Barnadesia Mutis ex Linnaeus

Ferreira, Paola De Lima, Antonelli, Alexandre & Groppo, Milton, 2021, Touch me carefully: a step towards understanding morphological diversity in the South American spiny sunflowers (Compositae, Barnadesioideae), Phytotaxa 518 (2), pp. 109-142 : 116-117

3. Barnadesia Mutis ex Linnaeus f. (1782: 55)

Type: — Barnadesia spinosa L. f.

Arching shrubs or trees, up to 20 m tall, multistemmed. Stems erect, decumbent or sometimes scandent, much-branched, lenticelate, cylindrical or rarely flat, with or without scales imbricate at base, glabrous, tomentose, villous, velutinous, sericeous, rarely dendritic, unarmed or with axillary spines in pairs or fascicles, straight or curved, convergent or divergent, glabrous, villous or pubescent at base and becoming glabrous towards the apex. Leaves alternate or in fascicles, sessile to petiolate, persistent or deciduous, blade elliptical, oblanceolate, or obovate, chartaceous to coriaceous, pale or lustrous, glabrous to densely villous, tomentose or sericeous on both surfaces, apex mucronate or spiny, margin flat, glabrous or ciliate, base acute or obtuse; venation eucamptodromous or actinodromous with 3 suprabasal nerves. Capitulescence terminal or axillary, monocephalous, corymbose or racemose. Capitula 9 or 16-flowered, heterogamous or homogamous, radiate, sessile to pedunculate, involucre cylindrical, turbinate or campanulate, 6–14-seriate, phyllaries scarious to coriaceous, green, brown, reddish-brown to purple, erect or reflexed, glabrous or pubescent, ovate grading to lanceolate, apex unarmed, mucronate or spiny, margin flat, ciliate. Receptacle flat, pilose. Flowers dimorphic or isomorphic. Ray flowers 8 or 13, bisexual, corolla subbilabiate (4+1), 5-lobed, red to purple, pink or rarely white, externally villous, internally glabrous or sericeous on throat, rarely sericeous on tube. Disc flowers 1 or 3, bisexual or pistillate by the androecium atrophy or staminate by the suppression of the style (only in Barnadesia subgenus Bacasia ), corolla tubular (5+0, 4+0, 3+0), subbilabiate (4+1, 3+1) or ligulate (5+0), 3–5- lobed, purple, pink or rarely white, externally and internally glabrous or villous. Anthers 5 (ray flowers) or 3–5 (disc flowers), apical appendage lanceolate to obtuse, basal appendage obtuse or shortly sagittate, ecaudate, ecalcarate, stamens inserted at the throat (ray flowers) or at the base or between the base and the throat (disc flowers), filaments glabrous, free, partially connate or totally connate into a staminal tube, (ray and disc flowers), or rarely connate in disc flowers, (only in B. spinosa ) Style cylindrical, purple. Cypsela (ray flowers) cylindrical or rarely turbinate, villous, or (disc flowers) turbinate, densely villous. Pappus (ray flowers) plumose, shorter than the corolla length, or (disc flowers) plumose, barbellate or setaceous, when barbellate or setaceous usually recurved at maturity, glabrous to villous, equal to or shorter than the corolla, bristles whitish. Pollen lophate, radially symmetric or asymmetric, smooth (Suessy & Urtubey 2007). Chromosome number = 12, 24, 25, 31, ca. 48, ca. 50, 62 ( Heiser 1963, Olsen 1980, Strother & Panero 1994, Stuessy et al. 1996).

Fig. 4 ; 10c .

Distribution and habitat: — Barnadesia is the third largest genus of Barnadesioideae , comprising 19 species distributed mainly along the Andes from Colombia to Argentina, with the highest number of species found in Peru ( Urtubey 1999, Hind 2001, Stuessy et al. 2009). The species occur at the edges of tropical and subtropical mixed (deciduous-evergreen) montane forests, and gallery forests; there are also records in the Páramos (alpine meadows) and in the Puna (highlands and steppe), c. 3,700 m a.s.l. One species, Barnadesia caryophylla (Vell.) S.F. Blake occurs in Brazil, in the Cerrado (Brazilian savanna) and in the Atlantic Forest (tropical or subtropical evergeen-semideciduos forest) ( Saavedra 2020).

