonal, subacute, light green, 0.6–0.7 mm long, 0.5–0.6 mm wide. Stamens: filaments transparent, 0.2 mm diam., 0.4–0.5 mm long, anthers ovoid, 0.4–0.45 mm long, 0.25–0.3 mm wide, bright red at first, then white. Normal fruit globose, 0.7–0.8 mm long, 0.6–0.7 mm diam., brown, covered of white 0.05 mm long whitish papillae up to distal third of fruit, style conical, light brown. Modified fruit, 2–2.5 mm long, 1.2–1.5 mm diam., bright green to brownish, minutely and densely covered with white 0.1–0.2 long trichomes, style prominent, bright green, 0.2–0.25 mm long, 0.3–0.4 mm diam., stigma dark, ripening from February to July.

Distribution and habitat

Plants grow from 1500 to 1600 m of the middle course of the Saña River valley, in the remnants of montane forest, mostly epiphytic.

Phenology

Inflorescences appear from October to March; fruits ripen from November to April.

Etymology

The epithet recalls the hairy surface of the modified fruits of this species, from the Latin pilus (hair) and Greek καρπός (fruit).

Notes

This specimen at first was considered a probable hybrid because it shows intermediate features between P. emarginulata and P. cacaophila. However, although those two species share their habitat to some extent they are found at lower altitudes and never close to P. pilocarpa. Instead, this species appears within the range of P. vivipara, but it seems not to be related to it. Another interesting fact is that most fruits are modified to form hairy galls. The etiology of these galls remains unknown.

This species belongs to Peperomia subg. Micropiper (Miq.) Miq. (Frenzke et al. 2015).

Additional specimen examined

Peru, dept. Cajamarca, Prov. San Miguel, dist. La Florida: Road from Monteseco to Chorro Blanco, 1545 m, 6°50'51.3"S, 79°06'23.5"W, 10 Feb 2023, G. Pino & L.E.Alomía 3830, (USM 333267).

12. Peperomia riosaniensis Hutchison ex Pino, Samain & L.E. Alomía, sp. nov.

urn:lsid:ipni.org:names:77317653-1

Fig. 8A–H

Type material

Peru, [dept. Lambayeque, prov. Chiclayo, dist. Oyotún]: road to Hacienda Taulis, ca. 80 km up the Río Saña from the Pan American Highway, 500 m, [06°51'23"S, 79°09'31"W], 28 Aug 1964, P. C. Hutchison 6308 (holotype: UC[1298471]!, isotypes US [2485139]!, USM [Not found]).

Figure 8.

Peperomia riosaniensis Pino, Samain & L.E. Alomía, sp. nov. A, B plant in habitat C plant ex situ D early stages of development E detail of the leaves F young plant in cultivation G spadix in anthesis H mature spadix with fruits I comparison of the fruits of P. riosaniensis (above) and P. palmiformis (below).

Diagnosis

Perennial, succulent terrestrial herb similar to P. palmiformis Pino & Samain, differs in inflorescences that have wider panicles, with more (30–60 compared to 16–40) and shorter spadices (1–5 cm long compared to 2–8 cm long), leaves attached all along the stem, slightly longer (6–9 cm compared to 5–7 cm), more succulent, canaliculate instead of flat, with longer petioles (2 cm compared to 0.7 cm), lamina wider at proximal third (1.2–1.8 cm compared to 0.5–1.2), margin entire distally instead of serrate, fruits larger (0.9 × 0.6 mm compared to 0.4 × 0.3 mm), widely ovoid instead of narrowly ovoid.

Description

Perennial, succulent, terrestrial herb, living in gorges or the shade of other plants, (4–) 9–18 cm tall, up to 50 cm tall when flowering. Roots fibrous, light gray, 0.7–0.4 mm diam., 2–4 cm long. Stem mainly erect or shortly decumbent at base, straight, terete, slightly puberulous, 0.7–1.2 cm diam. at the base, gradually tapering to 4–8 mm at the apex, bright green, brownish at the base, with reniform leaf scars spirally displayed every 3–8 mm, 1.5–2 mm × 3–4 mm, brownish; very rarely branching dichotomously from the distal 10 cm, but mostly with 4–6 branches from the base, resulting in a caespitose plant. Leaves alternate, glabrous, spirally inserted, present all along the stem or in the distal third of the main stem or branches under dry conditions, young leaves smaller, narrower, very succulent, induplicate or canaliculate, older leaves less succulent; petiole reniform in section, gradually continuing with leaf, 0.5–2 cm long, 2.5–3 mm wide, 1.5–2 mm thick, lamina convex to induplicate, narrowly obovate, (3–) 6–9 cm long, 1.2–2.4 mm thick, (1.4–)2–3 cm wide at distal third, (1.4–)1.8–3 cm wide at the middle, (1–)1.4–1.8 cm wide at proximal third, apex subacute, distal third slightly recurved, base cuneate; adaxially bright glossy green, convex to slightly induplicate, obscurely 3–5-palmatinerved, nerves lighter colored; canaliculate or markedly induplicate, only the central nerve is visible in young leaves; margin entire; abaxially pale green, obscurely 3–5-palmatinerved, central nerve subcarinate, the others rather depressed. Inflorescence is an open terminal panicle with 30–60 alternate spadices emerging solitary or from secondary axes with 2–7 spadices, each born from 10–20 nodes appearing 5–10 cm from the base of the central axis; each node with a basal lanceolate acute bract 0.4–2 cm long, 0.3–0.7 cm wide, bright green; central axis 30–40 cm long, 3–6 mm diam. at the base, gradually tapering to 1 mm, terete, longitudinally furrowed, very light green; peduncle terete or slightly funnel-shaped distally, 3–10 mm long, 0.7–1.2 mm diam., each with a 2 mm long, 0.5 mm wide oval bract at the base, deciduous; rachis 1–5 cm long, 1.2–1.6 mm diam., light green. Floral bracts broadly oval to round, subacute, light green, 0.5–0.6 mm diam. Stamens, filaments 0.4 mm long, 0.2 mm diam., transparent, anthers white ovoid, 0.4–0.5 × 0.3 mm. Fruit an ovoid berry, 0.85–0.9 mm long, 0.55–0.6 mm diam., bright brown to olive green, minutely papillate, pedicel inconspicuous, dry anthers attached, style prominent, ovoid, bright brown, 0.15–0.2 mm long, 0.15–0.2 mm diam., stigma lighter colored.

