Monograph |
Corresponding author: Tatiana Erika Boza Espinoza ( tatianaerika@gmail.com ) Academic editor: Hugo de Boer
© 2022 Tatiana Erika Boza Espinoza, Michael Kessler.
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:
Boza Espinoza TE, Kessler M (2022) A monograph of the genus Polylepis (Rosaceae). PhytoKeys 203: 1-274. https://doi.org/10.3897/phytokeys.203.83529
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We present a monograph of the high Andean tree genus Polylepis (Rosaceae), based on a species concept considering morphological, climatic and biogeographic distinctness as indicators of evolutionary independence. In total, we recognize 45 species of Polylepis, grouped in five sections. Polylepis sect. Sericeae is represented by 15 species in four subsections, P. sect. Reticulatae by seven species, P. sect. Subsericantes by three species, P. sect. Australes by two species and P. sect. Incanaee by three subsections with 18 species. We describe seven new species, one from Colombia (P. frontinensis), one from Ecuador (P. simpsoniae) and five from Peru (P. acomayensis, P. fjeldsaoi, P. occidentalis, P. pilosissima and P. sacra). Three species from Peru (P. albicans, P. pallidistigma and P. serrata) are re-instated as valid species. Two taxa from Bolivia (P. incanoides and P. nana) are elevated from subspecies to species rank. The morphology, habitat, distribution, ecology and conservation status of each species are documented. We also provide an identification key to the species of the genus and general introductions on taxonomic history, morphology, evolution, ecology and conservation.
Andes, morphology, new taxa, taxonomy
Polylepis may well be the most emblematic tree genus of the central and northern Andes. It often occurs in an otherwise treeless landscape, forming the highest forests in the Western Hemisphere at elevations of over 4800 m. Polylepis forests contain unique biodiversity and provide habitats for a wide range of Andean plants and animals. However, they also provide crucial ecosystem services for the people living in the Andes: clean water, protection against erosion, firewood, fodder and medicinal plants, amongst others. On the other hand, it has been estimated that over 90% of the natural cover by Polylepis forests has been lost over millennia of human land use, so that the majority of species are considered to be of conservation concern. For this reason, studies on the physiology, ecology and conservation of the genus Polylepis have led to the publication of hundreds of scientific papers and reports and to the establishment of numerous conservation areas and projects to safeguard the last forest remnants.
Against this background, the poor taxonomic knowledge of the genus is troublesome. Although the first monograph of the genus was already published in 1911, to date, there is still no consensus on the number of species and their delimitation, greatly hampering conservation and management of the species and forests. The reasons for the taxonomic chaos in the genus are to be found in the high phenological variability of the individual species, coupled with great similarity between species, extensive hybridization and gene flow between species, polyploidization and, possibly, even apomictic reproduction (asexual seed reproduction). Due to these reasons, standard species concepts are difficult to apply to the genus, so that different authors have taken different approaches, arriving at different species circumscriptions. The last comprehensive monograph of the genus was published in 1979 and, due to the numerous discoveries since then, there was a strong need for a modern taxonomic treatment of the genus.
In this study, we undertook this effort by combining 35 years of field and herbarium experience of the co-author, MK, with the fresh view of PhD student TB (co-author). Together, we have studied all, but three species (P. longipilosa, P. occidentalis, P. quadrijuga) of the genus in the field and have applied a novel species concept for Polylepis which is based on a combination of morphological, ecological and biogeographical distinctness. Our species concept is narrower than previous concepts and accounts for the evolutionary independence of geographically disjunct forms that were previously combined in broadly defined species. As a result, we here recognize 45 species (up from 26 in the last global count published in 2006), including seven newly-described species. We consider that this classification more accurately reflects the evolutionary variability of Polylepis than previous classifications and will provide a workable background for the conservation and management of Polylepis forests and associated biota. Naturally, every new classification must stand the test of practicability and we look forward to this in the future.
The species concept used here is the general lineage concept of
Species delimitation and identification in Polylepis relies on populations instead of single herbarium specimens, because there is phenotypical variability even within a single plant (e.g., leaf size, texture, shape and indumentum). This variability depends on the growing stage of a branch and its exposure (sun or shade). Between individuals of a species, phenotypic variability is even more pronounced depending on genetic background, as well as growing and microhabitat conditions, with individuals from sheltered and humid sites having larger leaves with more leaflets, less dense hair cover and longer inflorescences with more flowers.
We also, for the first time, provide a formal sectional and subsectional infrageneric classification of the genus to facilitate communication. This classification is based on morphological differentiation supported by climatic niches and ploidy levels. In this sectional classification, we clustered species, based on morphological similarity (number of lateral leaflet pairs, indument type of the lower leaflet surfaces, stipule sheaths and/or fruits, leaflet apex shape and fruit shape). This morphologically based separation is broadly supported by biogeography (with most species of a section or subsection replacing each other geographically), similarity in climatic niches and similarity in ploidy levels. In the two larger, more variable sections, we also applied a subsectional classification to group similar species. The orthography of sectional and subsectional names has been designated in accordance with the ‘Shenzhen Code’ (
The sections defined by us largely correspond to the informal groups as outlined by
Our study is based primarily on the examination of the external morphology of herbarium specimens, supplemented with observation of rehydrated material, photographs and living plants in the field. Field studies by TB were conducted in Peru (Ancash, Arequipa, Ayacucho, Cajamarca, Cerro de Pasco, Cusco, Huancavelica, Junín, Lima, Moquegua, Puno and Tacna) and Ecuador (Azuay, Chimborazo, Loja, Napo and Pichincha). MK has studied Polylepis in Venezuela (Mérida), Colombia (Antioquia), Ecuador (Azuay, Napo, Pichincha), Peru (Ancash, Arequipa, Cuzco, Lima and Puno), Bolivia (Chuquisaca, Cochabamba, La Paz, Oruro, Potosí, Tarija) and Argentina (Córdoba, Jujuy, Salta, Tucumán). Together, we have studied all, but three species of Polylepis in the field. Voucher specimens were collected according to standard herbarium techniques. Photographs of plants and associated notes were taken in the field.
All the herbarium specimens representing each species were carefully observed and those spanning the morphological variation and geographical distribution range were chosen for measurements. Characters were measured from corresponding positions on mature, reproductive plants in order to minimize variation due to developmental differences. We have chosen characters for measurements, based in part on those that have been previously used to differentiate species within the genus Polylepis (
We combined occurrence records, based on herbarium specimens with field locations and observations from previous studies from multiple sources into a comprehensive, relational database. This database comprises approximately 3,250 quality-checked records of all species of Polylepis. All coordinates were cleansed and georeferenced if needed. We created the distribution maps using QGIS 2.18.14.
We extracted Mean Annual Precipitation (MAP) and Mean Annual Temperature (MAT) as climatic variables from the global climatic model CHELSA version 1.2 (
We based our conservation assessment on the International Union for Conservation of Nature guidelines (
The genus Polylepis was described by
Along with the circumscription of the genus,
In the first modern revision of the genus Polylepis,
Accordingly, prior to the present study, there were 88 names available in Polylepis, of which 27 were recognized as valid at species level. In our study, we consistently applied a narrow species concept (see Material and Methods), re-instating four species to species level and describing 11 species as new. Seven of these new species are described here, while four have been described in collaboration with colleagues, namely P. argentea (
In his revision of the genus,
The infrageneric classification adopted here includes sections partly corresponding to the three informal and meaningful groups previously defined, based on morphology and supported by climatic niches and ploidy levels. We propose the five sections Sericeae, Reticulatae, Subsericantes, Australes and Incanaee (Table
Alignment of the species of Polylepis according to the infrageneric classifications of
Species |
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This study | ||
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Section | Group | Group | Section | Subsection | |
P. lanuginosa | Dendracaena | unnamed / Latifoliatae | Sericeae | Sericeae | Lanuginosae |
P. multijuga | Plurijugae | ||||
P. longipilosa | — | Pauta | |||
P. pauta | Annulatipilosae | ||||
P. serrata | Plurijugae | ||||
P. albicans | unnamed | Sericeae | |||
P. argentea | — | ||||
P. canoi | — | ||||
P. frontinensis | — | ||||
P. humboldtii | — | ||||
P. loxensis | — | ||||
P. ochreata | Annulatipilosae | ||||
P. sericea | |||||
P. pepei | — | Pepea | |||
P. rodolfo-vasquezii | — | ||||
P. hieronymi | Subtustomentosae | Reticulatae | |||
P. microphylla | unnamed | Reticulatae | |||
P. occidentalis | — | ||||
P. quadrijuga | Supranitidae | ||||
P. reticulata | |||||
P. simpsoniae | |||||
P. weberbaueri | |||||
P. australis | unnamed* | Incanaee | Australes | ||
P. neglecta | Gymnopodae | — | |||
P. flavipila | — | Subsericantes | |||
P. pilosissima | — | ||||
P. subsericans | — | Sericeae / Incanaee | |||
P. acomayensis | — | Incanaee | Incanaee | Racemosae | |
P. incarum | — | ||||
P. lanata | — | ||||
P. pacensis | — | ||||
P. racemosa | unnamed* | ||||
P. sacra | — | ||||
P. triacontandra | Paucijugae | ||||
P. besseri | unnamed | Besseria | |||
P. crista-galli | Paucijugae | ||||
P. pallidistigma | |||||
P. rugulosa | |||||
P. subtusalbida | — | ||||
P. fjeldsaoi | — | Incanaee | |||
P. incana | Paucijugae | ||||
P. incanoides | — | ||||
P. nana | — | ||||
P. tarapacana | Paucijugae | ||||
P. tomentella |
All species of Polylepis are woody plants growing as trees, multi-stemmed trees or shrubs (Fig.
Growth habit of Polylepis species: tree growth form: A P. microphylla, Chimborazo, Ecuador B P. fjeldsaoi, Lucanas, Peru C P. rugulosa, Moquegua, Perú D P. sacra, Mantanay, Cusco, Peru E P. acomayensis, Paruro, Cusco, Peru F P. pilosissima, Lima, Perú G P. simpsoniae, Cajas, Azuay, Ecuador; shrubby growth form H P. pallidistigma, Azángaro, Puno, Peru I P. tarapacana, Santa Rosa, Puno, Peru J P. microphylla, Chacan, Cusco, Peru. Photographs A E. Bastidas B, C, E E.G. Urquiaga F D, F–I T.E. Boza E.
The bark of Polylepis is one of the characteristic features of the genus. Indeed, the name Polylepis is derived from the Greek words poly (many) and lepis (layers, skins), referring to the shredding, multilayered bark that is common to all species of the genus. The bark can be made up of more than 100 such layers (
The multi-layered, shredding bark characteristic of all species of the genus A P. multijuga Cajamarca, Peru B P. hieronymi cultivated at Zurich Botanical Garden C P. humboldtii Chimborazo, Ecuador D P. pepei La Paz, Bolivia E P. incana Papallacta, Ecuador F P. pauta Ecuador G P. sericea Colombia H P. canoi Junin, Peru. Photographs A, E E.G. Urquiaga F. B, C, F T.E. Boza E. D A. Fuentes G A. Möhl H H.R. Quispe.
The branching of Polylepis is sympodial (
Branching patterns of Polylepis: petioles remaining on the branches A P. rugulosa Moquegua, Peru F P. incana Napo, Ecuador; leaves closely clustered at the top of the branches B P. pallidistigma Puno, Peru E P. rodolfo-vasquezii Huancavelica, Peru; twisted stems and branches C P. humboldtii Chimborazo, Ecuador D P. sacra Cusco, Peru. Photographs A–D T.E. Boza E. E G. Vargas F E.G. Urquiaga F.
Another characteristic feature of the genus is the growth of the two stipules fused around the branch, forming a sheath. The sheaths are congested at the ends of the shoots and shaped like tubes nested inside each other (Fig.
Stipule sheaths congested at the ends of the shoots A P. multijuga Cajamarca, Peru B P. reticulata Azuay, Ecuador C P. ochreata Pichincha, Ecuador D, E P. fjeldsaoi Ayacucho, Peru F P. weberbaueri Ancash, Peru G P. subsericans Cusco, Peru. Photographs A, D–F E.G. Urquiaga F. B, C, G T.E. Boza E.
The types and density of hairs represent some of the most important taxonomic features within the genus, although there are significant fluctuations especially in the length and density of the hair within species (Fig.
Lower leaflet surfaces of Polylepis species with different types of hairs. Lanate: A P. lanata B P. serrata. Pannose: C P. besseri D P. rugulosa. Sericeous: E P. rodolfo-vasquezii F P. sericea. Tomentose: G P. microphylla H P. reticulata. Pilose: I P. flavipila J P. pilosissima. Villous: K P. acomayensis. Puberulous: L P. neglecta. Strigose: M P. subericans. Photographs by T. E. Boza E.
In some species, glandular hairs are intermixed with the longer hairs, often tinting the latter ones yellowish, as in P. flavipila and P. incana. In other species, only glandular hairs are found on some organs. In the extreme case of P. tarapacana, the resin forms a thick, translucent layer on the upperside of the leaflets.
The imparipinnate leaves provide some of the most important taxonomic features within the genus, especially since their features are often correlated with those of the inflorescences and fruits. In addition, they can be determined, based on vegetative material. Important features are:
Leaflet sizes in Polylepis: A P. microphylla; 0.3–0.7 × 0.2–0.5 cm B P. rodolfo-vasquezii; 0.9–1.1 × 0.4–0.6 cm C P. tarapacana; 0.7–0.8 × 0.3–0.4 cm D Polylepis fjeldsaoi; 1.2–2.1 × 0.6–0.7 cm E P. incana; (1.4–)1.8–2.7 × 0.4–0.7 cm F P. subsericans; (1.3–)1.7–2.8 × 0.5–0.7 cm G P. canoi; (2.4–)3.4–3.9 × (0.8–)1.1–1.5 cm H P. humboldtii;1.8–2.8 × 0.6–0.9 cm I P. multijuga; 2.9–3.6(–5.4) × 1.1–2.0 cm. Photographs A, D, E, I E.G. Urquiaga F. B G. Vargas F T.E. Boza E. G H. Huaylla H E. Bastidas.
The inflorescences are simple clusters, rarely branched, generally long and pendulous as in P. multijuga (15.4–36.0 cm; 47–83 flowers), P. ochreata (8.1–17.4 cm; 21–49 flowers) and P. serrata (7.6–17.3 cm; 16–35 flowers). In other species, the inflorescences are more reduced, in the extreme to the axillary region of the leaves, such as in P. microphylla (3.8–5.3 cm; 1–3 flowers), P. pepei (1.2–3.5 cm; 3 flowers) and P. rodolfovasquezii (0.9–1.1 cm; 1 flower) (Fig.
Leaves showing leaflet shapes, margins and apices in the sections and subsections of Polylepis: section Sericeae: A P. multijuga; elliptic, serrate, obtuse (subsect. Lanuginosae) B P. ochreata; elliptic, entire to slightly serrate, emarginate (subsect. Sericeae) C P. pauta, elliptic, crenate, emarginate (subsect. Pauta) D P. rodolfo-vaquezii, elliptic, entire, emarginate (subsect. Pepea). Section Reticulatae: E P. microphylla, broadly elliptic, entire, deeply emarginate F P. reticulata, elliptic to obovate, entire or slightly crenate, deeply emarginate. Section Australes: G P. australis, elliptic, serrate, emarginate. Section Incanaee: H P. besseri, obovate, crenate, obtuse or emarginate and I P. pallidistigma, elliptic, crenate, round or emarginate (subsect. Besseria) J P. tarapacana, obovate, entire or very slightly crenate, obtuse or acute and K P. incana, elliptic to obovate, crenate, obtuse to emarginate (subsect. Incanaee) L P. acomayensis, narrowly obovate, crenate, round to emarginate and M P. sacra, obovate, crenate, emarginate (subsect. Racemosae) N P. flavipila, obovate, crenate, acute or emarginate. Section Subsericantes: O P. subsericans, narrowly elliptic, entire to slightly serrate, round or emarginate. Photographs A, E, K, L E.G. Urquiaga F. B–G, M, O T.E. Boza E. H M. Kessler J A. Domic D G. Vargas.
The flowers of Polylepis are hermaphroditic and have a number of adaptations to wind pollination: the petals are missing, the 3–4 sepals are mostly green or rarely red, nectar or fragrances are missing, the stamens are exerted and the styles are multilobed. Previous taxonomic work has rarely taken into account flower features. However,
The fruits are indehiscent achenes that envelop the single carpel with only one ovule. The surface has differently-shaped protuberances including irregular flattened ridges (as in section Subsericantes), flattened spines (in sections Sericeae and Reticulatae), thick wings and ridges (in sections Incanaee and Subsericantes) (Fig.
Fruit type in Polylepis: A P. reticulata: variable numbers and placement of flattened spines B P. pauta: with variable numbers and placement of flattened spines C P. flavipila: irregular flattened ridges with a series of spines D P. subsericans: irregular flattened ridges with a series of spines E, F P. australis: 2–3 irregular and pronounced thin wings. Photographs by T. E. Boza E.
The evolutionary history of Polylepis is poorly understood. This is linked to the complex taxonomy of the genus, which has led to highly variable classifications over time (see Taxonomic history) and which, in itself, is due to the recent and presumably ongoing radiation of the genus coupled with hybridization and polyploidization. We here review these aspects with respect to their relevance for the taxonomic treatment of the genus.
Polylepis is wind pollinated. Earlier studies suggested low dispersal distances in the range of a few tens of meters (
The fruits of Polylepis are poorly adapted for long-distance dispersal. Some species have long thin spines that appear to be adapted for ectozoochoric dispersal, whereas others have thin wings designed for wind dispersal (
A further potential unknown complication in Polylepis may be the occurrence of apomixis (asexual seed production). Apomixis leads to taxonomic complications because the scarcity of sexual reproduction leads to the formation of numerous clonal lineages that are evolutionarily partly independent (
Hybridization is widespread in the plant kingdom and is also well known in Rosaceae (
For Bolivia, also based on morphological intermediacy in mixed species forests,
Besides these hybridization events between extant taxa, there is also indirect evidence for speciation via hybridization.
Summarizing, based on morphological traits, hybridization in Polylepis appears to be common, ranging from cases where species occur sympatrically, but where no putative hybrids have yet been found (
Direct molecular evidence of hybridization in Polylepis is still lacking. However, both
Changes in ploidy level are an important macro- and microevolutionary processes (
The following account of ploidy levels in Polylepis is based on
Overview of the available data on genome sizes, chromosome numbers and guard cell sizes in species of Polylepis. Where possible, literature records were assigned to this taxonomy, but a few data points (especially from the hybrid zone at Mojanda, Ecuador) had to be excluded because they could not be unambiguously assigned to a species. Depending on data source, we report the mean ± standard deviation, only the mean or a range. Data sources: 1.
Species | Genome size | Chromosome number | Guard cell length | Inferred ploidy levels (2n = /x =) | |||
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Voucher/locality | Size (pg) | Voucher/locality | N | Voucher/locality | Length (µm) | ||
Section Sericeae | |||||||
Subsection Lanuginosae | |||||||
P. lanuginosa | 3 ind. Sangay, Ec 6 | 1.42 ± 0.13 | diploid / x = 6 | ||||
6 ind. Zhud, Ec 2 | 38–42 | ||||||
Laegaard 55036 1 | 10.8 ± 1.4 | ||||||
Laegaard 102637 1 | 11.8 ± 1.6 | ||||||
P. multijuga | Boza 3070 9 | 11.0 ± 2.0 | diploid / x = 6 | ||||
Boza 3074 9 | 11.2 ± 1.7 | ||||||
Boza 3076 9 | 12.8 ± 2.3 | ||||||
Subsection Pauta | |||||||
P. longipilosa | Jaramillo 10862 9 | 10.3 ± 1.6 | diploid / x = 6 | ||||
P. pauta | 2 ind. Oyacachi, Ec 6 | 3.21 ± 0.04 | tetraploid / x = 12 plus aneuploids; perhaps also diploid / x = 6 | ||||
2 ind. Oyacachi, Ec 6 | 3.37 ± 0.18 | ||||||
25 ind. Papallacta, Ec 2, 5 | 67–83 | ||||||
16 ind. Papallacta, Ec 6 | 72 | ||||||
15 ind. Cayambe-Coca, Ec. 3 | 68–77 | ||||||
Kessler 2749 1 | 12.5 ± 1.8 | ||||||
Laegaard 102327 1 | 16.5 ± 2.8 | ||||||
P. pauta | Oyacachi, Ec 8 | 14.4 ± 2.5 | |||||
Papallacta, Ec 8 | 12.3 ± 1.9 | ||||||
Papallacta, Ec 9 | 12.7 ± 1.9 | ||||||
Papallacta, Ec 9 | 12.7 ± 2.1 | ||||||
Papallacta, Ec 9 | 16.6 ± 1.6 | ||||||
P. serrata | Cult. Göttingen 1 | 1.57 ± 0.11 | Cult. Göttingen 1 | 10.6 ± 0.9 | diploid / x = 6 | ||
Cult. Göttingen 1 | 1.61 ± 0.11 | Cult. Göttingen 1 | 12.7 ± 1.8 | ||||
Subsection Sericeae | |||||||
P. albicans | Boza 3014 9 | 10.8 ± 1.3 | diploid / x = 6 | ||||
Frimer 44 9 | 13.5 ± 1.4 | ||||||
Renvoize 5074 9 | 12.2 ± 1.2 | ||||||
P. argentea | Cult. Göttingen 1 | 1.63 ± 0.15 | Cult. Göttingen 1 | 12.4 ± 2.4 | diploid / x = 6 | ||
Cult. Göttingen 1 | 1.67 ± 0.15 | Cult. Göttingen 1 | 13.0 ± 1.7 | ||||
Cult. Zurich 9 | 13.9 ± 1.7 | ||||||
Chevarria 1035 9 | 13.4 ± 1.2 | ||||||
Hanold 85 9 | 13.4 ± 1.2 | ||||||
P. canoi | Kessler 2880 1 | 14.3 ± 2.6 | diploid / x = 6 | ||||
P. frontinensis | Kessler 2772 9 | 11.5 ± 1.3 | diploid / x = 6 | ||||
Kessler 2776 9 | 13.0 ± 1.3 | ||||||
P. humboldtii | Carate 185 9 | 12.4 ± 1.4 | diploid / x = 6 | ||||
P. loxensis | 25 ind. Fierro Urco, Ec 2, 5 | 39–42 | |||||
Laegaard 19109 9 | 12.0 ± 1.5 | ||||||
Lewis 3804 9 | 11.4 ± 1.1 | ||||||
P. ochreata | 2 ind. Yanacocha, Ec 6 | 3.41 ± 0.09 | diploid / x = 6, tetraploid / x = 12, and hexaploid / x = 18; perhaps plus aneuploids or hybrids | ||||
2 ind. El Ángel, Ec 6 | 4.66 ± 0.57 | ||||||
8 ind. El Ángel, Ec 2, 5 | 37–40 | ||||||
9 ind. Yanacocha, Ec 2, 5 | 59–77 | ||||||
16 ind. Yanacocha, Ec 6 | 82 | ||||||
15 ind. Yanacocha, Ec 3 | 73–88 | ||||||
Molau 2536 9 | 10.7 ± 1.3 | ||||||
Laegaard 54474 9 | 11.9 ± 1.1 | ||||||
Romoleroux 1060 9 | 11.7 ± 1.0 | ||||||
Yanacocha, Ec 8 | 14.1 ± 2.3 | ||||||
P. sericea | Dorr 5220 9 | 13.3 ± 1.8 | diploid / x = 6 | ||||
Subsection Pepea | |||||||
P. pepei | Kessler 2795 1 | 10.9 ± 1.6 | diploid / x = 6 | ||||
Kessler 3386 1 | 11.8 ± 1.6 | ||||||
P. rodolfo-vasquezii | Cult. Göttingen 1 | 1.60 ± 0.07 | Cult. Göttingen 1 | 10.2 ± 1.5 | diploid / x = 6 | ||
Cult. Göttingen 1 | 1.70 ± 0.05 | Cult. Göttingen 1 | 10.8 ± 1.3 | ||||
Section Reticulatae | |||||||
P. hieronymi | Cult. Göttingen 1 | 1.52 ± 0.02 | Cult. Göttingen 1 | 12.6 ± 1.8 | diploid / x = 6 | ||
Cult. Göttingen 1 | 1.49 ± 0.04 | Cult. Göttingen 1 | 11.9 ± 2.0 | ||||
52 ind. Ar 7 | 1.45–1.57 | ||||||
Beck 9345 1 | 13.2 ± 1.8 | ||||||
Kessler 3123 1 | 11.2 ± 1.0 | ||||||
P. microphylla | Cult. Göttingen 1 | 1.53 ± 0.06 | Cult. Göttingen 1 | 13.9 ± 1.1 | diploid / x = 6 and tetraploid / x = 12 plus aneuploids | ||
Cult. Göttingen 1 | 1.53 ± 0.07 | Cult. Göttingen 1 | 14.3 ± 2.2 | ||||
2 ind. Ozongoche, Ec 6 | 2.03 ± 0.22 | ||||||
8 ind. Achupallas, Ec 2, 6 | 70–82 | ||||||
Galiano 1999 1 | 14.2 ± 1.8 | ||||||
Achupallas, Ec 8 | 10.7 ± 1.9 | ||||||
P. occidentalis | Diaz 2879 9 | 11.3 ± 1.2 | diploid / x = 6 | ||||
Diaz 4012 9 | 10.8 ± 1.0 | ||||||
Sánchez 10285 9 | 12.7 ± 1.6 | ||||||