Notes: —The species of the genus are classified in two subgenera (following Urtubey 1999): Barnadesia subg. Barnadesia , with 17 species from Colombia to Argentina and one species in Brazil ( B. caryophylla ), characterized by sessile or subsessile leaves disposed in fascicles, sessile or shortly pedunculate capitula, disc flowers with subbilabiate or ligulate corolla with pappus simple or barbellate; and Barnadesia subg. Bacasia , with two species distributed from Colombia to Bolivia, characterized by alternate and petiolate leaves, pedunculate capitula, disc flower with tubular corolla and plumose pappus.

Barnadesia is the type genus of the subfamily and can be distinguished from the other genera by the following set of characters: arching shrub or trees, radiate capitula, 8 or 13 ray flowers, 1 or 3 disc flowers, colorful corolla (pink, red, purple, white), ecalcarate and ecaudate anthers, obtuse apical appendage, free filaments, partial or connate into a staminal tube, the pappus of the disc flower usually setaceous or barbellate that recurves at maturity, and lophate pollen.

Iconography: — Chung (1965), Harling (1991), Urtubey (1999), Ferreyra (1995), Hind (2001).

Accepted species: — Barnadesia subgenus Barnadesia : 3.1 Barnadesia aculeata (Benth.) I.C. Chung ; 3.2 Barnadesia arborea Kunth ; 3.3 Barnadesia blakeana Ferreyra ; 3.4 Barnadesia caryophylla (Vell.) S.F.Blake ; 3.5 Barnadesia dombeyana Less ; 3.6.1 Barnadesia glomerata Kuntze var. glomerata ; 3.6.2 Barnadesia glomerata var. mucronata I.C. Chung ; 3.7 Barnadesia horrida Muschl. ; 3.8 Barnadesia jelskii Hieron. ; 3.9.1 Barnadesia lehmannii var. angustifolia I.C. Chung ; 3.9.2 Barnadesia lehmannii var. ciliata I.C. Chung ; 3.9.3 Barnadesia lehmannii Hieron var. lehmannii ; 3.9.4 Barnadesia lehmannii var. villosa (I.C. Chung) Urtubey ; 3.10 Barnadesia macbridei Ferreyra ; 3.11 Barnadesia macrocephala Kuntze ; 3.12 Barnadesia odorata Griseb. ; 3.13 Barnadesia polyacantha Wedd. ; 3.14 Barnadesia pycnophylla Muschl. ; 3.15 Barnadesia reticulata D. Don ; 3.16 Barnadesia spinosa L. f.; 3.17 Barnadesia woodii D.J.N. Hind. Barnadesia subg. Bacasia (Ruiz & Pav.) Urtubey : 3.18 Barnadesia corymbosa (Ruiz & Pav.) D. Don ; 3.19 Barnadesia parviflora Spruce ex Benth. & Hook. f.

