Paepalanthus pilosus (Kunth in H.B.K.) Kunth var. pilosus

Hensold, Nancy, 2016, The Andean Paepalanthus pilosus complex (Eriocaulaceae): a revision with three new taxa, PhytoKeys 64, pp. 1-57 : 33-40

5 a. Paepalanthus pilosus (Kunth in H.B.K.) Kunth var. pilosus Figs 1A-B ; 2D, F-J ; 3B ; 11A-D ; 12A-K

Paepalanthus pilosus (Kunth in H.B.K.) Kunth var. pilosus

Description.

Densely branched cushion plants, the cushions reportedly reaching 30 cm in diameter (Molau 3223) and up to 10 cm tall (Berry 4366). Leaves subulate, recurved, 7-16 (-20) mm long × (0.5-) 0.7-1.3 (-1.5) mm wide at midpoint; apex cuspidate-acute to short-aristate, the sharp tip evident in adaxial view; usually appressed pilose adaxially near tip (in Peru and Ecuador), the hairs smooth to roughened, in northern part of range usually with conspicuous long coarse scattered cilia to apex (rare in Peru and Ecuador), often early glabrate, bright green, texture chartaceous to rigidulous, the abaxial surface smooth or with nerves salient. Peduncles appressed-pilose when young especially at apex, often only 1.5-8 mm long and surpassed by the sheaths at anthesis, but frequently and variably elongating up to ten or more times this length (20-100 mm) in fruit; in Peru and Ecuador only rarely up to 50 mm at anthesis and strongly exsert (Cano 16840); the sheaths 3-15 mm long, scarious and glabrous or minutely tufted at apex, splitting apically into 2 or 3 triangular segments. Capitula 3-6 mm in diameter, often borne among the leaves at anthesis, frequently exsert in fruit, more rarely exsert at flowering. Involucral bracts equaling the capitulum to slightly surpassing it, the outer bracts 1.6-4.2 mm long, ovate to often triangular, pale gold or greenish especially along midvein, pilose on upper back to merely tufted or glabrate, the inner bracts more broadly ovate to triangular, sometimes tinged grayish-brown on shoulders; receptacle long-pilose. Trichomes of bract and sepal apices subacute to bulbous, obscurely tuberculate. Flowers ca.10-16 per capitulum, the pistillate peripheral, usually 1-2 (-3) times as many as the staminate flowers. Pistillate flowers: Pedicels 0.35-0.5 mm, fine or rarely wide and spongy, not thickened-rigid in fruit. Sepals elliptic to obovate or oblanceolate, apex acute to rounded, the base linear-ligulate to narrowly cuneate, 1.4-2.65 mm long, 0.3-1.1 mm wide at middle, 0.15-0.3 (-0.4) mm wide at base; usually tinged dusky brown above, rarely uniformly pale cream (Sagastegui 12242), short-tufted at apex, ciliate at upper margins, and usually pilose either side of the dorsal midvein; the basal half of the midrib hygroscopically thickened, spadiceous-brown in fruit and recurving when dry, the margins and distal half of the sepal remaining chartaceous, erect; sepals detaching from diaspore before dispersal. Petals oblanceolate-spatulate, acute to obtuse to emarginate, rarely (Wurdack 1616) broadly cuneiform; apex truncate-emarginate, apiculate; 1.5-2.35 mm long × 0.3-0.75 (-1.0) mm wide, ca. 2-6 times longer than wide, cream to brownish-tinged, pubescence similar to Paepalanthus caryonauta ; usually not thickening in fruit, dispersed with the fruit. Gynoecium with style base 0.5-1.05 mm, nectaries with stalks 0.3-0.6 mm long, glandular portion dark red to brown, 0.3-0.4 mm long, penicillate with stiff whitish papillae at upper margin (northernmost Peru and northwards), or 0.4-0.5 mm long, clavate-infundibular with stiff whitish sometimes elongate papillae distributed along the outer surface, densest at the rim (Cajamarca: Celendín, Amazonas: Chachapoyas); papillae rigidulous after anthesis; styles 0.8-1.1 mm long (Barbour 3427: 1.4-1.6 mm long), dark red-brown. Seeds 0.55-0.8 mm long, red-brown, reticulate with abundant white erect to suberect pseudotrichomes. Staminate flowers: Pedicels 0.2-0.5 mm. Sepals 1.2-2.45 mm, fused at base unequally, from very briefly to about 1/3 sepal length, obovate to usually spatulate, acute to rounded, narrowed to base and not thickening with age. Corolla 1.2-2.5 mm long, the anthophore 0.5-1.5 mm long, comprising 30 -65 % the corolla length, membranous and 0.1-0.2 mm diameter at base, broadening to 0.2-0.3 mm near apex; the corolla tube including lobes 0.5-1.65 mm long. Filaments similar to Paepalanthus caryonauta , usually dark red-brown between apex and the base of the corolla lobe, exsert ca.0.2-0.5 mm beyond the lobes. Nectaries reaching the sinuses of the corolla tube.