P. quadrijuga | Gradstein s.n 1 | 12.2 ± 1.7 | diploid / x = 6 | ||||
Gradstein s.n 1 | 12.3 ± 2.0 | ||||||
Olivares 570 9 | 14.3 ± 1.3 | ||||||
P. reticulata | 11 ind. Soldados, Ec 2 | 36–42 | diploid / x = 6 plus higher ploidy (hexaploid / x = 18?; in cultivated plants only?) | ||||
3 ind. Oyacachi, Ec (pl) 6 | ~118 | ||||||
Kessler 2746a 1 | 12.2 ± 1.4 | ||||||
Laegaard 102691 1 | 10.0 ± 0.9 | ||||||
Cajas, Ec 9 | 10.5 ± 1.8 | ||||||
Cajas, Ec 9 | 12.3 ± 1.6 | ||||||
Cajas, Ec 9 | 11.4 ± 1.4 | ||||||
P. simpsoniae | 3 ind. Sangay, Ec 6 | 1.40 ± 0.08 | diploid / x = 6 | ||||
25 ind. Zhud, Ec 2 | 37–42 | ||||||
2 ind. Sangay, Ec 6 | 38 | ||||||
Laegaard 102677 1 | 12.2 ± 1.0 | ||||||
Cajas, Ec 9 | 9.1 ± 1.1 | ||||||
P. weberbaueri | Acleto 364 1 | 12.3 ± 0.9 | diploid / x = 6 | ||||
Boza 3018 9 | 14.1 ± 1.1 | ||||||
Boza 3148 9 | 14.3 ± 1.5 | ||||||
Smith 9568 9 | 14.9 ± 1.1 | ||||||
Section Australes | |||||||
P. australis | Cult. Göttingen 1 | 2.98 ± 0.06 | Cult. Göttingen 1 | 16.7 ± 2.6 | tetraploid / x = 12 and diploid / x = 6 plus triploid/ x = 9 hybrids and hexaploid / x = 18 autopolyploid derivate | ||
Cult. Göttingen 1 | 3.03 ± 0.03 | Cult. Göttingen 1 | 17.4 ± 1.6 | ||||
261 indiv. 7 | 2.84–2.97 | ||||||
75 indiv. 7 | 1.44–1.54 | ||||||
24 indiv. 7 | 2.09–2.24 | ||||||
1 indiv. 7 | 4.15 | ||||||
Kessler 3350 1 | 18.9 ± 1.9 | ||||||
Lazaro 6695 9 | 17.6 ± 2.4 | ||||||
Lorentz 760 1 | 12.8 ± 1.4 | ||||||
Venturi 3010 9 | 12.2 ± 1.8 | ||||||
w/colector 2330 9 | 15.5 ± 2.5 | ||||||
Cult. Zurich 9 | 22.7 ± 2.7 | ||||||
P. neglecta | Cult. Göttingen 1 | 1.54 ± 0.06 | Cult. Göttingen 1 | ~80 | Cult. Göttingen 1 | 13.9 ± 1.1 | diploid / x = 6; perhaps also tetraploid / x = 12 |
Cult. Göttingen 1 | 1.55 ± 0.09 | Cult. Göttingen 1 | 14.3 ± 2.2 | ||||
Kessler 3531 1 | 13.6 ± 2.3 | ||||||
Kessler 3633 1 | 13.2 ± 2.0 | ||||||
Section Subsericantes | |||||||
P. flavipila | Boza 3163 9 | 17.9 ± 1.8 | tetraploid / x = 12 | ||||
Boza 3167 9 | 16.0 ± 1.5 | ||||||
Boza 3168 9 | 15.3 ± 1.6 | ||||||
P. pilosissima | Kessler 3426 1 | 18.0 ± 2.0 | tetraploid / x = 12 | ||||
Kessler 3591 1 | 17.2 ± 2.2 | ||||||
Boza 3023 9 | 17.3 ± 1.6 | ||||||
Cerrate 1265 9 | 15.6 ± 1.4 | ||||||
Gentry 638 9 | 16.2 ± 1.3 | ||||||
Kessler 3428 9 | 15.7 ± 1.1 | ||||||
P. subsericans | Cult. Göttingen 1 | 3.12 ± 0.18 | Cult. Göttingen 1 | 16.6 ± 2.2 | tetraploid / x = 12 | ||
Cult. Göttingen 1 | 3.21 ± 0.11 | Cult. Göttingen 1 | 17.3 ± 2.2 | ||||
Toivonen s.n 1 | 18.3 ± 1.5 | ||||||
Toivonen s.n 1 | 18.5 ± 2.4 | ||||||
Sylvester 428 9 | 13.9 ± 1.2 | ||||||
Sylvester 868 9 | 16.2 ± 1.7 | ||||||
Sylvester 1287 9 | 15.9 ± 1.8 | ||||||
Section Incanaee | |||||||
Subsection Racemosae | |||||||
P. acomayensis | Boza 3135 9 | 16.2 ± 1.8 | tetraploid / x = 12 | ||||
Boza 3141 9 | 15.1 ± 1.4 | ||||||
P. incarum | Jimenez 2716 1 | 18.3 ± 0.8 | tetraploid / x = 12 | ||||
Kessler 3465 1 | 17.2 ± 1.9 | ||||||
Jimenez 2716 9 | 17.6 ± 1.6 | ||||||
Kessler 13515 9 | 16.9 ± 2.1 | ||||||
Shepard 150 9 | 17.8 ± 2.2 | ||||||
P. lanata | Kessler 2851 1 | 19.6 ± 2.1 | tetraploid / x = 12 | ||||
Kessler 2962 1 | 18.8 ± 1.7 | ||||||
P. pacensis | Kessler 3028 1 | 15.9 ± 2.7 | tetraploid / x = 12 | ||||
Mendez & Arcienaga 14 1 | 17.7 ± 1.3 | ||||||
Kessler 14528 9 | 19.2 ± 1.7 | ||||||
Lopez & Bermejo 4 9 | 19.1 ± 1.9 | ||||||
Lopez & Bermejo 10 9 | 19.7 ± 2.0 | ||||||
P. racemosa (all pl) | 2 ind. Cotopaxi, Ec. 6 | 4.48 ± 0.19 | tetraploid / x = 12 to octoploid / x = 24, with many intermediate and aneuploid ploidy levels | ||||
2 ind. Cotopaxi, Ec. 6 | 2.63 ± 0.20 | ||||||
3 ind. Oyacachi, Ec. 6 | 4.57 ± 0.11 | ||||||
12 ind. Oyacachi, Ec 2, 5 | 80–82 | ||||||
2 ind. Oyacachi, Ec 2, 5 | 72–77 | ||||||
10 ind. Oyacaci, Ec. 6 | 62–80 | ||||||
Ferreyra 12418 1 | 18.0 ± 1.6 | ||||||
Papallacta, Ec 8 | 21.7 ± 3.8 | ||||||
Oyacachi, Ec 8 | 17.6 ± 2.6 | ||||||
Arce 161 9 | 17.2 ± 1.5 | ||||||
Arce 167 9 | 15.1 ± 1.3 | ||||||
Arce 207 9 | 13.8 ± 1.4 | ||||||
Bird 1384 9 | 16.1 ± 1.3 | ||||||
Boza 3020 9 | 16.7 ± 1.6 | ||||||
Boza 3030 9 | 15.1 ± 1.0 | ||||||
Boza 3031 9 | 14.2 ± 1.4 | ||||||
Boza 3119 9 | 18.0 ± 1.5 | ||||||
Ferreyra 3792 9 | 15.3 ± 1.6 | ||||||
Kenehira 5 9 | 15.9 ± 1.1 | ||||||
Kessler 14608 9 | 17.1 ± 1.1 | ||||||
Laegaard 20465 9 | 19.8 ± 2.0 | ||||||
Laegaard 22351 9 | 17.0 ± 1.9 | ||||||
Leiva 741 9 | 14.9 ± 1.2 | ||||||
Leiva 1090 9 | 16.4 ± 1.8 | ||||||
Nuñez 8117 9 | 16.3 ± 1.3 | ||||||
Renvoize 4847 9 | 17.7 ± 1.4 | ||||||
Sánchez Vega 5322 9 | 13.5 ± 1.1 | ||||||
Smith 11076 9 | 12.6 ± 1.0 | ||||||
Soukup 3498 9 | 16.7 ± 1.5 | ||||||
Stork 9972 9 | 16.6 ± 1.8 | ||||||
Tovar 2371 9 | 15.3 ± 1.4 | ||||||
Velásquez 12 9 | 16.3 ± 1.1 | ||||||
West 3787 9 | 13.2 ± 2.2 | ||||||
P. sacra | Cult. Göttingen 1 | 5.76 ± 0.26 | Cult. Göttingen 1 | 20.2 ± 3.3 | octoploid / x = 24; perhaps also tetraploid / x = 12 or intermediates | ||
Cult. Göttingen 1 | 5.72 ± 0.15 | Cult. Göttingen 1 | 16.7 ± 3.3 | ||||
Rosales 04 1 | 19.5 ± 0.8 | ||||||
Sylvester 644 9 | 15.6 ± 1.4 | ||||||
Sylvester 1262 9 | 22.1 ± 1.2 | ||||||
Sylvester 1270 9 | 15.8 ± 1.3 | ||||||
P. triacontandra | Cult. Göttingen 1 | ~80 | tetraploid / x = 12; perhaps also lower ploidy levels | ||||
Beck 4976 1 | 18.9 ± 1.9 | ||||||
Kessler 3420 1 | 20.4 ± 1.1 | ||||||
Steudel 427 9 | 13.5 ± 1.9 | ||||||
Steudel 431 9 | 14.1 ± 2.6 | ||||||
Steudel 433 9 | 18.0 ± 2.1 | ||||||
Subsection Besseria | |||||||
P. besseri | Kessler 2989 1 | 20.4 ± 2.8 | tetraploid / x = 12 or higher ploidy level | ||||
Kessler 2985 1 | 19.2 ± 1.9 | ||||||
P. crista-galli | Beck 9343 1 | 16.6 ± 1.4 | tetraploid / x = 12 | ||||
Kessler 3155 1 | 17.8 ± 2.1 | ||||||
P. pallidistigma | Boza 3005 9 | 17.2 ± 1.9 | tetraploid / x = 12 | ||||
Boza 3006 9 | 17.1 ± 2.2 | ||||||
Boza 3007 9 | 18.5 ± 1.9 | ||||||
Sylvester 1807 9 | 16.4 ± 1.4 | ||||||
Sylvester 1816 9 | 17.3 ± 1.9 | ||||||
Sylvester 1825 9 | 18.5 ± 1.9 | ||||||
P. rugulosa | Ferreyra 2594 1 | 16.8 ± 1.9 | tetraploid / x = 12 | ||||
P. subtusalbida | Beck 7395 9 | 22.1 ± 1.5 | tetraploid / x = 12 and higher ploidy level | ||||
Kessler 216 9 | 23.4 ± 2.4 | ||||||
Ritter 1196 9 | 15.9 ± 1.4 | ||||||
Subsection Incanaee | |||||||
P. fjeldsaoi | Mendoza 1019 9 | 11.9 ± 2.6 | diploid / x = 6 | ||||
Mendoza 1032 9 | 13.3 ± 1.4 | ||||||
Mendoza 1057 9 | 15.2 ± 1.7 | ||||||
P. incana | 3 ind. Sincholagua, Ec 6 | 1.99 ± 0.34 | mainly diploid / x = 6 but also hexaploid / x = 18 (in cultivated plants only?) | ||||
3 ind. Illinizas, Ec 6 | 1.60 ± 0.14 | ||||||
3 ind. Inga-Raya, Ec 6 | 1.67 ± 0.30 | ||||||
3 ind. Cayambe-Coca, Ec (pl) 6 | 1.42 ± 0.10 | ||||||
3 ind. Antisana, Ec (pl) 6 | 4.67 ± 018 | ||||||
16 ind. El Ángel, Ec 2, 5, 6 | (38 –) 42 | ||||||
6 ind. Illinizas, Ec 2, 5, 6 | 38 | ||||||
P. incana | 30 ind. Cayambe-Coca, Ec 2, 5, 6 | (39 –) 42 | |||||
15 ind. Inga-Raya, Ec 6 | 42 | ||||||
15 ind. El Inga, Ec 4 | 40–42 | ||||||
15 ind. Papallacta, Ec 4 | 41–42 | ||||||
15 ind. El Ángel, Ec. 4 | 40–42 | ||||||
Laegaard 102647 1 | 17.6 ± 2.0 | ||||||
Schmidt-Lebuhn 521 1 | 17.0 ± 2.3 | ||||||
Illinizas, Ec 8 | 9.7 ± 0.5 | ||||||
Boza 3066 9 | 15.4 ± 1.8 | ||||||
Boza 3095 9 | 13.1 ± 1.5 | ||||||
Laegaard 102282 9 | 18.6 ± 2.2 | ||||||
P. incanoides | Kessler 3288 1 | 16.4 ± 1.6 | tetraploid / x = 12 | ||||
Kessler 3293 1 | 18.3 ± 2.4 | ||||||
Beck 34512 9 | 15.3 ± 1.9 | ||||||
Kessler 2954 9 | 13.2 ± 0.7 | ||||||
P. nana | Cult. Göttingen 1 | 2.93 ± 0.05 | Cult. Göttingen 1 | 18.7 ± 1.7 | tetraploid / x = 12; also lower ploidy levels ? | ||
Cult. Göttingen 1 | 2.96 ± 0.05 | Cult. Göttingen 1 | 19.6 ± 1.6 | ||||
Kessler 3514 1 | 20.3 ± 2.6 | ||||||
Kessler 3642 1 | 19.5 ± 1.3 | ||||||
Kessler 3501 9 | 15.4 ± 1.8 | ||||||
Kessler 3518 9 | 13.1 ± 1.5 | ||||||
Kessler 3519 9 | 18.6 ± 2.2 | ||||||
P. tarapacana | Cult. Göttingen 1 | 3.02 ± 0.17 | Cult. Göttingen 1 | ~80 | Cult. Göttingen 1 | 17.4 ± 1.8 | tetraploid / x = 12 |
Cult. Göttingen 1 | 3.00 ± 0.16 | Cult. Göttingen 1 | 16.9 ± 2.3 | ||||
Kessler 3599 1 | 17.4 ± 2.2 | ||||||
Kumar 6 1 | 17.1 ± 1.1 | ||||||
Beck 9008 9 | 14.9 ± 2.8 | ||||||
Beck 19897 9 | 16.1 ± 1.1 | ||||||
Beck 32470 9 | 15.0 ± 1.2 | ||||||
Boza 3009 9 | 14.9 ± 2.3 | ||||||
Kessler 3599 9 | 19.7 ± 2.0 | ||||||
P. tomentella | 43 ind. Ar 7 | 2.90–3.01 | Kessler 3188 1 | 17.9 ± 1.8 | tetraploid / x = 12 | ||
Kessler 3368 1 | 18.7 ± 2.0 | ||||||
Boza 3107 9 | 16.6 ± 1.6 | ||||||
Boza 3110 9 | 13.9 ± 2.0 | ||||||
Boza 3111 9 | 15.3 ± 1.4 | ||||||
Kessler 3200 9 | 19.1 ± 2.5 |
Assigning ploidy levels to species of Polylepis can be based on two different base numbers.
Based on this approach, for guard cell length, based on comparison with chromosome counts and genome size measurements,
At present, for the 45 species of Polylepis, combined data on guard cell length, chromosome number and genome size are available for nine (20%) species, on guard cell length and genome size for another nine (20%) species and on guard cell length and chromosome number for three (7%) species, whereas for 24 (53%) species, only guard cell measurements are available (Table
Placing the ploidy levels in an evolutionary context, in section Sericeae, most species are diploid, but mixed di- and polyploidy is present in P. ochreata and P. pauta. These two species overlap in northern Ecuador where they hybridize extensively and it is conceivable that the polyploid condition stems from this hybridization, as polyploidy is often correlated with hybridization (
In section Reticulatae, again most species are diploid, but polyploidy occurs in P. microphylla and P. reticulata. Interestingly, at least in P. reticulata, polyploidy is only known only from cultivated plants (
In section Australes, P. australis has been well studied and includes di-, tri-, tetra- and hexaploids, most likely due to autopolyploidiation (
Sections Subsericantes and Incanaee mainly includes tetraploid species. These sections on average occur at higher elevations and in more arid environments than the other sections, which corresponds well with the polyploid condition, since polyploids are well known to be over-represented at high latitudes and elevations (
Finally, focusing on the taxonomic implications of ploidy levels in the genus, we found that, in some cases, closely related species have different ploidy levels, supporting their treatment as distinct species. For example, P. fjeldsaoi has previously been identified as P. tomentella (
Polylepis belongs to the tribe Sanguisorbeae DC., which is characterized by cup-shaped hypanthium that entirely encloses the carpel(s), resulting in a perigynous position of the flower (
The first attempt to study the evolutionary history of Polylepis using molecular methods was undertaken by Malin Kerr in an unfortunately largely unpublished PhD thesis (
The second attempt at a molecular phylogenetic reconstruction of Polylepis was undertaken by
The latest molecular study of the phylogeny of Polylepis was conducted by M. Claudia Segovia S. in a PhD thesis that also remains largely unpublished (
In conclusion, our understanding of the evolutionary history of Polylepis is still very incomplete. While morphological traits point to a plausible story of diversification and adaptation from humid cloud forests to arid high-elevation habitats, molecular data suggest a complex, reticulate evolutionary history. Additionally, while there is evidence that Polylepis is nested within Acaena, we refrain from merging both genera until a clearer picture of their evolutionary relationships emerges.
Based on all the above, we can conclude that there is ample gene flow between populations of Polylepis assigned to different species. Although species can be distinguished on morphological, biogeographical and ecological grounds, it is likely that gene flow between populations of different species in close proximity have more gene flow between them than geographically remote populations of the same species. At the same time, the presence of different ploidy levels in at least eight species of the genus suggests that there may be barriers to gene flow within species.
In such a situation, the classical biological species concept of species being reproductively and evolutionarily independent units is hardly applicable (
Polylepis may well be the ecologically best-studied Andean tree genus. This is because it reaches the highest elevations of any tree genus in the Andes and because Polylepis forests are among the most threatened ecosystems in the neotropics. In the following, we briefly review some aspects of the physiology, ecology, and biogeography of Polylepis as they are relevant for our monographic work.
Polylepis typically forms the uppermost forest belt in the tropical Andes, although some species also grow at lower elevations in mixed forest stands with other tree genera. Due to the high elevations and accordingly low temperatures at which these trees occur, they have been the focus of a number of ecophysiological studies aiming to understand the adaptations to low temperatures. However, since the 45 species inhabit a wide range of habitats, ranging from moderate to extremely high elevations and arid to superhumid environments, the different species express a wide range of physiological adaptations that go beyond only adaptations to low temperatures.
The ability to tolerate the low nocturnal temperatures that are typical of tropical mountains is, in some species, achieved by daily osmotic adjustments and supercooling down to -9 °C as in P. sericea or freezing tolerance as in P. australis and P. tarapacana (
Adaptations to drought conditions are also frequent in Polylepis and include small, thick leaflets with wax layers and sunken stomata to reduce transpirational water loss (
Photosynthetic rates of Polylepis range from 3 μmol·m-2·s-1 in P. tarapacana to 9 μmol·m-2·s-1 in P. australis (
Generally speaking, physiological traits in Polylepis are related either to the temperature or precipitation conditions at which they grow, revealing evolutionary specialization and adaptation of the species along environmental gradients.
Polylepis is wind-pollinated. Although most pollen is deposited at close distances to the trees (
The seeds of Polylepis are not well adapted for long distance dispersal. Species in section Sericeae have spines that allow them to be transported in the fur of animals, but nothing is known about dispersal distances. In many other species, the nutlets have spiny ridges that do not appear to be adapted to any specific dispersal type (
Polylepis seeds typically have low germination rates, possibly associated with dormancy (
The species of Polylepis are generally rather slow-growing, with radial growth rates typically in the order of 1 mm per year, although much variation exists (
Dendrochronological studies on P. tarapacana in Bolivia suggest tree longevity of at least 700 years, with precipitation being positively and high summer temperatures (which increase drought stress) negatively related to radial growth (
One the most conspicuous patterns in the distribution of Polylepis forests is that, frequently, they occur as isolated forest patches isolated from the closed treeline (
Current understanding of this so-called “Polylepis-problem” suggests that the natural vegetation pattern would be a grassland-forest mosaic, with increasing grassland contribution towards higher elevations (
Other habitat associations of Polylepis include a preference for cloud condensation belts in arid regions (
The upper limit of the growth of Polylepis forests has been a matter of some debate.
It is generally accepted that the genus Polylepis evolved in the Andes from the genus Acaena (
The arrival of humans increased fire frequencies, first by hunter-gatherers and later by agropastoralists with their livestock, leading to widespread reductions in Polylepis abundance (e.g.,
In southernmost Ecuador, at the border with Peru, Polylepis populations are currently restricted to tiny populations, but pollen evidence shows that Polylepis was much more common in the early to mid-holocene (about 11,000–4,000 bp) (
The development of human cultures in the Andes took an important step forward some 6,000 bp with the domestication of camelids (
Polylepis forests harbour unique biodiversity, including a number of highly specialized, often range-restricted and threatened bird species (
Polylepis forests also provide habitats for many plant species, including the world’s highest vascular epiphytes (
Fungal interactions are also important in Polylepis (
Polylepis forests represent one of the most endangered habitats in the high Andes (
Conservation of Polylepis forests is not only relevant for the trees themselves. The forests are rich in endemics species and represent hotspots of biodiversity (
In this context, the evaluation of the conservation status and the degree of threat to the species is necessary in order to successfully focus conservation action. Although the current online version of the IUCN Red List of Threatened Species (http://www.iucnredlist.org/) presents assessments of species of the genus Polylepis, it includes barely 15 species, with 14 species listed as “Vulnerable” (VU) and one as “Near Threatened”. Bearing in mind the novel taxonomic arrangement proposed here, we present a global assessment of the conservation status for the 45 species of Polylepis, applying the IUCN Criteria and Categories (Table
Conservation status of Polylepis species. Abbreviations: Ar = Argentina, Bo = Bolivia, Ch = Chile, Co = Colombia, Ec = Ecuador, Pe = Perú, Ve = Venezuela.
Species | Status | Criteria | Country | Conservation Areas |
---|---|---|---|---|
Section Sericeae | ||||
Subsection Lanuginosae | ||||
P. lanuginosa | EN | B1a+B2a, C1 | Ec | Cajas National Park |
P. multijuga | CR | A1, B1a+B2a, C1 | Pe | None |
Subsection Pauta | ||||
P. pauta | VU | A1, B1a+B2a, C1 | Ec | Cayambe-Coca National Park |
Antisana Ecological Reserve | ||||
P. longipilosa | CR | A2a, B1a+B2a, C1+C2 | Ec | El Angel Ecological Reserve |
P. serrata | VU | B1a+B2a, C1 | Pe | Río Abiseo National Park |
Manu National Park | ||||
Subsection Sericeae | ||||
P. albicans | VU | B1a+B2a | Pe | Huascarán National Park |
P. argentea | VU | B1a+B2a | Pe | Otishi National Park |
P. canoi | EN | B1a+B2a, C1 | Bo, Pe | Otishi National Park |
P. frontinensis | CR | B2ac, C2a | Co | Colibrí del Sol Private Reserve |
P. humboldtii | CR | B2a, C2 | Ec | Sangay National Park |
P. longipilosa | CR | A2a, B1a+B2a, C1+C2 | Ec | El Angel Ecological Reserve |
P. loxensis | CR | A2a, B1a+B2a, C2a | Ec | None |
P. ochreata | VU | B1a+B2a, C1 | Co, Ec | El Angel Ecological Reserve (Ec) |
Yanacocha Reserve (Ec) | ||||
P. sericea | VU | B1a+B2a | Co, Ve | Sierra Nevada National Park (Ve) |
Sierra de la Culata National Park (Ve) | ||||
Subsection Pepea | ||||
P. pepei | EN | A2a, B1a+B2a, C1, D1 | Bo, Pe | Madidi National Park (Bo) |
Carrasco National Park (Bo) | ||||
P. rodolfo-vasquezii | VU | B1a+B2a, C1 | Pe | Pui-Pui Protection Forest |
Section Reticulatae | ||||
P. hieronymi | VU | B1a+B2ac | Ar, Bo | Cordillera de Sama Biological Reserve (Bo) |
P. microphylla | EN | B1a+B2ab | Ec, Pe | Sangay National Park (Ec) |
Cordillera Huayhuash Reserved Zone (Pe) | ||||
P. occidentalis | EN | A1, B1a+B2a, C1 | Pe | None |
P. quadrijuga | VU | A2a, B1a+B2a, D2a | Co | Chingaza National Natural Park |
Cocuy National Park | ||||
Sumapaz National Natural Park | ||||
P. reticulata | VU | B1a+B2a, C1 | Ec | Cajas National Park |
Llanganates National Park | ||||
Pasochoa Ecological Reserve | ||||
Yacuri National Park | ||||
P. simpsoniae | EN | A1, B1a+B2a, C2a | Ec | Cajas National Park |
P. weberbaueri | VU | B1a+B2a | Pe | Huascarán National Park |
Section Australes | ||||
P. australis | LC | B1a+B2a | Ar | Quebrada del Condorito National Park |
P. neglecta | VU | A1,2a, B1a+B2a, C2a | Bo | None |
Section Subsericantes | ||||
P. flavipila | EN | B1a+B2a, C2a | Pe | Nor Yauyos-Cocha Landscape Reserve |
P. pilosissima | CR | A2a, B2a | Pe | Japani Private Conservation Area |
P. subsericans | VU | B1a+B2a | Pe | Vilcanota Private Conservation Areas Network |
Section Incanaee | ||||
Subsection Racemosae | ||||
P. acomayensis | EN | A2a, B1a+B2a, C2a | Pe | None |
P. incarum | CR | A1, B1a+B2a, C1+C2a | Bo, Pe | None |
P. lanata | VU | B1a+B2a, D2a | Bo | Carrasco National Park |
P. pacensis | EN | A2b, B1a+B2a, C1 | Bo | None |
P. racemosa | LC | B1a+B2a | Ec, Pe | None |
P. sacra | VU | B1a+B2a | Pe | Vilcanota Private Conservation Areas Network |
P. triacontandra | EN | A1, B1a+B2a, C1 | Bo, Pe | Apolobamba Integrated Management Natural Area (Bo) |
Subsection Besseria | ||||
P. besseri | VU | A1, B1a+B2a, C1 | Bo | None |
P. crista-galli | VU | A1, B1a+B2a, C2a | Ar, Bo | None |
P. pallidistigma | VU | B1a+B2a | Pe | None |
P. rugulosa | VU | B1a+B2a, C1 | Ch, Pe | Lauca National Park (Ch) |
Islunga National Park (Ch) | ||||
Las Vicuñas National Reserve (Ch) | ||||
Salinas y Aguada Blanca National Reserve (Pe) | ||||
P. subtusalbida | VU | A1, B1a+B2a | Bo | Tunari National Park |
Subsection Incanaee | ||||
P. fjeldsaoi | VU | B1a+B2a, C2a | Pe | None |
P. incana | LC | A1, B1a+B2a | Co, Ec, Pe | Cajas National Park (Ec) |
Illinizas Ecological Reserve (Ec) | ||||
El Angel Ecological Reserve (Ec) | ||||
Huascarán National Park (Pe) | ||||
Cordillera Huayhuash Reserve Zone (Pe) | ||||
P. incanoides | EN | A1+A2a, B1a+B2a, D1 | Bo | None |
P. nana | CR | A1+A2a, B1a, C1+C2a, D2a | Bo | None |
P. tarapacana | NT | A1+A2a, B1a+B2a, C1 | Ar, Bo, Ch, Pe | Sajama National Park (Bo) |
P. tomentella | LC | A1, B1a+B2a, C2a | Ar, Bo | None |
Most of the species of Polylepis are present in at least one protected area, but 16 species have no such conservation actions taken to date. However, actual protection of the species in conservation areas leaves much to be desired. In many cases, no specific conservation actions are taken, in others, extractive activities continue within protected areas. Even where reforestation schemes are undertaken, these are often counterproductive, since alien species of Polylepis are used which can hybridize with the native species. For instance, P. racemosa is not native to Ecuador, but has been widely planted and is already hybridizing with the native species.
More generally, there is mixed success of protected areas in conservations terms which may show the limitations of strictly conservationist approaches that fail to take into consideration the needs of local human population (
Polylepis racemosa Ruiz & Pavón.
Trees or shrubs 1–32 m tall; bark shredding, multilayered with thin exfoliating layers; branches twisted showing a striking arrangement of the leaves, young shoots with leaves usually all closely clustered at the top, causing a shrub-like growth, while the basal internodes stretch rather significantly afterwards. Leaves alternate, imparipinnate with 1–7 pairs of lateral leaflets, obtrullate in outline, 1.3–19.5 cm long, 0.6–10.7 cm wide; rachises lanate, pannose, sericeous, tomentose, villous or glabrous; point of leaflet attachment with a tuft of long hairs; stipular sheaths apically acute, truncate or spurred, glabrous to densely covered with lanate, tomentose or villous hairs on the other surface; leaflets elliptic, ovate or obovate in outline, 0.3–5.4 cm long, 0.2–2.0 cm wide; margins entire, revolute, crenate to serrate, apically acute to deeply emarginate, attenuate, cuneate or unequally cordate, upper leaflet surfaces glabrous or sparsely to densely lanate, pilose, sericeous or tomentose; lower leaflet surfaces covered with very short pannose hairs, pannose mixed with another type of hairs or sparsely to densely lanate, pilose, sericeous, strigose, tomentose or villous. Inflorescences axillar, simple rarely branched, upright or pendant to 36.0 cm long, bearing 1–83 flowers; floral bracts 2.1–15.8 mm long, narrowly triangular. Flowers hermaphroditic; sepals 3–4; stamens 5–27, anthers orbicular with a dense tuft of straight white hairs in the upper half; styles fimbriate, 0.9–4.9 mm long, ovary inferior; carpel 1, ovule 1. Fruit indehiscent achene, fusiform turbinate with protuberances, flattened-spines, irregular flattened ridges, thick wings and ridges or thin wings, glabrous to densely sericeous, tomentose or villous, 1.7–15.1 mm long, 1.3–10.1 mm wide including spines.
