Description.

Suffruticose plant, forming distant, rather solitary, erect stems. Rhizomes 6–10 × 0.2–0.3 cm, horizontal to oblique, glabrous. Stems 12–20 cm tall, usually simple, sometimes branched at the upper part, covered with adpressed, arachnoid trichomes, with leaves restricted to the upper part. Leaves imbricate, extending into a sheath-like base that bears arachnoid trichomes; leaf laminas 4.6–5 × ca. 0.9 mm, linear-acicular, aristate at the apex (arista up to 1 mm long, purplish), entire, terete in cross section, glabrous, unconspicuously nerved on both faces, somewhat fleshy, drying coriaceous, matte, papillose. Capitula radiate, erect, sessile, partially enclosed among the leaves. Involucres ca. 8.6 × 7.3 mm, cupuliform; involucral bracts ca. 13, 5.2–5.5 × 1.1–1.6 mm, rather acute at the apex, dark-purplish. Ray florets ca. 13; corollas ca. 12 × 2.3–2.5 mm, 4 to 5-veined, subentire to 3-toothed at the apex, conspicuously surpassing the involucre, white, purplish beneath. Disc florets ca. 50; corollas 3.8–4.2 mm long, pale yellow to creamy; style branches truncate with a crown of sweeping trichomes, yellowish. Achenes cylindrical, with scattered arachnoid trichomes (immature); pappus 3.1–3.3 mm long, barbellate, whitish. Chromosome number unknown. Fig. ​Fig.55.

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Figure 5.

Xenophyllum acerosumA habit B stem apical part and capitulum C detail of leaf surface D adaxial leaf surface E ray corolla and style F disc corolla and stamens (vertically sectioned, style removed) G disc corolla and style branches H style I immature achene with pappus. All details drawn from Prieto 280 (US). Illustration by Alice Tangerini.

Distribution and habitat.

Endemic to Ecuador (Azuay, Zamora-Chinchipe [expected]). It grows in marshy places of the paramo ecoregion, between elevations of 3050–3070 m (Fig. ​(Fig.66).

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Figure 6.

Distribution map of Xenophyllum acerosum (left hand) and X. humile (right hand).

Thus far, X. acerosum is only known from the type locality in the paramo of Yacumbi, which is located above Saraguro on the border between the Ecuadorian provinces of Azuay and Zamora-Chinchipe.

Phenology.

Collected in flower in September and November.

Etymology.

The adjective acerosus -a -um means needle-shaped, and it refers to the leaf shape of this species.

Notes.

Xenophyllum acerosum is unique in the genus in its distant, rather solitary, erect stems arising from long rhizomes and by its linear-acicular, aristate leaves. The stems only bear leaves in the upper part.

Description.

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. Stems 1–3 cm tall (aerial part), lanate. Leaves stellate-imbricate to subimbricate, extending into a sheath-like base that bears long silky trichomes, usually with the leaf lamina spreading at nearly 90° from the sheath-like base; leaf laminas 2.5–10.8 × 0.5–1.5 mm, linear, obtuse to truncate, usually callous-like tipped at the apex, entire, elliptical to almost terete in cross section, glabrous, 1-nerved above (sometimes barely visible), 1-nerved beneath (sometimes barely visible), fleshy, drying coriaceous, matte, finely papillose. Capitula radiate, erect, sessile to subsessile (rarely with a short peduncle up to 5 mm long). Involucres 4.6–9.3 × 2.9–9 mm, cupuliform; involucral bracts (8–)10 to 13, 3.1–6.1 × 1–2.1 mm, obtuse at the apex, greenish to dark-purplish. Ray florets (8–)10 to 13; corollas 4.2–12.2 × 0.4–2.1 mm, 4 to 7-veined, subentire to 3-toothed at the apex, conspicuously surpassing the involucre, white. Disc florets (8–)22 to 39; corollas 3.1–5 mm long, whitish, usually purple-tipped; style branches truncate with a crown of sweeping trichomes, purplish. Achenes 2–2.8 × 0.4–0.6 mm, cylindrical, 7 to 9-ribbed, glabrous; pappus 2.1–7.9 mm long, barbellate, whitish, rarely partially rose-colored. Chromosome number unknown. Figs ​Figs2A,2A, ​,77.

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Figure 7.

Xenophyllum humileA habit B stem apical part and capitulum C detail of involucral bract apex D adaxial leaf surface E ray corolla and style F style G disc floret (frontward bristles removed) H disc corolla, stamens, and style (vertically sectioned). All details drawn from Funk & Montezuma 11443 (US) except for A (drawn from Øllgaard & Balslev 8749, US) and G, H (drawn from Funk 8093, US). Illustration by Alice Tangerini.

Additional iconography.

Funk (1997a: 237, fig. 1A); Funk (1997b: 33, fig. 3B, C); Beltrán (2016: 358, fig. 2C, as photo).

Distribution and habitat.

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. ​(Fig.66).

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).

Phenology.

Flowering nearly all year round.

Etymology.

The epithet humile means low-growing, close to the ground.

Notes.

Xenophyllum humile (Kunth) V.A.Funk is a morphologically polymorphic species with rhizome-like stems covered with lanate indumentum and marcescent leaves, obtuse to truncate leaf apex (usually callous-like tipped), (8–)10 to 13 involucral bracts, and (8–)10 to 13 ray florets with white corollas. The leaves are usually bent backwards due to a sort of articulation at the inflexion point between the leaf lamina and the sheath-like base. It is also characterized by its glabrous achenes, although we studied a single specimen from El Cajas (Azuay, Ecuador) displaying achenes with indumentum (Ulloa & Minga 1393, HA).

The species shows high variability in leaf size and density, which led some authors to describe infraspecific taxa or even to propose distinct species. Werneria fontii Cuatrec. was described from Tolima (Cordillera Central, Colombia) mainly on the basis of the compact habit of the plants thriving in this area. These plants are certainly smaller than typical forms and display short leaves with the leaf lamina barely spreading from the sheath-like base. Specimens also collected in the paramos of Tolima were previously used by Weddell (1856) to describe a variety that he characterized as having flesher leaves than typical variety, which is in line with the remarks stated by Cuatrecasas (1935) under W. fontii. However, these small and compact plants are not exclusive from this region. We studied similar forms from Farallones de Cali (Cordillera Occidental, Colombia), Nariño (southern Colombia), Tungurahua (Ecuador), and from Ancash (central Peru). On the other hand, some specimens from the paramos of Chingaza and Sumapaz (Cordillera Oriental, Colombia) display unusual narrow leaves that Cuatrecasas named in sched. as W. humilis var. angustifolia (Cuatrecasas 25628, US-01269081; Cleef 1316, US-01324876). From label information, it can be understood that these plants grow almost submerged in the water, which would explain the singular morphology of these forms. The Bolivian populations, which represent the southern limit of distribution of X. humile, rather exhibit a morphology similar to the typical forms. The puzzling pattern of the different morphologies along the whole distribution area makes unadvisable the recognition of more than a single species.

Xenophyllum humile shows morphological affinities with X. funkianum, X. rigidum, X. roseum, and X. sotarense (see comments under these species). Some specimens from Cundinamarca (Colombia) have the rhizome-like stems covered with a dense lanate indumentum and with scarce leaves along them (e.g., Pedraza & Franco 452, COL; Cleef & Florschütz 5469, COL, US). They might be confused with X. crassum subsp. orientale (Cuatrec.) J.Calvo, although their distributions areas do not overlap and X. humile has shorter and narrower leaves, shorter involucral bracts, and smaller capitula. We also studied a few specimens containing material mixed with Werneria pygmaea Gillies ex Hook. & Arn., a strictly rosettiform species that can be readily differentiated because it grows in lax clumps or solitary plants and its leaves are acute to obtuse at the apex and have the leaf lamina barely spreading from the sheath-like base (vs. leaves obtuse to truncate at the apex or callous-like tipped, usually with the leaf lamina spreading at nearly 90° from the sheath-like base in X. humile); see Cuatrecasas & Idrobo 27005 (COL) and Soderstrom 1293 (US).

An isolectotype of the name Werneria humilis Kunth is kept at the Willdenow Herbarium. This specimen was sent by Humboldt to Willdenow as stated at the bottom right of the sheet. It is not numbered but contains individuals identical to those from the lectotype at P.

Regarding the name Werneria fontii, it should be noted that the three specimens at MA are numbered with the same herbarium number, i.e., MA-246359. The isolectotypes, therefore, are indicated as MA-246359b and MA-246359c in order to differentiate them from the lectotype.

Description.

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. Stems 1–1.5 cm tall (aerial part), sparsely pilose. Leaves densely stellate-imbricate, extending into a sheath-like base that bears scattered multicellular trichomes; leaf laminas 4–5 × 1–1.2 mm, linear, aristate at the apex (arista up to 0.5 mm long that falls as the leaves age), entire, thinly hyaline (in dried specimens), elliptical in cross section, glabrous, unconspicuously nerved above, 1-nerved beneath (barely visible), fleshy, drying coriaceous, matte, finely papillose. Capitula radiate, erect, sessile to subsessile. Involucres 4.9–5 × 4.8–5 mm, cupuliform; involucral bracts 12 to 13, 3.2–4 × 1.2–1.3 mm, obtuse at the apex, greenish. Ray florets ca. 13; corollas 6.3–6.5 × 0.8–1.2 mm, 2 to 4-veined, subentire to 2-toothed at the apex, conspicuously surpassing the involucre, white. Disc florets 14 to 15; corollas 3.3–4 mm long, whitish, usually purple-tipped; style branches truncate with a crown of sweeping trichomes, purplish. Achenes ca. 2.2 × 0.5 mm, cylindrical, 7 to 8-ribbed, glabrous; pappus 4.8–6.2 mm long, barbellate, whitish. Chromosome number unknown. Fig. ​Fig.2B2B.

Additional iconography.

Blake (1928: 496, fig. 1Q–S sub Werneria leucobryoides).

Distribution and habitat.

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. ​(Fig.99).

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Figure 9.

Distribution map of Xenophyllum sotarense (left hand, closed triangle), X. roseum (left hand, open circle), and X. funkianum (right hand).