Distribution and habitat

Plants grow from 450 to 600 m, the lowest and driest layer of the area studied, along the Saña valley between the towns of Oyotún and La Florida in the departments Lambayeque and Cajamarca, on rocky soil, in thickets shaded by shrubs and supplied with moisture from nearby water courses.

Phenology

Inflorescences appear from June to August; fruits ripen from July to September.

Etymology

The epithet was coined by Hutchison in 1967 after the Saña river, “Río Saña” in Spanish, which is Latinized to “riosaniensis” in the same way Rauhocereus riosaniensis was described by Backeberg. As Hutchison only labeled the herbarium sheets and never published this name it has been considered a nomen herbariorum (Mathieu 2007).

Notes

The closest species of Peperomia riosaniensis is P. palmiformis Pino & Samain (Pino et al. 2012) The web page “Peperomia.net” (http://www.peperomia.net/repertorysearch.asp) indicates that this taxon is merely a synonym of the latter. We compared both species in detail and the main differences between P. riosaniensis and P. palmiformis are resumed in Table 3.

Table 3.

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Comparison of the main differences between P. riosaniensis and P. palmiformis.

Features	P. riosaniensis	P. palmiformis
Height (vegetative in cm)	9–18	15–20
Distribution of leaves	All along the stem	Mainly on distal third.
Branches	Very common, basal, rarely dichotomous distally.	Rarely branched from the base, distal dichotomous branches.
Leaf (petiole)	0.5–2 cm long	3–7 mm long
	2.5–3 mm wide	1.2–1.8 mm wide
	1.5–2 mm thick	1–1.5 mm thick
Leaf (shape)	Canaliculate to induplicate, narrowly obovate. Distal third slightly recurved.	Mainly flat, occasionally slightly convex on both sides, very narrowly obovate.
Leaf (length in cm)	6–9	5–7
Leaf (thickness in mm)	1.2–2.4	0.5–3
Leaf (width)	2–3 cm wide at distal third	2–2.5 cm wide at distal third
	1.8–3 cm wide at the middle	1–2 cm wide at the middle
	1.4–1.8 cm wide at proximal third	0.5–1.2 cm wide at proximal third
Leaf (margin)	Completely entire.	Constantly serrate at distal third.
Inflorescence	Open panicle	Narrow panicle
	10–20 branches,	10–16 upright branches,
	30–60 spadices.	16–40 spadices.
Central axis	30–40 cm long	25–30 cm long
Central axis	3–6 mm diam. at base	3.5–5 mm diam. at base
Rachis of the spadix	1–5 cm long, 1.2–1.6 mm diam.	2–8 cm long, 1–1.4 mm diam.
Fruit (shape/color)	Ovoid, bright brown to olive green.	Narrowly ovoid, light olive green
Fruit (length in mm)	0.85–0.9	0.35–0.45
Fruit (diam. in mm)	0.55–0.6	0.25–0.35
Fruit (style)	Wide ovoid, bright brown,	Long ovoid to balaniform, bright orange,
	0.15–0.2 mm long,	0.25–0.30 mm long,
	0.15–0.2 mm diam.	0.22–0.28 mm diam.

While P. palmiformis grows from 800 to 900 m along the Utcubamba valley from Bagua to Pedro Ruiz in department Amazonas, Peru, P. riosaniensis grows 300 m lower. Nevertheless, the temperatures in the habitats of both species average 25–30 °C during the day and 18–22 °C at night, P. riosaniensis grows at 200 km distance from P. palmiformis, on the dry, xerophytic western slopes of the Andes, with 25 mm rain per month average in summer and 0–4 mm in winter (Clima.com: https://www.clima.com/peru/cajamarca/florida). Peperomia palmiformis lives on the flatter eastern slopes of the Andes, along the lower basin of the Utcubamba river, in a dry valley but already under the influence of the Amazon Forest, with nearly 60 mm rain per month in summer and 5 mm monthly in winter (Clima.com). Both species look similar from far away, but P. riosaniensis is slightly shorter when vegetative; its inflorescences have relatively wider panicles, with more horizontal branches and more spadices that are relatively shorter. Although leaves look very much alike when pressed, the new species has longer and thicker petioles, longer leaves that are not as narrow and succulent, wider at the proximal third, making its margin convex, compared to the concave margin of P. palmiformis at this segment. The new species also has induplicate or canaliculate leaves, adaxially slightly darker and glossier, frequently carinate beneath and slightly recurved at the distal third, compared to the straight, flat, mostly biconvex succulent leaves of P. palmiformis, mainly due to its more developed fenestra. The serrate distal margin of the leaves observed in P. palmiformis similar to P. erosa Hutchison ex Pino (Pino et al. 2012) has never been observed in the new species. Fruits are entirely different, more globular, larger, thicker, and papillate in P. riosaniensis, frequently with dry anthers attached, and lacking the conspicuous bright orange persistent basal pedicel. Peperomia palmiformis fruits also have a larger, bright orange style, with a bulge that is sometimes even thicker than the fruit itself. These morphological differences are supported by the fact that in a recent phylogenetic study, P. riosaniensis is located in a different branch from P. palmiformis that is more related to P. columella, P. ferreyrae, P. mathieui, and P. columnaris from the Utcubamba valley (Frenzke et al. 2015). All these species mentioned belong to Peperomia subgenus Fenestratae Pino (Frenzke et al. 2015).