1 | Lateral leaflets 1 pair | 2 |
– | Lateral leaflets 2–7 pairs | 21 |
2 | Lower leaflet surfaces densely pannose, hairs < 0.2 mm long | 3 |
– | Lower leaflet surfaces sparsely to densely pilose, sericeous, strigose, tomentose or villous, hairs 0.4–2.0 mm long | 10 |
3 | Leaflet margins crenate | 4 |
– | Leaflet margins entire or serrate | 6 |
4 | Stipular sheaths apically acute, outer sheath surfaces densely villous | P. incana |
– | Stipular sheaths apically truncate, outer sheath surfaces glabrous to densely villous | 5 |
5 | Leaflets obovate; upper leaflet surfaces and rachises tomentose; central Peru | P. fjeldsaoi |
– | Leaflets elliptic; upper leaflet surfaces and rachises villous; southern Peru | P. pallidistigma |
6 | Leaflets 0.7–1.2 cm long; inflorescences 0.7–1.5 cm long | 7 |
– | Leaflets 1.3–3.2 cm long; inflorescences 2.2–8.0 cm long | 8 |
7 | Leaflets 1.0–1.2 cm long; upper leaflet surfaces often with dark sheen, glabrous to sparsely villous; central Bolivia | P. nana |
– | Leaflets 0.7–0.8 cm long; upper leaflet surfaces rugose, glabrous, usually covered with a layer of yellowish resinous exudate; south-western Peru, north-western Chile, western Bolivia and north-western Argentina | P. tarapacana |
8 | Stipular sheaths with outer surfaces densely tomentose; 13–15 stamens per flower; fruits with 2–4 wide flattened hard irregular ridges, sparsely tomentose | P. crista-galli |
– | Stipular sheaths with outer surfaces glabrous to densely villous; 15–23 stamens per flower; fruits with 3–4 ridges with a variable number and placement of flattened spines, densely villous | 9 |
9 | Leaflets 0.5–0.7 cm wide, with 7–15 teeth per side; leaflet apices obtuse to slightly acute; upper leaflet surfaces glabrous; 5–7 flowers per inflorescence; 15–19 stamens per flower; Dpto. Cochabamba (Bolivia) | P. incanoides |
– | Leaflets 0.3–0.6 cm wide, with 5–10 teeth per side, leaflet apices round to emarginate; upper leaflet surfaces glabrous to sparsely villous; 4–5 flowers per inflorescence, 19–23 stamens per flower; Dptos. Potosi, Oruro, Chuquisaca, Tarija (Bolivia) and Jujuy (Argentina) | P. tomentella |
10 | Leaflet margins serrate | 11 |
– | Leaflet margins entire or crenate | 13 |
11 | Lower leaflet surfaces sparsely to densely tomentose without underlying short hairs; 7–21 flowers per inflorescence | P. racemosa |
– | Lower leaflet surfaces densely tomentose with a dense underlying layer of very short white hairs; 3–7 flowers per inflorescence | 12 |
12 | Leaflets 0.7–1.3 cm wide; leaflet apices acute; inflorescences 5.1–7.5(–8.9) cm long; Titicaca basin (Puno, Peru; La Paz, Bolivia) | P. incarum |
– | Leaflets 0.4–0.6 cm wide; leaflet apices obtuse to emarginate; inflorescences 1.8–3.7 cm long; Depts. Cochabamba and north-western Potosi (Bolivia) | P. subtusalbida |
13 | Leaflets 0.9–2.2 cm long; leaflet apices deeply emarginate without projection; Ecuador | P. reticulata |
– | Leaflets 0.9–3.3 cm long; leaflet apices round, obtuse, acute or emarginate with a mid-vein projection; Ecuador, Peru, Bolivia, Chile | 14 |
14 | Leaflet margins entire | 15 |
– | Leaflet margins crenate | 16 |
15 | Leaflets 0.9–1.1 × 0.4–0.6 cm; upper leaflet surfaces glabrous to sparsely sericeous; lower leaflet surfaces sparsely to densely sericeous; inflorescences 0.9–1.1 cm long, with 1 flower | P. rodolfovasquezii |
– | Leaflets (1.3–)1.7–2.8 × 0.5–0.7 cm; upper leaflet surfaces sparsely strigose; lower leaflet surfaces densely strigose; inflorescences (1.9–)2.5–4.9(–5.6) cm long, with 3–6 flowers | P. subsericans |
16 | Lower leaflet surfaces densely tomentose with a dense underlying layer of very short, white pannose hairs | 17 |
– | Lower leaflet surfaces densely pilose, villous or tomentose without pannose hairs | 18 |
17 | Upper leaflet surfaces smooth to slightly rugose; lower leaflet surface hairs 0.6–0.8 mm long; 7–9 flowers per inflorescence; central Bolivia | P. besseri |
– | Upper leaflet surfaces strongly rugose; lower leaflet surface hairs 0.8–1.0 mm long; 4–5 flowers per inflorescence; south-western Peru and north-western Chile | P. rugulosa |
18 | Upper leaflet surfaces sparsely to densely pilose; 3–5 flowers per inflorescence; 11–17 stamens per flower | 19 |
– | Upper leaflet surfaces glabrous to sparsely villous; 5–13 flowers per inflorescence; 19–23 stamens per flower | 20 |
19 | Lower leaflet surfaces densely pilose, hairs 0.5–0.6 mm long | P. flavipila |
– | Lower leaflet surfaces densely villous, hairs 1.0–1.2 mm long | P. pilosissima |
20 | Leaflet apices round to emarginate; upper leaflet surfaces sparsely villous; lower leaflet surfaces densely villous, hairs 0.9–11 mm long; inflorescences 2.0–4.0 cm long, with 5–7 flowers; south-central Peru | P. acomayensis |
– | Leaflet apices acute; upper leaflet surfaces glabrous; lower leaflet surfaces densely tomentose, hairs 0.4–0.8 mm long; inflorescences (4.9–)5.5–7.0(–9.5) cm long, with 11–13 flowers; southern Peru and northern Bolivia | P. triacontandra |
21 | Lateral leaflets 4–7 pairs | 22 |
– | Lateral leaflets 2–3 pairs | 38 |
22 | Lower leaflet surfaces glabrous to puberulous; fruits with 2–3(–4) irregular and pronounced, thin wings | P. neglecta |
– | Lower leaflet surfaces sparsely or densely lanate, sericeous, villous or tomentose, hairs 0.3–2.3 mm long; fruits with variable numbers of flattened or long, hard spines | 23 |
23 | Leaflets 0.3–0.7 cm long; leaflet apices deeply emarginate; inflorescences 3.8–5.3 cm long, with 1–3 flowers | P. microphylla |
– | Leaflets 1.1–5.4 cm long; leaflet apices slightly emarginate to emarginate; inflorescences 2.4–36.0 cm long, with 4–83 flowers | 24 |
24 | Lower leaflet surfaces sparsely or densely sericeous | 25 |
– | Lower leaflet surfaces densely lanate, villous or tomentose | 31 |
25 | Leaflet margin entire or serrate | 26 |
– | Leaflet margin slightly crenate to crenate | 28 |
26 | Leaflets (0.8–)1.1–1.5 cm wide; lower leaflet surface hairs 1.3–1.7 mm long | P. canoi |
– | Leaflets 0.4–0.9 cm wide; lower leaflet surface hairs 0.2–0.9 mm long | 27 |
27 | Leaflets broadly obovate; upper leaflet surfaces glabrous with few hairs on the mid-vein | P. loxensis |
– | Leaflets narrowly elliptic to elliptic; upper leaflet surfaces glabrous to sparsely sericeous | 28 |
28 | Lateral leaflet 3–4 pairs | 29 |
– | Lateral leaflet 4–7 pairs | 30 |
29 | Leaflet margins slightly crenate at the apex; inflorescences 3.9–6.6(–7.8) cm long, with 18–21 flowers; northern Peru | P. albicans |
– | Leaflet margins entire; inflorescences 13.0–17.9(–20.4) cm long, with 23–29 flowers; central Ecuador | P. humboldtii |
30 | Leaflets 1.6–3.0 cm long; leaflet margins entire to slightly serrate; lower leaflet surfaces of mature plants with hairs 0.3–0.5 mm long; north-western Ecuador and southern Colombia | P. ochreata |
– | Leaflets (1.1–)1.4–1.6 cm long, leaflet margins crenate; lower leaflet surfaces of mature plants with hairs 0.3–0.4 mm long; north-eastern Ecuador | P. pauta |
31 | Lower leaflet surfaces densely lanate or villous, the hairs 0.7–1.8 mm long | 32 |
– | Lower leaflet surfaces densely tomentose, the hairs 0.3–1.5 mm long | 35 |
32 | Leaflet margin entire to slightly crenate; northern Ecuador | P. longipilosa |
– | Leaflet margins serrate; northern Colombia, Peru | 33 |
33 | Leaflets 0.4–0.8 cm wide, obovate; leaflet apices slightly emarginate; 7–15 flowers per inflorescence; north-western Colombia | P. frontinensis |
– | Leaflets 0.8–2.0 cm wide, elliptic; leaflet apices acute or obtuse; 16–83 flowers per inflorescence; Peru | 34 |
34 | Leaflets 1.1–2.0 cm wide; leaflet apices obtuse; lower leaflet surfaces densely villous; inflorescences (15.4–)21.7–28.2(–36.0) cm long, with 47–83 flowers | P. multijuga |
– | Leaflets 0.8–1.0(–1.2) cm wide; leaflet apices acute; lower leaflet surfaces densely lanate; inflorescences (7.6–)9.5–13.3(–17.3) cm long, with 16–35 flowers | P. serrata |
35 | Leaflet apices moderately emarginate; upper leaflet surfaces sparsely tomentose; Bolivia and Argentina | P. hieronymi |
– | Leaflet apices deeply emarginate; upper leaflet surfaces glabrous; Colombia, Ecuador and Peru | 36 |
36 | Lateral leaflets (1–)2–3(–4) pairs; lower leaflet surface hairs 0.6–1.5 mm long; Ecuador | P. reticulata |
– | Lateral leaflets 3–5 pairs; lower leaflet surface hairs 0.3–0.9 mm long; Colombia and Peru | 37 |
37 | Leaflets 0.5–0.8 cm wide; lower leaflet surface hairs 0.3–0.6 mm long; inflorescences 2.4–6.7 cm long, with 4–12 flowers; northern Peru | P. occidentalis |
– | Leaflets 0.7–1.1 cm wide; lower leaflet surface hairs 0.7–0.9 mm long; inflorescences (6.0–)7.3–10.5(–12.3) cm long, with 11–19 flowers; north-eastern Colombia | P. quadrijuga |
38 | Lower leaflet surfaces glabrous to sparsely hispid, puberulous, or pannose, the hairs < 0.2 mm long | 39 |
– | Lower leaflet surfaces sparsely to densely lanate, sericeous, tomentose or villous, the hairs 0.3–2.5 mm long | 42 |
39 | Leaflets 0.5–0.8 cm wide; leaflet margins crenate, with 5–8 teeth per side; lower leaflet surfaces glabrous to sparsely villous; southern Peru | P. pallidistigma |
– | Leaflets 0.4–1.5 cm wide; leaflet margins serrate with 9–18 teeth per side; lower leaflet surfaces glabrous; Argentina and Bolivia | 40 |
40 | Leaflet margins with 12–18 teeth per side; leaflet apices acute; 14–27 flowers per inflorescence; Argentina and Bolivia | P. neglecta |
– | Leaflet margins with 9–13 teeth per side; leaflet apices obtuse to emarginate; 5–12 flowers per inflorescence | 41 |
41 | Leaflets elliptic, lower leaflet surfaces glabrous to sparsely hispid; fruits with 2–3 irregular and pronounced thin wings, glabrous; Argentina | P. australis |
– | Leaflets obovate, lower leaflet surfaces pannose; fruits with 2–4 wide flattened hard irregular ridges, sparsely tomentose; Argentina and Bolivia | P. crista-galli |
42 | Lower leaflet surfaces densely lanate, the hairs 1.3–2.5 mm long | 43 |
– | Lower leaflet surfaces sparsely to densely sericeous, tomentose or villous, the hairs 0.3–2.0 mm long | 45 |
43 | Leaflets elliptic; upper leaflet surfaces glabrous to sparsely lanate; lower leaflet surface hairs 1.5–2.5 mm long, yellowish; 13–16 flowers per inflorescence; Ecuador | P. lanuginosa |
– | Leaflets obovate to broadly obovate; upper leaflet surfaces sparsely lanate; lower leaflet surface hairs 1.3–1.5 mm long, whitish; 5–11 flowers per inflorescence; Peru and Bolivia | 44 |
44 | Leaflets (1.8–)2.2–2.7 × 0.9–1.4 cm; inflorescences (5.0–)6.1–4.9(–12.3) cm long; Bolivia | P. lanata |
– | Leaflets 1.6–2.6 × 0.6–1.1 cm; inflorescences 5.0–8.8 cm long; Peru | P. sacra |
45 | Lower leaflet surfaces densely sericeous, the hairs 0.2–1.7 mm long | 46 |
– | Lower leaflet surfaces sparsely to densely tomentose or villous, the hairs 0.4–2.0 mm long | 52 |
46 | Leaflets 0.8–1.3 × 0.2–0.7 cm; inflorescences 1.2–1.6(–3.5) cm long, with 3 flowers; southern Peru and Bolivia | P. pepei |
– | Leaflets 1.2–3.9 × 0.4–1.5 cm; inflorescences 3.3–20.4 cm long, with 5–29 flowers; Venezuela to Bolivia | 47 |
47 | Leaflets (2.4–)3.4–3.9 × (0.8–)1.1–1.5 cm; lower leaflet surface hairs 1.3–1.7 mm long, yellowish; central Peru to Bolivia | P. canoi |
– | Leaflets 1.2–2.8 × 0.4–1.0 cm; lower leaflet surface hairs 0.2–1.0 mm long, silky, whitish; Venezuela to central Peru | 48 |
48 | Lateral leaflets 3–5 pairs; lower leaflet surface hairs 0.2–0.6 mm long; Ecuador and north-western Peru | 49 |
– | Lateral leaflets 2–3 pairs; lower leaflet surface hairs 0.6–1.0 mm long; Venezuela, Colombia, central Peru | 51 |
49 | Leaflet margins entire; inflorescences 13.0–17.9(–20.4) cm long, with 23–29 flowers | P. humboldtii |
– | Leaflet margins slightly crenate at the apex or serrate; inflorescences 3.5–12.2 cm long, with 9–27 flowers | 50 |
50 | Leaflets elliptic; leaflet margins slightly crenate at the apex; north-western Peru | P. albicans |
– | Leaflets broadly obovate; leaflet margins serrate; southern Ecuador | P. loxensis |
51 | Leaflets (1.9–)2.4–2.6 × 0.5–0.7 cm; upper leaflet surface sparsely to dense sericeous; inflorescences 7.2–8.1 cm long; central Peru | P. argentea |
– | Leaflets 1.8–2.1 × 0.8–1.0 cm; upper leaflet surfaces glabrous; inflorescences 3.3–4.5 cm long; Venezuela and Colombia | P. sericea |
52 | Leaflet margins serrate | 53 |
– | Leaflet margins entire to crenate | 55 |
53 | Lateral leaflets 3–4(5) pairs; lower leaflet surface densely villous; Colombia | P. frontinensis |
– | Lateral leaflets 1–2 pairs; lower leaflet surfaces sparsely to densely tomentose; Ecuador, Peru and Bolivia | 54 |
54 | Leaflets (2.3–)3.1–3.9 × (0.7–)0.9–1.5 cm, apices round; inflorescences (4.2–)5.0–9.4(–11.7) cm long, with 7–21 flowers; Ecuador and Peru | P. racemosa |
– | Leaflets 0.9–1.6 × 0.4–0.6 cm, apices obtuse to emarginate; inflorescences 1.8–3.7 cm long, with 3–4 flowers; Bolivia | P. subtusalbida |
55 | Leaflet apices acute, round, obtuse or emarginate; lower leaflet surfaces densely tomentose or villous with a dense underlying layer of very short pannose hairs; Peru and Bolivia | 56 |
– | Leaflet apices moderately to deeply emarginate; lower leaflet surfaces densely tomentose without short pannose hairs; Ecuador to Argentina | 59 |
56 | Lower leaflet surface hairs 0.9–1.1 mm long; inflorescences 2.0–4.0 cm long; central Peru | P. acomayensis |
– | Lower leaflet surface hairs 0.4–0.9 mm long; inflorescences 3.6–10.0 cm long; southern Peru to Bolivia | 57 |
57 | Leaflets (2.0–)2.6–3.3 × 0.7–1.1 cm, narrowly elliptic; 11–13 flowers per inflorescence; Dptos. Puno (Peru) and La Paz (Bolivia) | P. triacontandra |
– | Leaflets 1.4–2.4 × 0.6–11 cm, narrowly obovate; 5–11 flowers per inflorescence; La Paz and Cochabamba (Bolivia) | 58 |
58 | Lateral leaflets 1–2(–3) pairs; stipular sheaths apically truncate; rachises and lower leaflet surfaces densely villous; Dpto. Cochabamba (Bolivia) | P. besseri |
– | Lateral leaflet 2–3 pairs; stipular sheaths apically acute; rachises and lower leaflet surfaces densely tomentose; Dpto. La Paz (Bolivia) | P. pacensis |
59 | Leaflets (0.9–)1.3–1.6 × (0.4–)0.7–1.1 cm, broadly ovate; Ecuador | P. simpsoniae |
– | Leaflets 0.9–2.2 × 0.4–1.1 cm, elliptic to obovate; Colombia to Argentina | 60 |
60 | Leaflets narrowly obovate; upper leaflet surfaces sparsely tomentose; Bolivia and Argentina | P. hieronymi |
– | Leaflets narrowly elliptic to obovate; upper leaflet surfaces glabrous; Colombia to Peru | 61 |
61 | Lower leaflet surface hairs 0.3–0.6 mm long; Peru | 62 |
– | Lower leaflet surface hairs 0.6–1.5 mm long; Colombia and Ecuador | 63 |
62 | Lateral leaflets 3–5 pairs; leaflets 1.1–1.9 cm wide; inflorescences 2.4–6.7 cm long | P. occidentalis |
– | Lateral leaflets 2–3 pairs; leaflets 1.6–2.1 cm wide; inflorescences 8.6–9.7 cm long | P. weberbaueri |
63 | Lower leaflet surface hairs 0.7–0.9 mm long; inflorescences (6.0–)7.3–10.5(–12.3) cm long, with 11–19 flowers; north-eastern Colombia | P. quadrijuga |
– | Lower leaflet surface hairs 0.6–1.5 cm long; inflorescences (2.3–)4.3–13.8 cm long, with 5–9 flowers; Ecuador | P. reticulata |
Trees or shrubs; lower leaflet surfaces with sericeous, lanate or villous hairs; fruits with a variable number and placement of flattened, thin or short spines, densely sericeous or villous.
Polylepis sericea Wedd.
The sectional epithet Sericeae is a plural adjective agreeing in gender with Polylepis. This section, already defined as a group by
The majority of species in this section are morphologically clearly distinct. Probably the two most similar species are P. pepei and P. rodolfovasquezii, which only differ in a few, partly overlapping characters. They might be treated at subspecies level, but as detailed in the Introduction, we decided not to accept infraspecific taxa because of the difficulty of deciding at which level to discriminate between species- and subspecies-level differentiation. Table
Alignment of the species of the Polylepis sect. Sericeae according to
|
|
|
This study |
---|---|---|---|
P. albicans | P. sericea | P. sericea | P. albicans |
P. ochreata | P. ochreata | ||
P. hypargyrea | P. sericea | ||
P. sericea | |||
P. argentea | |||
P. frontinensis | |||
P. humboldtii | |||
P. loxensis | |||
— | P. canoi | P. canoi | |
P. coriacea | P. lanuginosa | P. lanuginosa | P. lanuginosa |
P. lanuginosa | |||
P. lehmannii | |||
P. multijuga | P. multijuga | P. multijuga | P. multijuga |
P. annulatipilosa | P. pauta | P. pauta | P. pauta |
P. pauta | |||
P. stuebelii | |||
— | P. longipilosa | ||
P. serrata | P. serrata | ||
— | P. pepei | P. pepei | P. pepei |
— | — | P. rodolfo-vasquezii | P. rodolfo-vasquezii |
Within section Sericeae, we recognize four subsections, based on their morphological distinctness, as follows: subsection Lanuginosae (two species) with lanate or villous lower leaflet surfaces and densely villous fruits; subsection Pauta (three species) with 4–6 lateral leaflet pairs and lanate or sericeous lower leaflet surfaces; subsection Sericeae (eight species) with sericeous lower leaflet surfaces (except P. frontinensis) and fruits with flattened spines; and subsection Pepea (two species) with 1–2 lateral leaflet pairs, emarginate leaflet apices and densely sericeous, slightly twisted fruits with short spines.
Many species of this section differ markedly in their climatic niches (Figs
Box plots showing the climatic niches of the species of the subsections Lanuginosae, Pauta and Pepea in relation to Mean Annual Temperature (MAT) (A) and Mean Annual Precipitation (MAP) (B). The ends of each box represent the upper and lower quartiles and the median is indicated with a bold line inside the box; the whisker lines extend to the highest and lowest observations, except when observations are higher or lower than the interquartile range (i.e. outliers), in which case they are indicated by a dot. Box plots that share the same lowercase letters within each subsection are not significantly different at p = 0.05. Vertical lines represent subsectional divisions.
Focussing on the individual subsections, the two species in subsect. Lanuginosae show minor ecological differences and replace each other geographically. In subsect. Pauta, P. longipilosa and P. pauta from northern Ecuador have quite similar climatic niches and replace each other geographically, whereas P. serrata from Peru grows under substantially higher temperatures and higher precipitation. The two very similar species of subsect. Pepea have identical niches and complementary geographical distributions. These species clearly form a vicariant species pair, suggesting allopatric speciation after geographical isolation. Finally, in subsection Sericeae, there are major differences among almost all species, with only P. albicans and P. humboldtii having similar climatic niches, but these are geographically well separated. Indeed, all species of this subsection are geographical vicariants, except for P. argentea and P. canoi, which broadly overlap geographically, but have quite different niches, with P. argentea growing under colder and drier and P. canoi under warmer and more humid conditions. These marked ecological differences between species show that they are evolutionarily and ecologically independent lineages and support their treatment as separate species.
Trees; 3–6(–7) lateral leaflet pairs; lower leaflet surfaces lanate or villous; fruits with flattened spines, densely villous.
Polylepis lanuginosa Kunth.
The subsectional epithet Lanuginosae is a plural adjective agreeing in gender with Polylepis.
Polylepis lehmannii Hieron. Bot. Jahrb. Syst.20: Beibl. 49: 29. 1895. Type. Ecuador. Azuay: west of Cuenca, Lehmann 6487 (holotype: B destroyed; isotype: F!).
Polylepis coriacea Bitter Bot. Jahrb. Syst. 45: 603. 1911. Type. Ecuador. Chimborazo: Valley of Pangor Spruce s.n (holotype: W, photos at F!, MO!, US!).
Ecuador. Chimborazo: near Calpi “ad radicem montis Chimborazo”, June 1903, Humboldt & Bondpland 2191 (holotype: P!; isotypes: P!, photo at F!).
Trees 3–8 m tall. Leaves slightly congested at the branch tips, imparipinnate with 2–3 pairs of lateral leaflets, obtrullate in outline, (4.5–)5.2–7.7 × 3.4–4.5 cm; rachises densely villous, points of leaflet attachment with a tuft of long, lanate hairs; stipular sheaths apically acute with spurs, densely lanate on the outer surfaces; leaflets elliptic in outline, second pair from the terminal leaflet the largest, one of this pair 1.7–2.8 × 0.7–1.4 cm; margin crenate with 8–9 teeth, apically emarginate, basally unequally cordate; upper leaflet surfaces glabrous or sparsely lanate; lower leaflet surfaces densely lanate with yellowish hairs 1.5–2.5 mm long. Inflorescences branched at the base or simple, pendant, (4.3–)6.2–9.5(–13.0) cm long, bearing 13–16 flowers; floral bracts 3.8–5.5 mm long, narrowly triangular, densely lanate on the outer surface; rachises villous. Flowers 5.8–7.5 mm diam.; sepals 4, ovate, green, densely sericeous outside; stamens 13–15, anthers orbicular, with a dense tuft of straight white hairs on the upper half; styles fimbriate, 1.9–2.3 mm long. Fruits turbinate, with variable numbers and placement of flattened spines, densely villous; 4.4–6.0 × 6.1–7.4(–9.8) mm including spines. Diploid.
Polylepis lanuginosa is endemic to central and southern Ecuador (Azuay, Bolívar, Cañar and Chimborazo) (Fig.
Based on 17 collecting localities, the estimated EOO is 6,910 km2 and the occupied habitat or AOO is 96 km2. It is protected within Cajas National Park. The species was categorized as VU B1+2c by
Polylepis lanuginosa is most similar to P. multijuga, with which it shares the leaflet shape and hair density. However, it differs from it and all other members of the genus by its branched inflorescences. Further, P. lanuginosa has crenate leaflets 1.7–2.8 cm long, whereas P. multijuga has serrate leaflets 2.9–5.4 cm long. In P. lanuginosa, the hairs on the lower leaflet surfaces are densely lanate, whereas in P. multijuga, they are densely villous. Polylepis lanuginosa is also morphologically similar to P. canoi, but differs in its elliptic and shorter (1.7–2.8 cm long) leaflets (versus leaflets obovate and 2.4–3.9 cm long). Additionally, P. lanuginosa has shorter styles (1.9–2.3 mm long) than P. canoi (2.4–3.8 mm long).