Phenology.

Flowering from July to December.

Etymology.

The epithet sotarense refers to the Sotará Volcano, the locus classicus of this species. It is located in southern Colombia on the border between the departments of Cauca and Huila.

Notes.

Xenophyllum sotarense is characterized by its linear, 4–5 mm long, densely stellate-imbricate leaves, which are typically aristate at the apex. The arista measures up to 0.5 mm long and falls as the leaf ages. The sheath-like base bears scattered multicellular trichomes.

It is morphologically close to X. humile, especially to those small and compact plants with leaf laminas barely spreading from the sheath-like bases (see comments under it). To avoid misidentifications the following characters should be carefully studied: leaf apex (aristate or clearly acute when the arista falls in X. sotarense vs. obtuse to truncate in X. humile) and stem indumentum (sparsely pilose in X. sotarense vs. lanate in X. humile). Xenophyllum sotarense also shows morphological affinities with X. roseum. The involucre length (4.9–5 mm in X. sotarense vs. 10–12 mm in X. roseum), the ray corolla length and color (6.3–6.5 mm, white in X. sotarense vs. 11.8–19 mm, pink in X. roseum), and the achene indumentum (glabrous in X. sotarense vs. white-villous in X. roseum) are useful characters for differentiating them.

The lectotype of Werneria sotarensis Hieron. at MA consists in a single sterile fragment barely informative and no isolectotypes appear to be extant; the duplicate at B was destroyed in 1943 (F0BN015821). For that reason, an epitype collected in the locus classicus has been designated. It is noteworthy that the plants from this locality are restricted on the volcano summit and co-occur with X. humile, a species much more abundant in the area that was also found at lower elevations.

The isotypes of Werneria leucobryoides S.F.Blake located at the Ecuadorian herbaria Q and QPLS show that the collector of this material was the Italian naturalist Enrico Festa instead of Luis Sodiro as the protologue indicates. This correction has been adopted since these specimens are the only ones that bear a label handwritten by Sodiro. It is feasible to think that the collector information was corrected on the specimens belonging to Sodiro’s personal herbarium but omitted on the duplicates to be distributed.

Description.

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. Stems 1.5–2 cm tall (aerial part), lanate. Leaves stellate-imbricate, extending into a sheath-like base that bears long silky trichomes, usually with the leaf lamina spreading at nearly 90° from the sheath-like base; leaf laminas 4.6–10.6 × 1.3–1.5 mm, linear, obtuse, callous-like tipped at the apex (the young ones usually bearing a quickly deciduous arista up to 0.5 mm long), entire, elliptical to almost terete in cross section (rather flat when dried), glabrous, unconspicuously nerved above, 1-nerved beneath (barely visible; in dried specimens the nerves are noticeable on both faces), fleshy, matte, papillose. Capitula radiate, erect, sessile. Involucres 10–12 × 7.9–8 mm, cupuliform; involucral bracts ca. 13, 5.5–7.6 × 1.6–2 mm, acute at the apex, greenish, purple-edged. Ray florets 11 to 13; corollas 11.8–19 × 2.5–3 mm, 4-veined, subentire to 3-toothed at the apex, conspicuously surpassing the involucre, pink. Disc florets 28 to 34; corollas 5.7–8 mm long, yellowish; style branches penicillate, yellowish. Achenes cylindrical, white-villous with trichomes ca. 0.5 mm long (immature); pappus ca. 7 mm long, barbellate, whitish. Chromosome number unknown. Fig. ​Fig.88.

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Figure 8.

Xenophyllum roseumA habit B capitulum C adaxial leaf surface D detail of leaf surface E ray corolla and style F disc corolla and style branches G disc corolla and stamens (vertically sectioned, style removed) H style I immature achene with pappus. All details drawn from Jameson s.n. (US) except for A (drawn from Funk & Montezuma 11441, US). Illustration by Alice Tangerini.

Distribution and habitat.

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. ​(Fig.99).

Xenophyllum roseum is a narrow endemic species only known from El Cajas National Park (Azuay) and the Culebrillas Lagoon (Cañar). A single collection from Loja (Jameson s.n., US) would also support its presence in this province. However, the label information does not provide a precise locality (“province of Loxa”), and therefore, it has not been represented on the map. New collections are required to confirm that X. roseum thrives in Loja.

Phenology.

Flowering from August to March.

Etymology.

The adjective roseus -a -um means rose-like, rose-colored, pink-colored. It refers to the characteristic pink ray corollas of this species.

Notes.

Xenophyllum roseum is well defined by its linear, stellate-imbricate leaves that usually have the leaf lamina spreading at nearly 90° from the sheath-like base, and by the obtuse leaf apex that usually bears a quickly deciduous arista when young. The achenes are covered with a white-villous indumentum and the ray corollas are pink. This latter character makes this species unique within the genus. The color is very noticeable on living plants but in old dried specimens it might be less obvious, and therefore, somewhat equivocal for identification purposes.

It is morphologically similar to X. humile and X. sotarense. From the former species, it differs in the ray corolla length and color (11.8–19 mm, pink in X. roseum vs. 4.2–12.2 mm, white in X. humile), disc corolla length and color (5.7–8 mm, yellowish in X. roseum vs. 3.1–5 mm, whitish in X. humile), and achene indumentum (white-villous in X. roseum vs. glabrous in X. humile). The deciduous arista that usually adorns the young leaves of X. roseum also helps to differentiate it from X. humile. With respect to X. sotarense, the differences are detailed under this latter species. Among the aforementioned species, X. roseum overlaps its distribution area with that of X. humile.

Type.

Ecuador. Chimborazo: Mt. Chimborazo area, at the end of Polylepis road and beginning of hike to Polylepis forest, 1°31’50’’S, 78°52’55’’W, 4233 m, 20 Apr 2018, V.A. Funk & J.M. Bonifacino 14059 (holotype: US!; isotypes: MO!, QCA!).

Description.

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. Stems 2–3 cm tall (aerial part), arachnoid. Leaves imbricate, extending into a sheath-like base that bears arachnoid trichomes; leaf laminas 5.3–7.8 × 0.8–0.9 mm, linear, rather acute, callous-like tipped at the apex, entire, elliptical in cross section, glabrous, unconspicuously nerved above, 1-nerved beneath (only visible in the lower third), fleshy, shiny, papillose. Capitula radiate, erect, sessile. Involucres 9–11 × 5–7 mm, cupuliform; involucral bracts 13 to 14, 4.7–6.9 × 1–1.7 mm, acute at the apex, dark-burgundy. Ray florets 12 to 13; corollas 8.9–11.6 × 2.3–3 mm, 4-veined, subentire to 3-toothed at the apex, conspicuously surpassing the involucre, white, somewhat purplish beneath. Disc florets 20 to 23; corollas 5–5.6 mm long, yellowish; style branches truncate with a crown of sweeping trichomes, yellowish. Achenes cylindrical, white-villous (immature); pappus 3.9–6.2 mm long, barbellate, whitish. Chromosome number unknown. Fig. ​Fig.1010.

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Figure 10.

Xenophyllum funkianumA habit B habit with capitula C stem apical part D detail of leaf apex E capitulum F ray floret (frontward bristles removed) G disc corolla and style branches H stamen. All details drawn from Funk & Bonifacino 14059 (US). Illustration by Alice Tangerini.

Additional iconography.

Calvo and Funk (2020: 32, fig. 2, as photo).

Distribution and habitat.

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. ​(Fig.99).

The species is known from a few collections near the Chimborazo Volcano. Its distribution area partially overlaps with those of X. humile and X. rigidum. See Calvo and Funk (2020) for further details.

Phenology.

Flowering from April to July.

Etymology.

The epithet honors the American botanist Vicki A. Funk (1947–2019), who described the genus Xenophyllum and greatly contributed to the understanding of the family Compositae.

Notes.

Xenophyllum funkianum is distinguished by its creeping rhizome-like stems 20–35 cm long, the straight linear leaves extending into a sheath-like base that bears arachnoid trichomes, the dark-burgundy sessile involucres with 13 to 14 involucral bracts, the 12 to 13 ray florets with white corollas somewhat purplish beneath, and by having white-villous achenes.

The morphologically closest species is X. rigidum. However, they can be differentiated by the leaf lamina size (5.3–7.8 × 0.8–0.9 mm in X. funkianum vs. 12.6–13.5 × 2.3–2.4 mm in X. rigidum), involucre size (9–11 × 5–7 mm in X. funkianum vs. 11.5–12.8 × 7.2–11.3 mm in X. rigidum), number of disc florets (20 to 23 in X. funkianum vs. 38 to 41 in X. rigidum), and by the way of growing of the rhizome-like stems (distinctly horizontal in X. funkianum vs. rather erect in X. rigidum). Xenophyllum funkianum is, furthermore, a smaller plant and the capitula are not enclosed among the leaves as in X. rigidum. Another similar species is X. humile; they differ in involucre length (9–11 mm in X. funkianum vs. 4.6–9.3 mm in X. humile), achene indumentum (white-villous in X. funkianum vs. glabrous in X. humile), and habit (rather creeping in X. funkianum vs. forming dense mats or hummocks in X. humile). Moreover, X. funkianum differs in having the leaf lamina barely spreading from the sheath-like base (vs. leaf lamina usually spreading at nearly 90° from the sheath-like base in X. humile).

Description.

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. Stems 2.5–4 cm tall (aerial part), lanate. Leaves stellate-imbricate, extending into a sheath-like base that bears long silky trichomes; leaf laminas 12.6–13.5 × 2.3–2.4 mm, linear, obtuse at the apex, entire, elliptical in cross section, glabrous, unconspicuously nerved above, 1-nerved beneath (only visible in the lower half), fleshy, drying coriaceous, rather matte, papillose. Capitula radiate, erect, sessile, completely enclosed among the leaves. Involucres 11.5–12.8 × 7.2–11.3 mm, cupuliform; involucral bracts ca. 13, 6.9–8.5 × 1.7–2 mm, rather acute at the apex, dark-purplish. Ray florets ca. 13; corollas 9.6–11 × 1.5–1.9 mm, 4 to 6-veined, subentire to 3-toothed at the apex, conspicuously surpassing the involucre, white. Disc florets 38 to 41; corollas 5.1–6.2 mm long, whitish, usually yellow-tipped; style branches truncate with a crown of sweeping trichomes or slightly penicillate, purplish. Achenes 2.9–3.8 × 1–1.1 mm, cylindrical, white-villous with trichomes ca. 1 mm long (ribs invisible); pappus 6.8–8.4 mm long, barbellate, whitish. Chromosome number unknown. Fig. ​Fig.1111.

