Senecio melanandrus (Wedd.) J.Calvo, A.Granda & V.A.Funk, 2019

Calvo, Joel, Granda, Arturo & Funk, Vicki A., 2019, New combinations and synonyms in discoid caespitose Andean Senecio (Senecioneae, Compositae), PhytoKeys 132, pp. 111-130 : 113-119

1. Senecio melanandrus (Wedd.) J.Calvo, A.Granda & V.A.Funk comb. nov. Figs 1 , 2 , 3A, B , 4A, B

Werneria melanandra Wedd., Chlor. Andina 1: 88. 1856. Type: Bolivia. La Paz: ravin de Chuquiaguillo, 1851, H.A. Weddell s.n. (lectotype, designated by Rockhausen (1939) as “type”, pg. 284: P [P04319315]). Epitype, designated here: Bolivia. La Paz: am Chacaltaya (30 km von La Paz), 4800 m, Feb 1908, O. Buchtien 1589: US [US00622639]; isoepitype: US [US00622640].

Senecio humillimus var. melanolepis Wedd., Chlor. Andina 1: 104. 1856. Type: Bolivia. La Paz: Larecaja, viciniis Sorata, ad lacum Yuriguana, prope Anilaya, Ancumpampa, prope Ancohuma, 3800-5000 m, Apr 1860, G. Mandon 108 (neotype, designated here: GH [GH00012144]; isoneotypes: P [P03730752, P04370980], S [S-R-10871]), syn. nov.

Senecio vegetus var. lobatus Cabrera, Notas Mus. La Plata, Bot. 18(89): 222. 1955. Type: Bolivia. La Paz: Ingavi, Miriquiri, 4200 m, 10 Mar 1921, E. Asplund 2866 (holotype: S [not located, Arne Anderberg in litt.]), syn. nov.

Senecio pucapampaensis H. Beltrán, Arnaldoa 15: 212. 2009. Type: Peru. Huancavelica: Pucapampa, debajo de Chonta, 4500-4600 m, 9 May 1958, O. Tovar 2959 (holotype: USM-00277274), syn. nov.

Senecio sykorae Montesinos, PhytoKeys 39: 6. 2014. Type: Peru. Moquegua: General Sánchez Cerro, Yunga, E of Yunga, on the peaks of Perusa mountain, 16°11'08"S, 70°38'14"W, 4802 m, 13 Apr 2012, D. Montesinos & F. Calisaya 3805 (holotype: USM s.n.; isotype: HUSA n.v.), syn. nov.

Senecio tassaensis Montesinos, PhytoKeys 39: 11. 2014. Type: Peru. Moquegua: General Sánchez Cerro, Ubinas, cumbre nevada del cerro Pirhuani Querala, 4650 m, 16°09'S, 70°43'W, 7 Apr 2011, D. Montesinos 3103 (holotype: HUSA n.v.; isotypes: MOL n.v. [not located, likely never sent], USM-247549), syn. nov.

Senecio canoi P. Gonzáles, Montesinos & Ed. Navarro, Anales Jard. Bot. Madrid 72(2): 1. 2015. Type: Peru. Puno: Carabaya, Corani, Minaspata, arriba de Chacaconiza, 14°01'57"S, 70°41'54"W, 4999 m, 14 Apr 2014, P. Gonzáles 2989 (holotype: USM n.v.), syn. nov.

Senecio vegetus sensu Cabrera (1955, 1985), non Weddell (1856).

Description.

Caespitose perennial herb. Leaves 4-15 mm long, 1.2-2.6 mm wide, linear-oblong to spatulate, apex acute to obtuse, base narrowed, margins entire, crenate or dentate, conduplicate downwards (rarely flat), glabrous, with marginal trichomes on the narrowed base or densely pilose, somewhat fleshy, greenish or glaucous. Capitulum discoid, solitary, terminal, sessile or subsessile; involucre 5-8 mm long, 3.7-9 mm wide. Involucral bracts 11-16, oblong-lanceolate, 3.8-4.9 mm long, 0.9-1.8 mm wide, partially fused at the base, smooth, glabrous or with trichomes on the abaxial surface ca. 0.7 mm long, dark purple- or blackish-tipped. Supplementary bracts (calyculus) 2 –4(– 6), linear to slightly spatulate, 4.2-7.5 mm long, 0.5-1 mm wide, smooth, three-quarters to as long as the involucral bracts, with trichomes (rarely glabrous), dark purple- or blackish-tipped. Disc florets 20-45, 4.3-6.3 mm long, 0.8-1.1 mm wide, 5-lobed, conspicuously differentiated in a distinct tube and campanulate limb, whitish. Anther bases auriculate, clearly acute, dark purple to blackish; filament collar balusterform. Style branches truncate with a crown of sweeping hairs, dark purple to blackish. Achenes 2.1-2.2 mm long, ca. 0.5 mm wide, brownish, covered by dense indumentum of obtuse whitish myxogenic twin trichomes ca. 0.2 mm long; pappus 5-6 mm long, barbellate, whitish. Chromosome number: unknown.