Ecuador. — Azuay: Chaucha, Angas on western slope of western cordillera (due west of Cuenca), 02°55'S, 079°25'W, 3400 m, 05 January 1981, Balslev 1507 (AAU!, NY, QCA!, S); Angas “Parroquia chaucha” colecciones en margenes de Río Angas, 3400 m, 02 August 1983, Jaramillo 5464 (AAU!, NY, QCA!); 5468 (AAU!); 5478 (AAU!, MO!, NY, QCA!); Cuenca, Area Nacional de Recreación Cajas, collection made along Río Patul from the Comunidad Baute/Laguna Patul (watershed of Río Patul), 02°33'S, 079°21'W, 3500–4200 m, 05 February 2001, Clark 6227 (QCA!, QCNE, US!); Molleturo, on the road from Las Cajas National Park to Molleturo, about 10 km from Molleturo, 02°50'S, 079°20'W, 3400 m, 19 September 1983, Brandbyge 42264 (AAU!, MO!, QCA!); Cuenca-Molleturo road ca. 11 km W of pass in Las Cajas, 02°48'S, 079°18'W, 3350 m, 01 May 1992, Lægaard 102637 (AAU!, QCA!); Páramo de Cajas, W of Cuenca, along new road, ca. 14 km W of pass, 02°48'S, 079°17'W, 3450 m, 31 March 1985, Lægaard 53932 (AAU!); Páramo de Cajas, W of Cuenca, along new road, ca. 20 km W of pass, 02°48'S, 079°17'W, 2900 m, 31 March 1985, Lægaard 53934 (AAU!, MO!, QCNE); Páramo de las Cajas, W of pass, 02°46'S, 079°15'W, 2500 m, 26 August 1985, Lægaard 55036 (AAU!, MO!, QCA!); carretero Cuenca-Molleturo-Naranjal, 4.2 km de Molleturo, desvío a Río Blanco 16.4 km, 02°48'40"S, 079°23'07"W, 3630 m, 15 January 2003, Ulloa 1203 (HA, MO!, US!); Sayausi, Area Recreacional Las Cajas, 02°49'S, 079°07'W, 3740–4070 m, Romoleroux 1192 (AAU!); Zhud, at Panamericana, app. 3 km S of Zhud, 02°29'S, 079°00'W, 2800 m, 02 May 1992, Lægaard 102697 (AAU!, QCA!); límite del parque nacional, 3359 m, 20 April 2012, Barba BOP236 (QCA!); Río Blanco, Curiquinga, 3645 m, 05 May 2001, Calle 1 (QCA!); Cuenca-Molleturo road, 49 km NW of Cuenca, 26 July 1982, Clemants 2184 (AAU!, NY, QCA!); El Chorro ca. 6 km above Molleturo on road to Cuenca, 2800–2900 m, 07 March 1985, Harling 22858 (AAU!, GB, MO!, QCA!); Molleturo, 2600–2700 m, 31 October 1988, Harling 25539 (GB, MO!); Descente occidentale du páramo de Cajas vers Molleturo, 3350 m, 14 April 1988, Huttel 1021 (QCA!); Vallée du río Angas, à 1 Km au-dessus du hameau d’Angas, zone trés humide, 3300 m, 10 May 1988, Huttel 1112 (QCA!); Vertiente del Pacífico, 3200 m, 07 July 1995, León 3601 (QCA!); Área recreacional Cajas, 3470 m, 21 September 2000, Lizarzaburu 25 (QCA!); 3500 m, Romoleroux 408 (NY); Parque Nacional Cajas, carretera Soldados, 3260 m, 04 April 2007, Romoleroux 4461 (QCA!); 3040 m, 04 April 2007, Romoleroux 4466 (QCA!); Vía Soldados Angas, al frente del caserío Angas, 3321 m, 19 August 2008, Romoleroux 5030 (QCA!); Carretera Soldados-Angas, 3040 m, 04 April 2007, Romoleroux GPI4466 (QCA!); Cajas, found along path from Cochapamba to Molleturo, 3500–3600 m, 22 July 1999, Smeets 559 (QCA!); 2670–3275 m, Steyermark 52599 (F!); 3160 m, Valencia 458 (QCA!). — Bolívar: carretera Guaranda-Santiago-Totoras, 3000–3150 m, 21 February 1987, Romoleroux 269 (AAU!, QCA!). Cañar: Molleturo, along Páramo road to Manu W of Cañar, W of pass, 02°33'S, 079°02'W, 3300–3700 m, 20 June 1988, Lægaard 71563, 71565, 71569 (AAU!, QCNE); Molleturo, Páramo de Cajas, W of Cuenca, aong new road, ca. 11 km W of pass, 02°48'S, 079°17'W, 3200 m, 30 March 1985, Lægaard 53911 (AAU!, MO!, QCA!); Zhud, along Panamericana, 4 km S of Zhud, 02°28'S, 079°00'W, 3000 m, 26 August 1985, Lægaard 55030 (AAU!, MO!); along a paved road to Carshao, ca. 15 km off the Panamerican highway, 02°29'S, 079°00'W, 3180 m, 10 June 1999, Sklenar 7115 (AAU!); carretera entre Dacur y Gun, 2250 m, 08 October 1999, Bonifaz 3974 (QCA!); North rim of the valley of the río Cañar, between Tambo and Suscal, 3000 m, 23 April 1944, Camp E-2773 (F!, NY, VEN); colecciones entre Zhud, Joyagshi, 3500 m, 31 December 2007, Jaramillo 26120 (QCA!); 3300–3700 m, Romoleroux 1563 (QCA!); 3270 m, Romoleroux 384 (QCA!); 3100 m, Romoleroux 387 (NY, QCA!); carretera entre Zhud y El Tumbo, 3011 m, 06 December 2007, Romoleroux 4681 (QCA!); Oeste de Cañar, Km 10.5, Cerro Caucay, 3450 m, 27 April 1988, Romoleroux 588 (AAU!, MO!); Rose 2389 (NY). Chimborazo: Columbe, road Pallantanga-Riobamba, 01°55'S, 078°50'W, 2400–2900 m, 01 April 1993, Romoleroux 1565 (AAU!, QCA!); Juan de Velasco, Pangor-Tepeyac, 01°48'S, 078°52'W, 3200 m, 09 February 1983, Brandbyge 42059 (AAU!, MO!);, 3300 m, 03 May 1983, Brandbyge 42153 (AAU!, MO!, QCA!); Colta (Cajabamba)–Pallatanga, km 27, 01°50'S, 078°53'W, 2880 m, 21 May 1990, Jørgensen 91822 (AAU!); Km 64–68 on road Cumandá-Cajabamba, at Río Pangor, 01°55'S, 078°54'W, 2750–2800 m, 08 April 1985, Lægaard 54125 (AAU!, MO!, QCA!), 54128 (AAU!, QCA!); road Pallatanga–Cajabamba, 32 km from Pallatanga, 01°51'S, 078°53'W, 3000 m, 28 August 1976, Øllgaard 8959 (AAU!, MO!, NY, S). Penipe, Parroquia Puela, Palictahua, 01°31'05"S, 078°29'43"W, 2600 m, 20 January 1997, Estudiantes ESPOCH 701 (CHEP); Colta, Pangor, puente del Río Agua Dulce, 3000 m, 31 January 2007, Caranqui 1659 (QCA!); Colta. Pangor, Achín alto, 3140 m, 21 May 2013, Caranqui 2290 (QCA!); Comunidad de Tauris, Zona Zagin, 3700–3900 m, 03 September 2009, Cárate 1202 (QCA!); Comunidad de Ambrosio Lazo, Quebrada de Cumbo, 3374 m, 07 June 2009, Cárate 623, 624 (QCA!); Páramo cerca de la comunidad Yerba Buena, Cantón Pallatanga, 3374 m, 19 July 2012, Peyre 315 (QCA!); 2800–3200 m, Romoleroux 373 (NY, QCA!); Vía Cajabamba–Pallatanga (km 24 desde la Y), entrada a Pangar, 3298 m, 27 December 2011, Romoleroux 5696 (QCA!); Vía Cajabamba-Palatanga (km 24 desde la Y), entrada al Pangar, 3298 m, 27 December 2011, Romoleroux 5697 (QCA!); Carretera Alausi-Cañar, km 16, localidad Achupallas, 3400 m, 26 April 1988, Romoleroux 574 (AAU!, QCA!); Cañar, 2 km al sur de Zhud, 2850 m, 27 April 1988, Romoleroux 584 (AAU!, QCA!); 3250 m, Romoleroux 591 (QCA!), W Andes of Cuenca, Lehmann 6487 (F!).
Peru. Cajamarca: at Chugur near Hualgayoc, 2700–3000 m, May 1904, Weberbauer 4098 (holotype: G!; isotypes: MOL!; photos at F!, MO!).
Trees 5–15 m tall. Leaves slightly congested at the branch tips, imparipinnate with 5–7 pairs of lateral leaflets, obtrullate in outline, (11.0–)14.6–19.5 × 6.2–9.1(–10.7) cm; rachises densely lanate, points of leaflet attachment with a tuft of long, lanate hairs; stipular sheaths apically acute with spurs, densely lanate on the outer surfaces; leaflets elliptic in outline, second pair from the terminal leaflet the largest, one of this pair 2.9–3.6(–5.4) × 1.1–2.0 cm; margin serrate with 6–10 teeth, apically obtuse, basally unequally cordate; upper leaflet surfaces glabrous or sparsely villous; lower leaflet surfaces densely villous with whitish hairs 0.9–2.3 mm long. Inflorescences pendant, (15.4–)21.7–28.2(–36.0) cm long, bearing 47–83 flowers; floral bracts 7.6–9.7 mm long, narrowly triangular, densely villous on the outer surface; rachises villous. Flowers 6.4–7.5 mm diam.; sepals 4, ovate, green, densely lanate outside; stamens 7–13, anthers orbicular, with a dense tuft of straight white hairs on the upper half; styles fimbriate, 1.9–3.8 mm long. Fruits turbinate, with variable numbers and placement of irregular spines, densely villous; 4.5–9.6 × 6.0–10.1 mm including spines. Diploid.
Polylepis multijuga is restricted to northern Peru (Fig.
Based on 22 localities, the EOO for Polylepis multijuga is estimated at 17,200 km2 and the AOO at 116 km2. The species was categorized as VU (B1+2c, D2) in the World List of Threatened Trees (
Polylepis multijuga is similar to P. canoi but has 5–7 pairs of elliptic leaflets, whereas the latter has 2–4 pairs of obovate leaflets. Polylepis multijuga also has longer inflorescences (15.4–36.0 cm) with 47–83 flowers (P. canoi 8.2–14.5 cm, 12–26 flowers). Polylepis multijuga is also similar to P. ochreata, but differs by having larger and broader leaflets (2.9–5.4 × 1.1–2.0 cm versus 1.6–3.0 × 0.5–0.7 cm) and densely villous lower leaflet surfaces (densely sericeous in P. ochreata). Polylepis ochreata also has shorter inflorescenses (8.1–17.4 cm) with fewer flowers (21–49).
Peru. Amazonas: Chachapoyas, Dist. Leymebamba, surroundings of La Esperanza, 06°49'04"S, 077°43'01"W, 3200–3300 m, 27 June 2010, Glenn 411 (CAS, COL!, F!, K, MO!, P!); Dist. Leymebamba. Río El Jardín, 06°55'52"S, 077°43'09"W, 3370 m, 30 June 2009, Gruhn 173 (MO!); Dist. Leymebamba, a 2 Km de la Laguna de Los Cóndores, ruta hacia Leymebamba, 2700–2950 m, 18 August 1998, Quipuscoa 1329 (MO!); Quintecocha. Dist. Leymebamba, vicinity of guard cabin at Quintecocha, 06°51'33"S, 077°42'15"W, 3134 m, 12 July 2008, Rothrock 239 (BRIT, MO!); Balsas, Chuquillurco, ruta a Calla Calla, 3400 m, 06 October 2001, Sánchez 11018 (MO!); Chachapoyas-Cajamarca road, jalca de Calla-Calla, 30–37 km from Leimebamba Natural grassland, ‘Jalca’, and ‘ceja de selva’ just leaving the pass entering the Marañon valley, 06°50'S, 077°50'W, 3500–3600 m, 04 September 1983, Smith 5037 (MO!, USM!); Leimebamba, Oseres, 06°58'05"S, 077°39'57"W, 2542 m, 22 May 2015, Vega 257 (HAO, MO!); road Balsas to Chachapoyas, upper eastern Calla-calla slopes descending from pass, 3000–3300 m, 02 June 1998, Weigend 98/330 (USM!). Luya, Distr. Conila-Cohechan, 06°16'25"S, 078°00'10"W, 3050 m, 23 August 2012, Bussmann 17289 (MO!); Colcamar, 3200–3300 m, 24–26 June 1948, Pennell 15632 (USM!). Cajamarca: Chota, Bosque de Pagaibamba (Ocshawilca), al oeste del Chorroblanco, entre Huambos y Querocoto, 2500 m, 18 October 1987, Sánchez 4588 (F!); Paccha, al O de Chadin, 3650 m, 22 July 1993, Sánchez 6586 (F!). Cutervo, Gruta de San Andres, 2200 m, 15 July 1990, Llatas 2749 (F!, MO!). Hualgayoc, Dist. de Chugur, 06°43'08"S, 078°42'58"W, 3222–3568 m, 12 August 2009, Castillo 786 (USM!); Hacienda Taulis, 13 km beyond Palmito junction towards La Playa, 2900 m, 02 September 1964, Hutchison 6463 (MO!, USM!); Chugur, sobre la ruta de Perlamayo, 2950–3000 m, 20 March 1988, Sánchez 4681 (AAU!, F!). San Miguel, El Prado, Hacienda Taulis, 06°59'02"S, 078°58'21"W, 3398 m, 01 June 2015, Boza 3029; 3070; 3071; 3072; 3073; 3074; 3075; 3076; 3077; 3078; 3079; 3080 (USM!, Z!); Quishuarpampa (Agua Blanca), 2900 m, 04 July 1986, Mostacero 1201 (F!, MO!); Quishuarpampa (El Tingo–Jalca de las Estacas), 07°21'00"S, 077°50'00"W, 2950 m, 12 May 1977, Sagástegui 8833 (MO!); Millán (El Tingo–Taulis), 3000 m, 20 June 1980, Sagástegui 9536 (F!, MO!, USM!); Sobre el desvio a Tongot, entre Quilcate bajo y Catilluc, 3050 m, 13 September 1991, Sánchez 5762 (MO!). Santa Cruz, Distr. Pulán, parte baja de la Quebrada Cocan, ladera Oeste, 3280 m, 02 November 2001, Sánchez 11112 (MO!); Dist. Pulan, La Palma, 2800 m, 12 February 2007, Santa 927 (USM!); ad Chugur versus Hualgayoc, 2700–2900 m, 1901–1929, Weberbauer 4098 (G, MO!). La Libertad: Bolivar, District Uchumarca, Páramo in surroundings of Vira Vira/Lagunas La Quinua, 07°00'12"S, 077°45'07"W, 3670 m, 17 May 2011, Bussmann 16931 (MO!). Huicungo, Dist. Uchumarca, 06°59'30"S, 077°43'07"W, 3140 m, 02 November 2012, Paniagua 8642 (MO!). Lambayeque: Ferrenafe, Dos Puentes, arriba de Incahuasi, 2900–3000 m, 09 July 1987, Ferreyra 20908 (USM!); road Incahuasi to Sinchihual and Tungula, 06°12'07"S, 079°17'57"W, 2897 m, 24 November 2014, Weigend 9660 (USM!).
Trees; 4–6 lateral leaflet pairs; lowers leaflet surfaces lanate or sericeous; fruits with flattened or thin spines, densely villous.
Polylepis pauta Hieron.
The subsectional epithet Pauta is a noun in apposition ['Nomen vernaculum: Pauta'; fide: Hieron., Bot. Jahrb. Syst. 21(3): 314. 1895].
Ecuador. Carchi: Cantón Montúfar, Loma El Corazón (Bretaña), al sureste de Huaca, al este de la Colonia Huaqueña, Río Minas, 00°35'N, 077°42'W, 3200–3500 m, 9 Apr 1989, Tipaz 35 (holotype: QCA!; isotypes: AAU!, MO!).
Trees 5–10 m tall. Leaves strongly congested at the branch tips, imparipinnate with (4–)5–6 pairs of the lateral leaflets, obtrullate in outline, (3.8–)4.3–7.3 × 2.4–4.5 cm; rachises densely sericeous, points of leaflet attachment with a tuft of long, straight whitish to yellowish hairs; stipular sheaths apically truncate, densely sericeous in the upper surface; leaflets narrowly to ovate in outline, second pair from the terminal leaflet the largest, one of this pair 1.4–2.2 × 0.4–0.5 cm; margins entire to slightly crenate with 6–7 teeth, apically slightly emarginate seemingly acute by the prolongation of hairs, basally unequally cordate; upper leaflet surfaces glabrous; lower leaflet surfaces densely lanate with whitish silky hairs 1.1–1.6 mm long. Inflorescences pendant, (6.8–)11.1–16.6 cm long, bearing 19–29 flowers; floral bracts 6.1–9.4 mm long, narrowly triangular, glabrous on the outer surface; rachises densely sericeous. Flowers 4.8–5.5 mm diam.; sepals 4, ovate, green, glabrous outside; stamens 8–10, anthers orbicular, with a dense tuft of straight white hairs on the upper half; styles fimbriate, 1.6–2.0 mm long. Fruits turbinate, with variable numbers and placement of flattened spines, densely sericeous; 3.9–6.7 × 4.3–7.5 mm including spines. Diploid.
Polylepis longipilosa is restricted to north-western Ecuador (Carchi) (Fig.
The EOO for Polylepis longipilosa is estimated as 17,689 km2, the AOO is assessed at 60 km2 and it is known from eight locations. It occurs in Reserva Ecológica El Angel. However, the Andean Forest and páramos at Carchi have, in recent years, come under increasing threat from timber cutting and forest burning and advancement of the agricultural frontier, which has contributed to the fragmentation and destruction of high Andean ecosystems (
The populations of Polylepis from Carchi on the southern slopes of Volcán Chiles and on the road between Maldonado and Tulcán have previously been identified either as P. ochreata (
There are three distinctive collections of the P. pauta/sericea complexes that
Ecuador. Carchi: Tulcán, carretera Túlcan-Tufiño-Maldonado-Chical col. en km 12 de Tufiño, cerca de las lagunas, 00°48'N, 077°55'W, 3900 m, 23 April 1993, Freire & Andersen 2547 (AAU!); road Tulcán-Maldonado, near Volcán Chiles, 00°48'N, 077°56'W, 3850–4000 m, 16 August 1985, Laegaard 54965 (AAU!, MO!, QCA!), 54967A, 54967B, 54967C (AAU!), 54967D, 54967E, 54967F (AAU!, QCA!); along the road from Tulcán to Volcán Chiles, 3900 m, 6 October1995, Sklenár & Kosteckova 1412A (QCA!); camino Tufiño, sitio Agua Hediondas, en la base del Volcán Chiles, límite con Colombia, 00°48'N, 077°54'W, 3500 m, 8 November 1993, Palacios 11847 (AAU!, MO!, QCNE); carretera entre Tulcán y Maldonado, faldas del Volcán Chiles, punto más alto del cruce de carretera, 00°45'N, 077°59'W, 3800 m, 19 May 1991, Palacios & Rubio 7349 (AAU!, MO!); southern slopes of Volcan Chiles, 00°49'N, 077°57'W, 3600 m, Ramsay 911 (QCA!, QCNE); route de Tufiño a Maldonado, 10 km après Tufiño, zone très humide, 3850 m, 06 July 1988, Huttel 1390 (QCA!); carretera San Gabriel-Shután alto, 3500 m, 25 March 1989, Jaramillo-Asanza 10862 (QCA!); comuna La Esperanza, páramo de El Artezón, sector Monte Redondo, 3789 m, 18 September 2007, Salgado 220B, 239A (QCA!).
Polylepis annulatipilosa
Bitter, Bot. Jahrb. Syst. 45: 596. 1911. Type. Ecuador. Pichincha: Andes of Quito, Jameson 16 (lectotype, designated by
Polylepis stuebelii Hieron., Bot. Jahrb. Syst. 21: 313. 1896. Type. Ecuador. Napo: E slope of Cerro Quilindaña near Bambasacha, 3700 m, Stübel 204 (holotype: B destroyed; photos at F!, MO!, NY!, US!).
Ecuador. Pichincha: “Corredor Machai”, 3900 m, Oct 1871, Stübel 232a (holotype: B destroyed; photos at F!, GH!, MO!).
Trees 2–12 m tall. Leaves strongly congested at the branch tips, imparipinnate with 4–5(–6) pairs of the lateral leaflets, obtrullate in outline, 3.2–4.9 × 2.2–3.0 cm; rachises sparsely sericeous, points of leaflet attachment with a tuft of long, straight whitish hairs; stipular sheaths apically acute with spurs, glabrous to sparsely sericeous (adult) or densely sericeous (juvenile) in the upper surface; leaflets elliptic in outline, second pair from the terminal leaflet the largest, one of this pair (1.1–)1.4–1.6 × 0.5–0.6 cm; margin crenate with 4–6 teeth, subcoriaceous, apically emarginate, basally unequally cordate; upper leaflet surfaces glabrous or sparsely sericeous with few hairs on the mid-veins; lower leaflet surfaces sparsely sericeous with whitish hairs 0.4–0.9 mm long. Inflorescences pendant, (8.1–)12.6–14.3(–19.3) cm long, bearing 9–15(–21) flowers; floral bracts (9.1–)10.0–12.2 mm long, narrowly triangular, densely sericeous on the outer surface; rachises densely villous. Flowers 6.0–7.4(–9.2) mm diam.; sepals 4, ovate, green, densely sericeous outside; stamens 9–15, anthers orbicular, with a dense tuft of straight white hairs on the upper half; styles fimbriate, 2.2–3.0 mm long. Fruits turbinate, with variable numbers and placement of flattened spines, densely sericeous; (2.6–)3.4–5.5 × 3.3–6.0(–8.2) mm including spines. Tetraploid, aneuploid; perhaps also diploid.
Polylepis pauta occurs in the north-eastern Cordillera Oriental of Ecuador (Fig.
The EOO for Polylepis pauta is estimated as 1,590 km2, the AOO is assessed at 132 km2 and it occurs at 14 locations. Large populations occur within Cayambe-Coca National Park and Antisana Ecological Reserve, but there is genetic evidence for a loss of genetic diversity due to forest fragmentation (
An outstanding feature of P. pauta are the differences in leaflet number, shape and indument between young and adult plants. On young plants, leaves look very similar to those of P. longipilosa, whereas as the plants mature, the leaves become almost glabrous and have 4–5 lateral leaflet pairs.
Ecuador. Cotopaxi: 2 km S of Paso de la Virgen on road Quito-Baeza, 00°20'S, 078°13'W, 3850–4000 m, 16 May 1984, Lægaard 52133 (AAU!, QCA!). Imbabura: González Suárez, Lagunas de Mojanda, Laguna Negra, 00°08'N, 078°15'W, 3700 m, 22 September 1990, Øllgaard 98194 (AAU!); vía hacia la laguna de Mojanda, 00°08'N, 078°15'W, 3500–3700 m, 02 November 1987, Romoleroux 475 (AAU!, QCA!). Otavalo, road from Otavalo to lagunas Mojanda, ca. 3 km before the lakes, 00°10'N, 078°17'W, 3500–3700 m, 23 October 1983, Balslev 4450 (AAU!, QCA!). Quiroga, Cotacachi, Reserva Ecológica Cotacachi-Cayapas, 00°18'N, 078°22'W, 3300–3350 m, 02 March 1992, Peñafiel 1091 (MO!). Tocachi, Laguna Grande de Mojanda, 15 km S of Otavalo, 00°08'N, 078°16'W, 3750 m, 14 May 1985, Eriksen 59359 (AAU!); 59374 (AAU!). Mojanda, Tomauco, 3309 m, 05 June 2008, Salgado 428 (QCA!); Mojanda, Tomauco, 3274 m, 05 June 2008, Salgado 458 (QCA!); Páramo de Mojanda, on the SW slope of the peak Nudo de Mojanda, 4130 m, 06 November 2007, Sklenar 10746 (QCA!). Napo: Oyacachi, 0°12'S, 78°8'W, 3680 m, 08 April 2012, Homeier 4948 (QCA!); N of Volcán Los Puntos, 00°12'S, 078°10'W, 4200 m, 27 July 1985, Lægaard 54756A (AAU!); N of Volcán Los Puntos, 00°12'S, 078°10'W, 3850–3900 m, 28 July 1985, Lægaard 54756B (AAU!); N of Volcán Los Puntos, 00°12'S, 078°10'W, 3850–3900 m, 28 July 1985, Lægaard 54756C (AAU!); 54756D (AAU!); 54756E (AAU!); 54756F (AAU!); 54761 (AAU!, MO!); Reserva Ecológica Oyacachi, 00°13'49"S, 078°08'44"W, 3915 m, 16 December 2008, Romoleroux 5346 (MO!, QCA!). Papallacta, Oyacachi, 0°18'6"S, 78°8'28"W, 3970 m, 10 June 2009, Homeier 4191 (QCA!); Páramo de Papallacta, 00°20'S, 078°10'W, 12 January 2015, Kessler s.n (Z!); Pifo-Papallacta, 3–5 km E of Paso de La Virgen, Páramo-swamp, 00°21'S, 078°11'W, 3700–3900 m, 09 June 1992, Lægaard 103115 (AAU!, GOET!, QCA!); 3 km E of Paso de la Virgen on road Pifo-Papallacta, 00°20'S, 078°11'W, 3950–4050 m, 02 June 1985, Lægaard 54442 (AAU!, QCA!); 54443 (AAU!); 54444 (AAU!, MO!); 54446 (AAU!); 54447 (AAU!, MO!, QCA!); 54448 (AAU!, MO!, QCA!); 54449 (AAU!, QCA!); 54450 (AAU!, MO!); 54451 (AAU!, MO!); 54452 (AAU!, MO!); along road Pifo-Papallacta, E of Paso de la Virgen, 00°21'S, 078°11'W, 3750–3850 m, 21 June 1985, Lægaard 54558B (AAU!); 54558C (AAU!); 54558D (AAU!); 54559 (AAU!, MO!, QCA!); 54560 (AAU!, MO!); 54561 (AAU!); road Quito–Baeza, 7–8 km NW of Laguna de Papallacta (Páramo de Guamaní), 00°19'S, 078°08'W, 3800 m, 20 July 1976, Øllgaard 8156 (AAU!, MO!, NY); Reserva Ecológica Oyacachi, 00°17'46"S, 078°08'49"W, 3927 m, 20 September 2008, Romoleroux 5194 (MO!, QCA!); carretera Quito–Baeza, páramo above Papallacta, 00°21'S, 078°10'W, 3400–3700 m, 28 May 1987, van der Werff 9638 (AAU!, GB, MO!). Pintag, Paso de Guamaní, quebrada, about +-4 km E Paso de Guamaní, on road to Papallacta, 00°20'S, 078°20'W, 3900 m, 26 March 1967, Sparre 15029 (AAU!, S). Quijos, Parroquia Papallacta, 00°21'S, 078°11'W, 3700 m, 28 May 1990, Cerón 10054 (MO!); Reserva Ecológica Antisana, carretera Pifo-Baeza, Páramo de la Virgen, 00°20'S, 078°12'W, 3960 m, 23 November 1998, Freire 2852 (AAU!, ILLS, MO!, QCNE); Reserva Ecológica Antisana, carretera Pifo–Baeza, Páramo de la Virgen, 00°23'S, 078°12'W, 3730 m, 24 November 1998, Freire 2870 (ILLS, MO!, QCNE); Reserva Ecológica Antisana, Páramo de Guamaní, carretera Pifo-Papallacta, La Virgen, 00°20'S, 078°12'W, 4140 m, 24 July 1998, Vargas 1946 (AAU!, ILLS, MO!, QCNE); carretera Quito-Tena via Baeza km 52, 3820 m, 03 August 1984, Dodson 14832 (MO!); 8 kms de la población de Oyacachi, siguiendo el sendero hacia Cochapamba, 3500 m, 12 March 1991, Gavilánez 462 (QCA!); carretera Oyacachi-Papallacta, colecciones a 11 km de la Laguna de Loreto, 3800 m, 27 April 1998, Guerrón 343 (QCA!); Papallacta, 3400–3600 m, 16 August 1990, Jaramillo 11832 (AAU!, MO!); Papallacta, 3400 m, 17 August 1990, Jaramillo 11842 (MO!); Páramo de Guamaní, road Quito Papallacta, 4000 m, 04 March 1979, Kieft 228 (QCA!); 3 km E of Paso de la Virgen on road Pifo-Papallacta, 3951–4050 m, 06 February 1985, Lægaard 54452 (QCA!); along road Pifo-Papallacta, E of Paso de la Virgen, 3750 m, Lægaard 54558 (QCA!); along road Pifo-Papallacta, E of Paso de la Virgen, 3750–3850 m, 21 June 1985, Lægaard 54560 (QCA!); N of Volcán Los Puntos, 3850 m, Lægaard 54756 (QCA!); N of Volcán Los Puntos, 3850–3900 m, 28 July 1985, Lægaard 54757 (QCA!); N of Volcán Los Puntos, 3851–3900 m, 28 July 1985, Lægaard 54758 (QCA!); 54759 (QCA!); Oyacachi, Yarupaccha, 3620–3680 m, 16 January 1996, Navarrete 1449 (QCA!); Reserva Ecológica Oyacachi, 3940 m, 28 January 2007, Romoleroux 4282 (QCA!); 4297 (QCA!); Reserva Ecológica Oyacachi, 3895 m, 23 February 2007, Romoleroux 4340 (QCA!); Reserva Ecológica Oyachachi, 3465 m, 08 March 2008, Romoleroux 4751 (QCA!); Páramo de Guamaní, alrededores de la laguna de Papallacta, 3900–4000 m, 06 December 1987, Romoleroux 491 (AAU!, NY, QCA!); Reserva Ecológica Oyachachi, 3929 m, 13 September 2008, Romoleroux 5167 (QCA!); 3681 m, 06 February 1985, Romoleroux 5168 (QCA!); 3917 m, 14 April 2009, Romoleroux 5475 (QCA!); 3880 m, 16 May 2009, Romoleroux 5489 (QCA!); Reserva Ecológica Oyacachi, 3560 m, 2 February 2007, Romoleroux A4321 (QCA!); Páramo de la Virgen, 3904 m, 29 September 2004, Salgado 1 (QCA!); about 3 km W of Oyacachi, 3550 m, 27 March 1996, Ståhl 2278 (QCA!); crescit prope Bambasacha in declivibus orientalibus mentis Quilindaña sitis, 3700 m, s.d., Stübel 204 (B, F!, MO!, NY, US!). Pichincha: Cayambe, carretera Cayambe-Hda. Piamonte-Patapamba, 00°02'S, 078°04'W, 3700 m, 04 December 1993, Freire 2606 (AAU!, QCA!). Papallacta, Along road Pifo-Papallacta, E of Paso de la Virgen, 00°21'S, 078°11'W, 4200–4300 m, 20 June 1985, Lægaard 54558A (AAU!); Pichincha-Napo, base del Volcán Antisana, entrada por Pintag hacia laguna Micacocha, campamento de EMAP, 00°27'S, 078°10'W, 4000–4100 m, 09 October 1990, Romoleroux 1117 (AAU!, QCA!); Pichincha-Napo, base del Volcán Antisana, entrada por Pintag hacia laguna Micacocha campamento EMAP, 00°27'S, 078°10'W, 4000–4100 m, 09 October 1990, Romoleroux 1118 (AAU!, QCA!); Páramo de la Virgen, camino antiguo, 0°20'S, 78°12'W, 3938 m, 20 September 2004, Salgado 3A (QCA!). Pifo, Mount Guamaní, 0°20'S, 78°33'W, 3600–3800 m, 15 September 1939, Asplund 8767 (QCA!); 2 km west of La Virgin on the road from Pifo to Papallacta, 00°17'S, 078°12'W, 3950–4050 m, 20 May 1984, Brandbyge 42638 (AAU!, MO!); Pifo-Papallacta (new road) app. 1 km W of Paso de la Virgen, 00°19'S, 078°13'W, 3700 m, 16 April 1992, Lægaard 102327 (AAU!, GOET!); 2 km S of Paso de la Virgen on road Quito-Baeza, 00°20'S, 078°13'W, 4000–4200 m, 19–20 May 1984, Lægaard 52134 (AAU!); 52135 (AAU!); 52138 (AAU!, MO!); 52162 (AAU!); 52176 (AAU!, QCA!); road Pifo–Papallacta, near Paso de la Virgen, 00°19'S, 078°13'W, 4000–4100 m, 13 March 1985, Lægaard 53849 (AAU!, MO!, QCA!); road Pifo-Papallacta, 3 km W of Paso de la Virgen, 00°18'S, 078°14'W, 3700–3900 m, 07 August 1985, Lægaard 54901A; 54901B; 54901C; 54901D; 54902AA; 54902K; 54902M; 54902P; 54902S; 54902U; 54902W; 54902Y (AAU!); at Paso de la Virgen, 00°18'S, 078°12'W, 4000–4050 m, 28 November 1985, Lægaard 55729 (AAU!, GOET!, MO!); carretera Quito–Papallacta, 1 km al este de la cumbre (La Virgen), 00°20'S, 078°15'W, 3800 m, 06 October 1986, Neill 7378A (AAU!, MO!, QCA!); 2 km al E de la cumbre de la carretera Pifo-Papallacta (La Virgen), 00°20'S, 078°15'W, 3900 m, 28 November 1987, Neill 8018 (AAU!, GB, MO!, QCA!, QCNE); Vía Baeza, 1 km antes del cruce de la Virgen, 00°18'S, 078°12'W, 3950 m, 01 March 1989, Palacios 3994 (AAU!, MO!); carretera Quito-Papallacta km 40–53, 00°16'S, 078°15'W, 3300–3800 m, 27 December 1992, Romoleroux 1507 (AAU!, QCA!); 00°21'S, 078°13'W, Romoleroux 353 (QCA!); Páramo de Guamaní, on the left side of the road Quito-Papallacta, 0°19'S, 78°12'W, 4000 m, 28 June 1997, Sklenár 2019 (QCA!). Quito, Parroquia Pifo, carretera Quito–Baeza, Páramo de la Virgen, 00°14'S, 078°20'W, 3500–3900 m, 25 April 1992, Cerón 18792 (MO!); Jameson 16 (MO!); road from Quito via Pifo to Papallacta, 00°34'S, 078°19'W, 3950 m, 04 July 2014, Kessler 14602; 14603; 14604; 14605 (Z!); Pass on Quito-Papallacta road, 3800–3900 m, 06 April 1991, Kessler 2750 (GOET!); 2755 (GOET!); Páramo de Guamaní, carretera Pifo-Papallacta, Km 27, 00°19'S, 078°12'W, 3960 m, 13 June 1990, León 1149 (QCA!); Baeza-Quito km 53, 00°20'S, 078°12'W, 4200 m, 08 July 2002, Schmidt-Lebuhn 378 (GOET!, QCA!). Tabacundo, at highest pass on road Mojanda-Tabacundo, 00°07'N, 078°15'W, 4030 m, 08 April 2001, Lægaard 21538A; 21538B (AAU!). Tocachi, Páramo de Mojanda, at Laguna Negra and S-side of Laguna Grande, 00°08'N, 078°16'W, 3800 m, 14 May 1985, Lægaard 54316B (AAU!, MO!, QCA!); 54330 (AAU!, MO!); 54333 (AAU!, MO!, QCA!); 54336 (AAU!); 54346 (AAU!, MO!, QCA!); Lagunas Mojanda, 00°07'N, 078°16'W, 3800 m, 30–31 Jul 1992, Palacios 10210 (AAU!, MO!); 10239 (AAU!, MO!); Lagunas de Mojanda, ca. Laguna Grande, 00°08'N, 078°16'W, 3700–3800 m, 01 June 1988, Romoleroux 654 (AAU!, QCA!); 3400–3500 m, Acosta-Solís 8379 (F!); 3700–4000 m, s.d., Asplund 18244 (S); alrededores de la Laguna Grande de Mojanda Cajas, 3960 m, 27 February 1999, Jaramillo 20986 (QCA!); the pass on Quito-Papallacta road, 3800–3900 m, 06 April 1991, Kessler 2749 (GOET!, MO!); 2750 (GOET!, MO!); 2753 (GOET!, MO!); 2754 (GOET!, LPB, MO!); Páramo de la Virgen, 3100 m, 01 November 2006, Muñoz 4 (QCA!); Laguna grande de Mojanda-Cajas, 3800 m, 19 September 2011, Pérez 5117 (QCA!); 3960 m, 01 August 1975, Little 22 (MO!).