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Figure 11.

Xenophyllum rigidumA habit B capitulum C adaxial leaf surface D ray floret (pappus removed) E detail of ray corolla apex F style of ray floret G disc floret (frontward bristles removed) H style of disc floret I stamen. All details drawn from Asplund 8391 (US) except for A (drawn from Grubb et al. 569, NY), C (drawn from Asplund 17334, NY), and G–I (drawn from Anthony & Tate 316, US). Illustration by Alice Tangerini.

Additional iconography.

Funk (1997b: 33, fig. 3D).

Distribution and habitat.

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. ​(Fig.1212).

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Figure 12.

Distribution map of Xenophyllum rigidum (left hand), X. crassum subsp. crassum (right hand, closed triangle), and X. crassum subsp. orientale (right hand, open circle).

Phenology.

Flowering from April to November.

Etymology.

The epithet rigidum means stiff, inflexible and it describes the leaves of this species.

Notes.

Xenophyllum rigidum is readily distinguished by its tough, linear, and apically obtuse leaves, as well as by its white-villous achenes. The rhizome-like stems are very robust and covered with a matted lanate indumentum where the leaf bases are sunked in. Another characteristic feature of this species is that the capitula are completely enclosed among the leaves.

Xenophyllum rigidum is morphologically close to X. funkianum, X. crassum (S.F.Blake) V.A.Funk subsp. crassum (see comments under these taxa), and X. humile. From the latter species, it differs in having larger leaves (12.6–13.5 × 2.3–2.4 mm vs. 2.5–10.8 × 0.5–1.5 mm in X. humile), longer involucral bracts (6.9–8.5 mm vs 3.1–6.1 mm in X. humile), higher number of disc florets (38 to 41 vs. (8–)22 to 39 in X. humile), and white-villous achenes (vs. glabrous in X. humile).

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 Hind and Jeffrey (2001) for further details on Humboldt and Bonpland’s material in the Willdenow Herbarium.

Description.

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. Stems 1.5–4 cm tall (aerial part), lanate. Leaves stellate-imbricate, extending into a sheath-like base that bears long silky trichomes, usually with the leaf lamina spreading at nearly 90° from the sheath-like base; leaf laminas 6.5–27.6 × 1.2–2.2 mm, linear, obtuse, usually callous-like tipped at the apex, entire, flat to elliptical in cross section, glabrous, 1-nerved above, 1-nerved beneath, fleshy, matte or shiny. Capitula radiate, erect, sessile to subsessile (rarely with a short peduncle up to 8 mm long). Involucres 10.2–19.7 × 7.8–15.8 mm, cupuliform; involucral bracts 11 to 21, 6–14.9 × 1.6–2.2 mm, rather acute at the apex, greenish to dark-purplish. Ray florets 13 to 21; corollas 13–19.7 × 1.7–3.4 mm, 4 to 6-veined, subentire to 3-toothed at the apex, conspicuously surpassing the involucre, white. Disc florets 50 to 95; corollas 6.6–8.1 mm long, yellow; style branches truncate with a crown of sweeping trichomes, yellowish, purple-tipped. Achenes 2.6–4 × 0.8–1 mm, cylindrical, 6 to 8-ribbed, glabrous; pappus 7.8–19 mm long, barbellate, whitish. Chromosome number unknown.

Description.

Stems 2.5–4 cm tall (aerial part), lanate. Leaves stellate-imbricate, extending into a sheath-like base that bears long silky trichomes, usually with the leaf lamina spreading at nearly 90° from the sheath-like base; leaf laminas 14.7–27.6 × 1.2–2.2 mm, linear, obtuse, usually callous-like tipped at the apex, entire, flat to elliptical in cross section, glabrous, 1-nerved above, 1-nerved beneath, fleshy, rather matte. Capitula radiate, erect, sessile to subsessile. Involucres 12.6–19.7 × 11–15.8 mm, cupuliform; involucral bracts 16 to 21, 8.6–14.9 × 1.5–2.2 mm, acute at the apex, greenish to dark-purplish. Ray florets 16 to 21; corollas 14.5–19.7 × 2.9–3.4 mm, 4 to 6-veined, subentire to 3-toothed at the apex, conspicuously surpassing the involucre, white. Disc florets ca. 95; corollas 6.5–8.1 mm long, yellow. Achenes 2.6–4 × 0.8–1 mm, cylindrical, 6 to 8-ribbed, glabrous; pappus 14–19 mm long, barbellate, whitish. Chromosome number unknown. Fig. ​Fig.1313.

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Figure 13.

Xenophyllum crassum subsp. crassumA habit B capitulum C adaxial leaf surface D ray floret (frontward bristles removed) E style of ray floret F disc floret (frontward bristles removed) G stamen H style of disc floret. All details drawn from Funk & Gavilanes 11070 (US). Illustration by Alice Tangerini.

Additional iconography.

Funk (1997b: 33, fig. 3A, sub X. crassum).

Distribution and habitat.

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. ​(Fig.1212).

Phenology.

Flowering nearly all year round.

Etymology.

The adjective crassus -a -um means thick and it refers to the robust rhizome that this species displays.

Notes.

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 X. crassum subsp. orientale by the leaf length (14.7–27.6 mm vs. 6.5–12.7 mm in subsp. orientale), number and length of the involucral bracts (16 to 21, 8.6–14.9 mm vs. 11 to 13, 6–8.2 mm in subsp. orientale), number of ray florets (16 to 21 vs. 13 to 15 in subsp. orientale), number of disc florets (ca. 95 vs. ca. 50 in subsp. orientale), and pappus length (14–19 mm vs. 7.8–8 mm in subsp. orientale). Both taxa have a well defined distribution area. The typical subspecies grows along the Cordillera Central and Cordillera Occidental in Colombia and extends southwards to central Ecuador. In contrast, X. crassum subsp. orientale is restricted in the northern part of the Cordillera Oriental of Colombia.

The habit of X. crassum subsp. crassum also resembles that of X. rigidum, but X. crassum subsp. crassum differs in having longer and narrower leaves (14.7–27.6 × 1.2–2.2 mm vs. 12.6–13.5 × 2.3–2.4 mm in X. rigidum), higher number of involucral bracts and ray florets (16 to 21 vs. ca. 13 in X. rigidum), and by displaying glabrous achenes (vs. white-villous in X. rigidum). Their distribution areas only overlap in the Antisana Volcano (Napo, Ecuador).

Description.

Stems 1.5–2.5 cm tall (aerial part), lanate. Leaves stellate-imbricate, extending into a sheath-like base that bears long silky trichomes, usually with the leaf lamina spreading at nearly 90° from the sheath-like base; leaf laminas 6.5–12.7 × 1.2–2 mm, linear, obtuse, usually callous-like tipped at the apex, entire, flat to elliptical in cross section, glabrous, 1-nerved above, 1-nerved beneath, fleshy, rather shiny (in dried specimens). Capitula radiate, erect, sessile to subsessile (rarely with a short peduncle up to 8 mm long). Involucres 10.2–12.2 × 7.8–10.6 mm, cupuliform; involucral bracts 11 to 13, 6–8.2 × 1.6–2.1 mm, acute to obtuse at the apex, greenish to dark-purplish. Ray florets 13 to 15; corollas 13–13.2 × 1.7–2.6 mm, 4 to 5-veined, subentire to 3-toothed at the apex, conspicuously surpassing the involucre, white. Disc florets ca. 50; corollas 6.6–6.9 mm long, yellow. Achenes cylindrical, glabrous (immature); pappus 7.8–8 mm long, barbellate, whitish. Chromosome number unknown.

Distribution and habitat.

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. ​(Fig.1212).

Phenology.

Flowering nearly all year round.

Etymology.

The adjective orientalis -is -e means eastern, which refers to the further eastern distribution of this taxon in relation to the typical subspecies. It occurs in the northern part of the Cordillera Oriental of Colombia (Eastern Ranges).

Notes.

Xenophyllum crassum subsp. orientale is characterized by the leaf length (6.5–12.7 mm), the number of involucral bracts (11 to 13), and the number of ray florets (13 to 15). In dried specimens, the leaves are rather shiny. This taxon was synonymized with X. crassum subsp. crassum by Ávila et al. (2016) but no discussion supporting such taxonomic decision was provided. The aforementioned morphological differences against the typical subspecies, in addition to the fact that their distribution areas are well defined and do not overlap, lead us to maintain the treatment by Cuatrecasas (1980).

It should be noted that X. crassum subsp. orientale has been confused with Werneria pygmaea, a species that grows in the same habitat. Indeed, we detected several specimens that contain mixed material, including an isotype (see comments below). Xenophyllum crassum subsp. orientale has erect, robust rhizome-like stems whereas W. pygmaea is a strictly rosettiform species not forming hummocks. Further differences are detailed in Calvo et al. (in press). Some confusion with X. humile is also likely (see comments under it).

The presumed isotype of W. crassa subsp. orientalis Cuatrec. kept at COL (COL-000005514) comes from “extremo S en Peralonso, 3100 m”, a locality slightly different from that indicated in the protologue, i.e., “vertiente norte, 3700−3800 m” [as “3600−3800 m” on the labels]. It is excluded from the type material as it does not belong to the same gathering. On the other hand, it has to be noted that the isotype at BC contains an individual corresponding to W. pygmaea (above on the right hand, with fragments of Disterigma sp. (Ericaceae) at the base).

Description.

