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Discoid caespitose Andean Senecio 111
New combinations and synonyms in discoid caespitose
Andean Senecio (Senecioneae, Compositae)
Joel Calvo1, Arturo Granda2, Vicki A. Funk3
1 Instituto de Geografía, Facultad de Ciencias del Mar y Geografía, Ponticia Universidad Católica de Valpa-
raíso, Avenida Brasil 2241, 2362807 Valparaíso, Chile 2 Herbario del Departamento Académico de Biología,
Facultad de Ciencias, Universidad Nacional Agraria La Molina, Av. La Molina s/n, apartado 12-056, Lima
12, Perú 3 US National Herbarium, Department of Botany, Smithsonian Institution, Washington D.C., USA
Corresponding author: Joel Calvo (calvocasas@gmail.com)
Academic editor: Peter de Lange|Received 25 July 2019|Accepted 2 September 2019|Published 3 October2019
Citation: Calvo J, Granda A, Funk VA (2019) New combinations and synonyms in discoid caespitose Andean Senecio
(Senecioneae, Compositae). PhytoKeys 132: 111–130. https://doi.org/10.3897/phytokeys.132.38534
Abstract
e names Werneria melanandra and W. pygmophylla are transferred to the genus Senecio. ey belong to
the group of the discoid caespitose Andean Senecio, specically to the subgroup with blackish anthers and
style branches and whitish corollas. e recognition of S. digitatus as a distinct species is also discussed.
Within the framework of the mentioned group, the names S. casapaltensis and S. macrorrhizus are lecto-
typied, S. humillimus var. melanolepis is neotypied, an epitype is designated for the name W. melanan-
dra, and nine new synonyms are proposed. An updated comprehensive dichotomous key including all
discoid caespitose Senecio species from Bolivia and Peru is provided.
Keywords
Asteraceae, Bolivia, Chile, dichotomous key, Peru, taxonomy, Werneria
Introduction
e discoid caespitose Andean Senecio L. species have traditionally been placed within
S. subser. Caespitosi (O. Hom.) Cabrera & S.E. Freire (Freire et al. 2014). is infra-
generic group was conceived for embracing the strictly caespitose species but also suf-
frutescent plants. As circumscribed by Cabrera et al. (1999), it includes ca. 50 species
from southern South America. e infrageneric classication of Senecio at the subserial
PhytoKeys 132: 111–130 (2019)
doi: 10.3897/phytokeys.132.38534
http://phytokeys.pensoft.net
Copyright Joel Calvo et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0),
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Joel Calvo et al. / PhytoKeys 132: 111–130 (2019)
112
rank has been proposed for the Argentinian species, which are reasonably well-known
(Freire et al. 2014). In Bolivia and Peru, by contrast, the understanding of the genus is
poorer; the infrageneric classication remains barely resolved and several species remain
misplaced. is is the case for two species that were hitherto recognized as members of
Werneria Kunth, a genus morphologically similar to Senecio that can be dierentiated
by the combination of the following characters: involucral bracts fused at the base, ab-
sence of genuine supplementary bracts (calyculus), achene trichomes not myxogenic,
rosettiform habit rather than caespitose. However, some Senecio species sometimes also
have the involucral bracts partially fused at the base or do not have supplementary
bracts. Such overlapping means that historically some species have been interchange-
ably treated as Senecio or Werneria, depending on the authors’ concepts. is is the case
with S. wernerioides Wedd., a species that Grisebach (1874) and Kuntze (1898) trans-
ferred to Werneria but that is presently widely accepted as a heterotypic synonym of S.
breviscapus DC. (Cabrera 1985; Beck and Ibáñez 2014; Freire et al. 2014). Similarly, S.
repens var. macbridei (Cuatrec.) Cabrera was initially described at the specic rank as W.
macbridei Cuatrec. In Chile, Ricardi and Marticorena (1964) described S. psteri Ri-
cardi & Martic., a species that has been recently synonymized with Xenophyllum esqui-
lachense (Cuatrec.) V.A. Funk [≡Werneria esquilachensis Cuatrec.] (Calvo et al. 2018).