Additional iconography.

Beltrán (2008: pg. 216, fig. 2, sub S. pucapampaensis ); Montesinos-Tubée (2014: pg. 7, fig. 2; pg. 13, fig. 4B, sub S. sykorae ); Montesinos-Tubée (2014: pg. 12, fig. 3; pg. 13, fig. 4C, sub S. tassaensis ); Montesinos-Tubée et al. (2015: pg. 2, fig. 1; pg. 3, fig. 2, sub S. canoi ).

Distribution and habitat.

Bolivia (Cochabamba, La Paz, Oruro, Potosí) and Peru ( Apurímac, Arequipa, Ayacucho, Cusco, Huancavelica, Moquegua, Puno) ( Fig. 5 ). It grows in exposed places mainly in the subhumid and dry puna ecoregions, at elevations of 3800-5100 m.

Phenology.

Flowering mainly from January to June, although some flowering specimens have been collected in November.

Etymology.

The epithet melanandrus means having dark or black stamens, which describes a striking character of this species.

Discussion.

This species is transferred to Senecio on the basis that it has genuine supplementary bracts (calyculus), the involucral bracts are not clearly fused at the base, it displays a caespitose habit with short stems, and it has myxogenic achene trichomes. Furthermore, its morphologically most similar species are currently treated as Senecio members: i.e., S. digitatus , S. madidiensis J. Calvo & A. Fuentes, S. pygmophyllus (see new combination below), and S. scorzonerifolius Meyen & Walp. All the names included in the synonymy were also described within the genus Senecio .

Senecio melanandrus is a highly variable species that has been variously interpreted. The poor condition of the type material probably helped to maintain the uncertainty surrounding the application of this name. Weddell (1856) described the leaves as "integerrimis vel nonnullis dente triangulari, [ …] glabriusculis vel inconspicue ciliolatis" [entire or with a few triangular teeth, rather glabrous or inconspicuously ciliate]. Several years later Rockhausen (1939), who published the first comprehensive taxonomic revision of the genus Werneria , stated that the leaves have "marginibus obsolete glanduloso-ciliolata" [margins scarcely glandular-ciliate]. On the basis of our studies, this species displays an unusually wide variability in leaf margin and indumentum of leaves and involucre, which is reflected in the number of names included in the synonymy. The leaf margin may be entire, crenate or dentate, variability that can be even found in the same individual. Likewise, the leaf indumentum varies from densely pilose ( Fig. 3A, B ) to almost glabrous ( Fig. 4A, B ). In Bolivia, the pilose forms are common although some glabrescent specimens are found near Nevado Sajama (Liberman 821, LPB, US) and in northern La Paz Department (Menhofer 1901, US). The glabrous forms also ap pear in the Peruvian regions of Huancavelica, northern Ayacucho, and northern Puno, and were recently treated under the names S. pucapampaensis H. Beltrán and S. canoi P. Gonzáles & al. [see Calvo and Fuentes (2018)]. These glabrous plants usually have dentate leaves, but forms exist with rather subentire leaves ( Gonzáles 3568, USM). The dentate, pilose forms that are frequently found in Bolivia were described as S. tassaensis Montesinos on the basis of a single collection from Moquegua (southern Peru). From the same region, a form with almost entire, glabrous leaves was named S. sykorae Montesinos. This form was also collected near the Bolivian locality of Ulla Ulla (Menhofer 1901, US). This puzzling distribution pattern and a continuum of intermediates sug gest that these forms do not deserve taxonomic recognition. Despite this variability, S. melanandrus is well characterized by supplementary bracts that are almost as long as the involucral bracts, the leaf lamina narrowed at the base, the blackish anthers and style branches, the whitish corollas, and by its myxogenic achene trichomes. The apex of the involucral bracts is usually remarkably dark-colored. Indeed, the epithet melanolepis of Weddell’s varietal name, here included in the synonymy, explicitly refers to this character, i.e., having black scales (involucral bracts). It is noteworthy that the anther bases were hitherto described as obtuse; however, they are auriculate and clearly acute.