Phenology.

In Central America, mostly collected January to May, and in July, coinciding with the main dry season, and possibly a shorter second dry season (veranillo), respectively ( Grayum et al. 2004). In the climatically wet paramos of Colombia and Venezuela, peak flowering is June to December, generally the wet season ( Rangel-Ch. 2000; Cuello et al. 2010; Vareschi 1970), though probably coinciding with short dry periods, as observed by Pedraza-Peñalosa et al. (2005). In Peru, flowering and fruiting mostly from July to September, during a pronounced dry season; in Ecuador, also during predominantly dry months from September to November (to January), fruiting through April.

Distribution.

Costa Rica (Cerro Talamanca): Cartago, Limón, San José. Panama: Bocas del Toro. Colombia (Eastern Cordillera, Sierra Nevada de Santa Marta, possibly Serrania de Perijá, also rare in Western and Central Cordilleras): Antioquia, Bogotá D.C., Boyacá, Cauca, Cundinamarca, Magdalena, Norte de Santander, Santander. Venezuela (Cordillera de Merida and Ramal de Guaramacal): Apure, Lara(?), Merida, Táchira, Trujillo. Ecuador: Azuay, Loja, Morona-Santiago, Zamora-Chinchipe(?). Peru (to 7° S lat.): Amazonas, Cajamarca, Lambayeque, Piura. (Fig. 6 )

Habitat and ecology.

Common and characteristic ground cover (rasante) or cushion species of wet páramo, rarely subpáramo, on boggy organic soils, or at pond and bog margins but not in standing water ( Schmidt-Mumm and Vargas-Rios 2012); often associated with Sphagnum , usually growing amongst shrubs or bunchgrass, in Colombia frequently also with bamboo ( Chusquea tessellata Munro) or Espeletia spp. ( Rangel-Ch. et al. 1997; Rangel-Ch. and Ariza 2000; Vargas and Rivera 1991). It is reported by some observers as growing at the base of bunchgrass ( Franco-Rosselli et al. 1986) or shrubs ( Weberbauer 1945) or under rock ledges, but is also reported in full sun. In Ecuador and the western cordillera of Peru, it is occasionally reported from montane forest. Elevation (2700-) 3000-4000 (-4400) m.

Paepalanthus pilosus is a commonly cited species in phytosociological analyses of North Andean paramo, usually under the name Paepalanthus karstenii ( Rangel-Ch. et al. 1997). It is reported as a "characteristic and dominant species" of the alliance Paepalantho karsteni - Chusquioni tessellatae in wet páramo east of Bogotá ( Rangel-Ch. and Ariza 2000). It is tolerant of disturbance and may even have a minor role in succession, being reported by several collectors as locally abundant on wet banks and roadsides, and in regenerating burnt paramo. Cleef (1981) noted that it is one of the first species after Castilleja to colonize burnt bunchgrass páramo. Cuello and Cleef (2009), in Venezuela, found it occurring abundantly in Sphagnum mats around the margins of an old lakebed undergoing succession to bunchgrass páramo, and suggested that this new azonal alliance ( Sphagno -recurvi - Paepalanthion -pilosi Cuello & Cleef) was associated with disturbance by wildlife fragmenting the Sphagnum mat. Vargas and Rivera (1991, quoted by Rangel-Ch. et al. 1997) found Paepalanthus pilosus in paramo disturbed by grazing cavies and rabbits. One collector reported that the plant forms an abundant ' necromasa ' with good water-retaining properties relevant to peat formation (Bernal 1647).