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. Stems 1–2 cm tall (aerial part), glabrous. Leaves densely imbricate, extending into a glabrous sheath-like base; leaf laminas 3.3–7.6 × 1.7–2.6 mm, triangular, acute at the apex, denticulate in the upper half, rather flat in cross section, glabrous, 1-nerved above (barely visible), 1-nerved beneath, rather fleshy, matte. Capitula radiate, erect, sessile to subsessile. Involucres 8.8–10.8 × 5.7–10 mm, cupuliform; involucral bracts 13 to 15, 4.8–6.7 × 1.8–2.2 mm, acute at the apex, greenish to dark-purplish. Ray florets 14 to 21; corollas 7.4–12.9 × 1.6–2.3 mm, 3 to 4-veined, subentire to 2-toothed at the apex, conspicuously surpassing the involucre, white. Disc florets 35 to 48; corollas 5–6.2 mm long, yellowish; style branches truncate with a crown of sweeping trichomes, purplish. Achenes 2.8–3 × ca. 1 mm, cylindrical, ca. 7-ribbed, glabrous; pappus 5–8.8 mm long, barbellate, whitish to partially purple-colored. Chromosome number unknown. Fig. ​Fig.1414.

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Figure 14.

Xenophyllum marcidumA habit B stem apical part and capitulum C capitulum D adaxial leaf surface E uppermost leaf F ray corolla and style G disc corolla and style branches H disc corolla and stamens (vertically sectioned, style removed) I style J achene with pappus. All details drawn from Funk & Bernal 11285 (US) except for A (drawn from Stafford 749, K) and C–E, H (drawn from Macbride & Featherstone 2475, US). Illustration by Alice Tangerini.

Additional iconography.

Blake (1928: 496, fig. 1H–J sub Werneria marcida, fig. 1N–P sub W. sedoides); Funk (1997a: 237, fig. 1B); Beltrán (2016: 358, fig. 2B, as photo).

Distribution and habitat.

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. ​(Fig.1515).

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Figure 15.

Distribution map of Xenophyllum marcidum (left hand) and X. ciliolatum (right hand).

Phenology.

Flowering from February to September.

Etymology.

The adjective marcidus -a -um means withered. It probably refers to the fact that a large proportion of the stem bears marcescent leaves and the green ones are restricted to the distal part.

Notes.

Xenophyllum marcidum is characterized by its triangular, densely imbricate leaves. Among the Xenophyllum members forming dense mats or hummocks, this species is readily distinguishable due to the leaves are denticulate in the upper half, acute at the apex, and rather flat in cross section. The capitula are radiate and have 14 to 21 ray florets with white, considerably long corollas.

Because of the marginally denticulate leaves, on dried specimens X. marcidum might be confused with X. ciliolatum, a species that has a partially overlapping distribution area. They can be easily differentiated by the habit (forming dense mats or hummocks in X. marcidum vs. forming clumps of erect or decumbent stems in X. ciliolatum), leaf lamina shape (triangular in X. marcidum vs. linear in X. ciliolatum), number of involucral bracts (13 to 15 in X. marcidum vs. ca. 8 in X. ciliolatum), number of ray florets (14 to 21 in X. marcidum vs. 8 to 11 in X. ciliolatum), and by the ray corolla length and color (7.4–12.9 mm, white in X. marcidum vs. 6.2–7.3 mm, yellow in X. ciliolatum).

Description.

Suffruticose plant, forming clumps of erect or decumbent stems. Rhizomes 5–8 × 0.3–0.5 cm, horizontal to oblique, glabrous. Stems 4–7 cm tall, simple or branched, glabrous, with leaves usually clustered in the upper part. Leaves imbricate, erect (not adpressed to the stem), extending into a glabrous sheath-like base; leaf laminas 4.9–8.6 × 0.6–1.2 mm, linear, rather acute at the apex, denticulate, elliptical to slightly curved forwards in cross section, glabrous, 1-nerved above (barely visible), 1-nerved beneath (sometimes purple-colored), rather fleshy, matte. Capitula radiate, erect, sessile. Involucres 8.9–12.4 × 4–7.3 mm, narrowly cupuliform; involucral bracts ca. 8, 4.1–7.7 × 1.4–2.5 mm, acute at the apex, greenish to dark-purplish. Ray florets 8 to 11; corollas 6.2–7.3 × 0.8–1.2 mm, 3-veined, subentire to 2-toothed at the apex, not surpassing the involucre, yellow. Disc florets 11 to 28; corollas 5–7.3 mm long, yellow; style branches truncate with a crown of sweeping trichomes, yellowish. Achenes 2–3.3 × 1–1.2 mm, cylindrical, 7 to 8-ribbed, glabrous; pappus 6.2–7.4 mm long, barbellate, whitish. Chromosome number unknown.

Iconography.

Beltrán (2016: 358, fig. 2E, as photo).

Distribution and habitat.

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. ​(Fig.1515).

Phenology.

Flowering from February to November.

Etymology.

The adjective ciliolatus -a -um is the diminutive of ciliatus -a -um, which describes the denticulate leaf margin of this species.

Notes.

Xenophyllum ciliolatum is one of the species displaying yellow ray corollas not surpassing the involucre. It is characterized by having erect or decumbent stems 4–7 cm tall, linear leaves 4.9–8.6 mm long with denticulate margin, ca. 8 involucral bracts, and 8 to 11 ray florets. The leaves are erect (not adpressed to the stem) and tend to be clustered upwards.

The morphologically closest species is X. juniperinum, known from southern Peru (Tacna), northern Chile, and western Bolivia (Oruro and Potosí). They differ in habit (erect or decumbent stems 4–7 cm long in X. ciliolatum vs. erect stems 8–15 cm long in X. juniperinum), leaf disposition (clustered up the stem and erect in X. ciliolatum vs. uniformly arranged along the stem and adpressed in X. juniperinum), leaf lamina shape (linear in X. ciliolatum vs. linear-triangular in X. juniperinum), leaf margin (denticulate in X. ciliolatum vs. minutely, irregularly denticulate in X. juniperinum), and involucral bract length (4.1–7.7 mm in X. ciliolatum vs. 2.8–3.4 in X. juniperinum). Their distribution areas do not overlap. Xenophyllum ciliolatum also shows morphological affinities with X. marcidum (see comments under it) and X. weddellii. From this latter species, X. ciliolatum differs in having denticulate leaf margin (vs. entire in X. weddellii), yellow ray corollas not surpassing the involucre (vs. white and conspicuously surpassing the involucre in X. weddellii), and disc corollas 5–7.3 mm long (vs. 4.5–5.2 mm long in X. weddellii). These species co-occur in southern Peru (Arequipa, Moquegua, Puno).

The lectotype designation of the name Werneria ciliolata A.Gray by Calvo et al. (2017) should be rather attributed to Funk (1997a) although she used the term “holotype”. It is here corrected to lectotype because it cannot be established that the author only used this element and that the gathering is represented by a single specimen (McNeill 2014).

Description.

Suffruticose plant, forming clumps of erect stems. Rhizomes 5–10 × 0.4–0.6 cm, horizontal to oblique, glabrous. Stems 8–15 cm tall, usually simple, glabrous, with leaves uniformly arranged along it. Leaves imbricate, adpressed to the stem, extending into a glabrous sheath-like base; leaf laminas 5–7 × 1.1–2 mm, linear-triangular, rather acute at the apex, minutely, irregularly denticulate, obtusely triangular in cross section, glabrous, 1-nerved above (barely visible), 1-nerved beneath, fleshy, drying coriaceous, matte. Capitula radiate, erect, sessile. Involucres 7–7.7 × 4–6.2 mm, narrowly cupuliform; involucral bracts 8 to 11, 2.8–3.4 × 1.8–2.8 mm, acute at the apex, dark-purplish. Ray florets 8 to 13; corollas 4.5–5.5 × 0.4–0.7 mm, unconspicuously veined, subentire to slightly 3-toothed at the apex, not surpassing the involucre, yellow. Disc florets 28 to 36; corollas 5.5–5.7 mm long, yellow; style branches truncate with a crown of sweeping trichomes, yellowish. Achenes 3.4–3.9 × 0.8–1 mm, cylindrical, 7 to 8-ribbed, glabrous; pappus 5.2–9 mm long, barbellate, whitish. Chromosome number unknown. Figs ​Figs2C,2C, ​,1616.

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Figure 16.

Xenophyllum juniperinumA habit B stem apical part and capitula C, D uppermost leaves and capitulum (notice leaf variability) E adaxial leaf surface F ray floret (frontward bristles removed) G ray floret (pappus removed) H disc floret (frontward bristles removed) I stamen J style. A, B drawn from Werdermann 1164 (GH), C–E (drawn from Werdermann 1164, CAS), and F, G (drawn from Werdermann 1164, US). Illustration by Alice Tangerini.

Additional iconography.

Blake (1928: 496, fig. 1K–M sub Werneria lycopodioides); Beltrán (2016: 358, fig. 2A, sub X. lycopodioides, as photo); Calvo et al. (2017: 228, fig. 1, as photo).

Distribution and habitat.

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. ​(Fig.1717).

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Figure 17.

Distribution map of Xenophyllum juniperinum (left hand), X. weddellii (right hand, closed triangle), and X. incisum (right hand, open circle).

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.

Phenology.

Flowering from April to December.

Etymology.

It is named after the genus Juniperus L. (Cupressaceae) due to the leaves of this species resembling those of some species belonging to the mentioned genus.

Notes.

Xenophyllum juniperinum forms clumps of erect stems 8–15 cm long, which bear imbricate leaves that are adpressed to the stem and uniformly arranged along it. The leaves are somewhat broadened at the base and the margin is minutely, irregularly denticulate. The yellow ray corollas not surpassing the involucre is a distinctive character of this species, which is shared with X. ciliolatum (see comments under it for comparison purposes).

Although the syntypes of the name Werneria juniperina Hieron. were apparently destroyed at B in 1943, it should be mentioned that Rockhausen (1939) had the opportunity to study them and designated the collection Stübel 106 as “typus”. See Calvo et al. (2017) for further details on the neotypification.

Description.