Selected specimens examined: — ARGENTINA. Jujuy: Capital, Lozano-Tiraxi, 11 November 1978, S.A. Renvoize 3439 (B; 3.12). Salta: Capital, Castellanos, Arroyo Castellanos 5 Km al W del puente a Yacone, 1500–1800 m, 15 September 1988, L.J. Novara 8098 (B, 3.12). Santa Victoria, Los toldos, Quebrada del Artillera, 4–7 km al E de la Municipalidad, 1500–1700 m, 17 March 1986, L. Novara 5225 (B; 3.12). Tucumán: Tafí Viejo, Cumbres de San Javier, Ruta Provincial 340, 04 November 1978, S.A. Renvoize 3340 (B; 3.12). BOLIVIA. Cochabamba: Ayopaya, Independencia, in the second valley north of La Mina on road to Sailapata, c 48 km N of Independencia, 2800 m, 16°56.35’S, 66°57.05’W, 15 May 2002, J.R.I. Wood 18477 (K; 3.6.1). Carrasco, Siberia-Cochabamba, 2800 m, March 1959, Cárdenas 5736 (US; 3.12). Carrasco, Dianpampa, 2160 m, 17°40’53’’S, 64°40’55’’W, 09 September 2003, S. Duran 6 (MO; 3.18). Carrasco, Siberia, Chua Kocha, 2195 m, 17°48’56’’S, 64°42’27’’W, 22 June 2003, E. Fernández 1943 (MO; 3.6.2). Cochabamba, Quebradas von Tiraque, 2900 m, 13 November 1928, J. Steinbach 8723 (MO, K; 3.11); Chapare road, 36 km from Cochabamba, roadside, 12.000 ft (c. 3.557 m alt.)., 22 August 1964, W.J. Badcock 296 (K; 3.11). Tapacari, kilo 86 (near Sarari) on the road from Cochabamba to Oruro, 4000 m, 22 June 1997, J.R.I. Wood 12323 (K, 3.11). Chuquisaca: Luis Calvo, comunidad Chuya Yaco, a 3 km aproximadamente al Sur-este de la comunidade de Chuya Yaco, 1239 m, 19°45’12’’S, 63°52’36’’W, 20 October 2005, A. Lliully 461 (MO; 3.18). Sud Cinti, ca. 11 km al NW de la Comunidad de Las Abras caminho hacia la comunidade de El Palmar, pendiente superior cerro Alto de Rosas, 862 m, 21°02’24’’S, 64°13’08’’W, 10 June 2005, R. Lozano 1380 (MO; 3.18). La Paz: Franz Tamayo, Parque Nacional Madidi entre Piñalito y Cuchiwani, 2140 m, 14°30’44’’S, 68°16’28’’W, 21 July 2012, A.F. Fuentes 17401A (MO; 3.18). Inquisivi, along the trail between Loma El Abra and Cerro Negro Kkota, a 6 km hike, ca. 6 km N from Inquisivi, 2900–3000 m, 16°50’S, 67°08’W, 22 August 1988, M. Lewis 881108 (MO; 3.6.2). Murillo, valle del río Zongo, 21.1 km al N de la cumbre, 3200 m, 16°09’S, 68°07’W, 04 April 1987, J.C. Solomon 16453 (MO, K; 3.14). Saavedra, Rio Comata valley, on descent from Charazani on road to Apolo, 1600 m, 06 August 2000, J.R.I. Wood 16594 (K; 3.17). Potosí: Charcas, Toro Toro, Siti Wilta caminho de Toro Toro a San Pedro de Buena vista, 3418 m, 18°07.99’S, 65°50.25’W, 27 February 2003, J.R.I. Wood 19251 (K; 3.14). Santa Cruz: Comaropa, Jungas de San Mateo, 2500 m, 22 October 1928, J. Steinbach 8400 (K; 3.6.2). Florida, Comunidad de Bella Vista, 1600–1900 m, 18°14’S, 63°39’W, 18 June 2006, L. Arroyo 3406 (K; 3.18). Samaipata, Bella Vista, sendero Ecologico el Cañadon, 1487 m, 42°80’38’’S, 79°83’15.4’’W, 08 September 2005, D. Villarroel 151 (MO; 3.18). Vallegrande, Huasacañada, 5 km al S de la ciudad de Vallegrande, 2050 m, 18°31.5’S, 64°5.8’W, 05 August 1991, I.G. Vargas 1071 ( US; 3.12). Tarija: Entre Ríos, Tarija road, c.km. 40, 19 July 1964, W.J. Badcock 273 (K, 3.12). BRAZIL. Goiás: Silvania, Área