Conservation status.

The status of this widespread variable taxon, reported as a colonizer tolerant of disturbance, is presumed to be of Least Concern ( IUCN 2014).

Taxonomic history.

No type material is found for Eriocaulon pilosum Kunth in the herbarium P-Bonpl. in Paris ( Stauffer et al. 2012). The lectotype specimen chosen from Kunth’s herbarium (B) includes two individuals of Paepalanthus pilosus , with one short-pedunculate individual of Paepalanthus dendroides mounted between them. The middle plant is a good match for the type of Eriocaulon dendroides Kunth, described from the same locality. Kunth’s concepts of the two species were based partly on the shorter peduncles in Paepalanthus pilosus , but this character varies in both species. Kunth also differentiated Eriocaulon pilosum by leaves rigid, “pilose-ciliate,” with a sharp apex, and involucral bracts ovate, acute, while Eriocaulon dendroides was described as having leaves acuminate, membranous and glabrous, and involucral bracts obovate. These characters are adequate to distinguish the two elements on the sheet and to justify exclusion of the middle plant from the type material.

Paepalanthus pilosus has suffered confusing taxonomic treatment over time. Körnicke (1863) recognized Paepalanthus pilosus and Paepalanthus dendroides (both described from Bogotá) as well as Paepalanthus selaginoides ( Popayán) as distinct taxa, distinguishing Paepalanthus selaginoides by the near obsolete peduncles, and Paepalanthus pilosus from Paepalanthus dendroides by the robust scattered cilia of the leaf margin. Ruhland (1903) synonymized all three under Paepalanthus pilosus with the claim that these diagnostic characters were too variable, sometimes even within specimens, an impression perhaps fostered by the mixed sheet of Paepalanthus dendroides and Paepalanthus pilosus from Kunth’s herbarium. At the same time, Ruhland erected an additional new species, Paepalanthus karstenii , also from near Bogotá, distinguished from Paepalanthus pilosus by the "leaf indument and apex," the involucral bracts broad and glabrous abaxially, and "a different form of the perianth." Inexplicably, he also described capitula as 2-3 mm wide in Paepalanthus pilosus versus 6-8 mm wide in Paepalanthus karstenii , which accords neither with the type of Paepalanthus pilosus (capitula 6.5 mm) nor earlier descriptions. How Ruhland thought the leaf indument, apex, or perianth in Paepalanthus karstenii differed from that of Paepalanthus pilosus is not clear from his description, leaving only the key character of bract pubescence, which also varies widely within species. In fact the broad (obovate) subglabrous bracts seen in the type of Paepalanthus karstenii are more typical of Paepalanthus dendroides as recognized by both Kunth and Körnicke. The identity of Paepalanthus karstenii needs further study (see Doubtful Taxa).

Moldenke (1975b) ostensibly followed Ruhland, treating Paepalanthus dendroides as a synonym of Paepalanthus pilosus , and distinguishing Paepalanthus karstenii by the "involucral bracts glabrous on the outer surface," but his use of the names in annotations (ca.1930's-1980's) doesn’t correlate with bract pubescence or shape. In his pattern of annotations, Moldenke revived the appropriate distinction between Paepalanthus dendroides and Paepalanthus pilosus , but confused the nomenclature, mostly annotating typical Paepalanthus pilosus as Paepalanthus karstenii , while applying the name Paepalanthus pilosus to Paepalanthus dendroides and occasional long-pedunculate individuals of Paepalanthus pilosus . This convention was followed by later authors (e.g., Cleef 1981, Madriñán and Zapata 2001). Huft’s treatment ( 1994) was similar but treated Central American Paepalanthus pilosus as Paepalanthus kupperi rather than Paepalanthus karstenii . Hensold and Hammel (2003), in a treatment of Costa Rican species, re-established Paepalanthus dendroides as a distinct taxon and corrected application of the name Paepalanthus pilosus to accord with the original concept of Kunth and Körnicke, which includes both Paepalanthus kupperi and most material determined as Paepalanthus karstenii .