The name Senecio vegetus var. lobatus Cabrera, here synonymized with S. melanandrus , was included by Cabrera (1985) in the synonymy of S. vegetus (Wedd.) Cabrera. Cabrera (1955, 1985) described this latter species as having silky-pubescent achenes. We had the opportunity of studying some of the specimens that he examined and they indeed correspond to S. melanandrus (i.e., Beck 7952, Mandon 108, Menhofer 2013, Weberbauer 7491). Cabrera’s interpretation of S. vegetus (≡ S. humillimus var. vegetus Weddell) might be explained by the fact that one of the syntypes (P [P01816588]) contains mixed material and some plants certainly correspond to S. melanandrus (the individual on the left hand and likely the fragment at the right hand below). The syntype P [P01816917], not containing admixtures, shows plainly glabrous plants with the leaves entire and obtuse at the apex. Therefore, and in disagreement with Cabrera, we consider that S. vegetus and S. melanandrus correspond to distinct taxonomic entities. The former belongs to the subgroup with yellowish anthers, style branches, and corollas whereas the latter is a member of the subgroup displaying blackish anthers and style branches and whitish corollas. However, it is important to point out that the accurate taxonomic position of S. vegetus remains uncertain. Because of the number of involucral bracts, the leaf morphology, the yellowish corollas, and the presumed glabrous achenes, we believe that this taxon is related to S. gamolepis Cabrera. Additional studies are needed to establish its correct taxonomic position. For the time being, we prefer not including it in the key.

Our efforts to locate the type material of S. humillimus var. melanolepis Wedd. were unsuccessful. In fact, all the collections cited in the protologue that were located correspond to the other varieties described by Weddell. For that reason, we selected as the neotype a Mandon collection that perfectly matches the diagnosis provided by Weddell. Moreover, it was identified as S. humillimus var. melanolepis by Schultz Bipontinus [see Mandon (1865)], which supports our interpretation of this taxon. The specimen P03730757 is excluded because it contains mixed material.

The holotype of the name S. canoi should be housed at USM ( Montesinos-Tubée et al. 2015); however, it was not located. The paratype Gonzáles 3429 (USM) was also not located at USM. As a result, we studied the collections Gonzáles 3428 and Gonzáles 3441 (USM), both collected around the locus classicus on the same day as the paratype. Likewise, the holotype of Senecio vegetus var. lobatus appears to be missing (Arne Anderberg in litt.). Cabrera indicated as paratype the collection Mandon 108, which is here selected as neotype for S. humillimus var. melanolepis .

Finally, in order to remove any uncertainty surrounding the application of this name, and considering that the conditions of the type material are deficient for a proper study of the diagnostic characters, we consider it appropriate to designate an epitype. The selected specimen is Buchtien 1589 (US00622639) from Chacaltaya, a mountain not far from the locus classicus of W. melanandra . A duplicate was found at US.

Selected specimens examined.