Suffruticose plant, forming clumps of rather decumbent stems. Rhizomes 7–10 × 0.5–0.8 cm, horizontal to oblique, glabrous. Stems 3–8 cm tall, simple or branched, glabrous, with leaves rather uniformly arranged along it. Leaves imbricate, extending into a glabrous sheath-like base; leaf laminas 3.4–7(–11.8) × 1.3–1.6 mm, linear-subulate, acute at the apex, entire, obtusely triangular in cross section, glabrous, unconspicuously nerved above, 1-nerved beneath (barely visible), fleshy, matte. Capitula radiate, erect, sessile. Involucres 7–9 × 5–7 mm, cupuliform; involucral bracts 8 to 12, 3.2–5.9 × 1.4–2.3 mm, acute to obtuse at the apex, greenish, sometimes purple-edged. Ray florets 9 to 15; corollas 5.9–8 × 1–1.2 mm, 3 to 4-veined, subentire at the apex, conspicuously surpassing the involucre, white, rarely pale purplish beneath. Disc florets 24 to 32; corollas 4.5–5.2 mm long, whitish to creamy, sometimes with the tube purplish; style branches truncate with a crown of sweeping trichomes, purplish. Achenes 3.8–4 × 0.9–1 mm, cylindrical, 8 to 9-ribbed, glabrous; pappus 6.3–7.5 mm long, barbellate, whitish. Chromosome number unknown.

Iconography.

Beltrán (2016: 358, fig. 2F, as photo).

Distribution and habitat.

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. ​(Fig.1717).

Phenology.

Flowering nearly all year round.

Etymology.

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.

Notes.

Xenophyllum weddellii is a species forming clumps of rather decumbent stems with linear, entire, glabrous leaves. It has 8 to 12 involucral bracts and 9 to 15 ray florets with white corollas that conspicuously surpass the involucre. On living plants, the color of the leaves is typically glaucous green.

The leaf apex is acute but we studied a specimen showing two lateral divisions near the apex (Moreira-Muñoz & Diazgranados 2625, SGO). It is identified as X. weddellii because the remaining characters perfectly fit with this species. Two emerging protrusions near the apex were also observed in a few other collections, which indicates that Moreira-Muñoz’s collection can be considered as an extreme of variability.

This species is morphologically similar to X. ciliolatum (see comments under it) and X. incisum. From this latter species, it mainly differs in the leaf lamina shape (linear-subulate, tapering upwards in X. weddellii vs. linear-oblong, similar width from base to apex in X. incisum) and leaf apex (acute in X. weddellii vs. 3-notched with the lobes rather truncate at the apex in X. incisum). It is interesting to mention that both species have similar ecological requirements and they usually thrive near the banks of the salt lagoons and moist valley bottoms. They could be referred to as allopatric species since their distribution areas are well defined and do not overlap.

The single original collection that we found comes from the Huasco Lagoon (Tarapacá, Chile). However, Philippi (1891) indicated additional sites in the protologue, i.e., “Machuca” and “regione inter Copacoya et Inacaliri”. These localities are located further south in the region of Antofagasta, where this species appears to be absent. In contrast, X. incisum is not rare in this region and, indeed, the provenance of the type material is “Inter Copacoya et Inacaliri”. The lack of original material of X. weddellii from these localities and the fact that we did not study any collection from there suggests that Philippi might had confused both species.

The holotype designation of the name Werneria weddellii Phil. has been corrected to lectotype according to McNeill (2014). In addition, the name Werneria decumbens Hieron. is synonymized with X. weddellii as suggested by Funk (1997a). Since the original material was likely destroyed at B in 1943, a neotype is designated. The provenance of the selected collection perfectly matches the protologue information.

Description.

Suffruticose plant, forming clumps of rather decumbent stems. Rhizomes 4–8 × 0.4–0.6 cm, horizontal to oblique, glabrous. Stems 4–8 cm tall, simple or branched, glabrous, with leaves rather uniformly arranged along it. Leaves imbricate, extending into a glabrous sheath-like base; leaf laminas 3.2–7.3 × 1.4–1.9 mm, linear-oblong, 3-notched at the apex, sometimes with the central lobe wider than lateral ones (rarely subentire), entire, obtusely triangular to slightly curved forwards in cross section, glabrous, unconspicuously nerved above, 1-nerved beneath, fleshy, matte; leaf lobes 0.5–1 × 0.5–0.6 mm, obtuse to truncate. Capitula radiate, erect, sessile. Involucres 6.4–8.3 × 4.6–6.1 mm, cupuliform; involucral bracts 8 to 9, 3.1–4.3 × 1.1–2.6 mm, obtuse at the apex, greenish. Ray florets 8 to 13; corollas 6.2–8.9 × 0.6–1.2 mm, 2 to 4-veined, subentire to 3-toothed at the apex, conspicuously surpassing the involucre, white. Disc florets 26 to 37; corollas 4.4–5 mm long, pale yellow to creamy; style branches truncate with a crown of sweeping trichomes, purplish. Achenes 2.3–2.4 × 0.8–0.9 mm, cylindrical, ca. 8-ribbed, glabrous; pappus 4.8–8 mm long, barbellate, whitish. Chromosome number unknown. Fig. 19C.

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Figure 19.

Detail of leaves and capitulum AXenophyllum lorochaqui, drawn from Castellanos s.n. (BA) BX. poposa, drawn from Cárdenas 362 (US) CX. incisum, drawn from Funk et al. 13078 (US). Illustration by Alice Tangerini.

Additional iconography.

Cabrera (1978: 473, fig. 200G, H, sub Werneria incisa).

Distribution and habitat.

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. ​(Fig.1717).

Phenology.

Flowering from October to March.

Etymology.

The epithet incisum means sharply and deeply cut into, which describes the leaf apex of this species. However, its leaf apex is rather notched (shallowly cut into).

Notes.

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.

Xenophyllum incisum is somewhat variable in leaf apex shape. Typical forms have 3-notched leaf apex, usually with the central lobe wider than lateral ones, but in some specimens the apex is barely notched or even subentire (e.g., Moreira-Muñoz 298, CONC; Arroyo et al. 97315, CONC). In these specimens, however, the leaf apex is truncate and slightly curved forwards as typically in the species. Similarly, we studied one specimen having the leaf apex with each lobe 3-notched (Villagrán et al. 9321, CONC).

This species has morphological affinities with X. lorochaqui J.Calvo & V.A.Funk, X. weddellii (see comments under these species), and X. poposa. With regard to the latter species, the main and unequivocal character that allows a proper identification is the stem indumentum (glabrous in X. incisum vs. arachnoid in X. poposa). The characters concerning the leaf apex have traditionally been used for discriminating among these species, however, they are barely useful because X. poposa also shows high variability in leaf apex (see further comments under it). A rather useful character is the leaf section, which is obtusely triangular to slightly curved forwards in X. incisum vs. terete or almost so in X. poposa. The sheath-like base is also more broadened in X. incisum.

The holotype designation by Funk (1997a) is corrected to lectotype because the gathering indicated by Philippi in the protologue is not represented by a single specimen. Moreover, a second-step lectotypification is made because two specimens of this gathering were found at SGO.

Description.

Suffruticose plant, forming clumps of erect stems (rarely dense mats). Rhizomes 4–10 × 0.3–0.7 cm, horizontal to oblique, glabrescent. Stems 2–15 cm tall, simple or branched, arachnoid, usually with functional green leaves restricted to the upper part. Leaves imbricate, extending into a sheath-like base that bears arachnoid trichomes; leaf laminas 2.5–7.1 × 0.5–1.1 mm, linear, rounded to truncate or 2 to 3-notched at the apex (usually white callous-tipped), entire, rather terete in cross section, glabrous, unconspicuously nerved on both faces, fleshy, matte; leaf lobes (when present) 0.6–1.5 × 0.3–0.4 mm, rounded to truncate, usually white callous-tipped. Capitula radiate, erect, sessile. Involucres (3.3–)5.6–8 × (2.7–)3–5.3 mm, narrowly cupuliform; involucral bracts 8 to 9, (1.1–)1.6–3.9 × 0.7–2.1 mm, acute to obtuse at the apex, greenish to partially dark-purplish. Ray florets 8 to 11; corollas 3.6–8.9 × 0.5–1.2 mm, 4-veined (sometimes unconspicuous), subentire to 3-toothed at the apex, conspicuously surpassing the involucre, white. Disc florets 7 to 18; corollas 3.2–5.9 mm long, whitish to creamy (rarely purplish); style branches truncate with a crown of sweeping trichomes, purplish. Achenes 2.8–2.9 × 0.7–0.8 mm, cylindrical, ca. 7-ribbed, glabrous; pappus (2.3–)4.1–5 mm long, barbellate, whitish to partially rose-colored. Figs ​Figs18,18, 19B.

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Figure 18.

Xenophyllum poposaA habit B stem apical part C capitulum D adaxial leaf surface E abaxial leaf surface F ray corolla and style G ray corolla and style (notice staminodes) H disc corolla I disc corolla and stamens (vertically sectioned, style removed) J achene with pappus. All details drawn from Funk et al. 13138 (US) except for A, B, D (drawn from Venturi 2964, US). Illustration by Alice Tangerini.

Additional iconography.

Cabrera (1978: 473, fig. 200I, J, sub Werneria poposa); Freire and Ariza-Espinar (2014: 227, sub X. incisum var. pubescens A–H, 228, X. poposa A–G); Beltrán (2016: 358, fig. 2D, as photo, 359, fig. 3E, sub X. incisum, as photo).

Distribution and habitat.

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. ​(Fig.2020).

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Figure 20.

Distribution map of Xenophyllum poposa (left hand), X. lorochaqui (right hand, closed triangle), and X. rosenii (right hand, open circle).

Phenology.

Flowering nearly all year round.

Etymology.

The epithet poposa is the vernacular name of this plant as Philippi (1891) stated in the protologue “ubi Poposa vocabatur” [called poposa]. He also added “infusum plantae contra dolores colicos propinant” [for relieving stomach cramps and colic] referring to its medicinal properties. It is important to underline that poposa is a noun in apposition not to be declined. The correct epithet is, therefore, poposa instead of the widely misapplied poposum (Funk 1997a; Freire and Ariza-Espinar 2014; Beltrán 2016).

Notes.