de influência da AHE Corumbá IV, 23 May 2003, A.A. Santos 2053 (CEN; 3.4). Serranópolis, RPPN Pousada das Araras, 18°26’22’’S, 51°59’43’’W, L.F. Souza 2980 (HJ; 3.4). Mato Grosso: Juína, ca 40 km WNW (em linha reta) de Juína, 11°18’S, 59°05’W, 06 July 1997, V.C. Souza 18251 (UEC; 3.4). Minas Gerais: Carandaí, Pedra do Sino Hotel Fazenda, BR 040 Km 6, Trilha da matinha atraz da cas., 1000–1200 m, 15 July 2005, N.F.O. Mota 286 (BHCB; 3.4). Lima Duarte, arraial de Conceição do Ibitipoca, Parque Estadual do Ibitipoca e Pedra do Gavião, 20 May 2001, R.M. Castro 393 (CESJ; 3.4). Santana do Garambeu, Trecho do Alto Rio Grande adjacente a zona urbana de Santana de Garambeu, 1100–1150 m, 21°36’12.6’’S, 44°07’25.3’’W, 08 June 2001, A. Salino 7007 (BHCB; 3.4). São Paulo: Cunha, 20 April 1939, J. Kiehl 3711 ( US; 3.4). São José do Rio Preto, Mirassol, 07 August 1991, N. Taroda-Ranga s.n. (SJRP; 3.4). COLOMBIA. Boyacá: Cordillera Oriental, Sierra Nevada del Cocuy, Quebrada de San Paulino próximo Alto Ritacuva, 3500 m, H.G. Barclay 7305 (MO; 3.16). Cauca: Puracé, márgen derecha Río Vinagre, corregimiento Chibchiguará, 3250 m, 02°23’47’’ S, 76°50’21’’ W, 15 January 1991, R. Ruiz 1232 (MO; 3.16). Cudinamarca: Above Guasca, 10.000 ft. (c. 3058m), 16 December 1938, E.K. Balls 85767 (K; 3.16). Salto de Tequendama, 25 km westsouthern of Bogotá, western margin Cordillera Oriental, 17 June 1950, S. Galen Smith 1091 (K; 3.16). Meta: Puerto López, 3000 m, 12 August 1046, M. Jaramillo Mejia 402 (US; 3.16) Nariño: Near top of divide on Tuquerres-Ricaurte road, roadside subparamo, 3000 m, 22 November 1981, Al. Gentry 34833 (MO; 3.16). Putamayo: Valle de Sibundoy, Sibundoy, ca. 2200 m, M.L. Bristol 858 ( US; 3.16). Quindío: Salento, Paraje Cocora, orilla del río Quindío, 2470 m, 11 June 1983, S. Royos 203 (MO; 3.19). Risaralda: La Pastora, Reserva Ucumari, 2610 m, 11 October 1989, D. Rangel 5406 (MO; 3.19). Santander: Páramo de las Puentes, above La Baja, 3500–3700 m, 25 January 1927, E.P. Killip 18250 (US; 3.16). Valle del Cauca: Tuluá, Corregimiento Santa Lucía, finca San Luis, 2800 m, 22 September 1984, W. Devia 724 (MO; 3.16). ECUADOR. Azuay: 8–10 km. S of Cumbe on the road to Loja, 9050–9400 ft. (c. 2758–2865 m), 22 January 1979, R.M. King 7762 (MO; 3.2). Bolívar: Simiatug, Hacienda Talahua, 3200 m, 28 Apri; 1939, C.W. Penland 547 (US; 3.2). Carchi: Wooded hills about 5 miles south of Tulcán, 2500 m, 10 August 1923, A.S. Hitchcock 21012 ( US; 3.16). Cañar: 20 km S of Cañar on the road to Cuenca, along railroad tracks, 9000 ft. (c. 2743m), 21 January 1979, R.M. King 7742 (MO; 3.2). Cotopaxi: Corazón, E.F. André K-271 (K; 3.2). Imbabura: Cotacachi, Hda. La Florida, 1900–2500m, 00°23’S, 78°28’W; 27 August 1992, A. Alvarez 614 (QCA; 3.19). Guayas: Guayaquil Canton, Bosque Protector Cerro Blanco, carretera a Salinas, km 15, 350 m, 02°10’S, 79°58’W, 15 August 1991, D. Rubio 1790 (MO; 3.9.4). Loja: Old road to Catamayo (La Toma)-Loja, km 7, c. 18 km from Catamayo, 2600 m, 04°00’S, 79°16’W, 21 May 1997, G.P. Lewis 3329 (MO; 3.1). Cordillera de Las Lagunitas, Amaluza-Jimbura-Zumba, km 