BOLIVIA. Cochabamba: Arque, Cruce Ventilla, 17°46'S, 66°40'W, 17 May 1981, O. Murgia 276 (LPB); cordillera del Tunari, cumbres del cerro Tunari, 17°17'S, 66°23'W, 25 Mar 1990, G. Navarro 653 (BOLV); Tapacarí, arriba rancho Wacakhariña, 3 km al NE de Japo K’asa (km 125 Cbba-Oruro), 17°39'S, 66°45'W, 9 Mar 1995, H.U. Pestalozzi 446 (BOLV); Tiraque, P.N. Carrasco, cordillera Juno, 17°18'S, 65°41'W, 18 Mar 2001, M. Zárate & D. Méndez 1087 (LPB); La Paz: Murillo, La Paz 32 km hacia Unduavi, 16°19'S, 68°2'W, 3 Apr 1983, S.G. Beck 7952 (LPB); Murillo, camino La Paz-Lambate, cerca Apacheta entrando al desvío hacia el Illimani, 2 km entrando hacia Milla Milla, 16°34'S, 67°52'W, 6 Apr 2012, S.G. Beck 32782 (LPB); Murillo, La Paz subiendo el valle Kaluyo hasta el albergue ecoturístico Pampalarama, 16°19'S, 68°4'W, 22 Mar 2009, S.G. Beck 33091 (LPB); Murillo, subiendo el valle de Irpavi hasta Palcoma, subiendo el río Hati Jahuira, 16°25'S, 67°57'W, 26 Apr 2013, S.G. Beck 34141 (LPB); Los Andes, above cumbre (pass) on rd. through Hichu-Kkota valley on rd. to mina La Fabulosa, 21 km from base of lag. Khara Kkota, 16°10'S, 68°20'W, 29 Apr 1995, V.A. Funk 11406 (US; the duplicate at LPB corresponds to Werneria apiculata Sch. Bip.); Murillo, Zonga valley, laguna Pata Kkota, 1.5 km S of pass, 16°18'S, 68°7'W, 11 Apr 1995, V.A. Funk & N. Bernal 11270 (LPB, US); Murillo, nev. Huayna Potosí, E slopes above rd., 16°17'S, 68°8'W, 12 Apr 1995, V.A. Funk & N. Bernal 11284A (US); Franz Tamayo, Canhuma (Ulla-Ulla), subiendo al cerro Laramani, 15°0'S, 69°6'W, 22 Jan 1983, X. Menhofer 1901 (US); Franz Tamayo, estancia Okaria (Ulla-Ulla), 15°3'S, 69°6'W, 24 Feb 1983, X. Menhofer 2013 (LPB); Murillo, 3.4 km N of Milluni on road to Zongo, 16°18'S, 68°7'W, 25 Apr 1985, J.C. Solomon & M. Moraes 13440 (LPB, US); Ingavi, cantón Jesús de Machaca, comunidad Titicani-Tacaca, a 20 km de Guaqui, 16°41'S, 68°49'W, 8 Apr 1989, X. Villavicencio 457 (LPB); Oruro: Eduardo Abaroa, Challapata, comunidad Churacani, cerca a la laguna Chullumpiri, 18°55'S, 66°40'W, 1 Apr 2018, M. Guzmán 125 (LPB); Sajama, nevado de Sajama, sur del cerro Jasasuni [Asa-asuni], 18°11'S, 68°55'W, 18 Mar 1984, M. Liberman 821 (LPB, US); Sajama, cantón Sajama, 18°10'S, 68°55'W, 17 Feb 1998, F. Loza de la Cruz 315 (LPB); Potosí: cordillera Kari Kari, aprox. 3.2 km arriba de la laguna San Sebastián, 19°37'S, 65°41'W, 13 Feb 2019, J. Calvo & M. Zárate 7872 (BOLV); José M. Linares Lizarazu, comunidad Alkatuyo, cerro Ichurata, 53 km SE de Potosí, 14 km al N de la escuela de Alkatuyo, 19°53'S, 65°33'W, 22 Jan 1994, F. Marino 309 (LPB). PERU. Apurímac: Antabamba, Juan Espinoza Medrano, paraje Ccanccahuane a 18 km al S de la comunidad campesina de Mollebamba, zona Minaminayoc, 14°29'S, 72°52'W, 5 Jun 2017, B. Espinoza-Prieto 534 (USM); Arequipa: pr. Chivay, ladera S del nevado Huarancante, 15°45'S, 71°32'W, 1 Apr 2005, C. Aedo & A. Galán 11022 (MA, USM); Castilla, Orcopampa, minas de Poracota, cerca a quebrada Faculla, 15°14'S, 72°32'W, 20 Apr 2011, H. Beltrán 7112 (USM); La Unión, Huaynacotas, Sarajorepampa, 15°1'S, 72°47'W, 18 Mar 2011, D. Montesinos 2949 (USM); Ayacucho: Huanca Sancos, Putajasa, 14°6'S, 74°14'W, 24 Feb 2002, A. Cano et al. 11963 (USM); Huanta, mt. Razuhuilca, 12°52'S, 74°9'W, 4-6 Feb 1926, A. Weberbauer 7491 (CONC, F); Cusco: Chumbivilcas, Santo Tomás, compañía minera Azuca (borde departamental Cusco-Apurímac), 14°35'S, 72°25'W, 13 Apr 2011, H. Beltrán 7032 (USM); Velille, Uchucarco, cerca a Soracocha, 14°26'S, 71°44'W, 23 Apr 2015, P. Gonzáles 3600 (USM); Velille, Uchucarco, cerca a Soracocha, 14°26'S, 71°44'W, 23 Apr 2015, P. Gonzáles 3601 (USM); cordillera de Vilcanota, cuenca de la laguna Sibinacocha, cerro Rititica, 13°45'S, 71°4'W, 5 Mar 2019, R.I. Meneses et al. 6968 (LPB); Huancavelica: Huaytará, Pilpichaca (abra Apacheta), 13°20'S, 74°44'W, 4 Jul 2010, A. Cano, W. Mendoza & A. Delgado 19680 (USM); Huachocolpa, alrededores de la unidad minera Caudalosa, 13°4'S, 75°0'W, 23-31 Mar 2015, P. Gonzáles 3568 (USM); Castrovirreyna, cordillera between Pisco and Ayacucho, 13°16'S, 75°18'W, May 1910, A. Weberbauer 5446 (F, GH); Moquegua: General Sánchez Cerro, Ubinas, S of Pillone, 16°10'S, 70°49'W, 24 Mar 2013, D. Montesinos 4023 (USM); General Sánchez Cerro, Ubinas, Matazo, 16°10'S, 70°49'W, 28 Mar 2015, D. Montesinos 4242 (USM); General Sánchez Cerro, Ubinas, Querala, 16°10'S, 70°49'W, 2 Mar 2018, D. Montesinos 5918 (USM); Puno: just W of abra on unpaved track, ca. 17 km from Puno-Ananea rd., 14°41'S, 69°41'W, 16 Mar 2014, V.A. Funk, M. Diazgranados & E. Cochachin 13184 (US, USM); Carabaya, Corani, Chacaconiza, 14°1'S, 70°40'W, 14 Jan 2015, P. Gonzáles 3428 (USM); Carabaya, Corani, Chacaconiza, 14°3'S, 70°40'W, 14 Jan 2015, P. Gonzáles 3441 (USM); Carabaya, Corani, Chacaconiza, 14°3'S, 70°40'W, 14 Jan 2015, P. Gonzáles 3444 (USM); Carabaya, alrededores de Condena, 13°46'S, 70°38'W, 9 Nov 2017, H. Trinidad 4192 (USM).