Xenophyllum poposa forms clumps of erect stems (rarely dense mats) characterized by having arachnoid indumentum. When the stems are very short and barely protrude from the soil, then, it tends to develop a mat-forming habit. The leaves are rather terete with the sheath-like base weakly broadened at the base. The leaf apex is usually undivided, rounded to truncate, and callous-tipped, however, plants having 2 to 3-notched apex are also found. It has 8 to 9 involucral bracts, 8 to 11 ray florets (with white corollas surpassing the involucre), 7 to 18 disc florets, and glabrous achenes. It is noteworthy that we observed a few ray florets with staminodes (Fig. 18G).

The type material of this species shows plants with clearly truncate and callous-tipped leaf apex, which probably led Rockhausen (1939) to relate a collection displaying notched leaf apex to X. incisum and describe it as Werneria incisa var. pubescens Rockh. He differentiated the new variety from the typical one by the longer leaves and the pubescent-lanate sheath-like bases. A detailed study of the lectotype (CORD-00005636) reveals that the stem bears arachnoid indumentum and that the sheath-like bases are not conspicuously broadened. These characters are distinctive of X. poposa rather than X. incisum. Moreover, this latter species is restricted to the edges of salt lagoon edges and desert plains with a certain humidity. The type material of Rockhausen’s variety was collected in the upper part of the Famatina Range (La Rioja, Argentina), which better matches the habitat of X. poposa (exposed slopes). On this basis, and because we believe that the glabrousness is a diagnostic character of X. incisum, we do not follow Rochkausen’s treatment as previous botanists did (Cabrera 1948; Freire and Ariza-Espinar 2014). Otherwise, we consider it appropriate to recognize W. incisa var. pubescens as part of the variability encompassed by X. poposa. Our decision is also supported by the fact that we studied further specimens corresponding to X. poposa that display notched leaf apex. These populations appear very scattered from southern Peru to northwestern Argentina, namely in Moquegua, Oruro, Potosí, Tucumán, Catamarca, and La Rioja. In Moquegua some specimens are strikingly tiny, barely protruding from the soil, and exhibit 2 to 3-notched leaf apex (Beltrán 7736, USM; Funk et al. 13155, US, USM). Due to field work in this region we can state that these forms grow mixed with plants displaying undivided, rounded to truncate leaf apex. It also deserves to be underlined that the different leaf morphologies can even be found within the same individual. The specimen Funk et al. 13155 (US), for example, has undivided, 2-notched, and 3-notched leaves. In Potosí (Bolivia), we visited another population composed of midsize plants showing 3-notched leaves where each lobe can be, in turn, 2-notched (Calvo & Zárate 7869, BOLV, LPB; Cárdenas 362, US [see Fig. 19B]). No individuals with undivided leaves were found there. On the other hand, the specimens from Famatina (La Rioja, Argentina) are quite robust plants having stems 11–13 cm long and 2 to 3-notched leaves. Based on the study of herbarium specimens, we also can conclude that the populations having notched leaf apex from Oruro, Tucumán, and Catamarca grow mixed with or close to the typical plants. All these examples demonstrate that a great intrapopulation and interpopulation variability exists in terms of both leaf apex and habit. Our efforts for trying to propose an infraspecific classification in pursuit of recognizing the distinct morphologies failed because of the intermediate forms and the lack of defined distribution patterns. Therefore, we believe that X. poposa should be treated as a highly variable species.

The distribution area of this species partially overlaps with that of X. incisum in the region comprised between Antofagasta (Chile), Potosí (Bolivia), Salta, and Jujuy (Argentina). However, they occur in different ecological niches as already mentioned above. The morphological differences between them are commented under X. incisum. Likewise, X. poposa might be confused with X. lorochaqui (see notes under it).

Regarding nomenclatural issues, it is noteworthy that Funk’s typification of the name Werneria poposa Phil. has been narrowed to a single specimen because two duplicates of the same gathering were found at SGO (Turland et al. 2018, ICN Art. 9.17). The specimen SGO-000006433 has therefore been selected as the second-step lectotype because it is most complete and best preserved.

The original material of Werneria lorentziana Hieron. comprises the syntypes Stübel 107 and Stübel 117, both apparently destroyed. Rockhausen (1939) designated as “typus” the collection Stübel 117 but indicated the locality that in the protologue corresponds to the number Stübel 107. Since these specimens are not extant anymore and we did not find pictures of them at F, such mismatch cannot be elucidated. Later, Freire and Ariza-Espinar (2014) misguidedly designated as lectotype the specimen Lorentz & Hieronymus 117 (CORD). Although the collector number coincides with one of the syntypes, this element is not part of the original material and has to be treated as a neotype.

The typification of W. incisa var. pubescens should be attributed to Cabrera and Freire (1999) although they used the term “isotipo”. With this action the authors presumed that a duplicate was kept at B but the specimen probably was destroyed during the Second World War. On this basis, the term isotype is corrected to lectotype (Turland et al. 2018, ICN Art. 9.10). Accordingly, the later lectotypification by Freire and Ariza-Espinar (2014) becomes superfluous.

Type.

Argentina. Catamarca: El Cajón, Negroara, 15 Jan 1914, L. Castillón 3365 (holotype: LIL-26677!; isotypes: BM s.n.!, BR s.n.!, US-00622893!, W-334!).

Description.

Suffruticose plant, forming clumps of erect stems. Rhizomes 5–10 × 0.6–0.8 cm, horizontal to oblique, glabrous. Stems 10–20 cm tall, branched, glabrous, usually with leaves restricted to the upper part. Leaves imbricate, extending into a sheath-like base glabrescent or with evanescent arachnoid trichomes; leaf laminas 7.9–11.5 × 2.3–2.7 mm, linear, broadened at the apex, 3-notched at the apex, with the central lobe longer than lateral ones, entire, curved forwards in cross section, glabrous, unconspicuously nerved on both faces, fleshy, matte; central leaf lobe 1–1.6 mm long (lateral ones 0.5–0.8 mm long), 1.3–1.6 mm wide at the maximum width point, entire or barely notched, obtuse. Capitula radiate, erect, sessile. Involucres 10.3–10.9 × 6.6–8.7 mm, cupuliform; involucral bracts ca. 13, 5.9–7 × 1.4–2.5 mm, obtuse at the apex, greenish. Ray florets 26 to 39; corollas 8.3–9.2 × 1.1–1.4 mm, 4-veined, subentire to 3-toothed at the apex, conspicuously surpassing the involucre, white. Disc florets 40 to 57; corollas 4.2–5.5 mm long, yellowish; style branches truncate with a crown of sweeping trichomes, yellowish. Achenes 2.5–3.1 × 0.7–0.9 mm, cylindrical, 8 to 9-ribbed, glabrous; pappus 4.3–6.1 mm long, barbellate, whitish. Chromosome number unknown. Figs 19A, ​,2121.

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Figure 21.

Xenophyllum lorochaquiA habit B stem apical part and capitulum C adaxial leaf surface (left hand) and vertical leaf profile (right hand) D capitulum E ray floret (pappus removed) F disc corolla G stamen H style. All details drawn from Castellanos s.n. (BA) except for A, B (drawn from Díaz s.n., GH). Illustration by Alice Tangerini.

Distribution and habitat.

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. ​(Fig.2020).

Phenology.

Flowering from January to March.

Etymology.

The epithet lorochaqui is the vernacular name of this plant and it means parrot’s foot (“loro” from Spanish: parrot; “chaqui” from Quichua: foot). It probably responds to the resemblance of the leaves with a parrot’s foot.

Notes.

Xenophyllum lorochaqui can be identified by the glabrous erect stems 10–20 cm tall (usually only bearing leaves in the upper part), leaf lamina 7.9–11.5 mm long, leaf apex 3-notched with the central lobe entire or barely notched and longer than the lateral ones, involucre with ca. 13 involucral bracts, and by displaying 26 to 39 ray florets with white corollas.

This species shows morphological affinities with X. incisum, X. dactylophyllum, and X. poposa. The differences against X. incisum are the leaf lamina length (3.2–7.3 mm vs. 7.9–11.5 mm in X. lorochaqui), the length of the leaf apex lobes (similar among them vs. central lobe longer than lateral ones in X. lorochaqui), the involucre length (6.4–8.3 mm vs 10.3–10.9 mm in X. lorochaqui), the involucral bract number (8 to 9 vs. ca. 13 in X. lorochaqui), and the ray floret number (8 to 13 vs. 26 to 39 in X. lorochaqui). With regard to X. dactylophyllum, the leaf apex is at least 9-divided (finger-like) with the primary division extending deeper than the subsequent ones, whereas in X. lorochaqui the leaf apex is 3-notched with the central lobe longer than lateral ones. From X. poposa, the distinguishable characters are the stem indumentum (arachnoid in X. poposa vs. glabrous in X. lorochaqui), the leaf lamina length (2.5–7.1 mm in X. poposa vs. 7.9–11.5 mm in X. lorochaqui), the involucral bract number (8 to 9 in X. poposa vs. ca. 13 in X. lorochaqui), and the ray floret number (8 to 11 in X. poposa vs. 26 to 39 in X. lorochaqui). Among these species, the distribution area of X. lorochaqui partially overlaps with that of X. incisum and X. poposa.

Description.

Suffruticose plant, forming clumps of erect stems. Rhizomes 6–10 × 0.4–0.5 cm, horizontal to oblique, glabrous. Stems 4–8 cm tall, simple or branched, glabrous, usually with leaves restricted to the upper part. Leaves imbricate, extending into a glabrous sheath-like base; leaf laminas 7–11 × 3–4.5 mm, linear-spatulate (broadened upwards), 2-forked at the apex (incision depth ca. 1/3 of the total length), entire, flat to elliptic or almost terete upwards in cross section, glabrous, 1-nerved above (barely visible), 1-nerved beneath, fleshy, matte; leaf lobes 2.3–4.5 × 1–1.5 mm, acute to obtuse. Capitula radiate, erect, sessile. Involucres 11–13.2 × 8–10 mm, cupuliform; involucral bracts 11 to 12, 5.6–7.2 × 1.9–3.2 mm, rather acute at the apex, greenish, purple-edged. Ray florets ca. 18; corollas ca. 9.2 × 1.7 mm, 3 to 4-veined, subentire to 3-toothed at the apex, conspicuously surpassing the involucre, white. Disc florets ca. 50; corollas 5–5.1 mm long, yellowish; style branches penicillate, purplish. Achenes ca. 3.5 × 0.9 mm, cylindrical, 6 to 8-ribbed, glabrous; pappus 8–10 mm long, barbellate, whitish. Chromosome number unknown.