Moldenke did not specify distinguishing characters for his other species here placed in synonymy of Paepalanthus pilosus , nor were these names widely used. Of his new infraspecific taxa, Paepalanthus karstenii f. corei was said to differ from that species by the peduncles only 1-2 cm in length, and Paepalanthus karstenii var. minimus by both leaves and peduncles shorter. The type of the former is in fact Paepalanthus dendroides , though Moldenke most commonly applied the name to Paepalanthus pilosus and Paepalanthus caryonauta .

Paepalanthus espinosianus and Paepalanthus loxensis were first treated in synonymy of Paepalanthus pilosus by León-Yánez and Hensold (1999), and Paepalanthus subsessilis by Hensold (2008). Paepalanthus selaginoides was originally placed in synonymy of Paepalanthus pilosus by Ruhland (1903) and is clearly closest to that species, but represents the only record of the species from the Central Cordillera of Colombia. Pending closer examination of the type, its placement here is provisional.

Discussion.

Paepalanthus pilosus in Peru and southern Ecuador is characterized by the usually dwarf flowering peduncles with capitula subsessile at anthesis, and the outer involucral bracts greenish or with a green midvein and often longer than wide. In this area, soft hirsutulous pubescence of the upper leaf surface near the apex, similar to that found in Paepalanthus dendroides , is common. From Costa Rica to Colombia and Venezuela, long robust scattered cilia along the distal margin are frequent and diagnostic when they occur (hence the species epithet; cf. Fig. 2D ), but rare in our area. These cilia can often be detected even in older glabrate leaves due to the persistent enlarged basal cells. From Paepalanthus dendroides , Paepalanthus pilosus may also be distinguished by the relatively narrow elliptic to oblanceolate petals, and the prominent dark rigid nectaries; and from Paepalanthus caryonauta by the usually larger capitula, the sharply acute to aristate leaf tips, and (in the typical variety) by the form of the fruiting calyx. (See Table 1 .)

Habit varies from a dense compact mat pressed horizontally by collectors (as in the type of Paepalanthus espinosianus Moldenke), to a rounded cushion with branch lengths of a few centimeters, pressed and mounted vertically (as in the type of Paepalanthus loxensis Moldenke) The superficial difference in aspect of these two forms caused Moldenke to ally the former species with Paepalanthus karstenii , but the latter species with Paepalanthus glaziovii Ruhland (subsect. Dichocladus ). This character almost certainly varies in response to habitat, as Heilborn (1925) noted in the vegetatively similar species Plantago rigida Kunth, which forms rounded cushions on wet sites and flat mats on drier sites. Leaves may be ascending to strongly reflexed and appressed to the ground, the latter perhaps a response to drier or sunnier conditions. An example are the collections Hernandez-Schmidt 1330 and 1432 from the same locality in Cundinamarca, the former reported on rocks in full sun, with leaves reflexed close to ground, the latter on saturated soil, with leaves erect.

Peduncle length is highly variable, as already noted by Ruhland (1903), but much of this variation may be developmental. Actively flowering capitula are usually borne on peduncles only a few millimeters long, barely emergent from the sheath, and in vested by the sharp cuspidate tips of subtending foliage leaves. Dramatic peduncle elongation often occurs later in fruit, as shown by specimens bearing both subsessile flowering capitula at stem tips, and fruiting capitula lower on the stem on peduncles up to ten times as long (e.g., Larsen 237, Fig. 11D ). The delayed timing of peduncle elongation may afford wind and frost protection during anthesis, while later enhancing fruit dispersal. Peduncle elongation doesn’t always accompany fruiting or may be minimal, but even in specimens with many subsessile fruiting capitula still embedded in the leaf mat, occasional remains of much longer peduncles can be found (e.g. Barbour 3427, Sagastegui 12242). Notably, the only specimen from Peru and Ecuador with peduncles already elongate (2-5 cm) at anthesis was also the only specimen reportedly collected from "montane forest," where it was said to be rare (Cano 16840). This plant may represent a taxonomically distinct variant, but for now is treated as an environmental form. In Colombian populations of the Eastern Cordillera, early peduncle elongation may be more common.