Figures

Figure 1. Senecio melanandrus. Habit (drawn from Buchtien 1589). Illustration by Alice Tangerini.

Figure 2. Senecio melanandrus A-C variability of leaves D capitulum E, F supplementary bracts G involucre H style I achene and floret J stamen. All details drawn from Weberbauer 5446 except for A (drawn from Calvo & Zarate 7872), C (drawn from Montesinos 3103), and G (drawn from Buchtien 1589). Illustration by Alice Tangerini.

Figure 3. A, B Senecio melanandrus (pilose form) C, D Senecio pygmophyllus E, F Senecio digitatus A habit (Peru, Cusco, Sibinacocha; Meneses et al. 6968) B leaves (Bolivia, Potosi, Kari Kari; Calvo & Zarate 7872) C habit (Chile, Tarapaca, Colchane; Moreira-Munoz 2876) D leaves (Chile, Arica-Parinacota, Las Cuevas; Moreira-Munoz & Luebert 2470) E habit F leaves (Chile, Antofagasta, Pacana; Calvo 7926). Picture A by Jim Farfan B, E, F by Joel Calvo C, D by Andres Moreira-Munoz.

Figure 4. A, B Senecio melanandrus (glabrous form) C, D Senecio pygmophyllus A habit B leaves (Peru, Puno, pr. Ananea; Funk et al. 13184) C habit D leaves (Peru, Moquegua, pr. Anillune; Funk et al. 13153). Pictures by Mauricio Diazgranados.

Figure 5. Distribution map of Senecio melanandrus (black circle), S. pygmophyllus (red circle), and S. digitatus (blue circle).