Additional specimens were examined

Peru, dept. Lambayeque, prov. Chiclayo, dist. Oyotún: road from Oyotún to El Espinal, on the trail to waterfall Velo de la Novia, 515 m, 06°48'30.3"S, 79°11'08"W, 26 Jul 2020, G. Pino & L.E. Alomía 3237 (USM 330870); same road, 520 m, 06°48'27.4"S, 79°11'04.7"W, 26 Jul 2020, G. Pino & L.E. Alomía 3238 (USM 330871); Dept. Cajamarca, prov. Santa Cruz, dist. Catache: Road from El Espinal to La Florida, 582 m, 06°51'13.7"S, 79°09'16.5"W, 26 Jul 2020, G. Pino & L.E. Alomía 3230 (USM 330869); Road from Cayaltí to La Florida, 8.4 km from La Florida, 492 m, 06°50'51,6"S, 79°09'53,7"W, 1 Mar 2009, G. Mathieu & L. Symmank 2009-167 (BR, GENT, USM 258833!) Same road, 591 m, 06°51'13.4"S, 79°09'18.4"W, 28 Aug 2022, G. Pino et al. 3637 (USM 330882) prov. San Miguel, dist. La Florida, Puente el Papayo, on slopes at the borders of the roadside, 18 Jul 1982, S. Llatas Quiroz 861 (F 1931956, NY).

13. Peperomia rotundata Kunth. Nov. Gen & Sp. [Humboldt] 1: 67, tab.12. 1815.

Fig. 5H

Piper rotundatum (Kunth) Poir., Lam. Encycl. Meth. Suppl. 4: 468. 1816. Type: Based on Peperomia rotundata Kunth.

Piper villosum Rich. mss. in herb. Francov. in DC. Prodr. 16(1):442. 1869. Type: Based on Peperomia rotundata Kunth.

Type

Colombia, dept. Cauca: “Crescit in excelsis regni Novogranatensis, inter Pansitara et vallem Yacanocatu, alt. 980 hexap. Floret Novembri.” [Grows in elevations of the Kingdom of New Granade, between Pacitará and the Yacanocatu valley, at 1800 m, Flowers in November], Humboldt s/n, (holotype: B [lost?]; isolectotype: P!)

Distribution and habitat

Montane forests of Venezuela, Colombia, Ecuador, Peru, and Bolivia: rare, but uniformly distributed, terrestrial in shaded places growing on humus.

Note

This species belongs to Peperomia subg. Micropiper (Miq.) Miq. (Frenzke et al. 2015).

Specimen examined

Peru, dept. Cajamarca, prov. San Miguel, dist. Unión Agua Blanca: El Tingo, camino a Taulis, [on the way to Taulis] 3175 m, [06°59'26"S, 79°01'21"W], 18 Feb 2000, E. Rodríguez et al. 2366 (HUT 37499, F, M, MO).

14. Peperomia sagasteguii Pino, Samain & L.E. Alomía, sp. nov.

urn:lsid:ipni.org:names:77317654-1

Fig. 9A–G

Type

Peru, dept. Cajamarca, prov. Santa Cruz, dist. Catache: Monteseco, 1 Km above, on the path to Chorro Blanco, 1353 m, [06°51'17.8"S, 79°06'33.2"W], 16 Mar 1986, M.O. Dillon & A. Sagástegui 4315a (holotype: F [1978653]!).

Figure 9.

Peperomia sagasteguii Pino, Samain & L.E. Alomía, sp. nov. A plant in habitat in the dry period B plant ex situ in anthesis C plant in habitat with inflorescences D detail of the leaves E comparison of P. sagasteguii and P. trinervis at the beginning of anthesis F detail of immature spadix G mature spadix showing long pedicels and fruits. (Details: Above, mature fruit; Below, fruit standing on long pedicel).

Diagnosis

Perennial, semi-succulent, terrestrial, epiphytic, or saxicolous herb similar to P. trinervis, differs in stems more pubescent, dark red to olive green instead of bright red, leaves with a longer and wider petiole (7–17 long × 1.8–2 wide compared to 3–10 long × 1.2–1.5 wide), lamina more succulent, green adaxially and light green-pinkish abaxially instead of olive green with gray nerves adaxially and red-purple abaxially; spadices longer (4–20 cm compared to 10 cm long), but narrower.

Description

Perennial, semi-succulent, terrestrial, epiphytic, or saxicolous herb, living in the shade of other plants, 10–30 cm tall, up to 45 cm tall when flowering. Roots fibrous, around 1 mm diam., transparent-reddish, 2–6 cm long, emerging from basal nodes and stolons. Stem stoloniferous at the base, then assurgent, thicker below, straight erect, terete, dark red to olive green, 0.4–0.6 cm diam. at the base, gradually tapering to 1.4–1.6 mm at the apex, readily branched alternately, internodes 2.5–3 cm at the base gradually descending to 0.8–1 cm towards the apex, surface crisp-pubescent, trichomes whitish, around 0.5 mm long at base, slightly shorter upwards. Branches 5–9, irregularly alternate, sometimes pseudo-opposite below, emerging almost horizontally mainly from the distal half, 7–15 cm long in the vegetative state. Leaves alternate, elliptic, elliptic obovate to subrotundate; petiole terete, same surface and color as stem, 0.7–1.7 cm long, 1.8–2 mm wide, slightly channeled above; lamina slightly convex to flat, 2.5–4 cm long, 1–2 mm thick, 1.6–2.8 cm wide, apex acute to attenuately subacuminate, base cuneate to sub truncate; adaxially bright glossy green, convex, 3-palmatinerved, nerves depressed; margin entire; abaxially pale green to pinkish, 3-palmatinerved, nerves elevated. Young leaves can be pinkish adaxially. Both sides are puberulous. Inflorescence terminal in an umbel of 1–3 spadices and sometimes 1–2 individual spadices from upper leaf nodes, umbel with 1–3 basal bracts at the base and lateral spadices with opposite bracts at base similar to leaves but much smaller; peduncle terete, light green, 4–6 mm long, 1.2–1.8 mm diam.; rachis 4–15 (–20) cm long, 1.5–1.7 mm diam., light green. Floral bracts broadly oval to round, light green, flat, almost inconspicuous, 0.35–0.45 × 0.30–0.40 mm. Stamens, filaments short, anthers white ovoid, 0.3 × 0.4 mm. Fruita globose berry, borne on a 1–1.5 mm long, 0.25–0.3 mm diam. narrow-conical transparent pedicel; 0.7–0.8 mm long, 0.6–0.7 mm diam., brown, covered in white 0.05 mm long whitish papillae up to distal third of fruit, style conical, light brown, 0.15–0.2 mm long, 0.20–0.25 mm diam.