Iconography.

Cabrera (1948: 52, fig. 2, sub Werneria rosenii).

Distribution and habitat.

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. ​(Fig.2020).

Xenophyllum rosenii is hitherto known from very few gatherings, i.e., the type locality in Jujuy (Argentina) plus two collections from southern Sur Lípez (Bolivia). Jørgensen (2014) also indicated this species from Oruro (Bolivia) on the basis of a misidentified collection that actually corresponds to X. digitatum (Liberman 352, LPB).

Phenology.

Collected in flower in March and November.

Etymology.

It is named after Eric von Rosen (1879–1948), a Swedish aristocrat, explorer, and ethnographer.

Notes.

Xenophyllum rosenii is unique in the genus by its 2-forked leaves. The species has been confused with X. digitatum (e.g., Meneses 5446, LPB; Treviño 761, HSP; Tupayachi 736, US). A detailed study of the leaves highlights that these species can be straightforwardly discriminated from one another by the leaf apex (2-forked in X. rosenii vs. 3-forked in X. digitatum). The other characters appear to be useless because of a significant overlapping among them.

Since two specimens of the gathering Fries 862 were found at UPS, the lectotypification has been further narrowed to a single specimen by way of a second-step lectotype (UPS-V-833144).

Description.

Suffruticose plant, forming clumps of erect stems. Rhizomes 5–12 × 0.5–0.9 cm, horizontal to oblique, glabrous. Stems 10–13 cm tall, simple or branched, glabrous, usually with leaves restricted to the upper part. Leaves imbricate, extending into a glabrous sheath-like base; leaf laminas 7.7–11.5 × 2.8–6.7 mm, linear-spatulate (broadened upwards), 3-forked at the apex (incision depth 1/3–1/2 of the total length), entire, usually with some minute cilia in the lower half, flat to elliptic or almost terete upwards in cross section, glabrous or arachnoid above (evanescent), 1-nerved above (barely visible), 1-nerved beneath, fleshy, matte; leaf lobes 3.6–5.9 × 0.8–1.2 mm, entire or 2(–3)-divided, acute to obtuse, sometimes mucronate. Capitula radiate, erect, sessile to subsessile. Involucres 12.1–16.7 × 9.8–15.7 mm, cupuliform; involucral bracts 12 to 13, 6.3–9.3 × 2.4–3.9 mm, obtuse at the apex, greenish, purple-edged. Ray florets 21 to 36; corollas 9.2–14.1 × 1.4–1.9 mm, 4 to 5-veined, subentire to 3-toothed at the apex, conspicuously surpassing the involucre, white. Disc florets 49 to 63; corollas 5.5–7.2 mm long, yellowish; style branches penicillate, purplish. Achenes 3.5–3.9 × 1–1.1 mm, cylindrical, 7 to 8-ribbed, glabrous; pappus 6.7–8.9 mm long, barbellate, whitish to partially rose-colored. Chromosome number 2n = 104(±4) (Diers 1961). Fig. ​Fig.2D2D.

Additional iconography.

Weddell (1856: pl. 17D, sub Werneria digitata); Ricardi and Marticorena (1966: 26, fig. 6, sub W. pseudodigitata); Beltrán (2016: 359, fig. 3B, as photo); Gonzáles et al. (2016: 169, fig. 1H, I, sub X. pseudodigitatum, as photo).

Distribution and habitat.

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. ​(Fig.2323).

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Figure 23.

Distribution map of Xenophyllum digitatum (left hand) and X. pseudodigitatum (right hand).

Phenology.

Flowering nearly all year round.

Etymology.

The adjective digitatus -a -um refers to the shape of the leaves, which have divisions arranged like those of a bird’s foot (3-forked).

Notes.

Xenophyllum digitatum is characterized by its 3-forked leaves, where each lobe can be, in turn, 2 to 3-divided. The depth of the primary division reaches 1/3–1/2 of the total leaf length and the leaf margin usually have some cilia in the lower half. It has quite large involucre (12.1–16.7 × 9.8–15.7 mm) with 12 to 13 involucral bracts, 21 to 36 ray florets (with white corollas conspicuously surpassing the involucre), 49 to 63 disc florets, and glabrous achenes.

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 (Meneses 5446, LPB; Yager 4697, LPB), or even 3-divided (Humbert 30793, USM; Hutchinson 1212, CONC, F, GH, LE, NY, UC, US, USM). The leaf lobe apex is acute to obtuse, although populations with clearly mucronate apex also exist such as those from Ausangate in Cusco (e.g., Humbert s.n. [Vargas 12143], US, CUZ). The leaf lamina is usually glabrous but specimens with evanescent arachnoid indumentum on the adaxial surface are also found. Based on this latter character, Rockhausen (1939) described the plants with indumentum as Werneria digitata var. lanata Rockh. We consider it as part of the variability encompassed by X. digitatum, and accordingly, Rockhausen’s name is treated as a heterotypic synonym.

This species has been confused with X. pseudodigitatum (Rockh.) V.A.Funk and X. rosenii. For details on their morphological differences see the comments under the respective species.

Description.

Suffruticose plant, forming clumps of erect stems. Rhizomes 5–8 × 0.4–0.6 cm, horizontal to oblique, glabrous. Stems 5–12 cm tall, simple or branched, glabrous, usually with leaves restricted to the upper part. Leaves imbricate, extending into a glabrous sheath-like base; leaf laminas 6–9.6 × 2–3.8 mm, linear-spatulate (broadened upwards), 3-forked at the apex (incision depth 1/8–1/7 of the total length), entire, sometimes with some minute cilia in the lower half, flat to elliptic or almost terete upwards in cross section, glabrous, 1-nerved above (barely visible), 1-nerved beneath, fleshy, matte; leaf lobes 1.3–2.6 × 1–1.6 mm, acute to mucronate (rarely notched). Capitula radiate, erect, sessile. Involucres 7–8.4 × 5.2–8 mm, cupuliform; involucral bracts 10 to 11, 3.4–3.9 × 2–2.8 mm, obtuse at the apex, greenish. Ray florets 20 to 25; corollas 7.1–8.2 × 1–1.2 mm, 3 to 4-veined, subentire to 3-toothed at the apex, conspicuously surpassing the involucre, white. Disc florets 29 to 33; corollas 4.2–5.1 mm long, yellowish; style branches penicillate, purplish. Achenes 2.3–2.5 × 0.5–0.7 mm, cylindrical, 7 to 9-ribbed, glabrous; pappus 5.2–7.1 mm long, barbellate, whitish. Chromosome number unknown. Fig. ​Fig.2222.

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Figure 22.

Xenophyllum pseudodigitatumA habit B, C capitula at different stages of development D leaf E leaf apex variability F ray floret (frontward bristles removed) G ray floret (pappus removed) H disc floret (frontward bristles removed) I stamen J style. All details drawn from Ellenberg 4295 (US) except for B, C, F (drawn from Petersen & Hjerting 159, C). Illustration by Alice Tangerini.

Additional iconography.

Cabrera (1978: 473, fig. 200A–F, sub Werneria pseudodigitata); Funk (1997a: 237, fig. 1C); Freire and Ariza-Espinar (2014: 228, X. pseudodigitatum A–E).

Distribution and habitat.

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. ​(Fig.2323).

Phenology.

Flowering from January to May.

Etymology.

The epithet pseudodigitatum refers to the resemblance between this species and X. digitatum. It probably responds to the fact that both species have 3-forked leaves.

Notes.

Xenophyllum pseudodigitatum is characterized by its 3-forked leaves. The leaf lobes are 1.3–2.6 mm long, which represents 1/8–1/7 of the total leaf length, and they are acute to mucronate at the apex (rarely notched); on living plants, they are noticeably spread out. The involucre length ranges from 7 to 8.4 mm and has 10 to 11 involucral bracts. The radiate capitula are composed of 20 to 25 ray florets and 29 to 33 disc florets.

This is a narrow endemic species to northwestern Argentina that has erroneously been reported from northern Chile (Ricardi and Marticorena 1966; Freire and Ariza-Espinar 2014; Moreira-Muñoz et al. 2016) and southern Peru (Gonzáles et al. 2016). All these records are confusions with X. digitatum. In order to avoid further misidentifications, the following characters should be studied: leaf lobe length (1.3–2.6 mm in X. pseudodigitatum vs. 3.6–5.9 mm in X. digitatum), involucre length (7–8.4 mm in X. pseudodigitatum vs. 12.1–16.7 mm in X. digitatum), number of involucral bracts (10 to 11 in X. pseudodigitatum vs. 12 to 13 in X. digitatum), ray corolla length (7.1–8.2 mm in X. pseudodigitatum vs. 9.2–14.1 mm in X. digitatum), and number of disc florets (29 to 33 in X. pseudodigitatum vs. 49 to 63 in X. digitatum). Their distribution areas do not overlap.

Description.

Suffruticose plant, forming clumps of erect stems. Rhizomes 4–10 × 0.4–0.5 cm, horizontal to oblique, glabrous. Stems 11–19 cm tall, usually branched, glabrous, usually with leaves restricted to the upper part. Leaves imbricate, extending into a glabrous sheath-like base; leaf laminas 7.2–16 × 2.2–3.7 mm, linear-oblong, slightly broadened towards the apex, 3-notched at the apex, with the lateral lobes wider than central one, entire and shortly ciliate (cilia ca. 0.3 mm long), flat to slightly curved forwards in cross section, glabrous, 1-nerved above (barely visible), 1-nerved beneath, somewhat fleshy, matte; leaf lobes 0.6–1.5 × 1–1.6 mm (central one significantly smaller), rounded to subtruncate. Capitula radiate, erect, sessile. Involucres 10–12.6 × 8–10 mm, cupuliform; involucral bracts 13 to 14, 5.2–8 × 2–2.9 mm, acute to obtuse at the apex, greenish. Ray florets 20 to 21; corollas 10.4–11.9 × ca. 1.5 mm, 3-veined, subentire to 2-toothed at the apex, conspicuously surpassing the involucre, white. Disc florets 37 to 58; corollas 6–7 mm long, yellow; style branches truncate with a crown of sweeping trichomes or slightly penicillate, purplish. Achenes 4.8–5.4 × 1–1.2 mm, cylindrical, 7 to 8-ribbed, glabrous; pappus 8.7–9.8 mm long, barbellate, whitish. Chromosome number unknown.