Polylepis serrata var. parcipila Bitter, Bot. Jahrb. Syst. 45: 593. 1911. Type. Peru. Cusco: La Convencion, Yanamanche, between Cusco and Santa Ana, 3500–3800 m, Weberbauer 4954 (holotype: B destroyed; isotype: Vratisl).
Polylepis serrata var. psilanthera Bitter, Bot. Jahrb. Syst. 45: 593. 1911. Type. Based on Polylepis serrata Pilg.
Peru. Huanuco: Huamalics, southeast of Monzon, 3400–3500 m, 1903, Weberbauer 3354 (holotype: B destroyed; photos at MO!, US).
Trees 3–27 m tall. Leaves slightly congested at the branch tips, imparipinnate with 4–7 pairs of lateral leaflets, obtrullate in outline, (5.4–)6.7–8.7(–11.1) × (3.2–)3.9–5.7(–6.4) cm; rachises densely tomentose; points of leaflet attachment with a ring of short tomentose hairs around; stipular sheaths apically acute with spurs, densely lanate on the outer surfaces; leaflets elliptic in outline, second pair from the terminal leaflet the largest, one of this pair (1.8–)2.4–3.5 × 0.8–1.0(–1.2) cm; margin serrate with 4–5 teeth, apically acute, basally unequally cordate; upper leaflet surfaces glabrous or sparsely lanate mainly in the mid-vein depression; lower leaflet surfaces densely lanate with whitish hairs 0.7–1.2 mm long. Inflorescences pendant, (7.6–)9.5–13.3(–17.3) cm long, bearing 16–35 flowers; floral bracts 3.4–4.5 mm long, narrowly triangular, densely villous on the outer surface; rachises villous. Flowers 5.2–5.9 mm diam.; sepals 4, ovate, green, densely sericeous outside; stamens (4–)6–14, anthers orbicular, with a dense tuft of straight white hairs on the upper half; styles fimbriate, 1.2–2.3 mm long. Fruits turbinate, with variable numbers and placement of thin spines, densely villous; (3.8–)6.1–6.7 × 5.6–8.8 mm including spines. Diploid.
Polylepis serrata is distributed from San Martín to Cusco, Peru (Fig.
The EOO for Polylepis serrata is estimated as 68,454 km2, the AOO is assessed at 100 km2 and it occurs at 18 locations. In Peru, was categorized as Near Threatened (
This species is quite similar to P. pauta and, in fact, it was treated as a junior synonym by
Polylepis serrata also is morphologically similar to P. ochreata, with which it shares the number of leaflets (4–7 pairs). The most obvious differences between P. serrata and this species are the leaflet width, margin, apex and hair type and length, with P. serrata having elliptic leaflets 0.8–1.2 cm long, with acute apex and longer lanate hairs (0.7–1.2 mm) on the lower surface, whereas P. ochreata has narrowly elliptic leaflets 0.5–0.7 cm long, emarginate apex and short sericeous hairs (0.3–0.5 mm) on the lower surface.
Peru. Cusco: Calca, Lares Cuncani, 07 June 1991, Tupayachi 1505 (CUZ!). La Convención, Prov. Machupicchu, Chakimayu, 3235 m, 01 September 2002, Arce s.n (CUZ!, USM!); Batiyayoc 13°08'01"S, 072°19'45"W, 3705 m, 01 October 2002, Arce s.n (CUZ!); Dist. Santa Teresa, Uchuyillaspay, 13°07'23"S, 072°37'30"W, 3883 m, 22 September 2005, Huamantupa 7018 (CUZ!, MO!); Dist. Echarate, Huayopata, San Luis, 13°04'43"S, 072°23'25"W, 2800 m, 30 March 2006, Huamantupa 7526 (CUZ!, MO!, USM!); Potrero, Bosque de Ukumuriyoc, 3600 m, 01 October 2002, Palomino 1737 (QCA!); Dist. Huayopata, Panticalle, Abra Málaga 13°08'02"S, 072°19'41"W, 3690 m, 30 May 2006, Toivonen 88 (CUZ!); 89 (CUZ!); 90 (CUZ!); 91 (CUZ!); Cerca Canchayoc, 3600 m, 29 June 1967, Vargas 19872 (CUZ!); Canchayoc, 3650–4000 m, 10 January 1968, Vargas 20086 (CUZ!); Canchayoc, 3700 m, 23 April 1980, Vargas 23311 (CUZ!); Yanamanche Quellomayo, 3600–4000 m, 25 July 1944, Vargas 4457 (CUZ!). Paucartambo, Challabamba, Pumataki, 13°09'16"S, 071°38'33"W, 3671 m, 10 December 2014, Boza 3024 (USM!, Z!); Challabamba, between Acjanaco and Tres Cruces, 13°10'07"S, 071°37'58"W, 3450 m, 10 December 2014, Boza 3025 (USM!, Z!); Trocha Ericsson, Acjanaco, Parque Nacional Manu, 3250–3350 m, 01 September 1990, Cano 4041 (USM!); Qollatambo, Parque Nacional Manu, 3700–3800 m, 10 September 1990, Cano 4319 (USM!); Tres Cruces, Parque Nacional Manu, 3600–3700 m, 06 March 1991, Cano 4588 (USM!); Cerro Chapuyoc, Challabamba, ParqueNacional Manu, 3350–3450 m, 15 March 1991, Cano 4689 (USM!); Valle del Pilcopata, near Accanaco Pass, turnoff to Tres Cruces,13°13'S, 071°35'W, 3500 m, 15 December 1983, Foster 7548 (MO!, USM!); Acjanaco, Parque Nacional Manu, Trocha Ericsson, 3000–3200 m, 22 July 1991, Huapaya 221 (USM!); Region of Acanacu and the Cordillera or Tres Cruces, 3290–3500 m, 07 December 1978, Luteyn 6386 (AAU!, MO!, USM!); Pillahuata, alrededores, Tres cruces, 130 km de Cusco en el camino hacia Pilcopta, 13°05'S, 071°30'W, 2000 m, 04 April 1987, Núñez 7749 (CUZ!, MO!, USM!); Km 130 hacia Kosñipata; incluye Acjanacu, Pillahuata, parte alta del Parque Nacional del Manu y ceja de selva hacia Kosñipata, 13°05'S, 071°30'W, 2600 m, 30 October 1987, Núñez 8482 (CUZ!, MO!); Tres Cruces, Parque Nacional Manu, 3500–3700 m, 01 April 1989, Tovar 10081 (USM!); Abra de Acjanaco-Tres Cruces de Oro, carretera Acjanaco-Pillahuata, 13°07'S, 071°40'W, 3700 m, 13 November 1986, Tupayachi 44; Tupayachi 45 (CUZ!, MO!); Entre Paucartambo y Acjanacu, Abra de Acjanacu, 3500 m, 25 January 1960, Vargas 13130 (CUZ!); Hda. Pillco, Valle de Paucartambi, 2800 m, 12 April 1967, Vargas 19242 (CUZ!); Abra de Acjanacu, 3500 m, 20 June 1986, Vargas 24009 (CUZ!); Quebrada de Acjanacu, 3500 m, 11 December 1942, Vargas 3004 (CUZ!, MO!); Chacapampa, 1800–2000 m, 01 December 1950, Vargas 9909 (CUZ!, MO!). Quispicanchis, Marcapata; 176 km from Cusco on road to Maldonado, Marcapata remmant forest to Cocha, 13°25'S, 070°54'W, 3150 m, 08 March 1991, Núñez 13151 (CUZ!, MO!); entre Abra Walla Walla y Marcapata a 210 km de Cusco, 13°25'S, 070°54'W, 2800–4600 m, 21–25 April 1988, Núñez 9032 (CUZ!, IBE, MO!); Huaillai-Marcapata, junto al río Araza, 2900 m, 11 December 1943, Vargas 3765 (CUZ!); Marcapata, 15–16 February 1929, Weberbauer 7803 (A!, MO!). Urubamba, Pakaymayu, 13°14'11"S, 072°29'38"W, 3861 m, 01 June 2002, Arce s.n (USM!); entre San Luis y Abra Malaga, 13°06'S, 072°22'W, 3500 m, 16 October 2002, Lehnert 444 (GOET!); Machu Picchu’, in Urcoscancha, a pampa above the village of Palcay, 13°09'30"S, 072°31'53"W, 3645 m, 05 July 1982, Peyton 792 (MO!); Lado Oriental de Cumbre Málaga, 01 October 1984, Rivas s.n (USM!); Machupicchu, campamento (km 90), 13°11'17"S, 072°26'10"W, 3070 m, 02 August 2006, Toivonen 1 (CUZ!); Dist. Machupicchu, campamento (km 90), 13°11'17"S, 072°26'10"W, 3070 m, 02 August 2006, Toivonen 2 (CUZ!); Dist. Machupicchu, Pakaymayu, 13°14'9"S, 072°29'36"W, 3760 m, 24 August 2006, Toivonen 24, 25; 26; 27; 30 (CUZ!); Dist. Machupicchu. Pakaymayu, 13°14'9"S, 072°29'36"W, 3760 m, 13 September 2006, Toivonen 3760 (CUZ!). Huánuco: Pachitea, Dist. de Umari, Comunidad Campesina de San Marcos, 3400 m, 04 March 2010, Beltrán 6740 (USM!); 6760 (USM!); 01 July 1903, Weberbauer 3354 (B, MO!). Junín: Satipo, Cordillera Vilcabamba, Río Ene slope, near summit of divide, 11°39'36"S, 073°40'02"W, 3320 m, 14 June 1997, Boyle 4398 (USM!). San Martín: Mariscal Caceres, Primer derrumbe y laguna del pato y asociadas del P. N. del Río Abiseo, 3250–3450 m, 24 June 1996, Cano 7265 (USM!); Dist. de Huicungo, Parque Nacional Río Abiseo, Callejón Rojas, 3600–3700 m, 06 July 2011, Castillo 1026 (USM!); 959 (USM!); Dist. de Huicungo, zona de Alpamachay, 3200–3300 m, 14 June 2001, León 5224 (USM!); Río Abiseo National Park; forest on edge of Laguna de Chochos, NW corner of park, 07°00'S, 077°00'W, 3300 m, 19 May 1986, Young 3175 (F!, MO!, USM!); Río Abiseo National Park, jucture of Quebrada Misquichilca and Quebrada Quimar, 4 km SE of Condormarca, 07°00'S, 077°00'W, 3500 m, 05 June 1986, Young 3552 (F!, MO!, USM!); forest on the edge of Laguna de Chochos, Chochos, NW corner of Río Abiseo National Park, 07°S, 077°W, 3300 m, 17 July 1987, Young 4863 (MO!, USM!).
Trees; 2–7 lateral leaflet pairs; lower leaflets sericeous or villous; fruits with flattened spines, densely sericeous or villous.
Polylepis sericea Wedd.
The subsectional epithet Sericeae is a plural adjective agreeing in gender with Polylepis.
Peru. Ancash: Cordillera Blanca above Caraz, Jun 1903, Weberbauer 3229 (holotype: B destroyed; photos at F!, GH!, NY!).
Polylepis albicans Pilger A inflorescence B lower leaflet surface C upper leaflet surface D branch apex with young inflorescence E young inflorescence (A Boza & Urquiaga 3014 B Boza & Urquiaga 3013 C–E Boza & Urquiaga 3015). Scale bars: 1 cm (A, B, C); 2 cm (D); 0.4 cm (E). Photographs by E. G. Urquiaga F.
Trees 3–7(12) m tall. Leaves strongly congested at the branch tips, imparipinnate with 3–4 pairs of lateral leaflets, obtrullate in outline, 3.5–4.9 × (2.5–)2.8–3.4 cm; rachises densely sericeous, points of leaflet attachment with a tuft of long, straight hairs, with ferruginous resin at leaflet insertion; stipular sheaths apically acute with spurs, densely sericeous on the outer surfaces; leaflets elliptic in outline, second pair from the terminal leaflet the largest, one of this pair 1.4–2.0 × 0.4–0.7 cm; margin slightly crenate at the apex with 4–5 teeth, strongly revolute, coriaceous, apically emarginate, basally unequally cordate; upper leaflet surfaces glabrous or sparsely sericeous; lower leaflet surfaces densely sericeous with short silky hairs 0.3–0.5 mm long. Inflorescences pendant, 3.9–6.6(7.5) cm long, bearing 18–21 flowers; floral bracts 5.5–6.9 mm long, narrowly triangular, densely sericeous on the outer surface; rachises sericeous. Flowers 3.4–7.5 mm diam.; sepals 3–4, ovate, green, densely sericeous outside; stamens 7–18, anthers orbicular, with a dense tuft of straight white hairs on the upper half; styles fimbriate, 1.4–3.2 mm long. Fruits turbinate, with variable numbers and placement of flattened spines, densely sericeous; 3.1–5.6 × 2.3–5.6 mm including spines. Diploid.
Polylepis albicans occurs in north-western Peru in the Cordillera Blanca in Ancash and in the adjacent high Andes of La Libertad (Fig.
The Extent of Occurrence (EOO) for P. albicans is estimated as 13,028 km2, the area of occupancy (AOO) is assessed at 164 km2 and it is known from 24 locations. It occurs in Huascarán National Park which encompasses almost the entire Cordillera Blanca. However, illegal mining occurs within the Park, becoming a direct threat to the species. Polylepis albicans is subject to reforestation activities in Huascarán National Park (
This species was described by
To us, the reddish glands, thick leaf texture and often emarginated leaflet apices suggest that P. albicans may include some genetic elements from P. weberbaueri, with which it co-occurs, but this remains to be tested by molecular studies.
Peru. Ancash: Carhuaz, Sonquenua, Shilla, 4020 m, 21 December 1989, Arce & Sánchez 188 (MO); Valley of Río Marcará, 2.5 hours from Vicos on trail to Lejiacocha, 09°19'00"S, 077°31'00"W, 3600 m, 11 March 1964, Hutchison & Wright 4325 (F, MO, USM); Shacshicucho, 4050 m, 26 August 1978, Mostacero et al. 569 (MO); Huascarán National Park; Quebrada Ulta, north side of valley; S-facing, moderate to gentle slopes, 09°07'S,077°32'W, 3930 m, 29 July 1985, Smith 11410 (MO, USM); Huascarán National Park. N-side of main valley, Quebrada Honda, 09°18'S, 077°25'W, 4200 m, 03 October 1985, Smith et al. 11641 (F, MO, USM); Huascarán National Park, mouth of Quebrada Ishinca, 09°23'S, 077°29'W, 3880 m, 15 February 1985, Smith et al. 9597 (F, MO, USM). Huaraz, Quebrada Quillcayhuanca, 4200 m, 30 October 1989, Arce & Martel 163 (MO); Quilcayhuanca, 09° 29'53.8"S, 77°24'59.6"W, 3831 m, 08 November 2014, Boza & Urquiaga 3022 (USM, Z); Lance, 4500 m, 04 June 2015, Boza &Urquiaga 3144 (USM, Z); Llanganuco, 09°04'47"S, 077°38'36"W, 4445 m, 03 June 2015, Boza & Urquiaga 3145 (USM, Z); 3146 (USM, Z); 3147 (USM, Z); Ulta, 09°06'S, 077°32'W, 4300 m, 07 June 2015, Boza & Urquiaga 3152 (USM, Z); 3153 (USM, Z); Llaca, 09°26'S, 077°26'W, 07 June 2015, Boza & Urquiaga 3154 (USM, Z); 3155 (USM, Z); Boza & Urquiaga 3156 (USM, Z); 28 May 1982, Cerrate 7696 (MO, USM); Comprado en la feria de plantas medicinales de Huaraz, 07 July 1988, Cerrate 9123 (USM); Huascarán National Park, Quebrada Shallap, 09°30'S, 077°24'W, 3900 m, 20 February 1985, Smith et al. 9709 (F, MO, USM). Huari, Llanganuco, 4366 m, 29 November 2007, Lasermann I12 (USM); Huascarán National Park, southside of Quebrada Carhuazcancha, 09°28'S, 077°15'W, 4200 m, 06 May 1986, Smith et al. 12255 (MO, USM); Huascarán National Park, Quebrada Pachachaca, a lateral valley of Quebrada Rurichinchay, 09°27'S, 077°16'W, 3860 m, 12 June 1986, Smith et al. 12542 (F, MO); Huascarán National Park, Quebrada de Yuraccocha, a lateral valley of Quebrada Rurichinchay, 09°22'S, 077°17'W, 4300 m, 16 June 1986, Smith et al. 12737 (MO, USM); Acopalca, 09°20'25"S, 077°12'19"W, 3300 m, 11 August 2010, Xue-Jun 194 (USM). Huaylas, Paron, 09°02'13"S, 77°43'52"W, 3357 m, 07 November 2014, Boza & Urquiaga 3016 (USM, Z); Huascarán National Park, Quebrada Parón, 09°01'S, 077°43'W, 3760 m, 08 May 1985, Smith 10606 (MO); Huascarán National Park, 09°00'S, 077°41'W, 4200 m, 29 September 1985, Smith 11537 (MO, USM); Huascarán National Park, Parón Valley, 09°00'S, 077°42'W, 4150 m, 01 January 1985, Smith & Goodwin 8924 (AAU, F, MO, USM); Huascarán National Park, Parón Valley, 09°01'S, 077°43'W, 3700 m, 01 January 1985, Smith & Goodwin 8939 (MO, USM); Huascarán National Park, western flank of Cordillera Blanca, Alpamayo–Cashapampa trail, 08°53'S, 077°45'W, 3950 m, 13 March 1985, Smith & Valencia 10013 (MO, USM); Huascarán National Park, lower slopes of Cerro Pakla, 08°49'S, 077°57'W, 4300 m, 09 April 1986, Smith et al. 12055 (AAU, F, MO, USM); Huascarán National Park, Quebrada Santa Cruz at base of and entering Quebrada Artizonraju, 08°55'S, 077°36'W, 4800 m, 16 January 1985, Smith et al. 9298 (F, MO, USM). Yungay, Ruta Vaqueria–Portachuelo, 3900 m, 05 November 1989, Arce 165 (MO); Huaytajirca, en el Dist. de Yanama, procedencia Matca (Yanama), 16 December 1989, Arce & Abilio 186 (MO); 30 km, hacia arriba, Parque Nacional de Huascaran, 3850 m, 10 March 1983, Beck 7914 (GOET, MO); Llanganuco, 09°03'04"S, 77°36'42"W, 3852 m, 07 November 2014, Boza & Urquiaga 3013 (USM, Z); Llanganuco encima de Yungay, 4000 m, 27 June 1966, Ferreyra 16860 (MO); Llanganuco arriba de Yungay, 4200 m, 14 December 1967, Ferreyra & Blount 18727 (GOET, MO, USM); Llanganuco, arriba de Yungay, 3900 m, 22 October 1965, Ferreyra & Tryon 16503 (MO, USM); slopes below Laguna de LLanganuco in quebrada de Llanganuco ca. 25 km above Yungay, just above and below the lake, 4100 m, 27 June 1966, Gabriel & Schunke 3826 (A, F); Dist. Yungay, Laguna de Llanganuco, 3800 m, 17 February 1968, Gutiérrez 249 (F); Laguna de Llanganuco, 3800 m, 19 February 1968, Gutiérrez 249-AGR (MO); Quebrada Llanganuco, cerca de la laguna y el albergue, 3850 m, 04 July 1981, Peréz 62 (USM); Laguna Llanganuco, 3400 m, 01 November 1984, Sagástegui & Dillon 12315 (F, MO); near Laguna Llanganuco, 09°03'54"S, 077°38'00"W, 4300 m, 14 August 2002, Schmidt-Lebuhn 507 (USM); near laguna Llanganuco, 09°03'54"S, 077°38'00"W, 4300 m, 14 August 2002, Schmidt-Lebuhn 510 (USM); Huascarán National Park, Lake Llanganuco, 09°05'S, 077°39'W, 3860 m, 16 August 1984, Smith 8210 (MO); Huascarán National Park, Llanganuco sector, Quebrada Demanada, side valley to Nevado Pisco, 09°02'S, 077°37'W, 4250 m, 13 April 1985, Smith & Cautivo 10302 (MO, USM); Huascarán National Park, Quebrada Ranincuray, 09°00'S, 077°33'W, 3850 m, 11 January 1985, Smith et al. 9049 (AAU, F, MO, USM); Huascarán National Park, Morococha at largest lake, 08°55'S, 077°35'W, 4550 m, 14 January 1985, Smith et al. 9215 (AAU, F, MO, USM); Llanganuco P. N. Huascarán, 09°07'00"S, 077°37'00"W, 3475 m, 07 August 2010, Xue-Jun 25 (USM). Cordillera Blanca near Ingenio in upper Pumapampa Valley, 11°04'S, 077°36'W, 4350 m, 15 February 1987, Boertmann 53 (AAU); Quebrada Ishinca, Cordillera Blanca, 09°23'S, 077°28'W, 3950 m, 23 August 1988, Frimer & Nielsen 101 (AAU); Quebrada Matará in Quebrada Ulta, Cordillera Blanca, 09°07'S, 077°32'W, 4250 m, 03 September 1988, Frimer & Nielsen 104 (AAU); Quebrada Ulta, Cordillera Blanca, 09°06'S, 077°32'W, 4050 m, 02 September 1988, Frimer & Nielsen 107 (AAU); 108 (AAU); Quebrada Rurichinchay, Cordillera Blanca, 09°21'S, 077°18'W, 4000 m, 06 Oct 1988, Frimer & Nielsen 118 (AAU); 123 (AAU); Quebrada Rurec, Cordillera Blanca, 09°25'S, 077°17'W, 3950 m, 11 October 1988, Frimer & Nielsen 125 (AAU); Frimer & Nielsen 126 (AAU); Quebrada Carhuasccancha. Cordillera Blanca, 09°29'S, 077°15'W, 3900 m, 15 October 1988, Frimer & Nielsen 132 (AAU); Querada Paron, Cordillera Blanca (W of Paron), 09°00'S, 077°41'W, 4150 m, 18 August 1988, Frimer & Nielsen 42; 43; 44; 45; 59 (AAU); Quebrada Ishinca, Cordillera Blanca, 09°23'S, 077°28'W, 3950 m, 23 August 1988, Frimer & Nielsen 70; 71; 73 (AAU); Quebrada Ishinca, Cordillera Blanca, 09°23'S, 077°28'W, 3950 m, 23 August 1988, Frimer & Nielsen 74; 99 (AAU); road from Yungay to Yauya, vicinity of Lagunas Llanganuco, 09°02'S, 077°35'W, 3800 m, 10 July 1982, Gentry et al. 37376 (MO, USM); Llanganuco, 4377 m, 29 November 2007, Lasermann II/1 (USM); Cordillera Blanca, Laguna Paron, 30 km NE of Caraz in northern Huascaran National Park, 4100 m, 10 October 1988, Peterson s.n (MO); Cordillera Blanca, East of Yungay, Laguna de Llanganuco, 3800 m, 05 April 1988, Renvoize & Lægaard 5066 (AAU); Cordillera Blanca. 35 km east of Yungay, 4000 m, 05 April 1988, Renvoize & Lægaard 5074 (AAU, MO); 5075 (AAU); 40 km east of Yungay., 4350 m, 05 April 1988, Renvoize & Lægaard 5088A; 5088B (AAU); Llanganuco Valley, 09°00'S, 077°30'W, 1500 m, August 1959, Tothill 174 (F); 3700 m, 1901–1929, Weberbauer 3229 (B, MO); Parque Nacional Huascarán. Llanganuco, 11 July 1982, Zardini 1535 (MO). La Libertad: Sánchez Carrión, señal Huayllides, 07°53'S, 078°02'W, 4100 m, 21 August 1982, Smith 2278 (MO, USM).
Peru. Junín: Concepción, Dist. de Andamarca, a 2.5 km de la localidad de Alhuay, 11°41'30"S, 74°54'01"W, 4150 m, 10 Oct 2017, H.R. Quispe M. 85 (holotype: CUZ!; isotypes: USM!, Z!).
Polylepis argentea T.Boza & H.R. Quispe A flower B inflorescence C upper leaflet surface D lower leaflet surface E bark F branching patterns (A, C, D, F Boza & Urquiaga 3036 B, E Quispe 85). Scale bars: 3 mm (A); 3 cm (B); 1 cm (C, D). Photographs A, C, F T. E. Boza E. B, E H. R. Quispe D E. G. Urquiaga F.