Distribution and habitat

Plants have only been found from 1200 to 1500 m of the middle course of the Saña River valley, in the remnants of montane forest, epiphytic or on rocks, always in shaded places. Peperomia sagasteguii is endemic to the Saña River valley.

Phenology

Inflorescences appear from October to November; fruits ripen from January to March.

Etymology

This species is dedicated to Abundio Sagástegui (1932–2012), collector of the type, a notable botanist, who worked at both Universidad Nacional de Trujillo and Universidad Privada Antonio Orrego, author of many books and articles, director of the journals “Bulletin of the Society of Botany of the UNT” and Arnaldoa, author of four genera and 97 species (Rodríguez and León 2012).

Notes

Most collections were determined initially as P. trinervis Ruiz & Pav., a species that seems to be very close, (see Table 4). The main differences are stem color, which is dull dark red and greenish in P. sagasteguii compared to the bright red of P. trinervis. The indumentum of P. sagasteguii seems to be denser and longer, like that of P. rotundata. Leaf petioles of P. trinervis are shorter and of similar color to its stems. The leaf color of P. sagasteguii is bright green adaxially with depressed nerves, while P. trinervis has a high color contrast between the dark olive green and the bright silver gray of the nerves. Abaxially leaf color of P. sagasteguii is light green, while P. trinervis is dark red with light green nerves. Peperomia sagasteguii has thicker, more succulent leaves. Finally, spadices of P. sagasteguii are much longer. The type collection was determined initially as P. blanda (Jacq.) Kunth, which bears no similarity and later it was determined as P. rotundata Kunth by R. Callejas, but this sample lacks the constant decussate leaves and inflorescences characteristic of that species. Some branches and leaves can be pseudo-opposite by shortening of the stem, leading to this confusion.

Table 4.

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Comparison of the main differences between P. sagasteguii and P. trinervis. Data of P. trinervis taken from Trelease and Yuncker (1950) and from live cultivated plants.

Features	P. sagasteguii	P. trinervis
Height (cm)	10–30 (–45)	20–25 or more
Internodes (in cm)	2.5–3 below, 0.8–1 above	2–3 below, 1 above
Stem (in mm)	4–6 diam. at the base, gradually tapering to 1.4–1.6	4–6 diam. at the base, gradually tapering to 1.5–2
Stem (color)	Dull dark red to olive green	Bright dark red
Leaf (petiole in mm)	7–17 long, 1.8–2 wide	3–10 long, 1.2–1.5 wide
Leaf (shape)	Elliptic, elliptic obovate to subrotundate.	Elliptic, elliptic obovate or elliptic obovate.
Leaf (measure)	2.5–4 cm long, 1–2 mm thick, 1.6–2.8 cm wide	2–3.5 cm long, less than 1 mm thick, 1.2–2.5 cm wide.
Leaf (color)	Adaxially bright glossy green, abaxially pale green to pinkish. Young leaves can be pinkish adaxially	Adaxially dark olive green with nerves in gray, abaxially bright dark red, nerves in light green. Young leaves can be purplish
Inflorescence	Terminal in an umbel of 1–3 spadices and sometimes 1–2 individual spadices from upper leaf nodes	Terminal and from the upper nodes.
Rachis of the spadix	4–15 (–20) cm long, 1.5–1.7 mm diam.	10 cm long, 1.8–2 mm diam.

This species belongs to Peperomia subg. Micropiper (Miq.) Miq. (Frenzke et al. 2015).

Additional specimens examined

Peru, dept. Cajamarca, prov. Santa Cruz, dist. Catache: Alrededor de campamento, Bosque Monteseco, borde de trocha sobre troncos secos [Around the campsite, Monteseco Forest, border of track, on dry logs] 1500 m, [06°50'58.6"S, 79°05'53.7"W], 20 Jan 1989, S. Leyva G. 015 (F 2016269, USM 90066) Prov. San Miguel, dist. La Florida: Buenos Aires, La Florida, sobre troncos de Guaba [on trunks of Inga sp.] 1000 m, [06°52'27.3"S, 79°07'10"W], 22 Mar 1986, S. Llatas Quiroz & R. Palacios 1822 (F 1963941). Road from La Florida to Monteseco, just before entering the reserved zone, 1531 m, 6°51'00.7"S, 79°05'58.2"W, 10 Feb 2023, G. Pino & L.E. Alomía 3831 (USM 333270). dist. Niepos: Road from La Florida to Monte Seco (Naranjo), 5.8 km from La Florida, 1395 m, 06°53'25.5"S, 79°06'57.3"W, 1 Mar 2009, G. Mathieu & L. Symmank 163 (USM); Road from La Florida to Niepos, 1297 m, 06°53'06.9"S, 79°06'47.6"W, 28 Aug 2022, G. Pino et al. 3641 (USM 333268); Same road, 1393 m, 06°53'25"S, 79°06'57"W, 28 Aug 2022, G. Pino et al. 3644 (USM 333269).

15. Peperomia symmankii Pino & Samain, sp. nov.

urn:lsid:ipni.org:names:77317655-1

Fig. 10A–F

Type

Peru, dept. Cajamarca, prov. San Miguel, dist. La Florida: road from Cayaltí to La Florida, 731 m, 06°51'57.6"S, 79°08'32.3"W, 1 Mar 2009, G. Mathieu & L. Symmank 162 (holotype: USM[255772]!).

Figure 10.

Peperomia symmankii Pino & Samain, sp. nov. A plant in habitat B plants ex situ in several stages of development C plant in habitat in blossom D detail of the leaves E plant ex situ in anthesis with bracts. (Detail: mature panicle) F young spadix (Detail: Mature spadix with fruits).

