Corresponding author: Joel Calvo (
Academic editor: A.R. Magee
The Andean genus
Calvo J, Moreira-Muñoz A (2020) Taxonomic revision of the Andean genus
Number of
Species of the genus were traditionally treated under the morphologically similar genus
The taxonomic history of the group before the establishment of the genus
This study is mostly based on the revision of 1014 herbarium specimens (types included, duplicates excluded) kept at the following herbaria: A, AAU, B, BA, BM, BOLV,
A comprehensive synonymy of the genus
A general description of the genus and detailed species descriptions were prepared. For this, qualitative characters were studied with the aid of a binocular dissecting microscope, while quantitative characters were recorded using a Mitutoyo digital caliper, CD-15DC. A Zeiss Standard 16WL microscope was used for examination of the achene trichomes. Information concerning the habitat, elevation, and flowering period of each species was obtained from the herbarium specimen labels.
Accepted species are presented in an order that represents their morphological affinities. An index to the taxonomic names treated is provided in Appendix
Main habit types in
In contrast, the second habit type includes the species forming clumps of somewhat distantly spaced stems that are uniform in appearance along its whole length (Fig.
Diagram of measurements used in descriptions
The leaf morphology has a relevant taxonomic value within the genus [see treatment under
Species of
The ray florets are pistillate and fertile. The color of the ray corollas is usually white, sometimes slightly purplish on the abaxial surface near the apex. Three species have yellow or pale yellow ray corollas and
The disc florets are hermaphroditic. The corollas are tubular, 5-lobed at the apex, yellow, whitish, creamy, or purplish.
Further variability has been found in the morphology of the style branch apex. In all species examined it is truncate but differs on its ornamentation; some species bear a crown of sweeping trichomes (e.g.,
In
The pappus of all species is capillary, composed of barbellate, whitish to partially rose- or purple-colored bristles.
Suffruticose plants, forming mats, hummocks, or clumps of erect stems.
Colombia, Ecuador, Peru, Bolivia, northern Chile, and northwestern Argentina. The
The generic name
Distribution map of genus
1 | Leaves entire, undivided at the apex |
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– | Leaves notched, forked, or finger-like at the apex |
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2 | Ray corollas not surpassing the involucre, yellow |
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– | Ray corollas conspicuously surpassing the involucre, white or pink |
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3 | Leaves linear, denticulate, erect (not adpressed to the stem); involucral bracts 4.1–7.7 mm long; stems 4–7 cm long, erect or decumbent |
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– | Leaves linear-triangular, minutely, irregularly denticulate, adpressed to the stem; involucral bracts 2.8–3.4 mm long; stems 8–15 cm long, erect |
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4 | Ray corollas entirely pink |
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– | Ray corollas white, sometimes purplish beneath |
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5 | Leaf apices aristate (arista 0.5–1 mm long) |
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– | Leaf apices acute to obtuse |
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6 | Plants forming distant, rather solitary, erect stems; involucres ca. 8.6 × 7.3 mm; ray corollas ca. 12 mm long; disc florets ca. 50 |
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– | Plants forming dense mats or hummocks; involucres 4.9–5 × 4.8–5 mm; ray corollas 6.3–6.5 mm long; disc florets 14 to 15 |
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7 | Stems glabrous |
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– | Stems densely arachnoid |
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8 | Leaves triangular, denticulate in the upper half; involucral bracts 13 to 15; ray florets 14 to 21 with corollas 7.4–12.9 mm long; disc florets 35 to 48 |
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– | Leaves linear-subulate, entire; involucral bracts 8 to 12; ray florets 9 to 15 with corollas 5.9–8 mm long; disc florets 24 to 32 |
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9 | Achenes white-villous |
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– | Achenes glabrous |
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10 | Leaf laminas 5.3–7.8 × 0.8–0.9 mm; ray florets 20 to 23; capitula surpassing the leaves |