Trees 4–7 m tall. Leaves strongly congested at the branch tips, imparipinnate with 2 pairs of lateral leaflets, obtrullate in outline, (2.9)3.3–4.3 × (2.6–)3.3–4.3 cm; rachises densely sericeous, points of leaflet attachment with a tuft of long, straight hairs, sometimes with resin at leaflet insertion; stipular sheaths apically acute with spurs, densely sericeous on the outer surfaces; leaflets narrowly elliptic in outline, second pair from the terminal leaflet the largest, one of this pair (1.9)2.4–2.6 × 0.5–0.7 cm; margin entire, coriaceous, apically acute to slightly retuse, basally unequally cordate; upper leaflet surfaces almost glabrous with some hairs on the mid-veins to densely sericeous with silky hairs throughout; lower leaflet surfaces densely sericeous with silky hairs 0.6–0.9 mm long. Inflorescences pendant, 7.2–8.1 cm long, bearing 5–6(–9) flowers; floral bracts 4.5–5.6 mm long, narrowly triangular, densely sericeous on the outer surface; rachises sericeous. Flowers 7–9 mm diam.; sepals 3–4, ovate, green, densely sericeous outside; stamens 7–10, anthers orbicular, with a dense tuft of straight white hairs on the upper half; styles fimbriate, 2.7–4.4 mm long. Fruits turbinate, with variable numbers and placement of flattened spines, densely sericeous; 2.3–2.5 × 3.5–5.3 mm including spines. Diploid.
Polylepis argentea has been found in central Peru at La Mar (Ayacucho), Concepcion, Huancayo and Satipo (Junin) and La Convencion and Urubamba (Cusco) (Fig.
The estimated Extent of Occurrence (EOO) for P. argentea is 23,788 km2 and the Area of Occupancy is 40 km2. It is known from just eight locations, but several of these encompass forests of several square kilometers.
Polylepis argentea seems morphologically closest to P. sericea and P. canoi with which it shares similar lower leaflet surface hair type and density. The most obvious differences between P. argentea and these species is leaflet size, with P. argentea having leaflets of 1.9–2.6 × 0.5–0.7 cm, whereas P. canoi has leaflets of 2.4–3.9 × 0.8–1.5 cm and P. sericea of 1.8–2.1 × 0.8–1.0 cm. Further, P. argentea has shorter hairs (0.6–0.9 mm versus 1.3–1.7 mm) than P. canoi. In P. canoi, the hairs on the lower leaflet surfaces are yellowish and often most pronounced on the secondary veins, whereas in P. argentea, they are silky and more evenly distributed. Polylepis argentea has the upper leaflet surfaces with a few hairs on mid-veins whereas P. sericea has totally glabrous upper leaflet surfaces. Additionally, the inflorescence length and number of flowers per inflorescence differ between the species, with P. argentea having inflorescences 7.2–8.1 cm long with 5–9 flowers, compared with values of 3.3–4.5 cm and 9–15 flowers in P. sericea and 8.2–14.5 cm and 12–26 flowers in P. canoi. The three species can also be distinguished by the number of stamens and style length, with P. argentea having 7–10 stamens and styles 2.7–4.4 mm long, whereas the other two species have 13–15 stamens and styles 2.4–3.8 mm in P. canoi and 1.9–2.5 mm in P. sericea.
Polylepis argentea was first collected by B. Boyle during scientific expeditions carried out in 1997 and 1998 to the isolated Cordillera Vilcabamba where he recorded three species of Polylepis (
Peru. Ayacucho: La Mar, Dist. Tambo, Estera Community, sector Muyuorco, 12°54'19"S, 073°48'17"W, 3637 m, 29 June 2015, Boza 3036; 3096; 3097; 3098; 3099; 3100; 3101; 3102; 3103; 3104; 3105; 3106 (USM!, Z!). Cusco: La Convención, Dist. Huayopata Abra Málaga, 13°08'05"S, 072°19'18"W, 3802 m, 13 June 2015, Boza 3032; 3082; 3083; 3084 (USM!, Z!); Cuzco. Provincia La Convención, Bosque Qulcamachay, 4200 m, 01 October 2002, Palomino 2030 (QCA!); Dist. Huayopata, localidad Panticalle, Abra Málaga, 13°08'02"S, 072°19'32"W, 3725 m, 30 May 2006, Toivonen 84; 85; 86; 87 (CUZ!). Urubamba, Inkatambo, 13°18'06"S, 072°31'44"W, 4340 m, 01 September 2002, Arce s.n (USM!); Qésqa, 13°17'51"S, 072°24'57"W, 4000 m, 01 October 2002, Arce s.n (USM!); Abra Málaga, 13°08'43"S, 072°18'09"W, 4318 m, 01 October 2002, Arce s.n (CUZ!); Inkatambo 13°18'06"S, 072°31'44"W, 3840 m, 01 September 2002, Arce s.n (CUZ!); Dist. Ollantaytambo, Huaytampo, 13°10'47"S, 072°21'10"W, 3650 m, 07 November 2002, Calatayud 1035 (CUZ!, F!, MO!, USM!); Santuario Histórico Machu Pichu, camino Inca, Km 88–112, por puente Ruinas, 13°18'S, 072°07'W, 2000–4100 m, 20–21 June 1988, Núñez 9204 (MO!); Dist. Ollantaytambo, localidad Abra Málaga, 13°09'02"S, 072°18'09"W, 4230 m, 29 May 2006, Toivonen 15 (CUZ!); 16 (CUZ!); Dist. Ollantaytambo, localidad Huaytampo, 13°10'31"S, 072°21'03"W, 3800 m, 06 July 2006, Toivonen 95 (CUZ!); 96 (CUZ!). Junín: Concepcion, Andamarca, 11°41'30"S, 074°54'01"W, 2300 m, 14 June 2002, Martinez 18 (USM!); Dist. de Andamarca, a 2.5 km de la localidad de Alhuanay, 11°41'30"S, 074°54'01"W, 4150 m, 10 October 2017, Quispe 85 (CUZ!, USM!, Z!). Huancayo, Dist. de Santo Domingo de Acobamba, a 5 km de la localidad de Callanca, 11°45'43"S, 074°55'15"W, 4200 m, 12 October 2017, Quispe 87 (CUZ!, USM!, Z!). Satipo, Satipo/La Convencion Cordillera Vilcabamba Río Ene slope, near summit of divide, 11°39'30"S, 073°40'02"W, 3350 m, 07 June 1997, Boyle 4149 (USM!).
Peru. Cusco: La Convención, Cordillera del Vilcabamba, 30 km caminando de la Hacienda Luisiana y del Río Apurimac, 3400 m, 17 Jul 1968, T.R. Dudley 11180 (holotype: MO!; isotypes: NA, F!).
Trees 4–7(9) m tall. Leaves strongly congested at the branch tips, imparipinnate with 2–3(4) pairs of lateral leaflets, obtrullate in outline, (4.0–)7.9–9.4 × (4.2–)6.7–7.5 cm; rachises densely sericeous, points of leaflet attachment with a tuft of long, straight yellowish hairs, with ferruginous resin at leaflet insertion; stipular sheaths apically acute with spurs, glabrous in both surfaces; leaflets obovate in outline, second pair from the terminal leaflet the largest, one of this pair (2.4–)3.4–3.9 × (0.8–)1.1–1.5 cm; margin entire to slightly serrate with 4–6 teeth, coriaceous, apically slightly emarginate, basally unequally cordate; upper leaflet surfaces glabrous or with sparse sericeous hairs; lower leaflet surfaces densely sericeous with yellowish hairs 1.3–1.7 mm long. Inflorescences pendant, 8.2–14.5 cm long, bearing 12–17(26) flowers; floral bracts 7.0–15.8 mm long, narrowly triangular, densely sericeous on the outer surface; rachises sericeous. Flowers 7.8–11.2 mm diam.; sepals 3–4, ovate, green, densely sericeous outside; stamens 13–15, anthers orbicular, with a dense tuft of straight white hairs on the upper half; styles fimbriate, 2.4–3.8 mm long. Fruits turbinate, with variable numbers and placement of flattened spines, densely sericeous; 5.2 × 7.5 mm including spines. Diploid.
Polylepis canoi is distributed from the central-south-eastern Peruvian Andes to the central Bolivian Andes (Fig.
The EOO is estimated as 98,800 km2 and AOO as 84 km2. The species is known from 17 locations in Peru and Bolivia. In Peru, it has been categorized as EN (B1ab(iii)) (León-Yañez et al. 2006) and in Bolivia, as EN (B1ab(i,ii,iii)) (
Polylepis canoi seems morphologically closest to P. ochreata and P. sericea. However, it has obovate and larger (2.4–3.9 × 0.8–1.5 cm) leaflets than the other two species, which have elliptic and smaller (1.8–2.7 × 0.5–1.0 cm) leaflets. Additionally, P. canoi has longer hairs (1.3–1.7 mm) than the other two species (0.7–1.2 mm).
This species was treated as endemic to Peru by
Bolivia. Cochabamba: Chapare, Mayka Mayu, 17°12'S, 065°58'W, s.d., Hensen 2248 (BOLV, LPB, MO!, TEX); Maycamayu, ca. 60 Km N Sacaba, 17°12'S, 065°58'W, 3300 m, 11 August 1991, Kessler 2874 (GOET!); 2875 (GOET!); 2877 (GOET!); 2878 (AAU!); 2879 (GOET!, MO!); 2880 (GOET!). La Paz: Bautista Saavedra, Area Natural de Manejo Integrado Apolobamba, bajada de Waricunca, mas allá de Chaka, por el antiguo camino Sorapata-Apolo, 14°53'19"S, 068°47'04"W, 3550 m, 28 March 2009, Fuentes 13589 (BOLV, LPB, MA, MO!, USZ); Area Natural de Manejo Integrado Apolobamba, sector Chaka, bosque continuo al SE del campamento cerca de la cueva, por el antiguo camino Laji Sorapata-Apolo, 14°53'32"S, 068°47'12"W, 3461 m, 30 March 2009, Fuentes 13634 (LPB, MO!, QCA!, USZ); 13639 (BOLV, LPB, MO!, QCA!, USZ); Área Natural de Manejo Integrado Apolobamba. Bajada de Wuaricunca, más allá de Chaka, por el antiguo camino Hilo-Hilo – Apolo, 14°53'11"S, 068°47'04"W, 3550 m, 06 April 2009, Fuentes 13897 (BOLV, LPB, MA, MO!, QCA!, USZ); Area Natural de Manejo Integrado Madidi, Hilo Hilo. Sobre el Río Tumamayu en la localidad de Laji Sorapata, 14°53'14"S, 068°51'52"W, 4182 m, 10 April 2009, Loza 635A (LPB, MA, MO!); 645 (LPB, MO!, QCA!, USZ); Área Natural de Manejo Integrado Apolobamba, Hilo Hilo, Juchuy Queñua a medio día de Laji Sorapata, 14°54'52"S, 068°48'08"W, 3879 m, 16 April 2009, Loza 757 (LPB, MO!); 775 (BOLV, LPB, MA, MO!, USZ); 788 (LPB, MO!, QCA!, USZ); Chaka Machay(Laji), 14°53'S, 068°47'W, 3300 m, 14 September 2002, Zenteno 1507 (LPB). Franz Tamayo, Área Natural de Manejo Integrado Apolobamba, Keara bajo, 14°42'09"S, 069°04'35"W, 3500 m, 21 November 2007, Araujo 4078 (LPB, MO!); Área Natural de Manejo Integrado Apolobamba, Hilo Hilo, Chaka, sobre la senda hacia Amantala, 14°53'16"S, 068°47'16"W, 3576 m, 16 August 2009, Cayola 3417 (BOLV, LPB, MA, MO!, USZ); Parque Nacional Madidi, entre Queara y Mojos, sector Mosquito, 14°39'37"S, 068°57'54"W, 3400 m, 26 February 2008, Fuentes 12028 (BOLV, LPB, MO!, QCA!, USZ); Parque Nacional Madidi, Puina Viejo, ca. 3 km río abajo por camino al W del río, 14°34'58"S, 069°06'24"W, 3316 m, 21 June 2005, Fuentes 8549 (LPB, MO!); Parque Nacional Madidi, Hilo Hilo, arriba de la mina Kanupata en la localidad de Laji Sorapata, 14°52'28"S, 068°51'15"W, 4182 m, 11 April 2009, Loza 671 (BOLV, HSB, LPB, MA, MO!, NY, QCA!, USZ); Bosque de Queñuari, 14°54'31"S, 069°01'07"W, 4275 m, 28 September 2006, Palabral 489 (LPB); Senda Pelechuco-Mojo, sector Tambo Quemado, a media hora del campamento siguiendo senda Pelechuco Moxos, 14°41'03"S, 068°58'22"W, 3455 m, 01 May 2003, Paniagua 5710 (LPB, MA, MO!). Larecaja, bosque de a localidad de Hirola, pasando Lipichi, 15°26'41"S, 068°10'57"W, 3881 m, 05 November 2008, Palabral 705 (LPB). Murillo, 8 km after Palca on the road to Iquico, 4000 m, 10 November 1967, Vuilleumier 342 (MO!).
Peru. Cusco: La Convención, Cordillera de Vilcabamba, above Camp 7, ca. 30 km walking distance from Hacienda Luisiana and the Apurimac River, 12°30'S, 074°30'W, 3400 m, 17 July 1968, Dudley 11180 (F!, MO!, NA); usually on eastern slopes ca. 30 km walking distance NE from Hacienda Luisiana and the Apurimac River, 12°30'S, 073°30'W, 3400 m, 19 July 1968, Dudley 11221 (F!, USM!). Junín: Jauja, Dist. Molinos, Comunidad Curimarca, Jucha, 11°33'53"S, 075°18'58"W, 3893 m, 10 November 2016, Ames s.n (Z!). Satipo, Dist. de Pampa Hermosa, Comunidad de Toldopampa, Tasta, 11°26'08"S, 074°53'58"W, 3754 m, 04 October 2016, Ames s.n (Z!); Junin/Cusco Prov. Satipo/La Convención, Cordillera Vilcabamba. Río Ene, slope near summit of divide, 11°39'30"S, 073°40'02"W, 3350 m, 07 June 1997, Boyle 4151 (USM!). Puno: Limbani, Huancasayani, on road to Limbani just east of Abra Aricoma, 14°13'S, 069°42'W, 3750 m, 28 March 1987, Boertmann 129 (AAU!, QCA!); Huancasayani between Abra Aricoma and Limbani, 14°13'S, 069°42'W, 3750 m, 28 March 1987, Brandbyge 511 (AAU!).
This species differs from Polylepis quadrijuga
Colombia. Antioqui: Urrao, Páramo Frontino, 06°30'N, 76°07'W, 3400 m, 5 Sep 2000, J.A. Perez & N. Parra 1477 (holotype: MEDEL!)
Trees 4–8 m tall. Leaves only slightly congested at the ends of the branches, imparipinnate with 3–4(5) pairs or lateral leaflets, obtrullate in outline, (2.5–)3.3–5.2 × (1.8–)2.3–3.5 cm; rachises villous, point of leaflet attachment with a tuft of long, straight hairs, slightly resinous, stipular sheaths acute at the apex with spurs, densely sericeous on the outer surface; leaflets obovate in outline, second pair from the terminal leaflet the largest, one of this pair (1.1–)1.4–2.0 × 0.4–0.8 cm; margin serrate with 5–6 teeth, coriaceous, apically slightly emarginate, basally unequally cordate; upper leaf surfaces glabrous with few trichomes in the mid-vein depression; lower surfaces densely villous with hairs 1.4–1.8 mm. Inflorescences pendant, 6.3–10.6 cm long, bearing 7–15 flowers; floral bracts 4.5–5.4 mm long, narrowly triangular, sparsely villous on the outer surface; rachises sparsely villous. Flowers 7.5–8.2 mm diam.; sepals 3–4, ovate, densely villous outside; stamens 9–11; styles fimbriate, 2.8–3.2 mm long. Fruits turbinate, with variable number of spines, densely villous; 3.3–3.6 × 4.7–5.6 cm including spines. Diploid.
Polylepis frontinensis occurs in north-western Colombia in the Páramo Frontino, also called Páramo del Sol (Fig.
This species is named after the Páramo Frontino to which its distribution appears to be restricted.
Polylepis frontinensis is restricted to the upper humid montane cloud forest limit in the Páramo Frontino. Its estimated extent of occurrence (EOO) and area of occupancy (AOO) are 24 km2. The area of distribution of the species is largely unprotected, except for the private Reserva Colibrí del Sol which only includes a few hundred individuals of this species (M. Kessler, pers. obs.). In addition, there is evidence of clearance of Polylepis forests elsewhere in the páramo (
The populations of Polylepis from Páramo Frontino have previously been identified either as P. quadrijuga or P. sericea (e.g.,
Considering the morphological intermediacy of P. frontinensis with P. quadrijuga and P. sericea, we hypothesize that this species might be of hybridogenic origin between members of section Sericeae subsection Sericeae and section Reticulatae.
Colombia. Antioquia: Urrao, Vereda El Chuscal, sector Alto de las campanas, hacia La Laguna Campanas, 06°27'46"N, 076°07'37"W, 3830 m, 20 June 2013, Alvarez 84 (HUA!, MO!); Páramo del Sol, Sector Alto del Burro, 3600 m, 17 April 2011, Alzate 4168 (HUA!); Chical, Reserva ProAves “Colibrí del Sol” Páramo frontino, 06°26'22"N, 076°05'47"W, 3300 m, 04 February 2015, Kessler 14772; 14773; 14774; 14775; 14776; 14777 (Z!); Páramo de Frontino, Llano Grande, 3460 m, 06 January 1984, Londoño 51 (HUA!,MEDEL); Páramo de Frontino, 06°30'N, 076°07'W, 3400 m, 05 September 2000, Perez & Parra 1477 (MEDEL); Páramo de Frontino, 3450 m, 22 September 1994, Renteria 10555 (HUA!); Páramo de Frontino, Zona situada entre el 15 y la Esperanza, 2980–3680 m, 18 May 1985, Renteria 4038 (HUA!); Páramo de Frontino, sitio Llano grande, 06°27'24"N, 076°07'22"W, 3380 m, 10 September 1986, Roldán 315 (COL!, MO!, HUA!); Páramo frontino. Camino de Puente Largo al cerro Cuchilla de Frontino, 06°30'N, 076°07'W, 3600–3800 m, 19 July 1995, Sánchez 2244 (COL!, MEDEL); Páramo de Frontino. Camino entre Puente Largo y Llano Grande, 06°30'N, 076°07'W, 3550–3600 m, 20 July 1995, Sánchez et al. 2289 (MEDEL); Páramo El Sol, Vereda La Encarnación, 06°29'12"N, 076°06'33"W, 3518 m, 24 May 2014, Sarrazola 699 (HUA!). Páramo Frontino, near Llano Grande, 3450 m, 26 October 1976, Boeke 235 (MEDEL, MO!).
Ecuador. Chimborazo: Lagunas de Atillo, 02°08'S, 78°34'W, 3465 m, 17 Dec 2019, K. Romoleroux, T.E. Boza E. & E. Bastidas 6199 (holotype: QCA!; isotype: Z!).
Trees 4–12 m tall. Leaves strongly congested at the branch tips, imparipinnate with 3–4 pairs of the lateral leaflets, obtrullate in outline, 4.5–6.3 × 3.4–4.3 cm; rachises densely sericeous, points of leaflet attachment with a tuft of long, straight whitish hairs; stipular sheaths apically acute with spurs, densely sericeous in the upper surface; leaflets elliptic in outline, second pair from the terminal leaflet the largest, one of this pair 1.8–2.8 × 0.6–0.9 cm; margin entire, apically emarginate, basally unequally cordate; upper leaflet surfaces glabrous; lower leaflet surfaces densely sericeous with whitish hairs 0.2–0.4 mm. long. Inflorescences pendant, 13.0–17.9(–20.4) cm long, bearing 23–29 flowers; floral bracts 9.3–11.1 mm long, narrowly triangular, densely sericeous on the outer surface; rachises glabrous. Flowers 7.4–8.4 mm diam.; sepals 4, ovate, green, densely sericeous outside; stamens 9–15, anthers orbicular, with a dense tuft of straight white hairs on the upper half; styles fimbriate, 1.9–2.9 mm long. Fruits turbinate, with variable numbers and placement of flattened spines, densely sericeous; 3.3–5.1 × 3.1–7.4 mm including spines. Diploid.
Polylepis humboldtii is restricted to Chimborazo Province in Ecuador (Fig.
The AOO is estimated as 4 km2 and it has been collected at only two locations in Ecuador. Although it is protected within Sangay National Park, burning of the páramo grassland matrix likely affects the remaining Polylepis forest patches. Therefore, we assess P. humboldtii as Critically Endangered (B2a, C2).
Polylepis humboldtii seems morphologically closest to P. sericea with which it shares similar leaflet shape, margin, apex and upper and lower leaflet surfaces hairs type and density. The most obvious differences between these species are leaflet hair length, with P. humboldtii having shorter hairs than P. sericea (0.2–0.4 mm versus 0.7–1.0 mm) and longer inflorescences (13.0–20.4 cm) with more flowers (23–29) than P. sericea (3.3–4.5 cm, 9–15 flowers). Additionally, P. humboldtii occurs in central Ecuadorean Andes, whereas P. sericea is distributed from western Venezuela to central Colombia.
Ecuador. Chimborazo: Alausí, Achupallas, alrededores, 2°17'S, 78°39'W, 3300 m, 11 July 2013, Caranqui 2565 (QCA!); Lagunas de Atillo, 2°8'S, 78°34'W, 3465 m, 13 April 2009, Carate et al. 184; 185; 188 (QCA!).
Ecuador. Loja: Laguna Chinchilla, 03°36'20"S, 079°23'08"W, 3610 m, 21 Dec 2019, T.E. Boza E. & C. Medina 3185 (holotype: QCA!; isotypes: Z!, CUZ!).
Trees 4–10 m tall. Leaves strongly congested at the branch tips, imparipinnate with 3–4(–5) pairs of the lateral leaflets, obtrullate in outline, 2.6–3.6 × 2.1–3.2 cm; rachises densely sericeous, points of leaflet attachment with a tuft of long, straight whitish hairs; stipular sheaths apically truncate, densely sericeous in the upper surface; leaflets narrowly to broadly obovate in outline, second pair from the terminal leaflet the largest, one of this pair 1.2–1.6 × 0.5–0.8 cm; margin serrate at apex with 3–4 teeth, apically emarginate, basally unequally cordate; upper leaflet surfaces glabrous with few hairs on mid-depression; lower leaflet surfaces densely sericeous with whitish silky hairs 0.2–0.6 mm long. Inflorescences pendant, (3.5–)4.3–10.5(–12.2) cm long, bearing 9–27 flowers; floral bracts 5.5–6.3 mm long, narrowly triangular, densely sericeous on the outer surface; rachises densely sericeous. Flowers 4.4–5.2 mm diam.; sepals 4, ovate, green, densely sericeous outside; stamens 7–9, anthers orbicular, with a dense tuft of straight white hairs on the upper half; styles fimbriate, 1.7–2.0 mm long. Fruits turbinate, with variable numbers and placement of flattened spines, densely sericeous; 1.7–3.0 × 1.4–1.5 mm including spines. Diploid.
Polylepis loxensis is restricted to south-western Ecuador (Fig.
Polylepis loxensis is known from six locations with an EOO of 728 km2 and an estimated AOO of 32 km2. No conservation actions have been taken to date. The area is heavily grazed by cattle and horses, pine plantations occupy large extensions and a large proportion of the area is under gold mining concessions (
Polylepis loxensis is most similar to P. ochreata, with which it shares the emarginate leaflet apices and subcordate leaflet bases and similar dense, short, white silky hair on the lower leaflet surfaces. Indeed, they were treated as conspecific by
Ecuador. Azuay: Nabón, 3°28'20"S, 79°02'24"W, 2800–3300 m, 15 November 2008, Salgado 1419 (LOJA!); Loja: Loja, Fierro Urco, 03°36'20"S, 079°23'08"W, 3610 m, 19 December 2019, Boza & Medina 3184 (QCA!, Z!, CUZ!); Fierro Urco, Saraguro-Loja, km 12.4 turnoff towards Fierro Urco, km 23.8, 03°43'10"S, 079°19'18"W, 3840 m, 6 December 1994, Jørgensen et al. 1240 (AAU!, LOJA!, MO!); road San Lucas–Saraguro, km 9, turn off to Fierro Urco, 11 km to the pass, 03°43'03"S, 079°19'25"W, 3630 m, 4 November 2000, Jørgensen et al. 2228 (QCA!); ca. 10 km along road to Fierro Urco, 03°41'S, 079°01'W, 2850 m, 8 September 1998, Laegaard 19109 (AAU!, LOJA!, QCA!); Fierro Urco, grass Páramo 12 km to the left (northbound) from the Panamericana highway, 03°43'S, 079°19'W, 3600–3650 m, 9 June 1999, Sklenár & Laegaard 7096 (AAU!, GOET!); ca. km 12 along Páramo road to Fierro Urco, 03°41'S, 079°01'W, 3650 m elev., 9 June 1999, Laegaard & Sklenár 20279 (AAU!, LOJA!, QCA!); Páramo of Fierro Urco SW of Saraguro, 03°43'S, 079°19'W, 3500 m elev., 21 November 1996, Lewis et al. 2121 (AAU!); road Loja-Cuenca, km 50, track to Fierro Urco, km 5–7, 03°41'S, 079°17'W, 3150–3350 m elev., 25 October 1996, Lewis & Lozano2724 (AAU!,LOJA!, MO!, QCA!); road Loja-Saraguro, km 52, track to Fierro Urco, km 10, 03°42'S, 079°18'W, 3350–3450 m elev., 17 January 1997, Lewis et al. 2932 (AAU!, LOJA!, MO!); road Loja–Saraguro, 8.5 km N of San Lucas, track to Fierro Urco, km 11, 03°43'10"S, 079°19'18"W, 3550 m elev., 15 January 1998, Lewis & Hughes 3804 (AAU!, LOJA!, MO!, QCA!); Fierro Urco, 03°41'S, 079°22'W, 3700 m elev., 11 January 1995, P. Lozano 172 (LOJA!); Saraguro, Laguna Chinchilla, 03°36'20"S, 079°23'08"W, 3610 m elev., 21 December 2019, Boza & Medina 3186 (QCA!, Z!, CUZ!), cerro Chinchilla, parroquía Celén, 03°35'44"S, 079°20'17"W, 3000 m elev., 19 September 1984, Jaramillo 7332 (QCA!), 7335 (GB, QCA!); Laguna Chinchilla, 03°36'17"S, 079°23'49"W, 11 November 2008, Salgado et al. 1392; 1394 (LOJA!).
Polylepis ochreata var. integra Bitter, Bot. Jahrb. Syst. 45: 598, fig. 4. 1911. Type. Ecuador. Imbabura: Volcan Mojanda, Mar. 1901, Sodiro s.n. (holotype: FI n.v.; isotype: GOET!).
Polylepis subintegra Benoist, Bull. Soc. Bot. France 81: 326. 1934. Type. Ecuador. Pichincha: W slopes of Cerro Pichincha, Taurichupa, 4000 m, 28 Nov 1930, Benoist 3356 (holotype: P!).
Acaena ochreata Wedd., Chlor. And. 2: 240. 1855.
Ecuador. Pichincha: W slopes of Cerro Pichincha, 3600 m, May 1856, Jameson 73 (lectotype, designated by
Trees 2–10 m tall. Leaves strongly congested at the branch tips, imparipinnate with 4–7 pairs of the lateral leaflets, obtrullate in outline, (3.9–)4.4–7.0 × 2.9–4.7 cm; rachises glabrous to densely sericeous, points of leaflet attachment with a tuft of long, straight whitish hairs; stipular sheaths apically acute, glabrous to sparsely sericeous (adult) or densely sericeous (juvenile) in the upper surface; leaflets narrowly elliptic in outline, second pair from the terminal leaflet the largest, one of this pair 1.6–3.0 × 0.5–0.7 cm; margin entire to slightly serrate with 4–6 teeth, coriaceous, apically emarginate, basally unequally cordate; upper leaflet surfaces glabrous; lower leaflet surfaces densely sericeous with whitish silky hairs 1.3–2.0 mm long in juvenile plants and 0.3–0.5 mm long in adult plants. Inflorescences pendant, 8.1–15.5(–17.4) cm long, bearing (21–)26–49 flowers; floral bracts 5.9–12.8 mm long, narrowly triangular, densely sericeous on the outer surface; rachises sericeous. Flowers 6.6–9.0 mm diam.; sepals 4, ovate, green, densely sericeous outside; stamens 9–13, anthers orbicular, with a dense tuft of straight white hairs on the upper half; styles fimbriate, 2.1–2.6 mm long. Fruits turbinate, with variable numbers and placement of flattened spines, densely sericeous; 4.7–7.5 × 6.1–7.9 mm including spines. Diploid, tetraploid and hexaploid.