Diagnosis

Perennial, succulent, caulescent, terrestrial herb, similar to P. ricardofernandezii Pino & Samain, but shorter, less branched, and with more tortuous stems. Leaves are smaller (2.5–4.5 cm long compared to 4.5–8 cm long), more widely ovate, with an obtuse or rounded apex instead of acute-acuminate. Petioles are shorter (0.3–3.5 cm long compared to 2–4 cm), lighter in color, and attached closer to the cordate base, which is sometimes auriculate and overlapping compared to the truly peltate insertion of P. ricardofernandezii. The inflorescence is narrower and longer but otherwise very similar, with longer spadices (3–20 mm compared to 2–4 mm).

Description

Perennial, succulent, caulescent, terrestrial herb, 12–20 cm when vegetative, up to 5 cm when flowering, growing on soil among rocks at slightly shaded places, near water courses. Vegetative stem, 0.6–1.5 cm diam. At the base, light gray brownish, erect at first, then sometimes decumbent, rooting from the base, terete in the rainy period, slightly furrowed under dry conditions, with flat or projected spirally arranged round to reniform scars, 0.15–0.3 cm tall, 0.25–0.45 cm wide, distanced every 0.5–1 cm, corresponding to decurrent petioles. Leaves in young plants perennial, alternate, spirally attached, present at the distal 2–4 cm, glabrous; petiole reniform in cross-section, 0.5–3.5 cm long, 2–2.5 mm wide, erect or oblique upwards, light green or with a reddish blush, inserted almost at the base of the leaves, subpeltate at 1–2 mm from abaxial base; lamina widely ovate, 2–4.5 cm long, 2–4.5 cm wide at proximal third, 2.5–5 cm wide at the middle, 2.5–4 cm wide at distal third, apex obtuse, occasionally rounded, base cordate, sometimes auriculate and overlapping; adaxially dull green, flat to concave. 7-palmatinerved, nerves depressed; margin entire, undulated at distal half; abaxially very light green, sometimes reddish, nerves slightly carinate, darker green. A reproductive stem develops in the rainy season, from the apex the first time or from the next most distal node, with bracts similar to leaves, their lamina is first corrugate with undulate margins, and reddish abaxially, later they become larger, 4–7 cm long, 5–7 cm wide at proximal third, 3–6.5 cm wide at the middle, 4–5.5 cm wide at distal third, flat, adaxially bright green, abaxially very light green with nerves less conspicuous, apex acute to obtuse, base cordate, margins entire, petiole 4–7 cm long, light green. All bracts and inflorescences are deciduous after flowering. Inflorescence 2–5 (–10) panicles born alternately from the distal stem towards the apex, appearing from February to March. Each panicle is a raceme of 15–45 conferted, spirally arranged, horizontally inserted spadices around a vertical central axis, gradually opening from base to apex, longer and whiter at the base and shorter and greener distally. Central axis 10–30 cm long in total, 0.4–0.5 mm diam. at the base, gradually tapering to 0.9–1 mm, terete, light green to reddish, whitish towards the apex; peduncle terete or slightly funnel-shaped distally, 1–2 mm long, 0.3–0.4 mm diam., bright white; rachis 3–20 mm long, 0.4–0.5 mm diam., longitudinally furrowed, bright white. Floral bracts peltate, discoid, greenish white, 0.25–0.3 mm diam. Stamens white, pink when dry, filaments 0.2 mm long, transparent, anthers white ovoid, 0.15–0.20 mm. Ovary globose, white, 0.35–0.45 mm diam., stigma fimbriate. Fruit an ovoid berry, 0.75–0.85 mm long, 0.55–0.65 mm diam., olive green-brownish, minutely papillate, pedicel inconspicuous, style very broadly conical, white, stigma brown.

Distribution and habitat

Plants are rare and have been found only at four spots from 700 to 2400, semi-shaded on soil accumulated among huge rocks, on moist places near watercourses.

Phenology

Inflorescences appear from December to March; fruits ripen from February to April.

Etymology

The epithet is dedicated to Lars Symmank, a botanist from Dresden, Germany, who together with Guido Mathieu collected the specimen of this new species in the first expedition to Peru organized by Ghent University in 2009. He worked in several studies of subgenus Tildenia (Mathieu et al. 2011; Samain et al. 2011; Symmank et al. 2011).

Notes

The closest species to P. symmankii is P. ricardofernandezii from Piura. This species is mainly terrestrial, taller, and many branched with a bush-like appearance contrasted to the creeping habit of P. symmankii. The main difference is the leaf size and shape: P. symmankii has relatively smaller leaves, more widely ovate, with an obtuse to round apex, compared to the narrow ovate, acute-acuminate leaves of P. ricardofernandezii. Petioles of the new species are shorter, lighter in color, attached closer to the leaf base, which is truly cordate, even sometimes auriculate and overlapping, compared to the more peltate insertion in P. ricardofernandezii. The inflorescence of P. symmankii is narrower and longer but otherwise very similar, with slightly longer spadices (Table 5).

Table 5.

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Comparison of the main differences between P. symmankii and P. ricardofernandezii.

Features	Peperomia symmankii	Peperomia ricardofernandezii
Height (vegetative in cm)	12–20	10–50
Height (flowering, in cm)	25–45	15–60 or more
Petioles (surface and color)	Glabrous, light green or with reddish blush	Glabrous, burgundy red.
Petioles (length and insertion)	0.5–3.5 cm	2–4 cm
Petioles (length and insertion)	Subpeltate 1–2 mm from abaxial base	Subpeltate 2–3 mm from abaxial base
Leaves (shape and color)	Widely ovate, apex acute, base cordate; adaxially dull green, abaxially very light green, sometimes with a slight blush.	Narrowly ovate, apex acute to subacuminate, base subcordate; adaxially light green, abaxially reddish to burgundy red.
Leaves (size)	2–4.5 cm long	4.5–8 cm long
	2.5–4 cm wide at distal third	1.5–3 cm wide at distal third
	2.5–5 cm wide at the middle	2.5–6 cm wide at the middle
	2–4.5 cm wide at proximal third	2.5–5.5 cm wide at proximal third
Vegetative stem	Perennial, usually one branch from subterminal bud.	Perennial, 1–3 branches from subterminal bud.
Bracts (petiole)	4–5 cm long, light green, inserted subpeltate at 1–3 mm from abaxial base	2–10 cm long, red, inserted subpeltate at 2–5 mm from abaxial base
Bracts (shape and size)	Widely ovate	Ovate
	4–7 cm long	4.5–8 cm long, 2.5–6 cm wide,
	3–7 cm wide,	apex acuminate and clearly acute distally
	apex acute to obtuse	apex acuminate and clearly acute distally
Central axis of panicle (length in cm)	10–30	3.5–7
Rachis of spadix (length in mm)	3–20	2–4

This species belongs to Peperomia subg. Panicularia Miq. (Frenzke et al. 2015).