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– | Leaf laminas 12.6–13.5 × 2.3–2.4 mm; ray florets 38 to 41; capitula enclosed among the leaves |
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11 | Involucres 10.2–19.7 mm long; ray florets 13 to 21 with corollas 13–19.7 mm long; disc florets 50 to 95 |
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– | Involucres 3.3–9.3 mm long; ray florets 8 to 13 with corollas 3.6–12.2 mm long; disc florets 7 to 39 |
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12 | Leaf laminas spreading at nearly 90° from the sheath-like bases, 2.5–10.8 mm long, finely papillose; plants forming dense mats or hummocks (in paramo and humid puna) |
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– | Leaf laminas barely spreading from the sheath-like bases, 2.5–7.1 mm long, not papillose; plants forming clumps of erect stems, rarely dense mats (in subhumid, dry, and desertic puna) |
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13 | Capitula disciform |
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– | Capitula radiate |
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14 | Ray corollas not surpassing the involucre, pale yellow; capitula rather nodding; stems 30–60 cm tall |
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– | Ray corollas conspicuously surpassing the involucre, white; capitula erect; stems 2–24 cm tall |
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15 | Leaves 2 to 3-forked or finger-like at the apex |
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– | Leaves 3-notched at the apex |
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16 | Leaves finger-like at the apex (at least 9-divided) |
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– | Leaves forked (2 to 3-divided) |
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17 | Leaves 2-forked |
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– | Leaves 3-forked |
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18 | Leaf lobes 3.6–5.9 mm long, entire or divided; involucres 12.1–16.7 mm long; ray corollas 9.2–14.1 mm long; disc florets 49 to 63 |
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– | Leaf lobes 1.3–2.6 mm long, entire (rarely notched); involucres 7–8.4 mm long; ray corollas 7.1–8.2 mm long; disc florets 29 to 33 |
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19 | Stems arachnoid; leaf laminas 0.5–1.1 mm wide; disc florets 7 to 18 |
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– | Stems glabrous; leaf laminas 1.3–3.7 mm wide; disc florets 22 to 58 |
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20 | Ray florets 20 to 39; disc florets 37 to 58; involucral bracts 5.2–8 mm long |
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– | Ray florets 8 to 13; disc florets 22 to 37; involucral bracts 3.1–5.4 mm long |
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21 | Leaf margins entire and shortly ciliate; central lobe smaller than lateral ones; ray florets 20 to 21 with corollas 10.4–11.9 mm long; achenes 4.8–5.4 mm long |
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– | Leaf margins entire; central lobe larger than lateral ones; ray florets 26 to 39 with corollas 8.3–9.2 mm long; achenes 2.5–3.1 mm long |
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22 | Leaf laminas with evanescent arachnoid trichomes above; involucral bracts 10 to 13, 4.9–5.4 mm long; disc corollas 5.4–6 mm long |
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– | Leaf laminas glabrous; involucral bracts 8 to 9, 3.1–4.3 mm long; disc corollas 4.4–5 mm long |
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Suffruticose plant, forming distant, rather solitary, erect stems.
Endemic to Ecuador (Azuay, Zamora-Chinchipe [expected]). It grows in marshy places of the paramo ecoregion, between elevations of 3050–3070 m (Fig.
Thus far,
Distribution map of
Collected in flower in September and November.
The adjective
Suffruticose plant, forming dense mats or hummocks, with rhizome-like stems up to 15 cm long covered with lanate indumentum and old leaves, rather erect, simple or branched from the base.
Northwestern Colombia to northern Bolivia. Bolivia (La Paz), Colombia (Antioquia, Caldas, Cauca, Cundinamarca, Huila, Meta, Nariño, Quindío [expected], Risaralda, Tolima, Valle del Cauca), Ecuador (Azuay, Bolívar, Cañar [expected], Carchi, Chimborazo, Cotopaxi, Imbabura, Loja, Morona-Santiago, Napo, Orellana, Pichincha, Sucumbíos, Tungurahua, Zamora-Chinchipe), Peru (Amazonas [n.v.], Ancash, Cajamarca, Junín, La Libertad [expected], San Martín). This species grows in marshes, grasslands, rocky outcrops, and exposed places of the paramo and humid puna ecoregions, between elevations of (2700–)3000–4800 m (Fig.