Polylepis ochreata (Wedd.) Bitter A flowering branch B fruit C lower leaf surface D upper leaf surface E stipular sheaths F young leaves (A Romoleroux 350 B Laegaard 55665 C Romoleroux 5413 D Romoleroux 300 E Clark 5820 F Ericksen 59086). Scale bars: 6 cm (A); 6 mm (B); 4 cm (C, D). Photographs by E. G. Urquiaga F.
Polylepis ochreata is distributed in the Andes of Ecuador and in Nariño, southernmost Colombia (Fig.
The EOO is estimated at 7,525 km2 and AOO at 112 km2. The species is known from 16 locations. In Colombia, remnants of P. ochreata forests are under pressure by the expansion of potato cropland, so that the Cumbal population has been assessed as EN and Chile’s population as VU (
Described by Bitter in 1911, this species was synonymized under P. sericea by
Ecuador. Bolívar: Guaranda, Parroquia Salinas, recorrido entre los Arrayanes y Pambabuela, 01°22'06"S, 079°03'47"W, 3615 m, 10 Febrero 2005, Vargas López 4696 (AAU!, K, MO!, QCNE, US!). Carchi: Cumbal, 00°48'19"N, 077°53'03"W, 3500 m, Bensman 418 (MO!, WIS); Km 31 west of Tulcán on road to Maldonado, 00°52'N, 077°55'W, 3900 m, 21 June 1984, Todzia 2485 (MO!). La Libertad (Alizo), 00°45'N, 077°59'W, Asplund 17037 (S); Páramos de El Angel S of Volcán Chiles, 00°45'N, 077°58'W, 3850 m, 14 March 1985, Eriksen 59086 (AAU!, MO!). Maldonado, Volcán Los Chiles, along road 9 km W of Tufiño, 00°49'N, 077°57'W, 3500 m, 10 March 1992, Lægaard 101661 (AAU!, GOET, QCA!); Tufiño, Road Tulcán-Maldonado, near Volcán Chiles, 00°48'N, 077°56'W, 3850–4000 m, 16 August 1985, Lægaard 54966 (AAU!, MO!, QCA!); S slopes of volcán Chiles, 14–16 km W of Tufiño on road to Maldonado, 0–1 km S of the road, 00°47'N, 077°57'W, 3850–3900 m, 18 January 1988, Molau 2536 (AAU!, GB, MO!, QCA!); a 33 km de Tulcán, 00°48'N, 077°54'W, 3900 m, Romoleroux 173 (AAU!, QCA!); Carretera Tulcán-Tufiño-Maldonado, 00°47'N, 077°57'W, 3800–3900 m, 12 October 1986, Romoleroux 189 (AAU!, QCA!); Tulcán, 33.4 km W of Tulcán on road to Maldonado, Páramo de Chilos on Colombia border, 00°48'19"N, 077°53'03"W, 3900 m, 22 September 1979, Gentry 26342 (AAU!, MO!, QCA!). Cotopaxi: Toacaso, Quebrada Faldiguera, 00°41'S, 078°45'W, 3750 m, 16 February 1991, Jørgensen 93000 (AAU!, MO!, QCA!). Imbabura: Gonzalez Suarez, Laguna Mojanda, camino, forêt d’altitude, 00°08'N, 078°15'W, 2500 m, 01 February 1996, Billiet 6762 (BR, MO!). La Merced de Buenos Aires: at road Chauasqui–Merced de Buenos Aires, km 20, near pass, 00°33'N, 078°17'W, 3700–3850 m, 10 December 1984, Lægaard 53475 (AAU!, MO!, QCA!). Otavalo: forested path to Laguna Mojanda (La via antigua a Mojanda por el cementerio), 00°10'00"N, 078°15'00"W, 3800 m, 31 December 2000, Clark 5820 (QCA!, US). San Rafael, W slopes of Volcán Cayambe, 00°10'00"N, 078°15'00"W, 3700–3900 m, 27 July 1967, Sparre 17789 (AAU!, S). Napo: Nono, N-side of Volcán Pichincha above Hacienda Yanacocha, 00°07'S, 078°34'W, 3950–4050 m, 02 June 1985, Lægaard 54457 (AAU!, MO!, QCA!). Pichincha: along, Northern slopes of Cerro Corazón, 2–4 km W along on the road to Hacienda El Pongo, 00°28'S, 078°36'W, 3100–3200 m, 13 May 1979, Holm-Nielsen 18007 (AAU!, MO!); Corazón, 00°31'53"S, 078°39'36"W, 3260 m, Sodiro s.n. (AAU!); Lloa, Volcán Atacazo, W slope, 17 km from San Juan, 00°20'S, 078°38'W, 2850 m, 25 August 1980, Holm-Nielsen 25115 (AAU!); 25148 (AAU!); Volcán Atacazo, SW slope, km 19 from San Juan, 00°21'S, 078°39'W, 2900 m, 25 August 1980, Holm-Nielsen 25169 (AAU!); West-side of Volcán Atacazo, along drinkwater canal, 00°20'S, 078°38'W, 3700–3750 m, 11 August 1984, Lægaard 52639 (AAU!, MO!, QCA!); 52641 (AAU!); along drinkwater-canal on W-side of Atacazo, ca. 5 km S of Campamento, 00°20'S, 078°38'W, 3700–3800 m, 24 October 1984, Lægaard 53256 (AAU!); along drinkwater-canal on W-side of Atacazo, ca. 5 km S of Camparmento, 00°20'S, 078°38'W, 3750 m, 28 October 1984, Lægaard 53259 (AAU!); 53260 (AAU!); along drinkwater-canal on W-side of Volcan Atacazo, 00°20'S, 078°38'W, 3200 m, 24 November 1985, Lægaard 55665 (AAU!, GOET, MO!, QCA!); Volcán Atacatzo, 00°20'S, 078°37'W, 3500 m, Mille 364 (US); carretera Quito–San Juan–San José de la Victoria, 00°17'53"S, 078°38'20"W, 2900–3400 m, 24 December 1987, Zak 3265 (AAU!, GB, MO!); Nono, Camino Yanacocha NW of Volcan Pichincha, 00°05'S, 078°33'W, 3200–3800 m, 03 October 1981, Balslev 2049 (AAU!, MO!, NY, QCA!); 28 November 1930, Benoist 3356 (P); Yanococha, faldas noroccidentales, 00°07'S, 078°35'W, 22 March 1987, Jaramillo Asanza 9573 (AAU!, NY, QCA!); 9588 (AAU!, QCA!); N-side of Volcán Pichincha above Hacienda Yanacocha, 00°07'S, 078°34'W, 3800 m, 04 June 1985, Lægaard 54458 (AAU!, MO!); 54459 (AAU!); 54462 (AAU!, QCA!); 54463; 54467 (AAU!, MO!); 54474; 54476; 54477 (AAU!, MO!, QCA!); Carretera Quito-Nanegalito-Santa Ana del Tablón, desvío Hda Yanacocha km 1–10 desde el desvío, 00°07'S, 078°34'W, 3500–3600 m, 06 December 1992, Romoleroux 1495A (AAU!); Yanacocha, 3617 m, 28 November 2008, Romoleroux 5342 (QCA!); Yanacocha, sector La Despensa, 00°07'52"S, 078°35'06"W, 3837 m, 14 Febrero 2009, Romoleroux 5413 (MO!, QCA!); Reserva Yanacocha, Trocha “Inca” 1–600 m, 00°06'44"S, 078°34'24"W, 3536 m, 11 June 2011, Ulloa Ulloa 2171 (MO!, QCA!); carretera Quito-Nono-Tandayapa, desviación a Yanacocha en la localidad de Guanto-Pugro, en la hacienda “Alto Perú”, estribaciones N.O. del Volcán Pichincha, 00°05'S, 078°35'W, 3200–3300 m, 17 November 1987, Zak 2946 (AAU!, GB, MO!); Quito, SW-slopes of volcan Atacazo, 00°20'S, 078°35'W, 3650 m, 11 October 1984, Brandbyge 42817 (AAU!, MO!, QCA!); SW-slopes of volcán Atacazo, 00°20'S, 078°35'W, 3700–3800 m, 28 October 1984, Brandbyge 42837 (AAU!, MO!, QCA!); Volcán Pichincha, N slopes, road to Hda. Yanacocha from pass on Quito-Nono road, km 7–11.2, 00°07'S, 078°33'W, 3600–3500 m, 12 October 1991, Øllgaard 99187 (AAU!); Carretera a San Juan-Atacazo, km 1–12, 00°20'S, 078°35'W, 3700–4000 m, 02 September 1990, Romoleroux 1060 (AAU!, QCA!); Tocachi, 00°08'N, 078°16'W, 3260 m, Asplund 17103 (S); 00°08'N, 078°16'W, Benoist 4549 (S); NW side of Pichincha, 00°08'N, 078°16'W, Fagerlind s.n (S); 00°08'N, 078°16'W, Holmgren 664 (S); 00°08'N, 078°16'W, Jameson s.n (MO!); Páramo de Mojanda, at Laguna Negra and S-side of Laguna Grande, 00°08'N, 078°16'W, 3800 m, 14 May 1985, Lægaard 54316A (AAU!, QCA!); 00°08'N, 078°16'W, Romoleroux 1495 (AAU!, QCA!); 243 (QCA!); 245 (NY, QCA!); 305 (QCA!); 00°08'N, 078°16'W, 3700 m, Romoleroux 350 (QCA!).
Polylepis hypargyrea Bitter, Bot. Jahrb. Syst. 45: 600. 1911. Type. Venezuela. Páramo de la Culata, Sierra Nevada Moritz 1120 (holotype: B destroyed; isotypes: BM!; photos at F!, GH!).
Polylepis quindiensis Cuatrecasas, Revista Acad. Colomb. Ci. Exact. 4: 343 .1941. Type. Colombia. Caldas: Cordillera Central, W of Macizo del Quindio, Nevado del Ruiz, 3400–3500 m, 5 May 1940, Cuatrecasas 9327 (holotype: COL!; isotypes: BC!,US!).
Venezuela. Mérida: Sierra Nevada, 3500 m, Jun 1847, Funck & Schlim 1546 (lectotype, designated by
Trees 3–7(12) m tall. Leaves strongly congested at the branch tips, imparipinnate with 2–3(–4) pairs of lateral leaflets, obtrullate in outline, 3.9–4.2 × 2.5–3.8 cm; rachises glabrous, points of leaflet attachment with a tuft of long, straight whitish hairs; stipular sheaths apically acute with spurs, almost glabrous with some hairs at the edges on the outer surfaces and glabrous in the inner surfaces; leaflets elliptic in outline, second pair from the terminal leaflet the largest, one of this pair 1.8–2.1 × 0.8–1.0 cm; margin entire, coriaceous, apically emarginate to retuse, basally unequally cordate; upper leaflet surfaces glabrous; lower leaflet surfaces densely sericeous with whitish hairs 0.7–1.0 mm long. Inflorescences pendant, 3.3–4.5 cm long, bearing 9–15 flowers; floral bracts 4.1–6.4 mm long, narrowly triangular, densely sericeous on the outer surface; rachises sericeous. Flowers 4.2–8.1 mm diam.; sepals 4, ovate, green, densely sericeous outside; stamens 13–15, anthers orbicular, with a dense tuft of straight white hairs on the upper half; styles fimbriate, 1.9–2.5 mm long. Fruits turbinate, with variable numbers and placement of flattened spines, densely sericeous; 4.0–7.4 × 3.4–9.6 mm including spines. Diploid.
Polylepis sericea is found in two distinct geographic areas, the Cordillera de Mérida in the Andes of western Venezuela and the Cordillera Central of Colombia in Caldas, Quindio and Risaralda Departments (Fig.
The estimated Extent of Occurrence (EOO) for Polylepis sericea is 36,560 km2. The Area of Occupancy (AOO) is 100 km2. The species is known from 16 locations. It is protected in Venezuela within the Sierra Nevada and Sierra de la Culata National Parks, with some minor relicts in the highest areas of the Trujillo State, where more than 50% of the remnant forest of P. sericea are conserved (
In her seminal taxonomic revision of the genus Polylepis,
As defined by
Colombia. Caldas: Pereira, El Cisne, Laguna del Otúm, 04°46'N, 075°25'W, 3900–4200 m, 20 March 2009, Vargas 20063 (COL!). Villamaría, Cordillera central, vertiente occidental; cabeceras del río Otún, Laguna del Mosquito y plan del Villar, 04°58'N, 075°21'W, 3650–3750 m, 26 November 1946, Cuatrecasas 23257 (COL!); Cordillera Central, vertiente occidental, vert. sudoeste del Ruiz, El Prisco, páramos, 04°58'N, 075°22'W, 3500–3600 m, 05 May 1940, Cuatrecasas 9327 (COL!). Quindío: Salento, Vereda Cocóra; below Nevado del Quindio, 3800 m, 20–22 May 1989, Luteyn 12974 (MO!). Risaralda: Pereira, Cordillera central, en el paso de la Laguna del Otúm hacia la Quebrada Africa, 04°47'N, 075°24'W, 4300 m, 09 February 1980, Jaramillo 6276 (COL!).
Venezuela. Lara: Morán, Páramo del Jabon (Vertiente Oriental), 09°34'N, 070°06'W, 3100–3400 m, 02 November 1969, Cuatrecasas 28216 (MERF); Páramo Jabón, camino al páramo Cendé, 09°34'N, 070°06'W, 3000–3200 m, 30 December 1999, Riina 1036 (VEN). Mérida: Caracciolo Parra Olmedo, Páramo La Culata en quebrada, 08°46'43"N, 071°03'04"W, 3581 m, 07 October 2006, Bonifacino 2541 (VEN). Justo Briceño, Páramos de Laguna Grande, 08°48'N, 070°56'W, 21 January 1929, Pittier 13253 (MO!, VEN). Libertador, Parque Nacional Sierra Nevada. Loma Redonda Teleferico station and south, 08°33'N, 071°05'W, 4068 m, 20 May 1988, Dorr 5220 (AAU!); Pico Bolivar, 08°33'N, 071°02'W, 4200 m, 17 January 1968, Walter 443 (GOET!). Miranda, carretera hacia Piñango, Páramo Piedras Blancas, Dtto. Rangel, 09°00'N, 070°50'W, 3700 m, 03 March 1982, Aymard 1050 (MO!); Dist. Justo Briceño. Páramo y chirivital en la vertiente NW del Alto del Totumo, hoya del Río Chirurí, a 19.5 km de El Aguila por la carretera a Piñango, 08°51'N, 070°49'W, 3900–4000 m, 02 April 1982, Berry 3812 (MO!); 3844 (MO!, VEN); de El Aguila a Piñango, 08°56'24"N, 070°50'47"W, 3820 m, 03 August 2010, Grande 2565 (VEN). Pueblo Llano, Andes de Merida/Steilhang oberhalb Laguna Negra, 08°56'N, 070°41'W, 3500–3700 m, 01 August 1958, Schwabe s.n (GOET!); Andes de Merida, 08°56'N, 070°41'W, 4000 m, 01 January 1973, Schwabe s.n (GOET!). Rangel, Margenes del Río Chama, cerca de Apartadevos, 08°47'N, 070°51'W, 01 July 1971, Aristeguieta 7886 (MO!); Quebrada de la Mucuchache, SE de la entrada, 3600 m, 16 June 1981, Briceño 298 (VEN); Dist. Rangel, cascada SE of Laguna de Mucubaji and below Pico Mucuñuque, Parque Nacional Sierra Nevada, 08°48'N, 070°49'W, 3600–3800 m, 15 June 1988, Dorr 5524 (MO!, VEN); Sierra Nevada, 08°36'N, 070°53'W, 3800 m, 20 July 1934, Farenholtz 1833 (GOET!); Sierra Nevada, 08°36'N, 070°53'W, 4000 m, 27 July 1934, Farenholtz 1927 (GOET!); Quebrada Yoyo, 08°43'N, 070°49'W, 3880 m, 12 April 1930, Gehriger 73 (MO!, VEN); Distr. Rangel. Sierra Nevada de Santo Domingo, road between Laguna de Mucubaji and Laguna Negra, 08°47'N, 070°48'W, 3400 m, 03 July 1979, Kieft 87 (MO!, VEN); moraine at the head of the valley above L. Mucubají, on a small rocky cliff just above and east of the lowest falls, 08°47'N, 070°49'W, 3650 m, 21 July 1972, Loveless 1722 (MO!); Sierra Nevada, 08°36'N, 070°53'W, s.d., Moritz 1120 (MO!); La Nevada, 08°36'N, 070°53'W, 3352 m, 21 December 1904, Schlim 1546 (MO!); Berghange oberhalb Laguna Negra/Páramo, 08°46'N, 070°48'W, 3700 m, s.d., Schwabe s.n (GOET!); Páramo de Mucubají, Páramo vegetation around Cascadas along the trail to Laguna Negra Páramo, 08°46'49"N, 070°49'16"W, 3640 m, 12 October 2007, Sklenar 10240 (VEN); Caserio Mifafi, camino quebrada de río Chama-Caserio Mucumpis a través del páramo Piedra Blanca (entrada por la carretera Apartaderous-Pico Aguila), 08°48'N, 070°50'W, 14 August 1980, Stergios 2116 (MO!); Páramo seco y húmedo en el sector de Sto. Domingo de Mucubají los alrededores de la Laguna de Mucubají, 08°46'N, 070°49'W, 29 May 1986, Stergios 8378 (MO!). Santos Marquina, Sierra Nevada. Páramo alrededores de la Laguna Verde proximo Picos Humboldt y Bonpland, near edge of la LagunaVerde, 08°34'N, 070°59'W, 4000 m, 04 December 1959, Barclay 10034 (MO!); Cerro de Caballo, 08°32'N, 070°54'W, 3600–3850 m, 25 November 1959, Barclay 9816 (MO!); Sierra Nevada; alrededores de la Laguna Coromoto. Trail to Laguna Verde, 08°34'N, 071°00'W, 3300–3500 m, 03 December 1959, Barclay 9951 (MO!), Parque Nacional Sierra Nevada, Mérida, Páramo Media Luna, 300 m Westl der Teleferico-Station Loma Redonda, 3920 m, 10 January 1995, Berg 517 (VEN); Páramo del Aguila, 10 March 1951, Croizat 66 (VEN); alrededores inmediatos de la Laguna Brava (Páramo de la Laguna Brava), sector del Páramo de los Granates, Sierra de Santo Domingo, Cordillera de los Andes, 3300 m, 20 May 1971, López-Figueiras 8728 (VEN); Páramo, Los Colorados, 3900 m, 01 May 1988, López del Pozo 416 (VEN); Páramo, 3550 m, July 1988, López del Pozo 944 (VEN); Parque Nacional Sierra Nevada, Laguna Negra, 17 September 1998, Ramirez 5533 (VEN); Laguna Mucubají, above Los Apartaderos, 3625–3655 m, 21 July 1944, Steyermark 57513 (VEN); Laguna Negra, 3520 m, 18 May 1952, Varechi 962 (VEN). Tachira: Jauregui, Páramo Sumusica along the trail heading northwest from the mountain pass (road La Grita-San Jose de Bolivar), 08°01'31"N, 071°57'53"W, 3340 m, 17 October 2007, Sklenar 10356 (VEN). Trujillo: Boconó, Mun. Carache, P.N. Dinira, arriba de Mesa Arriba, debajo del Pico Cendé, ladera SO, 09°32'N, 070°07'W, 3200 m, 01 April 1999, Duno de Stefano 767 (MO!, VEN).
Shrubs or trees, 1–2 lateral leaflet pairs; lower leaflet surfaces lanate or sericeous; fruit slightly twisted with short spines, densely sericeous.
Polylepis pepei B.B. Simpson.
The subsectional epithet Pepea is a noun in apposition.
Bolivia. Cochabamba: 77 km after Chapare on the road to Todos Santos, 4200 m, 4 Jan 1968, Vuilleumier 465 (holotype: US!; isotypes: MO!, NY!, P!, TEX!,US!, VEN!).
Shrubs or trees 2–7(9) m tall. Leaves strongly congested at the branch tips, imparipinnate with 2 pairs of leaflets, obtrullate in outline, (1.3–)1.7–2.6 × 1.2–2.0 cm; rachises densely sericeous, points of leaflet attachment with a tuft of long; stipular sheaths apically truncate or with spurs, densely lanate on the outer surfaces; leaflets elliptic in outline, second pair from the terminal leaflet the largest, one of this pair 0.8–1.3 × 0.2–0.7 cm; margin entire, apically emarginate or tridentate due to a projection of the mid-vein, basally unequally cordate; upper leaflet surfaces sparsely to densely sericeous; lower leaflet surfaces densely sericeous with whitish hairs 0.6–0.9 mm long. Inflorescences upright, 1.2–1.6(–3.5) cm long, bearing 3 flowers; floral bracts 4.0–7.3 mm long, narrowly triangular, densely sericeous on the outer surface; rachises densely sericeous. Flowers 4.9–5.9 mm diam.; sepals 3–4, ovate, green, densely sericeous outside; stamens 5–9, anthers orbicular, with a dense tuft of straight white hairs on the upper half; styles fimbriate, 3.0–4.9 mm long. Fruits turbinate often slightly twisted, with variable numbers and placement of short spines, densely sericeous; 2.3–5.7 × 1.9–3.9 mm including spines. Diploid.
Polylepis pepei has been found in northern to central Bolivia and southern Puno (Peru) where it has been collected at one locality in San Antonio de Putina Province close to the border with Bolivia (Fig.
Polylepis pepei is known from 12 locations with an EOO of 35,111 km2 and an estimated AOO of 68 km2. Polylepis pepei was categorized as VU (A1c) in the World List of Threatened Trees (
Polylepis pepei is very similar to P. rodolfovasquezii. It differs by having two pairs of lateral leaflets (versus one pair in P. rodolfovasquezii) and longer inflorescences (1.2–1.6(–3.5) cm) bearing three flowers, whereas P. rodolfovasquezii has shorter inflorescences (0.9–1.1 cm) bearing just one flower. Additionally, P. pepei may be confused with P. subsericans and P. flavipila because they all share short leaflets and inflorescences. Polylepis pepei differs from these by having two pairs of lateral leaflets and sericeous hairs, whereas the other two species have one pair of lateral leaflets and strigose hairs in P. subsericans and pilose hairs in P. flavipila.
Bolivia. Cochabamba: Chapare, Km 74 Camino antiguo a los yungas del Chapare entrando por Aguirre, 3760 m, 24 April 1999, Mercado 2207 (MO!); 77 km. after Cochabamba on the road to Todos Santos, 4200 m, 04 January 1967, Vuilleumier 465 (MO!, NY, US!). Tiraque, El Ronco, ceja de monte yungena, 17°00'05"S, 065°39'20"W, 3930 m, 11 May 2005, Alcázar-Johansen 403 (BOLV); El Ronco, Ceja de monte yunguena, 17°00'05"S, 065°39'20"W, 3710 m, 11 May 2005, Johansen 403 (MO!). La Paz: Bautista Saavedra, Area Natural de Manejo Integrado Apolobamba, Hilo Hilo, a una hora y media de Pallalani en direccion a Laji Sorapata, sobre el camino, 14°52'40"S, 068°55'34"W, 4300 m, 06 April 2009, Loza 589 (LPB, MA, MO!, USZ); 590 (LPB, MO!, QCA!, USZ). Franz Tamayo, Parque Nacional Madidi, Queara, sector Quecara, Llantai Cunca, 14°39'01"S, 069°05'01"W, 21 April 2008, Fuentes 12687 (BOLV, CTES, HSB, LPB, MA, MO!, QCA!, USZ); Area Natural de Manejo Integrado Apolobamba, Keara, hacia el NW, 14°41'03"S, 069°05'35"W, 4151 m, 17 June 2005, Fuentes 8282 (LPB, MA, MO!, QCA!); Area Natural de Manejo Integrado Apolobamba, Waca Cocha, 4.7 km al SE de Keara, 14°43'47"S, 069°04'17"W, 18 June 2005, Fuentes 8341 (LPB, MO!, QCA!); Area Natural de Manejo Integrado Apolobamba, Hilo Hilo, frente a Pallalani, 14°52'49"S, 068°57'09"W, 4286 m, 05 April 2009, Loza 587 (LPB, MO!, QCA!, USZ); 588 (BOLV, LPB, MO!, QCA!, USZ); Parque Nacional Madidi, Queara nuevo, Chuñuña, queñual al N del pueblo, 14°41'04"S, 069°05'36"W, 4100 m, 09 April 2008, Paco 1 (BOLV, DAV, HSB, LPB, MA, MO!, USZ); Area Natural de Manejo Integrado Apolobamba, Queara nuevo Toilcacocha, 14°41'12"S, 069°05'17"W, 3930 m, 11 April 2008, Paco 80 (LPB, MA, MO!, QCA!, US!); Apolobamba, Puina, cerca de Queñuapata, 14°36'26"S, 069°05'52"W, 4365 m, 10 April 2008, Quisbert 801 (BOLV, LPB, MA, MO!, NY, USZ); 810 (LPB, MA, MO!, USZ); Apolobamba, entre la comunidad de Puina y cerro k’akepununa, 14°36'24"S, 069°05'47"W, 4458 m, 11 April 2008, Quisbert 821 (LPB, MA, MO!, USZ); 825 (BOLV, LPB, MA, MO!, QCA!, USZ); Apolobamba, Palomani, 14°34'58"S, 069°07'38"W, 4286 m, 12 April 2008, Quisbert 844 (BOLV, HSB, LPB, MA, MO!, QCA!, USZ); 848 (BOLV, LPB, MA, MO!, USZ). Inquisivi, 15 Km N Villa Victoria, ca. 15 km SE Quime, 17°06'S, 067°14'W, 4050 m, 05 December 1991, Kessler 3385 (AAU!, GOET!, MO!); 3386 (AAU!, GOET!, MO!). Murillo, entre Pongo y Unduavi, MIna 50, subiendo hacia la Mina SAn Luis, 3960 m, 28 October 1994, Beck 21532 (LPB); Pongo bajanado a los Yungas, del pueblo Pongo subiendo a los restos del bosque de Polylepis pepei, 16°19'32"S, 067°57'26"W, 3950 m, 10 January 2007, Beck 29771 (LPB); Valle del Zongo entrando arriba de Botijalca (Tiquimani) haia el Este, Umapalca media hora y entrando en Valle Latera, 16°12'S, 068°03'W, 4000 m, 31 January 2004, Beck 30014 (LPB); 14.8 km N of the pass at the head of The Zongo Valley, 16°13'S, 068°07'W, 3850–4050 m, 11 April 1987, Brandbyge 584 (AAU!); 854 (MO!); Valle del Río Zongo. 14.8 km al norte de la cumbre, 16°12'S, 068°07'W, 3900–4000 m, 20 February 1987, Solomon 16172 (LPB, MO!); 17.0 km al este de La Cumbre (vieja estación de ferrocarril) por el camino a Unduavi (4.2 km al oeste de Unduavi), 16°19'S, 067°55'W, 3350 m, 11 April 1988, Solomon 18267 (LPB, MO!). Nor Yungas, arriba de Unduavi subiendo aproximadamente 45 min hacia los bosques de Polylepis pepei, 16°18'S, 067°56'W, 4120 m, 13 September 2016, Escobari 78 (LPB). Sud Yungas, debajo de Unduavi, subiendo el valle de Cerromarca, 3450 m, 28 August 1988, Beck 14680 (LPB).
Peru. Puno: San Antonio de Putina, Tocko-Tocko, 14°43'56"S, 69°36'21"W, 4560 m, 10–12 June 1969, Vargas 21596 (CUZ!).
Peru. Junin: Satipo, Pampa Hermosa, rural community of Santa Rosa de Toldopampa, buffer area of the Bosque de Proteccion Pui-Pui, 4221 m, 11°29'33.5"S, 74°56'37.8"W, 21 Apr 2015, Valenzuela & Rojas 28873 (holotype: HOXA!; isotypes: MO!, USM!).