41, 3270 m, 4°44’55”S, 79°25’13”W, 23 November 1994, P.M. J ø rgensen 792 (MO; 3.1). Napo: Baeza, 2 km antes de Baeza, carretera Papallacta-Baeza parte alta de “Y”, 2000 m, 00°28’S, 77°54’W, 11–14 July 1992, R. Valencia 2461 (QCA; 3.19). Quijos, Serra Azul, Agrícola Industrial Río Aragón, Colinas del Tundal, 2200 m, 00°40’S, 77°54’W, 30 April 1992, A. Alvarez 300 (MO, QCA; 3.19). Cerca de baños termales, dos kms. Al N. de Papallacta, 22 March 1977, S. González 17 (MO; 3.2). Pichincha: Along road from Santo Domingo de los Colorados and Alóag, ca. 14 km W of Alóag, 2250 m, 00°26’54”S, 78°42’44”W, 01 August 2004, T.B. Croat 92841 (MO; 3.19). Quito, carretero Nono- Guarumos-Tandayapa, colleciones en sendero entre Guarumos y Cerro Guarumos, derecho de vía del Oleoducto de Crudos Pesados, 2400–2700 m, 00°03’S, 78°38’W, 01 September 2001, A.F. Fierro 3148 (MO; 3.19). Tabacundo, Quebrada Verdetola, km 2.8, 3000 m, 00°12’S, 78°23’W, 09 July 2000, B. Cuamacás 623 (MO; 3.8). Tambilho, 2700 m, 25 May 1939, E. Asplund 6439 (K; 3.2). Tungurahua: Bãnos-puyo road 6 km, South of Bãnos, trail leading disturbed wet montane forest, 2500 m, 01°22’S, 78°25’W, 10 December 1983, A. Barfod 49003 (QCA; 3.19). PERU. Ancash: Bolognesi, cerros al E. de Chiquian, 3700 m, 10 May 1950, E. Cerrate 523 (MO; 3.5). Amazonas: Chacapoyas, 28 kms along road from Leimebamba SW towards Celendín, 9700 ft. (c. 2956m), R.M. King 9254 (MO; 3.2). Bongará, banda frente a Pomacochas, en trocha, 2200–2300 m, 17 August 1963, R. Ferreyra 15201 (MO; 3.9.3). Mendoza, 1400 m, August 1963, F. Woytkowski 8121 (MO; 3.4). Partly along main road Jumbilla-Pedro Ruiz, partly along road to Tialango, 05°52’31”S, 76°46’36”W, 03 November 2012, H. van der Werff 24976 (MO; 3.9.3). Apurímac: Valley of Rio Colcachaca at Cotarusi-Colce ca. 1 km above junction w Rio Chalhuanca ca. 15 km (air) S of Chalhuanca, 3000–3100 m, 16 December 1962, H.H. Iltis 541 (US; 3.7). Ayacucho: Humanguilla, ca. 15 km before Ayacucho left of road Huanta-Ayacucho, 10.000 ft. (c. 3048m), 01 January 1962, S.G.E. Saunders 725 (K; 3.7). La mar, commonest of dry puna scrub along roadside from Ayacucho to San Francisco, shortly after crossing over the 2nd high pass, 34 km S of Tambo, c. 10 km from Rirconado at 3 am in the morning, 3500 m, 13°S, 74°W, 07 June 1968, T.R. Dudley 9053 ( US; 3.14). Cajamarca: Celendín, on roado to Celendín, 3 km east of Encañada, 3200 m, 16 May 1964, P.C. Hutchison 5120 (MO; 3.5). Celendín, cerca a Celendín, 3400–3500 m, 21 June 1963, R. Ferreyra 15032 (MO; 3.8). Contumazá, arriba de Trinidad, 2200 m, 06 July 1977, A. Sagástegui A. 8945 (MO; 3.9.3). Cutervo, entre Cutervo y Cochabamba, 2500–2600 m, 01 August 1946, R. Ferreyra 838 (MO; 3.8). entre Llama y Chongoyape, 2200–2500 m, 02 August 1946, R. Ferreyra 862 (MO; 3.9.3). Cusco: Cusco, road to Pisac, ca 1 km after ruins of Sacsayhuaman, 3700 m, 13°30’07”S, 71°58’50”W, 10 September 2002, M. Ackermann 262 (B; 3.7). Qorao, abra de Qoarao, 3817 m, 13°29’S, 71°56’W, 21 March 2003, W. Galiano 4793 (MO; 3.7). Urubamba, trilha para a lagoa Yanacocha, 