Other characters showing wide variation in Peru and Ecuador are involucral bract length, capitulum size, and flower and seed size. For example, the type of Paepalanthus loxensis (Loja) contrasts sharply with that of Paepalanthus espinosianus (Azuay) by its much smaller involucral bracts, capitula, flowers, and seeds. Indeed, in most plants from Azuay, in the Cerro Fierro-Urcu (Loja), and in some localities in Peru ( Jaén, Bagua, Celendín), outer involucral bracts are ca.2.6-4.0 mm, with the tips often surpassing the large capitula 4-6 mm in diameter (cf. Fig. 2H ). In contrast, most specimens from southern Loja, the western Cordillera of Peru, and southern Amazonas, Peru, have involucral bracts about 1.6-2.6 mm long and capitula 3-4 mm in diameter (Fig. 2I-J ). The length of the pistillate flower sepals was measured in 18 specimens of Peru and Ecuador and was imperfectly correlated with capitulum size. Small flowers (sepals 1.4-1.8 mm) were found in the small-capitate plants from southern Loja and northwestern Peru, and a few large-capitate plants of Azuay; all other specimens were large-flowered (sepals ca. 1.8-2.4 mm long). Seed size showed the strongest geographic correlation, with the smallest seeds observed in the small-flowered specimens from southern Loja and the western cordillera of Peru (0.53-0.63 mm, 5 samples), while elsewhere seeds were larger (0.67-0.8 mm, 8 samples). Among the large-flowered collections, those of Amazonas, Peru, south of the mouth of the Rio Utcubamba (ca. 6° S latitude) are distinguished by the unusually broad female petals (l/w ratio 1.8-2.9 vs. 3.5-7.0 elsewhere), broadly rounded to even truncate-emarginate at apex (Fig. 12I ), and by an ambiguous pattern of sepal thickening, both characters suggesting intermediacy with Paepalanthus pilosus var. leoniae , of San Martín. Also shared with that variety are the funnelform nectaries (Fig. 12I ) with unusually large, spreading papillae (also found in Sagastegui 12242, Celendín), in comparison to the commonly penicillate nectaries with small papillae found elsewhere.

Paepalanthus pilosus in Colombia requires further careful study. In the Eastern Cordillera, particularly in disturbed paramo near Bogotá, there are more pronounced morphological extremes in the species than in other parts of its distribution. Here, leaf lengths range up to 3 cm, capitula from 3.5-8 mm in diameter, peduncles are occa sionally strongly exsert at flowering, and involucral bract color and form are variable, ranging from nearly obovate-rounded to narrowly triangular, and pale gold to dark pigmented on the shoulders. Madriñán and Zapata (2001) indicate two sympatric elements for the Páramo Chingaza, one with ciliate leaves and peduncles exceeding the leaves in length (" Paepalanthus dendroides "), and one with leaves nearly glabrous and peduncles shorter (" Paepalanthus karstenii "). However type material of both Paepalanthus pilosus and Paepalanthus karstenii have prominent scattered cilia of the leaf margins, and both have variable peduncle lengths, which may be quite long in fruit. Judging from the photographs, both elements from Páramo Chingaza fit within my working concept of Paepalanthus pilosus , but the presence of two reproductively isolated crypto-taxa in this area cannot be ruled out. Some differences especially in leaf size, orientation and pubescence may be due to microhabitat, but polymorphism does seem more exaggerated in collections from the vicinity of Bogotá than from elsewhere, and collectors have often recognized two elements at one locality (e.g., Barclay 4031, 4057, Paramo de Guasca; Dwyer 8186, 8189, Paramo Cruz Verde; Rangel 4045, 4048, Boyacá, Duitama). For simplicity I have treated as Paepalanthus pilosus all forms with subulate, sharply cuspidate leaves and the flower and fruit structure described above, and consider robust spreading cilia of the distal leaf margin also diagnostic when present, as it is in Paepalanthus pilosus of Venezuela and Central America. This may however obscure a more complicated taxonomic picture.