The species is very common along the paramos of Colombia and Ecuador, but it becomes very scattered towards its southern limit. No collections have been studied from the region comprised between central Peru (Junín) and northern Bolivia (La Paz).
Flowering nearly all year round.
The epithet
The species shows high variability in leaf size and density, which led some authors to describe infraspecific taxa or even to propose distinct species.
An isolectotype of the name
Regarding the name
Suffruticose plant, forming dense mats or hummocks, with rhizome-like stems up to 5 cm long covered with sparse pilose indumentum and old leaves, rather erect, simple or branched from the base.
Southern Colombia to central Ecuador. Colombia (border Cauca/Huila), Ecuador (Chimborazo, Morona-Santiago, Napo, Tungurahua). It grows in rocky outcrops and scree slopes around the upper limit of vegetation of the superparamo ecoregion, between elevations of 4125–4800 m (Fig.
Flowering from July to December.
The epithet
It is morphologically close to
The lectotype of
The isotypes of
Suffruticose plant, forming dense mats or hummocks, with rhizome-like stems up to 6.5 cm long covered with matted lanate indumentum and leaf bases, rather erect, simple or branched from the base.
Endemic to Ecuador (Azuay, Cañar, Loja). It grows in moist grasslands and exposed places of the paramo ecoregion, between elevations of 3650–4275 m (Fig.
Distribution map of
Flowering from August to March.
The adjective
It is morphologically similar to
Ecuador. Chimborazo: Mt. Chimborazo area, at the end of
Suffruticose plant, forming lax mats, with rhizome-like stems 20–35 cm long covered with arachnoid indumentum and leaf base remnants resembling paleae, horizontal, creeping, simple or branched from the base.
Endemic to Ecuador (Bolívar, Chimborazo). This species grows in exposed places and on sandy soils of the dry superparamo ecoregion, at elevations of 4100–4300 m (Fig.
The species is known from a few collections near the Chimborazo Volcano. Its distribution area partially overlaps with those of
Flowering from April to July.
The epithet honors the American botanist Vicki A. Funk (1947–2019), who described the genus
The morphologically closest species is
Suffruticose plant, forming mats or hummocks, with rhizome-like stems up to 16 cm long covered with matted lanate indumentum and leaf bases, rather erect, simple or branched from the base.
Endemic to Ecuador (Bolívar, Chimborazo, Cotopaxi, Napo, Pichincha, Tungurahua). It grows in rocky outcrops and on sandy soils, rather dry, around the upper limit of vegetation of the superparamo ecoregion, at elevations of (3500–)3900–5100 m (Fig.
Distribution map of
Flowering from April to November.
The epithet
The specimen B-W-16432-01-0 kept at the Willdenow Herbarium is considered part of the original material and designated as isolectotype. Willdenow received the specimen from Humboldt as stated by Schlechtendal on the blue label at the bottom of the sheet. Although unnumbered, it contains an individual identical to those from the lectotype. See
Suffruticose plant, forming dense mats or hummocks, with rhizome-like stems up to 17 cm long covered with matted lanate indumentum and leaf bases, rather erect, simple or branched from the base.
1 | Leaf laminas 14.7–27.6 mm long; involucral bracts 16 to 21, 8.6–14.9 mm long; ray florets 16 to 21; disc florets ca. 95 |
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– | Leaf laminas 6.5–12.7 mm long; involucral bracts 11 to 13, 6–8.2 mm long; ray florets 13 to 15; disc florets ca. 50 |
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Central Colombia to central Ecuador. Colombia (Caldas, Cauca, Nariño, Risaralda, Tolima, Quindío, Valle del Cauca), Ecuador (Carchi, Chimborazo, Cotopaxi [expected], Morona-Santiago [expected], Napo, Pichincha, Tungurahua). It grows in marshes and moist grasslands of the paramo ecoregion, between elevations of 3125–4500 m (Fig.