Additional specimens examined

Peru, dept. Cajamarca, prov. San Miguel, dist. La Florida: El Papayo bridge over Saña River, East bank, 10 m to North, under the shade of Guadua angustifolia plants, on rocks, 731 m, 06°51'57.6"S, 79°08'32.3"W, 28 Aug 2022, G. Pino et al. 3639 (USM 333272). El Papayo, sobre materia orgánica de las rocas. [El Papayo, on rocks with the decayed organic matter] 850 m, [06°51'S, 79°08'W], 1 Feb 1986, S. Llatas Q. 1770 (HUT 22496); Road from Florida to Monteseco, 1095 m, 6°51'41.3"S, 79°07'08.9"W, 10 Feb 2023, G. Pino & L.E. Alomía 3832; (USM 333273). Dist. Bolívar: Bosque Oscuraná, Caserío el Nogal, Bosque Húmedo [Oscuraná Wet Forest, El Nogal Hamlet], 2400 m, [6°56'51.0"S, 79°08'04.0"W], Sep 9 2003, A. Juárez et al. s.n. (HLL 435, HUT 42219). Prov. Santa Cruz, dist. Catache: Carretera al Bosque de Monteseco, borde de Carretera. [Road to Monteseco forest, border of road], 1200 m, [06°51'21"S, 79°06'42"W], 20 Jan 1996, E. Rodríguez 750 (HUT 30979).

16. Peperomia vivipara Pino, Samain & L.E. Alomía, sp. nov.

urn:lsid:ipni.org:names:77317656-1

Fig. 11A–F

Type

Peru, dept. Cajamarca, prov. Santa Cruz, dist. Catache: Monteseco Forest. 1500 m, [06°50'58.6"S, 79°05'53.7"W], 19 Dec 1984, A. Sagástegui et al. 12396 (holotype: HUT [19867] !; isotype: F [2038843] !).

Figure 11.

Peperomia vivipara Pino, Samain & L.E. Alomía, sp. nov. A young plant in habitat B mature plant in habitat showing offsets C plant ex situ showing offsets D propagules of different sizes just detached from mother plant E detail of leaves F detail of a branch: adaxial side (left), abaxial side (right) G branch with terminal spadix H spadix in anthesis.(Detail: Above, before; and below, after development of stamens).

Diagnosis

Perennial, semi-succulent, caespitose herb similar to P. alata, but more succulent, with thicker and more stout branches, leaves are shorter, elliptic, acute based, instead of lanceolate and long acuminate. Leaf bases are obtuse or subtruncate instead of cuneate. After flowering, propagules are developed at the apex of old branches that readily wither to assure vegetative propagation.

Description

Perennial, semi-succulent, caespitose herb, growing terrestrial, epiphytic, or saxicolous in the shade of other plants, 30–45 cm tall. Roots fibrous, emerging from the base of plants, 0.7–1 mm diam., light brown. Stem stoloniferous at the base, from which 3–10 deciduous branches emerge, straight erect, sometimes in zig-zag towards the apex, terete at the base, then prominently winged towards the apex with a triangular section, dark olive green, slightly puberulous at the base, then glabrescent towards the apex, 0.7–1 cm diam. At the base, gradually tapering to 0.3–0.5 mm at the apex, rarely branched close to the base, internodes 2.5–5 cm at the base gradually descending to 0.5–0.7 cm towards the apex, wings prominent at the distal half of the stems, from below the node towards the node after the next, semitransparent, 1–2 mm tall. Leaves alternate, distichous, glabrous, elliptic to elliptic ovate; petiole short, decurrent to next opposite node, reniform in section, channeled above, reddish green-pinkish, 0.2–0.4 (–1) cm long, 1.5–2 mm wide; lamina 3–6.5 cm long, 0.5–1.5 mm thick, 1.8–3.2 cm wide, apex acute to very slightly acuminate, slightly recurvate, base sub truncate; adaxially bright glossy green in young leaves, then very dark army green and even bluish, flat to slightly canaliculate, 3–5-palmatinerved, nerves depressed; margin entire; abaxially very light green with a pink blush, 3-palmatinerved, nerves elevated. Inflorescence simple terminal or up to 3 individually from upper leaf nodes opposite to the leaves; peduncle terete to infundibuliform, bright green, 2–9 mm long, 1.4–1.5 mm diam.; rachis 3–6 cm long, 2–3 mm diam., light green. Floral bracts rotundate, light green, flat, almost inconspicuous, only darker green than rachis, 0.4–0.5 mm diam. Stamens, filaments transparent, 0.2 mm diam., 1.5–3 mm long, anthers white ovoid, 0.4–0.5 mm long, 0.3–0.4 mm wide. Fruit not seen. Propagule developing at the apex of the branch, 1–5 cm tall, persistent until the whole stem dries.

Distribution and habitat

Plants grow from 1500 to 1800 m of the middle course of the Saña River valley, in the remnants of montane forest, epiphytic or on rocks, always in shaded places.

Phenology

Inflorescences appear from December to March; fruits ripen from February to April.

Etymology

The epithet stands for the most striking feature of this plant, the production of offsets directly from the mother plant, bypassing germination, as a method of survival in extreme weather conditions, from the Latin viviparus (that brings forth its young alive).