3449 m, 13°18’01.7”S, 72°03’13.6’‘W, 11 January 2016, G. Cappatinta 54 (SPFR; 3.7). Huacanvelica: Paucartambo, 11.500 ft. (c. 3505m), 03 May 1939, B. Balls 6683 (K; 3.7). Huánuco: Ambo, Tomayaquichua, a caminho de Estanco, 3609 m, 76°08’52.0’’W, 10°03’52.9’’S, 27 January 2016, G. Cappatinta 49 (SPFR; 3.14). on open slope of ravine, 3 km east of Acomayo, 26 July 1946, F. Woytkowski 34335 (MO; 3.10). Huánuco, km 474 on Lima-Tingo Maria road, 1650 m, 02 June 1981, K. Young 632 (MO; 3.18). Llata, about 7000 ft. (c. 2133m), 21 August 1922, Macbride 2250 ( US; 3.5). Huaraz: Recuay, aos redores de Cátac, 12 February 2016, G. Cappatinta 48 (SPFR; 3.5). Junín: Tarma, entre La Merced y Quimiri, valle del Chanchamayo, 800–900 m, 27 June 1948, R. Ferreyra 3696 (MO; 3.4). San Luíz, 14 August 1944, C. Ridoutt 14564 (MO; 3.4). Yucapata, 1400 m, 16, July 1961, F. Woytkowski 6637 (MO; 3.4). Ica: Pisco, Arriba de Pisco, 2900 m, 20 March 1954, W. Rauh 386 (MO; 3.15). La Libertad: Otuzco, 9 km west of and below Shorey toward Trujillo, 3590 m, 11 August 1964, P.C. Hutchison 6289 (MO; 3.5). Lima: Canta, alreadores de Canta, 2900–2950 m, 16 March 1950, R. Ferreyra 6914 (MO; 3.15). Canta, cerca a San Juan, Valle de Chancay, 3000–3200 m, 29 May 1974, R. Ferreyra 18415 (MO; 3.5). Huarochirí, Monte de Zárate, encima de San Bartolomé, 2900 m, R. Ferreyra 10427 (MO; 3.3). Moyobamba, Mathews 1389 (K; 3.9.1); above Huamantanga and Obrajilla, Mathews 643 (K; 3.9.1). Pasco: Oxapampa, road from Paucartambo to Oxapampa, towards Tingo Chontabamba, km 82, 2280 m, 10°38’S, 75°25’W, 04 April 2001, M. Weigend 5485 (B; 3.18). Daniel Carrión, 16 km NE of Yanahuanca 1 km NW of Páucar, along fencerow, 3450, 01 August 1965, R. Bird 1258 (MO; 3.5). Piura: Huancabamba, Mitopampa (Huancabamba-Cuello del Indio), 2650 m, 22 July 1975, A. Sagástegui A. 8239 (MO; 3.8). Huancabamba, arriba de Palamba, 1300 m, 13 September 1981, A. Lopez M. 8802 (MO; 3.9.3). Puno: Carabaya, Ollachea, across San Gaban river from town, 23 August 1980, J.D. Boeke 3171 (MO; 3.18). Carabaya, 3 km South of Ollachea, roadside puna, 13 august 1980, J.D. Boeke 2997 ( US; 3.14). Sandia, San Juan del Oro, 1300 m, 06 August 1965, C. Vargas C. 16401 ( US; 3.4). Sandia, entre Sandia y Cuyocuyo, 3200–3300 m, 16 May 1966, R. Ferreyra 16740 (MO; 3.7). San Martín: Near to Moyobamba, 890 m, 22 June 1947, F. Woytkowski 35300 (MO; 3.4). Tumbes: Zarumilla, El Caucho, 300–400 m, September 1960, Maekava s.n. (MO; 3.9.3).

Figures

FIGURE 4. Barnadesia. A and E: Barnadesia pycnophylla. A. Habit. E. Divergent and fasciculate spines on stem. B: Barnadesia caryophylla. Capitulum. C: Barnadesia odorata. Capitulum. D: Barnadesia arborea. Capitula. Photos by Gari Ccana-Ccapatinta (A and E), Gustavo Shimizu (B), Claudia Martin (C), Carmen Ulloa (D).

FIGURE 10. Geographical distribution for the genera of Barnadesioideae. A: Dasyphyllum. B: Chuquiraga. C: Barnadesia (red dots) and Huarpea (blue triangle). D: Doniophyton (blue triangles); E: Archidasyphyllum (blue triangles); Fulcaldea (blue triangles)—these words (triagle) in plural (triangles).