Flowering nearly all year round.
The adjective
This taxon forms dense mats or hummocks and it is characterized by its linear, apically obtuse leaves, which are flat to elliptical in cross section and prominently 1-nerved on both faces. The rhizome-like stems are robust, long, and covered with a matted lanate indumentum that covers the leaf bases.
This subspecies can be differentiate from
The habit of
Endemic to Colombia (Arauca [expected], Boyacá, Norte de Santander, Santander). It grows in marshes and wet exposed areas of the paramo ecoregion, between elevations of 3200–4600 m (Fig.
Flowering nearly all year round.
The adjective
It should be noted that
The presumed isotype of
Suffruticose plant, forming dense mats or hummocks, with rhizome-like stems up to 8 cm long covered with old leaves, rather erect, simple or branched from the base.
Central Peru to northern Bolivia. Bolivia (La Paz), Peru (Ancash, Arequipa, Ayacucho, Huancavelica, Huánuco, Junín [expected], Lima, Moquegua [expected], Pasco [expected], Puno). It grows on exposed rocky slopes and cryoturbated soils of the subhumid and humid puna ecoregions, between elevations of 4550–5350 m (Fig.
Distribution map of
Flowering from February to September.
The adjective
Because of the marginally denticulate leaves, on dried specimens
Suffruticose plant, forming clumps of erect or decumbent stems.
Central Peru to central Bolivia. Bolivia (Cochabamba, La Paz, Oruro, Potosí), Peru (Arequipa, Ayacucho, Cusco, Huancavelica, Junín, Lima, Moquegua, Pasco [expected], Puno). This species grows in rocky outcrops, scree slopes, cryoturbated soils, and exposed grasslands of the subhumid and humid puna ecoregions, between elevations of 3900–5350 m (Fig.
Flowering from February to November.
The adjective
The morphologically closest species is
The lectotype designation of the name
Suffruticose plant, forming clumps of erect stems.
Southern Peru to northern Chile. Bolivia (Oruro, Potosí), Chile (Antofagasta, Arica-Parinacota, Tarapacá [expected]), Peru (Tacna). It grows in exposed places on rocky and sandy soils of the dry and desertic puna ecoregions, between elevations of 4000–5200 m (Fig.
Most populations of this species are located between eastern Tacna (Peru), northeastern Arica-Parinacota (Chile), and northwestern Oruro (Bolivia). However, a few collections are from further south (northeastern Antofagasta in Chile and western Potosí in Bolivia). Its presence in the Chilean region of Tarapacá is expected.
Distribution map of
Flowering from April to December.
It is named after the genus
Although the syntypes of the name
Suffruticose plant, forming clumps of rather decumbent stems.
Southern Peru to northern Chile. Bolivia (Oruro), Chile (Arica-Parinacota, Tarapacá), Peru (Arequipa, Moquegua, Puno, Tacna [n.v.]). It grows along the banks of marshes and salt lagoons of the subhumid and dry puna ecoregions, between elevations of 3800–4700 m (Fig.
Flowering nearly all year round.
This species honors H.A. Weddell (1819–1877), a British botanist (French by choice) and physician who devoted part of his life to the study of the South American flora.
The leaf apex is acute but we studied a specimen showing two lateral divisions near the apex (
This species is morphologically similar to
The single original collection that we found comes from the Huasco Lagoon (Tarapacá, Chile). However,
The holotype designation of the name
Suffruticose plant, forming clumps of rather decumbent stems.
Restricted to northern Chile, northwestern Argentina, and southwestern Bolivia. Argentina (Jujuy, Salta), Bolivia (Potosí), Chile (Antofagasta). This species grows along the banks of salt lagoons and desert plains with a certain humidity of the dry and desertic puna ecoregions, between elevations of (2600–)3500–4900 m (Fig.
Flowering from October to March.