Notes

The closest species to P. vivipara is P. alata Ruiz & Pav., a species of wide distribution in Mexico, Central America, and all countries of South America except Chile, Argentina, and Uruguay, in tropical forests at lower elevations. Plants are similar in habit and size, they share the zig-zag winged internodes, although P. vivipara is stouter, with thicker and more erect branches with longer internodes. The main difference is in leaves, which are shorter, elliptic with an acute short apex and an obtuse or almost truncate base in P. vivipara compared to the lanceolate, long acuminate, and acute-based leaves of P. alata. A further difference is that P. alata does not produce propagules at the distal stems. (Table 6) These plantlets, that drop readily and root close to the mother plant propagating the plants vegetatively, have not been seen in any other species of Peperomia and give the name to the epithet of the species.

Table 6.

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Comparison of the main differences between P. vivipara and P. alata.

Features	Peperomia vivipara	Peperomia alata
Height	30–45 cm	30 cm or more
Stem (diam. at base)	7–10 mm	3–8 mm
Internodes (distance)	2.5–5 cm	1–3 cm
Petioles (length and diam.)	2–10 mm long	3–20 mm long
Petioles (length and diam.)	1.5–2 mm diam.	0.8–2 mm diam.
Leaves (shape)	Elliptic to elliptic ovate,	Elliptic to elliptic-lanceolate,
	apex acute to slightly acuminate,	apex long acuminate,
	base obtuse, subtruncate	base acute, cuneate
Vegetative leaves (size)	3–6.5 cm long	4.5–15 cm long
Vegetative leaves (size)	1.8–3.2 cm wide	2–4.5 cm wide
Spadices (length)	3–6 cm	6–8 (–15) cm
Propagules at distal stem	Present	Absent

This species belongs to Peperomia subg. Micropiper (Miq.) Miq. (Frenzke et al. 2015).

Specimens examined

Peru, dept. Cajamarca, prov. Santa Cruz, dist. Catache: Monteseco, 5 km above, on the path to Chorro Blanco, 1500 m, [06°50'58.6"S, 79°05'53.7"W], 16 Mar 1986, M.O. Dillon & A. Sagástegui 4366 (F 1993688); Same place, on the path below the campsite, 1450 m, [06°51'10.8"S, 79°06'09.4"W], 19 Mar 1986, M.O. Dillon & A. Sagástegui 4432 (F 1993348); Monteseco, 3 km NE, 1750 m, 06°50'51.6"S, 79°06'14.7"W, 13 May 1987, J. Santisteban C. & J. Guevara B. 044 (HUT 24730, F1995346). Prov. San Miguel, dist. Niepos: Road from La Florida to Monte Seco (Naranjo), 14.5 km from La Florida, 1643 m, 06°54'04.8"S, 79°07'33.5"W, 1 Mar 2009, G . Mathieu & L. Symmank 165 (USM 258839); Road from Naranjo to La Florida, 1632 m, 06°54'00.2"S, 79°07'59.0"W, Jul 26 2020, G. Pino & L.E. Alomía 3214; Same road, 1613 m, 06°54'03.3"S, 79°07'33.5"W, 26 Jul 2020, G. Pino & L.E. Alomía 3215 (USM 330866); Idem, 1612 m, 06°54'03.3"S, 79°07'33.5"W, Feb 2 2023, G. Pino & L.E. Alomía 3833 (USM 333271); Idem, 1393 m, 06°53'25.3"S, 79°06'57.7"W, 26 Jul 2020, G. Pino & L.E. Alomía 3216 (USM 330867); Road from La Florida to Niepos, 1393 m, 06°53'25"S, 79°06'57"W, 28 Aug 2022, G. Pino et al. 3643 (USM 330886); Same road, 1613 m, 06°54'03.3"S, 79°07'33.5"W, 28 Aug 2022, G. Pino et al. 3645 (USM 330887).

Discussion

This research found 16 taxa of Peperomia in a radius of less than 25 km (Fig. 2). Previous studies had reported nine species of Peperomia in the protected area of Udima, north of the town of Monteseco (Sagástegui and Dillon 1991) and only four in the still unprotected forest La Oscurana in Bolívar (Juárez et al. 2005). It is noticeable that previous studies focus on “relict forests” that represent a sample of the former extension of the natural extents of these woodlands, but neglect to sample semi-arid environments that can also be rich in species, as we show in this article. This species richness could be explained by the influence of the AHZ, but the closest valleys to the north and south of Río Saña, also inside this zone have very scarce collections of this genus. Perhaps they deserve more research to verify if they share the same biodiversity.

We have observed that the highest diversity of Peperomia species occurs between 500 and 1600 m, although more rainfall and vegetation can be expected at higher altitudes. The species found in the upper layers (P. hispidula, P. rotundata. P. microphylla, P. galioides) have a broad distribution in the department of Cajamarca (Pino 2004) and even in the rest of the Americas. The fact that the endemic or rare species are concentrated in the middle range of mountain systems is an observation consistent with other floral studies (Vergara et al. 2017) but has no demonstrated explanation yet.

Another observation is that in the lower range, where more droughts threaten the diversity of species of Peperomia, we have found two species of subgenus Fenestratae and one of subgenus Panicularia. These subgenera are restricted to Ecuador and Peru, and they show an extreme adaptation to prolonged periods of drought through succulence. Other families adapted to this desertic environment, such as Cactaceae, – are also rich in diversity in this lower layer (Mischler 1988; Alomía 2020). Finally, our findings support the hypothesis that there are still many species of Peperomia in Peru awaiting their discovery.