Hybridization may be a factor contributing to the chaotic variation in Paepalanthus pilosus in the vicinity of Bogotá. Some material, such as the type of Paepalanthus karstenii (see Doubtful Taxa) looks intermediate with Paepalanthus dendroides , as to elongate peduncles and rounded involucral bracts. Typical Paepalanthus dendroides was originally described from Bogotá, but has not been collected in the vicinity since 1917, while it is still widespread elsewhere. In addition, probable hybrids between Paepalanthus pilosus and Paepalanthus caryonauta have been collected at the margins of the range of Paepalanthus pilosus , at Paramo Frontino (Antioquia), and on the eastern slope of Paramo Sumapaz, south of Bogotá (see Paepalanthus caryonauta discussion).

A final complication in the taxonomy of Colombian Paepalanthus pilosus is the presence in the Serranía del Perijá and vicinity of a closely related but more robust taxon, which may intergrade with it, discussed below under Paepalanthus sp. A.

Selected specimens examined

(of 122 total). COSTA RICA. Cartago: Cerro Asuncion , 3350 m, 19 Mar 1978, Wilbur 26123 (F) . Limón: Parque Nac. Chirripó, Valle Las Morrenas , 9°29'24"N, 83°29'24"W, 3500 m, E. Alfaro 417 (MO) . San José: Canton de Dota, Cerro Vueltas, 9°37'40"N, 83°51'10"W, 3150 m, 28 Mar 1994, Cascante et al. 212 (F, MO); Cerro Chirripo , 9°27'36"N, 83°29'24"W, 3400-3739 m, 27 Jul 1996, Gamboa R. et al. 490 (MO) . PANAMA. Bocas del Toro: Cerro Fabrega , 9°07'N, 82°52'40"W, 3100-3300 m, 6-8 Mar 1984, Davidse et al. 25303 (F, MO) . COLOMBIA. Antioquia: Mpio. Urrao, Paramo Frontino, near Llano Grande , 3450 m, 27 Oct 1976, Boeke & McElroy 265 (NY [01347937] p.p., mixed with Paepalanthus caryonauta × Paepalanthus pilosus at top center), Ibidem, 3320-3450 m, 2 Mar 1989, MacDougal & Roldán 4463 (MO) . Bogotá, DC: Entre Alto de las Cruces-La Viga y Diego Largo , 3380- 3320 m, 19 Jun 1939, Cuatrecasas 5553 (F); Paramo de Chisaca , 3660- 3720 m, 11 Sep 1961, Cuatrecasas & Jaramillo 25882 (F); Paramo de Cruz Verde , 3300-3500 m, 20 Sep 1917, Pennell 2074 (F, MO); Paramo de Sumapaz , 3750 m, 8 Aug 2003, S. Rios et al. 16 (UDBC [UDBC15910]) . Cerro Monserrate, 26 Apr 1951, Romero-Castaneda 2492 (F); Paramo de Chisaca , 3596 m, 15 Oct 1966, Soderstrom 1346 (MO); Usme, Predio El Refugio EAAB, alt. 3726 m, 2 Aug 2011, Trujillo 600 (UDBC [UDBC30513]) . Boyacá: Nevado del Cocuy, valle de Las Lagunillas 4000-4300 m, 12 Sep 1938, Cuatrecasas 1501 (F); Paramo de la Rusia , Duitama - Charalá, 3624 m, 20 Aug 1953, Langenheim 3508 (MICH); Duitama , Paramo de Belen , 06°01'35"N, 72° 57'08"W, 3900 m, 30 Sep 1986, Rangel et al. 4045 (MO) . Cundinamarca: Paramo de Guasca, 3000-3400 m, 13 Jun 1957, Barclay 4031 (MO); Represa de Neusa, 3500 m, 22 Jun 1957, Barclay 4150 (MO); Boqueron de Chipaque , 3350 m, 12 Jul 1957, Barclay 4489 (MO); Paramo de Choachi , 3400 m, 18 Jun 1959, Barclay 7757 (MO); Mpio. de Fomeque , Sector La Playa (Parque Nac. Natural Chingaza), 3180 m, 31 Aug 1991, Bernal et al. 1647 (COL [COL000066835]); La Calera, Paramo de Palacio , 3420-3500 m, 8 Dec 1959, Cuatrecasas et al. 25574 (F); Mpio. Subachoque, Paramo El Tablazo, 3400 m, 6 Oct 2003, M. Hernandez Schmidt et al. 1330 (COL [COL000034189]), 12 Dec 2003, M. Hernandez Schmidt 1432 (COL [COL000034188]); Sopó, 2700 m, 14 Feb 1951, Schultes 11590 (MO) . Magdalena: Sierra Nevada de Santa Marta, Laguna Chubdula, 10°55'N, 73°53'W, 3480 m, 29 Jul 1972, Kirkbride & Forero 1784 (F); Slopes of Cuchilla Chebachucua , valley of Rio Duriameina , 10°38'N, 73°38'W, 3500 m, 31 May 1977, S. White & Alverson 642 (MO) . Norte de Santander: Cucutilla , Páramo El Romeral, 3200-3800 m, 25 Feb 2002, Ortiz R. et al. 978 (COL [COL000066965]) ; Santander: Paramo del Almorzadero , 3600 m, 1 Jan 1960, Barclay 10394 (MO) . VENEZUELA. Apure: Distr. Paez , paramo de Tamá, 3100 m, 26 Jun 1973, Ruiz-Teran & López-Figueiras 8555 (MICH) . Lara / Trujillo: Parque Nacional Dinira, Paramo de Jabón, vista al Tocuyo , 09°33'55"N, 70° 06'18"W, 3000 m, 13 Aug 1999, Riina et al. 522 (F) . Merida: Distr. Rangel, Sierra de Santo Domingo, paramo de Mucubaji, 3560-3600 m, 19 Nov 1959, Barclay & Juajibioy 9565 (MO); Distr. Justo Briceño, Alto del Totumo , hoya del Rio Chiruri , a 19.5 km de El Aguila, 3900-4000 m, 11 Oct 1983, Berry 4229 (F); Distr. Campo Elias , San Jose de Acequias - Mucutuy, 3120 m, 22 Aug 1984, Berry 4366 (F, MO); Distr. Rangel, Paramo de Mucuchies , 3750 m, Sept 1952, Humbert 26300 (F); Distr . Miranda, Timotes - Paramito , 2285-3500 m, 24 Mar 1944, Steyermark 55727 (F); Distr. Libertador, La Aguada, 3475 m, 17 Dec 1969, Steyermark & Koyama 102359 (F, MO) . Táchira: Paramo de Tamá, Pata de Judio , 3100-3500 m, 19 Oct 1978, Luteyn et al. 5927 (MO) . Trujillo: Parque Nac. Guaramacal , 2800-3100 m, 15 Jun 2001, Dorr et al. 9005 (F) . ECUADOR. Azuay: Paramo de Matanga , 03°13'42"S, 78°57'14"W, 3340 m, 18 Jul 2006, Aedo et al. 13036 (MO); Gualaceo - Limón, side road at km 25.2, 03°00'S, 78° 40'W, 3540-3600 m, 12 Jan 2000, Jørgensen et al. 1851 (F, MO); Sigsig - Gualaquiza, km 28, 03° 11'48"S, 78°47'09"W, 3280-3330 m, 14 Nov 2000, Jørgensen et al. 2366 (F, MO) . Azuay / Morona-Santiago: Muletrack Sevilla de Oro-Mendez ( Paramo de Castillo ), 2°48'S, 78°36'W, 3250-3500 m, 16 Sep 1976, Øllgaard & Balslev 9557 (F) . Loja: Tambo de Savanilla, 18 Oct 1876, André K-1438 (F); Fierro Urco , Saraguro-Loja , 03°43'10"S, 79° 19'18"W, 3840 m, 6 Dec 1994, Jørgensen et al. 1241 (MO); Yangana - Valladolid, 2800-2900 m, 5 Sep 1985, Larsen & Dall 237 (MO), Ibid., 3150 m, 14 Nov 1997, Lewis & Klitgaard 3726 (MO) . PERU. Amazonas: Prov. Bagua, Cordillera Colán 3290 m, 9 Sep 1978, Barbour 3427 (F, MO); Prov. Chachapoyas , Leimebamba-Chilchos trail, 4 Jul 1977, Boeke 2133 (MO, USM n.v.); Prov. Luya , Distrito Camporedondo , Tullanya , 06°04'29"S, 78°18'44"W, 3320 m, 8 Dec 1996, R. Vásquez-M. & Rojas 21996 (MO); Prov. Chachapoyas, 3000 m, Weberbauer 4416 (F frag ex G); Molinopampa - Diosan pass, 3100-3300 m, 5 Aug 1962, Wurdack 1616 (F) . Cajamarca: Prov. Jaén, Sallique, Localidad El Páramo, 05°40'51"S, 79°19'00"W, 3400 m, 22 Jul 1998, Campos et al. 5334 (F, MO); Prov. Celendín, Jalca de Kumulca , 3300 m, 18 Aug 1984, Sagástegui et al. 12242 (F, MO) . Lambayeque: Ferreñafe, Cañaris, 3253 m, 22 Apr 2010, Chocce et al. 5688 (USM [photo]) ; Piura: Huancabamba, Carmen de la Frontera, Nueva York , 3160 m, 28 Jul 2006, Cano et al. 16840 (F, USM [photo]) .