The epithet
This species can be properly identified by the combination of the following characters: rather decumbent, 4–8 cm tall, glabrous stems, linear-oblong leaf lamina, 3-notched leaf apex with truncate lobes (rarely subentire), 8 to 9 involucral bracts, 8 to 13 ray florets with white corollas conspicuously surpassing the involucre, and glabrous achenes.
This species has morphological affinities with
The holotype designation by
Suffruticose plant, forming clumps of erect stems (rarely dense mats).
Detail of leaves and capitulum
Central Peru to northwestern Argentina. Argentina (Catamarca, Jujuy, La Rioja, Salta, Tucumán), Bolivia (Oruro, Potosí), Chile (Antofagasta, Arica-Parinacota, Tarapacá [expected]), Peru (Arequipa, Ayacucho, Cusco, Junín, Lima, Moquegua, Puno [expected], Tacna). It grows in rocky outcrops, grasslands, exposed slopes, and flat sandy pampas of the subhumid, dry, and desertic puna ecoregions, between elevations of (2600–)3000–5200 m (Fig.
Distribution map of
Flowering nearly all year round.
The epithet
The type material of this species shows plants with clearly truncate and callous-tipped leaf apex, which probably led
The distribution area of this species partially overlaps with that of
Regarding nomenclatural issues, it is noteworthy that Funk’s typification of the name
The original material of
The typification of
Argentina. Catamarca: El Cajón, Negroara, 15 Jan 1914,
Suffruticose plant, forming clumps of erect stems.
Endemic to Argentina (Catamarca, Jujuy, Salta, Tucumán). This species grows in exposed rocky slopes and on bare soils of the dry puna ecoregion, between elevations of 3500–5000 m (Fig.
Flowering from January to March.
The epithet
This species shows morphological affinities with
Suffruticose plant, forming clumps of erect stems.
Restricted to northwestern Argentina and southwestern Bolivia. Argentina (Jujuy, Salta [expected]), Bolivia (Potosí). It grows in wet meadows and rocky places of the dry puna ecoregion, between elevations of 4600–5200 m (Fig.
Collected in flower in March and November.
It is named after Eric von Rosen (1879–1948), a Swedish aristocrat, explorer, and ethnographer.
Since two specimens of the gathering
Suffruticose plant, forming clumps of erect stems.
Central Peru to northern Chile. Bolivia (Oruro, Potosí), Chile (Antofagasta, Arica-Parinacota, Tarapacá), Peru (Apurímac [expected], Arequipa, Ayacucho, Cusco, Huancavelica, Junín, Lima, Moquegua, Puno). It grows in wet rocky outcrops, marsh and stream edges, and on cryoturbated soils of the puna ecoregion, between elevations of 3700–5300 m (Fig.
Flowering nearly all year round.
The adjective
It is a variable species mainly concerning leaf lobe shape and leaf indumentum. Typical forms have entire leaf lobes but some specimens display 2-divided lobes (
This species has been confused with
Suffruticose plant, forming clumps of erect stems.
Endemic to Argentina (Jujuy, Salta). It grows in rocky slopes on rather wet soils of the dry puna ecoregion, between elevations of 3700–5125 m (Fig.
Distribution map of
Flowering from January to May.
The epithet
This is a narrow endemic species to northwestern Argentina that has erroneously been reported from northern Chile (
Suffruticose plant, forming clumps of erect stems.
Endemic to Peru (Ancash, Ayacucho, Huancavelica, Huánuco [expected], Junín, Lima). It grows in rocky outcrops, scree slopes, and on cryoturbated soils of the subhumid and humid puna ecoregions, between elevations of 3850–5000 m (Fig.
Distribution map of
Flowering from February to September.
The adjective
This is a distinctive species due to the shortly ciliate leaf margin and the 3-notched leaf apex (the lateral lobes being conspicuously larger than the central one). The leaf lamina is glabrous, linear-oblong, slightly broadened towards the apex, flat in cross section but tending to be curved forwards. It has 13 to 14 involucral bracts, 20 to 21 ray florets with white corollas, 37 to 58 disc florets, and glabrous achenes.