Acknowledgements

The present article is part of the research made for the Ph.D. Thesis – Review of subgenus Fenestratae (Peperomia, Piperaceae) of the first author, research inscribed by the Dirección de Posgrado, Facultad de Ciencias Biológicas, Universidad Nacional Mayor de San Marcos, Lima, Perú, by Dict. N° 149v-UPG-FCB-2021. We thank the authorities of Peru, (SERFOR, Ministry of Agriculture) for permission to collect material, (permit number 009-2009-AG-DGFFS-DGEFFS and 0124-2011-AG-DGFFS-DGEFFS), especially to Irma Hellen Castillo for her help and orientation; to the staff of USM Herbarium: Asunción Cano, Severo Baldeón, Hamilton Beltrán, Marissa Ocrospoma, Isabeu Cruz and Julio Torres. To Eric Rodríguez from HUT, for all his support, to Esteban Martínez, Emilio Pereyra, Luisa Infante, Merli Saavedra, Jorge Ernesto Romero Banda, Luis Alomía Díaz, Edith Collazos Balarezo, and Dannyel Alomía Collazos for assistance with fieldwork and logistics. To Carmen Martínez, Nicolás Cansinos and the staff of the Botanical Garden of the “Parque de Las Leyendas” for the facilities to cultivate live plants for this study. We are very grateful to both anonymous reviewers for their critical comments which have improved our manuscript. All photos by the authors except: 6D “Peperomia haematolepis” by Apipa is licensed under CC BY-NC https://www.inaturalist.org/taxa/1142412-Peperomia-haematolepis/browse_photos.

This research was funded by its authors.

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Supplementary material

Supplementary material 1

Localities of Peperomia species from the Saña River Valley, Peru

Guillermo Eloy Pino Infante, Marie-Stéphanie Samain, Luis Enrique Aarón Alomía Collazos

Data type: COL (excel document)

Explanation note: Database of localities of Peperomia species found at the Saña River Valley Peru, data obtained from Herbarium sheets, personal collections, or calculations based on data from Herbarium labels.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.

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﻿Abstract

﻿Resumen

Keywords

Palabras clave

﻿Introduction

﻿Materials and methods

﻿Study area

﻿Plant material

﻿Results

﻿Taxonomy

﻿Key to the species of Peperomia from the Saña River Valley

﻿1. Peperomia cacaophila Trel. & Yunck., Piperac. N. South Amer. 2(610). 1950.

Type

Distribution and habitat

Notes

Specimens examined

﻿2. Peperomia cymbifolia Pino, Pino, Peperomias de Cajamarca: 16–17, 50–51 f. 3 A–N. 2004.

Type

Distribution and habitat

Notes

﻿ Peperomia cymbifolia Pino var. occidentalis Pino & L.E.Alomía, var. nov.

Diagnosis

Type

Distribution and habitat

Phenology

Additional specimens examined

﻿3. Peperomia dolabriformis Kunth, Nov. Gen & Sp. [Humboldt] 1: 50, tab. 4. 1815.

Type

Distribution and habitat

Note

Specimens examined

﻿4. Peperomia emarginulata C.DC. DC. Prodr. 185. 16(1): 433. 1869.

Type

Distribution and habitat

Notes

Specimens examined

﻿5. Peperomia fraseri C.DC., J. Bot. 4: 134. 1866.

Type

Distribution and habitat

Specimens examined

Note

﻿6. Peperomia galioides Kunth, Nov. Gen & Sp. [Humboldt] 1: 58, tab. 17. 1815.

Type

Distribution and habitat

Note

Specimens examined

﻿7. Peperomia haematolepis Trel., Field. Mus. Pub. Bot. [Macbride] 13(2):52. 1936.

Type

Distribution and habitat

Notes

Specimen examined

﻿8. Peperomia hispidula (Sw.) A. Dietr., Sp. Pl. 1: 165. 1831.

Type

Distribution and habitat

Note

Specimen examined

﻿9. Peperomia inaequalifolia Ruiz & Pav., Flora Peruviana & Chilensis 1: 30, plat. 46 fig.a. 1798.

Type

Distribution and habitat

Note

Specimens examined

﻿10. Peperomia microphylla Kunth, Nov. Gen & Sp. [Humboldt] 1: 57, tab. 15, fig. 2. 1815.

Type

Distribution and habitat

Note

Specimen examined

﻿11. Peperomia pilocarpa Pino, Samain & L.E. Alomía, sp. nov.

Type

Diagnosis

Description

Distribution and habitat

Phenology

Etymology

Notes

Additional specimen examined

﻿12. Peperomia riosaniensis Hutchison ex Pino, Samain & L.E. Alomía, sp. nov.

Type material

Diagnosis

Description

Distribution and habitat

Abstract

Resumen

Introduction

Materials and methods

Study area

Plant material

Results

Taxonomy

Key to the species of Peperomia from the Saña River Valley

1. Peperomia cacaophila Trel. & Yunck., Piperac. N. South Amer. 2(610). 1950.

2. Peperomia cymbifolia Pino, Pino, Peperomias de Cajamarca: 16–17, 50–51 f. 3 A–N. 2004.

Peperomia cymbifolia Pino var. occidentalis Pino & L.E.Alomía, var. nov.

3. Peperomia dolabriformis Kunth, Nov. Gen & Sp. [Humboldt] 1: 50, tab. 4. 1815.

4. Peperomia emarginulata C.DC. DC. Prodr. 185. 16(1): 433. 1869.

5. Peperomia fraseri C.DC., J. Bot. 4: 134. 1866.

6. Peperomia galioides Kunth, Nov. Gen & Sp. [Humboldt] 1: 58, tab. 17. 1815.

7. Peperomia haematolepis Trel., Field. Mus. Pub. Bot. [Macbride] 13(2):52. 1936.

8. Peperomia hispidula (Sw.) A. Dietr., Sp. Pl. 1: 165. 1831.

9. Peperomia inaequalifolia Ruiz & Pav., Flora Peruviana & Chilensis 1: 30, plat. 46 fig.a. 1798.

10. Peperomia microphylla Kunth, Nov. Gen & Sp. [Humboldt] 1: 57, tab. 15, fig. 2. 1815.

11. Peperomia pilocarpa Pino, Samain & L.E. Alomía, sp. nov.

12. Peperomia riosaniensis Hutchison ex Pino, Samain & L.E. Alomía, sp. nov.

13. Peperomia rotundata Kunth. Nov. Gen & Sp. [Humboldt] 1: 67, tab.12. 1815.

14. Peperomia sagasteguii Pino, Samain & L.E. Alomía, sp. nov.

15. Peperomia symmankii Pino & Samain, sp. nov.

16. Peperomia vivipara Pino, Samain & L.E. Alomía, sp. nov.

Discussion

Acknowledgements

References