Figures

Figure 1. Cushion habit of Paepalanthus pilosus var. pilosus in Venezuela, November 2012. A Paepalanthus pilosus cushions with bunchgrasses, Paramo Batallon B Individual cushion, with graminoids emerging, Paramo Los Conejos. Photos by Serge Aubert, Station Alpine Joseph Fourier, France. Used by permission (www. cushionplants. eu).

Figure 2. Capitula and leaf tips of Paepalanthus pilosus and relatives A-E Fruiting capitula with diaspores; sepals detaching (A, D) or intact (C, E) A Paepalanthus dendroides B-C Paepalanthus caryonauta D Paepalanthus pilosus var. pilosus E Paepalanthus lodiculoides F-J Paepalanthus pilosus var. pilosus, involucral bract variation. Arrows = tips of the torn peduncle sheath. (A Davidse 28991 B Barclay 5176, Ecuador C Dudley 11060, Peru D Cuatrecasas 25574 E Ollgaard 9717 F Cuatrecasas 25882 G Cuatrecasas 5553 H Ollgaard 9557 I Cano 16840 J Jorgensen 2366.)

Figure 3. Vascular bundle morphology, adaxial side up. A Paepalanthus caryonauta B Paepalanthus pilosus var. pilosus C-E Paepalanthus dendroides F Paepalanthus huancabambensis midvein, with bundle sheath extensions. (A Valenzuela 8117 B Barbour 3427 C Leon 2683 D-E Leon 2243. F: Sagastegui 16799 D-F Mesophyll darkened for contrast.)

Figure 6. Distribution of Paepalanthus caryonauta and suspected hybrids, Paepalanthus pilosus var. pilosus, Paepalanthus pilosus var. leoniae, and Paepalanthus " Species A. "

Figure 11. Habits of Paepalanthus pilosus varieties and " Paepalanthus sp. A. " A-D Paepalanthus pilosus var. pilosus. (A Soderstrom 1346 B Sagastegui 12242 C Cano 16840 D Larsen 237.) E Paepalanthus pilosus var. leoniae (Leon 1597) F-G " Paepalanthus sp. A " (Cuadros 3732) F Habit G Capitulum. In D arrows indicate flowering (fl) and fruiting (fr) capitula.

Figure 12. Pistillate flowers and fruits of Paepalanthus pilosus varieties. A-K Paepalanthus pilosus var. pilosus (A-B Jorgensen 2366 C-F Cano 16840 G Cuatrecasas 25574 H MacDougal 4463 I Wurdack 1616 J Ollgaard 9557 K Boeke 2133) A-B Flower in anthesis with detail of gynoecial nectaries C Flower in anthesis D Gynoecium E Petal F Seed G Diaspore with detached calyx H Sepal (fr) I Diaspore releasing seed J-K Seeds after wetting L Paepalanthus pilosus var. leoniae. Mature diaspore, sepals and petals intact. (Leon 1597).