It has to be noted that we had problems in identifying
Suffruticose plant, forming clumps of erect stems.
Endemic to Peru (Junín, Lima). It grows in rocky outcrops of the puna ecoregion, at elevations of ca. 4600 m (Fig.
Collected in flower in May.
The epithet
This is a poorly known and collected species. Aside from the type material, we only studied one collection from southeastern Lima Department (
It might be confused with
Suffruticose plant, forming clumps of erect stems or cushions.
Central Peru to central Bolivia. Bolivia (Cochabamba, La Paz, Potosí [n.v.]), Peru (Ancash, Apurímac, Arequipa, Ayacucho, Cusco, Huancavelica, Huánuco, Junín, Lima, Moquegua, Pasco, Puno). This species grows in rocky outcrops, scree slopes, cryoturbated soils, and exposed grasslands of the subhumid and humid puna ecoregions, between elevations of (2800–)3400–5500 m (Fig.
Flowering nearly all year round.
The epithet
Another species with finger-like leaves is
Our efforts in locating the type material of
Shrubby plant.
Endemic to Peru (Arequipa, Cusco, Huánuco, Lima, Moquegua, Puno). It grows in rocky outcrops, scree slopes, and exposed grasslands of the subhumid puna ecoregion, between elevations of 3900–5200 m (Fig.
Distribution map of
Flowering from March to December.
The epithet
This is a shrub with 30–60 cm tall stems that have glabrous or floccose-lanate finger-like leaves, with at least 12 divisions at the apex. The divisions are similar in length, ca. 1.5 mm deep. It is characterized by its capitula that tend to be nodding as the plant ages, with 15 to 28 ray florets with pale yellow corollas that do not surpass the involucre. As in the case of
It should be mentioned that the elevation indicated on the isolectotype at F is 16000 feet, which slightly differs from that indicated on the lectotype (15500 feet). We believe that such mismatch is not a major reason for excluding it from the type material.
Suffruticose plant, forming clumps of erect or decumbent stems.
Southern Peru to central-western Bolivia. Bolivia (Oruro), Chile (Arica-Parinacota), Peru (Arequipa, Moquegua, Puno, Tacna). It grows in rocky outcrops, steep volcanic slopes, and open shrublands of the subhumid and dry puna ecoregions, between elevations of 3700–4950 m (Fig.
Flowering from February to June.
It is named after the Peruvian village of San Antonio de Esquilache (western Puno), where the type material was collected in 1937 by Dora Stafford.
This is the unique species within the genus displaying disciform capitula. They have ca. 10, peripheral, pistillate florets with a corolla reduced to a vestigial tube ca. 2.1 mm long or without corolla, and therefore, the style is exsert in its whole length or almost so. Another distinctive feature is the morphology of the leaves; they are 3-forked, abruptly narrowed at the base, and marginally scabrous-ciliate. Each lobe usually has 2 to 3 rough mucronate teeth. Any confusion with the other members of the genus is unlikely.
See
We are grateful to the curators and staff of the herbaria mentioned in the text. We would like to express our gratitude to the late Vicki Funk (US) for research support and for sharing worthy information and material that she compiled over a long period of time. Special thanks are also addressed to Carol Kelloff (US) for herbarium assistance, Alice Tangerini (US) for the nice illustrations, and Carlos Aedo (MA) for advice on taxonomic issues. We are indebted to Luvo Magoswana and to another anonymous reviewer for their insightful comments during the review process. The first author was a Short-Term Visitor Program Fellow at the Smithsonian Institution and was granted the Robinson Award of its Department of Botany. This work has been funded by FONDECYT from Chile by means of a postdoctoral fellowship to the first author (project N°3170270) and supported by the project N°1180211 of the second author.
Index to the taxonomic names treated, with the currently accepted names indicated in bold type.
sect.
sect.
subg.
List of exsiccatae.