Iconography.

Blake (1928: 496, fig. 1D, E sub Werneria decora); Beltrán (2016: 359, fig. 3C, as photo).

Distribution and habitat.

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. ​(Fig.2424).

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Figure 24.

Distribution map of Xenophyllum decorum (left hand, closed triangle), X. amblydactylum (left hand, open circle), and X. dactylophyllum (right hand).

Phenology.

Flowering from February to September.

Etymology.

The adjective decorus -a -um means handsome, elegant, decorous. Indeed, Blake (1928) pointed out in the protologue that it is “an attractive and very distinct species”.

Notes.

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.

Xenophyllum decorum has the leaf apex somewhat similar to that of X. incisum, however, their distribution areas are geographically very distant. In addition, these species differ in the leaf length (7.2–16 mm in X. decorum vs. 3.2–7.3 in X. incisum), leaf margin (shortly ciliate in X. decorum vs. not ciliate in X. incisum), number of involucral bracts (13 to 14 in X. decorum vs. 8 to 9 in X. incisum), number of ray florets (20 to 21 in X. decorum vs. 8 to 13 in X. incisum), and length of ray corollas (10.4–11.9 mm in X. decorum vs. 6.2–8.9 mm in X. incisum).

It has to be noted that we had problems in identifying Beltrán 7987 (USM). This specimen exhibits the characteristic flat and ciliate leaves of X. decorum but has the leaf apex strongly divided (ca. 2.5 mm deep) with each lobe entire or divided once again. It might be treated as an intermediate form between X. decorum and X. dactylophyllum or X. digitatum since these species co-occur in the region where Beltrán’s specimen was collected.

Description.

Suffruticose plant, forming clumps of erect stems. Rhizomes 10–12 × ca. 0.5 cm, horizontal to oblique, glabrous. Stems 4–7 cm tall, rather branched, glabrous, usually with leaves restricted to the upper part. Leaves subimbricate, extending into a sheath-like base that bears evanescent arachnoid trichomes; leaf laminas 4.3–10 × 1.3–1.4 mm, linear, barely broadened at the apex, 3-notched at the apex, entire, usually with some minute cilia near the base, flat to obtusely triangular in cross section, with evanescent arachnoid trichomes above, 1-nerved above (barely visible), 1-nerved beneath, somewhat fleshy, matte; leaf lobes 0.8–1.4 × 0.6–1 mm, obtuse. Capitula radiate, erect, sessile. Involucres 8.8–9.8 × 5.5–7 mm, cupuliform; involucral bracts 10 to 13, 4.9–5.4 × 1.2–1.8 mm, acute at the apex, greenish. Ray florets ca. 11; corollas 9–10.2 × ca. 1.2 mm, 4-veined, subentire to 2-toothed at the apex, conspicuously surpassing the involucre, white. Disc florets ca. 22; corollas 5.4–6 mm long, yellowish; style branches truncate with a crown of sweeping trichomes or slightly penicillate, yellowish. Achenes cylindrical, glabrous (immature); pappus ca. 5.6 mm long, barbellate, whitish. Chromosome number unknown.

Iconography.

Blake (1928: 496, fig. 1B, C sub Werneria amblydactyla).

Distribution and habitat.

Endemic to Peru (Junín, Lima). It grows in rocky outcrops of the puna ecoregion, at elevations of ca. 4600 m (Fig. ​(Fig.2424).

Phenology.

Collected in flower in May.

Etymology.

The epithet amblydactylum means having blunt fingers (“ambly-”: blunt, obtuse; “dactyl-”: finger). It refers to the leaf apex lobes of this species.

Notes.

Xenophyllum amblydactylum forms clumps of erect stems 4–7 cm tall that bear linear, 4.3–10 mm long leaves. The leaf apex is 3-notched and the lobes are small (0.8–1.4 × 0.6–1 mm) and connivent. The leaf lamina has arachnoid trichomes on the adaxial surface that are quicky evanescent. The involucre is 8.8–9.8 mm long and it is composed of 10 to 13 involucral bracts. The capitula are radiate and display ca. 11 ray florets with white corollas.

This is a poorly known and collected species. Aside from the type material, we only studied one collection from southeastern Lima Department (Beltrán 4174, USM) and a specimen that was bought as medicinal in a farmers market in the city of Lima (Yarupaitán 1499b, USM). Further collections are required in order to improve the understanding of its variability and properly describe the achenes.

It might be confused with X. dactylophyllum because their distribution areas overlap. However, several characters let anyone discriminate them from one another, i.e., leaf apex (3-notched in X. amblydactylum vs. at least 9-divided in X. dactylophyllum), leaf lobe length (0.8–1.4 mm in X. amblydactylum vs. primary division 2–3 mm in X. dactylophyllum), involucre length (8.8–9.8 mm in X. amblydactylum vs. 10.8–12.8 mm in X. dactylophyllum), ray floret number (ca. 11 in X. amblydactylum vs. 17 to 22 in X. dactylophyllum).

Description.

Suffruticose plant, forming clumps of erect stems or cushions. Rhizomes 5–15 × 0.5–0.8 cm, horizontal to oblique, glabrous. Stems 11–24 cm tall, branched, glabrous, usually with leaves restricted to the upper part. Leaves imbricate, extending into a sheath-like base glabrous or with long silky trichomes; leaf laminas 5.7–9.9 × 2.4–4.1 mm, spatulate (finger-like at the apex), at least 9-divided at the apex, with the primary division extending deeper than the subsequent ones, entire, sometimes with some minute cilia, elliptical to terete upwards in cross section, glabrous or floccose-lanate, 1-nerved above (barely visible), 1-nerved beneath, somewhat fleshy, matte; primary division 2–3 mm long, 3 to 6 divided, rounded to truncate, sometimes with some scattered cilia on the margin. Capitula radiate, erect, sessile. Involucres 10.8–12.8 × 8.2–9.5 mm, cupuliform; involucral bracts 11 to 20, 6.6–7.6 × 1.9–2 mm, obtuse at the apex, greenish, sometimes purple-edged. Ray florets 17 to 22; corollas 9.2–11 × 1.3–1.6 mm, 4-veined, subentire to 3-toothed at the apex, conspicuously surpassing the involucre, white. Disc florets 37 to 50; corollas 5.3–6.3 mm long, yellowish; style branches truncate with a crown of sweeping trichomes or slightly penicillate, purplish. Achenes 3.1–4.8 × 0.9–1.3 mm, cylindrical, 7 to 9-ribbed, glabrous; pappus 8.1–8.9 mm long, barbellate, whitish. Chromosome number 2n = 108(±4) (Diers 1961). Fig. ​Fig.2525.

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Figure 25.

Xenophyllum dactylophyllumA habit B stem apical part and capitulum C leaf D ray corolla and style E disc corolla and style branches F disc corolla and stamens (vertically sectioned, style removed) G style H achene with pappus. All details drawn from Funk & González-Quint 11372 (US). Illustration by Alice Tangerini.

Additional iconography.

Linares et al. (2010: 102, fig. 1, sub X. oscartovarii, as photo); Beltrán (2016: 359, fig. 3A, as photo).

Distribution and habitat.

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. ​(Fig.2424).

Xenophyllum dactylophyllum has been reported from the Bolivian department of Potosí on the basis of the collection Steinbach s.n. (Jørgensen 2014; Tropicos 2020). This material, which has not been located, was collected nearby Andacaba, a locality ca. 30 km south of the city of Potosí. If the identification is confirmed, it would represent the southern distribution limit of this species. The specimen US-00974139 most likely corresponds to a mislabeling because both the locality (San Carlos de Bariloche, Río Negro Province, Argentina) and the elevation (600 m) do not correspond to the known distribution area and altitudinal rank of this species.

Phenology.

Flowering nearly all year round.

Etymology.

The epithet dactylophyllum means finger-like leaves and it refers to the resemblance between the leaves of this plant and a hand.

Notes.

Xenophyllum dactylophyllum is characterized by having glabrous or floccose-lanate finger-like leaves, with at least 9 divisions at the apex. The divisions are usually arranged in multiples of three, the primary one extending deeper than the subsequent ones. The capitula are radiate, erect, with 17 to 22 ray florets with white corollas conspicuously surpassing the involucre.

Another species with finger-like leaves is X. staffordiae, which has occasionally been confused with X. dactylophyllum. They can be readily differentiated by the habit (11–24 cm tall in X. dactylophyllum vs. 30–60 cm tall in X. staffordiae), leaf divisions (at least 9-divided, with the primary division extending deeper than the subsequent ones in X. dactylophyllum vs. at least 12-divided, with all divisions similar in length in X. staffordiae), capitulum position (erect in X. dactylophyllum vs. rather nodding in X. staffordiae), and length and color of the ray corollas (9.2–11 mm, surpassing the involucre, white in X. dactylophyllum vs. 6.4–8.3 mm, not surpassing the involucre, pale yellow in X. staffordiae). Their distribution areas overlap from central to southern Peru. Other morphologically similar species are X. amblydactylum and X. lorochaqui (see comments under these species).

Our efforts in locating the type material of X. oscartovarii E.Linares, J.Campos, Nauray, Vicente Orell. & A.Galán were unfruitful. Due to the original description and the pictures included in the publication (Linares et al. 2010), we believe that it fairly corresponds to X. dactylophyllum. Oddly, this latter species was not mentioned in the taxonomic discussion of the presumed new species. In addition, we studied several specimens of X. dactylophyllum collected close to the type locality of X. oscartovarii.