Shrubs or trees 1–8 m tall. Leaves strongly congested at the branch tips, imparipinnate with 1 pair of lateral leaflets, obtrullate in outline, 1.4–1.6 × 1.5–2.0 cm; rachises glabrous, points of leaflet attachment with a tuft of long hairs; stipular sheaths apically with spurs, sparsely sericeous on the outer surfaces; leaflets elliptic in outline, second pair from the terminal leaflet the largest, one of this pair 0.9–1.1 × 0.4–0.6 cm; margin entire, apically emarginate with a projection of the mid-vein, basally unequally cordate; upper leaflet surfaces glabrous to sparsely sericeous; lower leaflet surfaces sparsely to densely sericeous with whitish hairs 0.8–1.0 mm long. Inflorescences upright, 0.9–1.1 cm long, bearing 1 flower; floral bracts 4.0–4.8 mm long, narrowly triangular, densely sericeous on the outer surface; rachises glabrous. Flowers 5.7–6.6 mm diam.; sepals 3, ovate, green, densely sericeous outside; stamens 9–10, anthers orbicular, with a dense tuft of straight white hairs on the upper half; styles fimbriate, 3.6–4.1 mm long. Fruits turbinate often slightly twisted, with variable numbers and placement of short spines, densely sericeous; 4.8–6.0 × 2.6–2.9 mm including spines. Diploid.
Polylepis rodolfovasquezii is distributed along the Eastern Cordillera of central Peru from San Martin to Cusco and Puno (Fig.
The estimated EOO is 164,207 km2 and AOO is 116 km2. The species is known from 19 locations. Although it is protected within the Pui-Pui Protection Forest in Junin, the species was categorized as VU for Peru (
Specimens of this recently described species were long identified as Polylepis pepei, based on their very similar morphology. However, P. rodolfovasquezii differs from P. pepei by having only one pair of lateral leaflets (versus two pairs) and shorter inflorescences (0.9–1.1 cm long) bearing just one flower (versus 1.2–1.6(–3.5) cm long bearing three flowers). When Polylepis rodolfovasquezii was described by
Polylepis rodolfovasquezii also resembles P. subsericans and P. flavipila. It differs from these in its shorter inflorescence (0.9–1.1 cm) bearing just one flower, whereas in P. subsericans, the inflorescences are 1.9–5.6 cm long with 3–6 flowers and in P. flaviplia 2.7–4.4 cm long with 3–5 flowers.
Peru. Cusco: La Convención, bosque de Mandor, 4200 m, 01 October 2004, Palomino 2043 (QCA!); Dist. Santa Teresa, Mountain edges on the lower Eastern portion of the Phachaq valley, Yanama, 13°17'11"S, 072°50'13"W, 4232 m, 28 April 2012, Sylvester 1451 (Z!); Dist. de Ollantaytambo, Mountain edges on the lower Eastern portion of the Phachaq valley, Yanama, 13°17'12”S, 072°50'13"W, 4211 m, 01 May 2012, Sylvester 1501 (Z!); Dist. de Santa Teresa, grazed slopes in the central Pacchaq valley on the East side of the river Yanama, 13°15'40"S, 072°50'17"W, 4268 m, 04 May 2012, Sylvester 1558 (Z!); Dist. Santa Teresa, Mountain edges on the lower Eastern portion of the Phachaq valley, Yanama, 13°17'12"S, 072°50'13"W, 4174 m, 05 May 2012, Sylvester 1564 (Z!); Dist. de Ollantaytambo, topmost forest found on the lower North side of the lower Phachaq valley, Yanama, 13°17'01"S, 072°50'01"W, 4566 m, 15 May 2012, Sylvester 1597 (Z!); 1598 (Z!). Urubamba, Q’esqa, 3960 m, 01 September 2002, Arce s. n. (USM!); Abra Malaga, 13°08'46"S, 072°18'14"W, 4284 m, 01 October 2002, Arce s.n (USM!); Paljay, 13°08'46"S, 072°18'14"W, 4177 m, 01 September 2002, Arce s.n (CUZ!); Chaupiwayco, 13°14'59"S, 072°29'10"W, 4290 m, 01 May 2002, Arce s.n (CUZ!); Piñasniocj, Panticalla pass, 3600 m, 15 July 1915, Cook 1241 (US!); 1837 (US!); Cañon above Peñas ruins towards Nevado Veronica, Peñas Cañon beyond Ollantaytambo on road to Abra Malaga, 4100 m, 26 August 1989, Driesch s.n (GOET!); Cumbre Malaga, 01 October 1984, Rivas s.n (USM!); Dist. de Ollantaytambo, Congunayoc; 3.5 km 175 South of the village Thastayoc, on SE facing slope facing towards Ollantaytambo, 13°10'26"S, 072°16'06"W, 4438 m, 09 March 2012, Sylvester 1392 (Z!); 13°10'24"S, 072°16'14"W, 4415 m, 09 March 2012, Sylvester 1393 (Z!); 13°10'26"S, 072°16'06"W, 4427 m, 10 March 2012, Sylvester 1396 (Z!); 13°10'22"S, 072°16'11"W, 4417 m, 10 March 2012, Sylvester 1397 (Z!); 13°10'25"S, 072°16'14"W, 4414 m, 10 March 2012, Sylvester 1398 (Z!); Machupicchu, Warmiwañuska, 13°14'21"S, 072°29'06"W, 4235 m, 13 September 2006, Toivonen 67; 68; 69; 70; 71; 72; 73; 74; 75; 80 (CUZ!); Ollantaytambo, Abra Malaga, 13°08'40"S, 072°17'51"W, 4340 m, 10 May 2006, Toivonen 82 (CUZ!); Dist. Machupicchu, Microcuenca Pacaymayo; laguna Pacaymayo, 13°13'48"S, 072°29'48"W, 3900 m, 26 June 2001, Tupayachi 5049 (CUZ!); Machupicchu Microcuenca Cusichaca, Sisaypampa Abra Palkay, 13°20'00"S, 072°30'44"W, 4100 m, 28 June 2001, Tupayachi 5155 (CUZ!); 4350 m, 01 July 1915, Bingham 2068 (US!). Junín: Concepcion, Dist. de Comas, localidad de Pomamanta, 11°44'24"S, 075°09'39"W, 4400 m, 23 August 2017, Quispe 76 (CUZ!, USM!, Z!). Satipo, Pampa Hermosa, Toldopampa, 11°29'34"S, 074°56'37"W, 4160 m, 02 August 2016, Boza 3169; 3170; 3171; 3172; 3173; 3174; 3175; 3176; 3177; 3178 (USM!, Z!); Pampa Hermosa. Toldopampa, 13°12'15"S, 075°20'22"W, 4131 m, 02 August 2016, Boza 3179 (USM!, Z!); 3180 (USM!, Z!); Dist. Pampa Hermosa, Comunidad Campesina Santa Rosa de Toldopampa, 11°29'34"S, 074°56'38"W, 4221 m, 21 April 2015, Valenzuela 28873 (HOXA, MO!, USM!). Puno: Limbani, Huancasayani on road to Limbani just east of Abra Aricoma, 14°13'S, 069°42'W, 3750 m, 28 March 1987, Boertmann 130 (AAU!); 512 (AAU!). San Martín: Mariscal Caceres, Dist. de Huicungo, Callejón de Corneadas, 07°57'46"S, 077°23'23"W, 3925 m, 11 June 2001, León 5153 (USM!); Dist. Huicungo, en pirca, debajo del camino de abra Ventanas y Laguna Colorada, 08°00'53"S, 077°23'30"W, 3924 m, 20 June 2010, León 5539 (USM!). San Martín, Dist. de Huicungo, cerca a Laguna Colorada, camino al abra Ventanas, 3900 m, 18 June 2001, León 5260 (USM!).
Trees or shrubs, lower leaflet surfaces tomentose; apices emarginate; fruits with variable numbers and placements of flattened, almost cylindrical or long spines, densely lanose, tomentose or villous.
Polylepis reticulata Hieron.
The sectional epithet Reticulatae is a plural adjective agreeing in gender with Polylepis. Section Reticulatae, first informally recognized by
Alignment of the taxa of the Polylepis sect. Reticulatae according to
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This study |
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P. brachyphylla | P. reticulata | P. reticulata | P. reticulata |
P. nitida | |||
P. reticulata | |||
P. occidentalis | |||
P. hieronymi | P. hieronymi | P. hieronymi | P. hieronymi |
P. microphylla | P. weberbaueri | P. microphylla | P. microphylla |
P. weberbaueri | P. weberbaueri | P. simpsoniae | |
P. weberbaueri | |||
P. quadrijuga | P. quadrijuga | P. quadrijuga | P. quadrijuga |
Many species of this section differ notably in the Mean Annual Temperature (MAT) of their climatic niches, with only P. quadrijuga and P. simpsoniae not being statistically different (Fig.
Polylepis hypoleuca (Weddell) Bitter, Bot. Jahrb. Syst. 45: 607. 1911.
Polylepis racemosa β hypoleuca Weddell, Chlor. Andina 2:238. 1857 [1861]. Basionym. Type. Bolivia. Tarija: between Tarija and San Luis, July-August 1846, Weddell 4607 (lectotype, designated by
Polylepis racemosa var. albotomentella Kuntze, Revis. Gen. Pl. 3: 77. 1898. Type. Argentina. Córdoba: Sierra de Córdoba, Los Gigantes, Kurtz 6926 (holotype: NY!).
Polylepis australis var. bijuga
Polylepis hieronymi
var. dolicholopha
Polylepis hieronymi var. saltensis Bitter, Bot. Jahrb. Syst. 45: 609. 1911. Type. Argentina. Salta: near Pampa Granda, pass “El Alizar”, 2400–2600 m, 1900, Nelson 12584 (holotype: S).
Bolivia. Tarija: Salinas, Cuesta de Polla, Valle del Tambo, June 1873, Lorentz & Hieronymus 938a (holotype: B destroyed; isotypes: G!, GOET!, NY!).
Trees 3–8(–25) m tall. Leaves slightly congested at the branch tips, imparipinnate with 3–4 pairs of lateral leaflets, obtrullate in outline, (3.3–)3.6–5.2 × 2.1–3.1 cm; rachises densely tomentose, points of leaflet attachment with a tuft of long, lanate hairs; stipular sheaths apically truncate with spurs, densely sericeous on the outer surfaces; leaflets narrowly obovate in outline, second pair from the terminal leaflet the largest, one of this pair (1.2–)1.5–2.1 × 0.6–1.0 cm; margin crenate with 5–7 teeth, apically emarginate, basally unequally cordate; upper leaflet surfaces sparsely tomentose; lower leaflet surfaces densely tomentose with whitish hairs 0.8–1.1 mm long. Inflorescences pendant, (4.5–)5.6–7.5(–8.1) cm long, bearing 13–25 flowers; floral bracts 3.2–6.3 mm long, narrowly triangular, densely lanate on the outer surface; rachises villous. Flowers 5.4–6.5 mm diam.; sepals 4, ovate, green, densely sericeous outside; stamens 9–19, anthers orbicular, with a dense tuft of straight white hairs on the upper half; styles fimbriate, 2.3–3.9 mm long. Fruits almost cylindrical, with variable numbers and placement of long spines, densely lanose; (4.1–)6.0–6.4(–8.3) × 3.5–7.0 mm including spines. Diploid.
Polylepis hieronymi occurs in Boliviano-Tucumanic forests at 1500–3450 m elevation (Fig.
Based on 28 collecting localities, the estimated EOO is 150,691 km2 and the AOO is 148 km2. It was categorized as VU (B1+2c) in the World List of Threatened Trees (
Sterile individuals of P. hieronymi can be confused with sterile plants of P. besseri, also because both species broadly overlap in distribution. Both species have quite similar leaflet shapes and texture, numbers of lateral leaflet pairs and densely tomentose lower leaflet surfaces. If no fruits are available (spiny in P. hieronymi, with broad ridges in P. besseri), they are best distinguished by the sericeous hairs on the stipular sheaths in P. hieronymi and tomentose hairs in P. besseri. Polylepis hieronymi also somewhat resembles P. neglecta in having 3–4 lateral leaflet pairs and relatively long inflorescences with many flowers. However, it has narrowly obovate leaflets with crenate margin and tomentose hairs 0.8–1.1 mm long, styles 2.3–3.9 mm long and spiny fruits, whereas P. neglecta has elliptic leaflets with serrate margins and glabrous to puberulous lower surfaces, shorter styles 1.5–2.2 mm long and winged fruits.
Argentina. Jujuy: Capital, Tiraxi, Alto Salviar, ladera E, 08 November 1989, Tupayachi s.n (MO!). Salta: Guachipas, Cuesta del La jar, 1600–1700 m, 07 February 1983, Novara 3142 (MO!).
Bolivia. Chuquisaca: Azurduy, Saliendo de Azurduy hacia el río Pilcomayo, 20°12'52"S, 064°26'37"W, 3114 m, 15 October 2007, Cervantes 187 (HSB, MO!); Belisario Boeto, Trayecto Villa Serrano hacia la comunidad de Tampa Mayu y Nuevo Mundo, 19°00'04"S, 064°18'55"W, 2297 m, 12 December 2007, Cervantes 157 B (HSB, MO!); 1 km S Nuevo Mundo on road to Padilla, 19°28'S, 064°10'W, 2200 m, 07 October 1991, Kessler 3306 (GOET!, LPB); 3307 (AAU!, GOET!, MO!); 3308 (AAU!, GOET!, MO!); 3309 (GOET!); 8 km SW Nuevo Mundo on road to Padilla, 19°25'S, 064°11'W, 2500 m, 07 October 1991, Kessler 3317 (GOET!); 3318 (GOET!); 3321 (GOET!); próximo a Lagunillas, 2240 m, 24 January 1988, Murguía 128 (LPB); Hernando Siles, subiendo de la Hacienda Guzman para el Abra, 20°17'16"S, 064°02'56"W, 1993 m, 24 December 2005, Peñaranda 22 (HSB, MO!, QCA!); Oropeza, Municipio de Yotala, Canton Huayllas. Comunidad Pitatorillas, 19°09'06"S, 065°20'48"W, 3374 m, 23 September 2007, Jiménez 376 (HSB, MO!, QCA!). Sud Cinti, Cerro Cobre Khasa, between Culpina and El Palmar, 20°48'S, 064°34'W, 3100 m, 21 September 1991, Fjeldså s.n (GOET!). Tomina, 25 km hacia Montegudo, 19°03'S, 064°16'W, 2400 m, 01 October 1983, Beck 9345 (BOLV, GOET!, LPB, MO!, NY); ca. 20 km SE Padilla on road to Monteagudo, 19°03'S, 064°16'W, 2450 m, 07 October 1991, Kessler 3319 (AAU!, GOET!); Trayecto Lima Bamba – EL Villar, 19°33'01"S, 064°19'55"W, 2553 m, 13 October 2007, Portal 138 (HSB, MO!). Santa Cruz: Mairana, within the Flora de la Region del Parque Nacional Amboro, but above the 700 m contour, 18°06'30"S, 063°57'00"W, 2000–2100 m, Nee 43429 (MO!, NY). Manuel Maria Caballero, Parque Nacional Amboró. San Juan del Potrero; entre Yunguillas y cabeceras del río Zapallar, 17°53'S, 064°25'W, 2300–2400 m, 12–13 May 1992, Vargas 1350 (MO!, NY, USZ). Vallegrande, between “Mataralcito” and “El Palmar” on road from Valle Grande to Tierras Nuevas, 17 km by air ESE of Valle Grande, 18°32'00"S, 065°57'00"W, 2150 m, 29 December 1988, Nee 37403 (NY); camino de Tierras Nuevas a Vallegrande, 18°30'27"S, 063°55'04"W, 2249 m, 31 July 2011, Parada-Gutierrez 3580 (MO!, USZ); camino hacia Khasa Monte, sobre la cima de la serrania, 18°38'09"S, 064°02'01"W, 2550 m, 04 August 2011, Parada-Gutierrez 3671 (MO!, USZ); camino del Cruce hacia Alto Seco, 18°44'45"S, 064°06'53"W, 2717 m, 08 July 2011, Parada-Gutierrez 3746 (MO!, USZ); Senegilla a 17 km de Vallegrande, 18°40'03"S, 064°01'55"W, 2400 m, 20 August 2012, Parada-Gutierrez 4828 (MO!, USZ); a 4 km al norte de Postrervalle sobre el camino a Mairana, 2000 m, 13 November 1999, Saldias 6192 (USZ). Vallegrande, 2363 m, 24 August 2008, Arroyo 4007 (QCA!); Meson at Samaipata, 2200 m, 01 March 1911, Herzog 1786a (GOET!). Tarija: Arce, 43 km hacia Padcaya, Huancanqui, 2500–2600 m, 20 November 1986, Beck 14080 (GOET!); cerca de Camacho, 2600 m, 17 December 1987, Beck 16067 (GOET!); bajando del Abra del Cerro Cabildo hacia el S via estancia Cabildo, 2350 m, 29 January 1988, Beck 16240 (GOET!); ca. 5 km W Padcaya, 21°54'S, 064°46'W, 2200 m, 18 September 1991, Kessler 3114 (AAU!, GOET!); 3646 (GOET!); detras de Padcaya, 2450 m, 23 January 1988, Liberman 1637 (GOET!); Municipio Padcaya, Cantón Emborozú, Reserva Natural Alarachi, recorrido a cima más alta de la Zona Alarachi, próximo al Cerro Yauparuna, 22°10'44"S, 064°36'33"W, 2260–2380 m, 16 September 2004, Serrano 4828 (MO!); 39.9 km S of jct. of road to Entre Rios, on road to Padcaya, 21°54'S, 064°41'W, 2100–2200 m, 29 April 1983, Solomon 10218 (LPB, MO!, NY). O’Connor, ca. 5 Km W Padcaya, 21°54'S, 064°46'W, 2200 m, 18 September 1991, Kessler 3113 (AAU!, GOET!, MO!); 3115 (GOET!, MO!); ca. 70 km on road from Tarija to Entre Rios, 21°26'S, 064°19'W, 2200 m, 20 September 1991, Kessler 3123; 3124 (AAU!, GOET!); 3125 (GOET!, MO!); 3660 (AAU!, GOET!, MO!); 21.1 km on road to entre Rios, 21°27'S, 064°20'W, 1900 m, 01 October 1983, Solomon 10918 (LPB, MO!). Valle del Tambo bei Tarija, 10 June 1973, Hieronymus 938 (GOET!); Cult. at Jardin Botanico La Paz 2000 from seeds, s.d., Kessler 12625 (GOET!). s.d., Cárdenas 3906 (US!); 3907 (US!); Salinas, Cuestas de Polla, in valle Tambo, June 1873, Hieronymus 938a (B, F!); s.d., Hieronymus 938a (B, MO!).
Polylepis microphylla var. polyarthotricha Bitter, Bot. Jahrb. Syst. 45: 612. 1911. Type. Goudot 1 (holotype: W).
Polylepis lanuginosa var. microphylla Weddell, Chlor. Andina 2:238.1861.
Ecuador. Chimborazo: Humboldt & Bonpland 3141 (holotype: P!; isotypes: F!, GOET! US!).
Shrubs and trees 1.5–8 m tall. Leaves strongly congested at the branch tips, imparipinnate with 3–6 pairs of lateral leaflets, obtrullate in outline, (1.3–)2.0–3.5 × (0.6–)1.2–1.5 cm; rachises densely tomentose often intermixed with twisted dark red hairs; stipular sheaths apically acute with spurs, densely tomentose on the outer surfaces; leaflets broadly elliptic in outline, second pair from the terminal leaflet the largest, one of this pair 0.3–0.7 × 0.2–0.5 cm; margin entire, apically deeply emarginate, basally unequally cordate; upper leaflet surfaces glabrous or sparsely tomentose mainly in the mid-vein depression; lower leaflet surfaces densely tomentose with whitish hairs 0.8–1.0 mm long. Inflorescences branched at the base or simple, pendant, 3.8–5.3 cm long, bearing 1–3 flowers; floral bracts 2.2–2.5 mm long, narrowly triangular, densely tomentose on the outer surface; rachises tomentose. Flowers 4.0–6.4 mm diam.; sepals 4, ovate, green, glabrous outside; stamens 9–11, anthers orbicular, with a dense tuft of straight white hairs on the upper half; styles fimbriate, 2.6–3.5 mm long. Fruits turbinate, with variable numbers of long spines, densely tomentose; 2.7–4.3 × 1.3–2.1 mm including spines. Diploid and tetraploid.
Polylepis microphylla occurs in small, isolated populations in the central Ecuadorian Andes on the slopes of Volcán Chimborazo, in north-western Peru in the high Andes of Cajamarca, the Cordillera Blanca and the adjacent Cordillera Huayhuash at the boundaries of Ancash and Lima and in Arequipa and Cusco (Fig.
The EOO is estimated as 407,902 km2 and AOO as 100 km2. The species is known from 18 locations. The two forests in Ecuador form a single genetic population (
Polylepis microphylla was included in P. weberbaueri by
Ecuador. Chimborazo: Alausí, camino Totoras-Charicando, 02°11'30"S, 078°42'18"W, 3500 m, 08 July 2004, Caranqui 1205B (CHEP); Shumit, Pucara Achupallas, 3986 m, 04 September 2009, Cárate 1200 (QCA!); carretera Alausí-Achupallas-Osogoche, 3300–3400 m, 10 August 1987, Romoleroux 372 (AAU!, NY, QCA!); Vía Achupallas-Osogoche, km 14.5, localidad Zula, 3600 m, 24 March 2001, Romoleroux 3995 (QCA!); Alausí-Achupallas, Páramo Parada 1, 3576 m, 17 March 2007, Romoleroux 4414 (QCA!); Alausí-Achullapas, 3655 m, 17 March 2007, Romoleroux 4435 (QCA!); Vía Totoras Achupallas, Lagunas de Osogoche, 3626 m, 08 November 2008, Romoleroux 5325 (QCA!); Vía desvío Osogoche-Achupallas, 3589 m, 28 December 2011, Romoleroux 5704 (QCA!); Vía Achupallas-lagunas de Osogoche, km 11.5, 3000 m, 26 April 1988, Romoleroux 576 (AAU!, MO!, QCA!); Vía Achupallas-lagunas de Osogoche km 11.5, 3650 m, 26 April 1988, Romoleroux 577 (AAU!, NY, QCA!); 3650 m, s.d., Romoleroux 578 (AAU!, NY, QCA!); Vía Achuapallas-Lagunas de Osogoche, km. 15, 3650 m, 26 April 1988, Romoleroux 579 (AAU!, MO!, NY, QCA!); 3650 m, Romoleroux 580 (AAU!, NY, QCA!); Vía Achupallas-Lagunas de Osogoche, km 15, 3650 m, 26 April 1988, Romoleroux 581 (AAU!, NY, QCA!); Vía Osogoche-Achupallas, 3610 m, 30 September 2016, Romoleroux 6126 (QCA!); Vía Osogoche-Achupallas, 3610 m, 30 September 2016, Romoleroux 6127 (QCA!); Vía Osogoche-Achupallas, 02°15'53"S, 078°42'09"W, 3610 m, 09 March 2017, Romoleroux 6148 (QCA!); Alausí-Achupallas, 3655 m, 17 March 2007, Romoleroux GPI4435 (QCA!), Quitesian Andes, s.d., Cothouy s.n (NY); 1857–1864, Spruce s.n (MO!).
Peru. Ancash: Huaylas, Caraz, Laguna Parón, flanco norte, 4170 m, 27 April 2013, Baldeón 7816 (USM!); Caraz, Laguna Paron, Irazábal 199 (CUZ!); Yungay, Parque Nacional de Huascaran, laguna Paron, 08°59'55"S, 077°41'26"W, 4200 m, 11 February 1997, Tupayachi 3271 (CUZ!). Cajamarca: San Miguel, San Miguel de Pallaques, road Agua blanca to Oyotum, Ponga la Mesa, 3500–3600 m, 14 October 2000, Weigend 2000/748 (F!, USM!). Cusco: Acomayo, Queuñayocpampa, 14°04'01"S, 071°35'08"W, 3940 m, 01 April 2003, Arce s.n (USM!); Rondocan, localidad Parara, 13°47'10"S, 071°45'08"W, 4085 m, Pfuro AS-133 (Z!). Calca, Pisac, 13°25'S, 071°51'W, 3400 m, 10 February 2003, Lægaard 22250 (AAU!). Cusco, Huacoto, 13°30'54"S, 071°51'19"W, 3960 m, 01 May 2003, Arce s.n (USM!); Chacan, 13°29'04"S, 071°59'26"W, 3805 m, 17 September 2014, Boza 3001; 3002 (USM!, Z!); Cuzco, Chacan camino al grupo arqueológico, borde de la microcuenca Chacan, 3600 m, 03 October 2000, Galiano 3999 (QCA!); Huacoto, 3991 m, Irazábal 204 (CUZ!); Chacan S of Cusco, 13°29'S, 072°00'W, 3900 m, 10 February 2003, Lægaard 22350 (AAU!, US!); San Jerónimo, localidad de Huacoto, 13°31'06"S, 071°51'32"W, 3940 m, 25 May 2006, Toivonen 19 (CUZ!); 20 (CUZ!); 21 (CUZ!); Chacan, 14 June 2006, Toivonen 98 (CUZ!); Chacan, 14 June 2006, Toivonen 99 (CUZ!); Ruinas de Pisac, 3200 m, 01 May 2002, Toivonen s.n (CUZ!); Chacan, 3600 m, 28 April 1993, Tupayachi 2280 (CUZ!). Quispicanchis, Dist. Huaro, Urpay, 13°41'01"S, 071°38'22"W, 3200 m, 01 November 2002, Galiano 4512 (AMAZ, CUZ!, HUT, MO!, MOL, USM!). Urubamba, Cusco. Prov. Urubamba, Yucay, Hatum Wayko, 3500 m, 09 September 2001, Herrera 4175 (QCA!); Yucay, 3833 m, Irazábal 207 (CUZ!); Yucay, cerro Turukuntur, 3750 m, 13 January 1991, Tupayachi 1460 (CUZ!). Lima: Cajatambo, Huayllapa. Dist. de Copa, cerro empinado a 115 km del pueblo, 3360 m, 13 May 2001, Callupe 1 (USM!).
Resembles the Ecuadorian species, P. reticulata Hieron. by having 3–5 lateral leaflet pairs and similar type and density of hairs, but differs by its shorter hairs (0.3–0.6 mm vs. 0.6–1.5 mm), shorter inflorescences (2.4–6.7 cm vs. 2.3–13.8 cm) and shorter styles (1.5–2.0 mm vs. 2.6–3.9 mm).
Peru: Piura: Huancabamba, Talanco, 2900 m, 5 Nov 1976, A. Sagastegui A.8635 (holotype: MO!; isotype: UMO!).
Trees 3–15 m tall. Leaves slightly congested at the branch tips, imparipinnate with 3–5 pairs of lateral leaflets, obtrullate in outline, 3.1–4.7 × 1.8–2.9 cm; rachises densely tomentose, points of leaflet attachment with a tuft of long, lanate hairs; stipular sheaths apically acute with spurs, densely lanate on the outer surfaces; leaflets elliptic in outline, second pair from the terminal leaflet the largest, one of this pair 1.1–1.9 × 0.5–0.8 cm; margin entire or slightly crenate at the apex with 3–6 teeth, apically emarginate, basally unequally cordate; upper leaflet surfaces glabrous; lower leaflet surfaces densely tomentose with whitish hairs 0.3–0.6 mm long. Inflorescences pendant, 2.4–6.7 cm long, bearing 4–12 flowers; floral bracts 6.8–7.6 mm long, narrowly triangular, densely lanate on the outer surface; rachises tomentose. Flowers 6.8–7.6 mm diam.; sepals 4, ovate, green, densely tomentose outside; stamens 9–15, anthers orbicular, with a dense tuft of straight white hairs on the upper half; styles fimbriate, 1.5–2.0 mm long. Fruits turbinate, with variable numbers and placement of flattened spines, densely villous; 3.8–4.1 × 5.8–7.1 mm including spines. Diploid.
Polylepis occidentalis is distributed in the high mountains of western Peru from Huancabamba (Piura) to Pataz (La Libertad) (Fig.
The species epithet “occidentalis” (Latin: western) refers to the distribution range occupying the western mountains in Peru.
The EOO for Polylepis occidentalis is estimated as 12,906 km2 and the AOO at 52 km2. It is known from 11 locations. It has been found in monospecific forests at the southern margins of the Huancabamba Andean Depression, which forms an important dispersal corridor for Andean tree species (
The populations of Polylepis section Reticulatae from western Peru have been previously identified either as P. reticulata (