Such disparate treatments highlight the taxonomic complexity of these groups and in-
dicate that some species are dicult to assign to one or another genus. In these cases, a
detailed study based on the aforementioned set of characters is needed. In addition, the
achene indumentum type appears to be a useful character for a proper identication.
In arid regions Senecio species with myxogenic trichomes (with mucilaginous proper-
ties when soaked in water) are common (Nordenstam et al. 2009; Mukherjee and
Nordenstam 2012). is character is also found in other genera within the tribe, e.g.,
Dauresia B. Nord. & Pelser, Dolichoglottis B. Nord., Euryops (Cass.) Cass. (Nordenstam
et al. 2009), but it has not been reported in Werneria. Indeed, most species belonging
to this genus have glabrous achenes or rarely scattered long trichomes near the base.
Only W. nubigena Kunth usually displays achenes with dense, villous indumentum. It
is composed of twin liform trichomes, ca. 0.7 mm long, with acute to subacute, asym-
metrical, usually forked apex, but does not exude mucilage when treated in water. On
this basis, the myxogenic trichomes appear to be absent in Werneria, and therefore, it
is another useful character to discriminate between the two genera.
Herein, we transfer W. melanandra Wedd. and W. pygmophylla S.F. Blake to the ge-
nus Senecio. Furthermore, and in disagreement with previous treatments (Rockhausen
1939; Cabrera 1949; Freire et al. 2014), we believe that W. pygmophylla and S. digitatus
Phil. correspond to two dierent taxonomic entities and we justify this here accord-
ingly. ese species belong to a group of discoid caespitose Andean Senecio with black-
ish anthers and style branches and whitish corollas but dier from one another in some
characters (see discussions below). Detailed illustrations and pictures are provided for
each species, as well as a dichotomous key including the discoid caespitose Senecio spe-
cies from Bolivia and Peru.
Discoid caespitose Andean Senecio 113
Materials and methods
is contribution is the result of an intensive review of the published bibliography and
the revision of herbarium specimens kept at BOLV, CONC, HSP, LPB, MA, MOL,
SGO, US, and USM. Additionally, digital herbarium specimens from LP and P were
studied; herbarium acronyms follow iers (2018). A light microscope was used for
examination of microcharacters. Field work was conducted in Bolivia, southern Peru,
and northern Chile.
Results
New combinations
1. Senecio melanandrus (Wedd.) J.Calvo, A.Granda & V.A.Funk, comb. nov.
urn:lsid:ipni.org:names:60479386-2
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, Ancump-
ampa, 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 (holo-
type: 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: Gen-
eral 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.; iso-
types: MOL n.v. [not located, likely never sent], USM-247549), syn. nov.
Joel Calvo et al. / PhytoKeys 132: 111–130 (2019)
114
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 at), glabrous, with marginal trichomes
on the narrowed base or densely pilose, somewhat eshy, greenish or glaucous. Ca-
pitulum 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 sur-
face ca. 0.7 mm long, dark purple- or blackish-tipped. Supplementary bracts (calycu-
lus) 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 orets 20–45, 4.3–6.3 mm long, 0.8–1.1 mm
wide, 5-lobed, conspicuously dierentiated in a distinct tube and campanulate limb,
whitish. Anther bases auriculate, clearly acute, dark purple to blackish; lament 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, g. 2, sub S. pucapampaensis);
Montesinos-Tubée (2014: pg. 7, g. 2; pg. 13, g. 4B, sub S. sykorae); Montesinos-
Tubée (2014: pg. 12, g. 3; pg. 13, g. 4C, sub S. tassaensis); Montesinos-Tubée et al.
(2015: pg. 2, g. 1; pg. 3, g. 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 eleva-
tions of 3800–5100 m.
Phenology. Flowering mainly from January to June, although some owering
specimens have been collected in November.
Etymology. e epithet melanandrus means having dark or black stamens, which
describes a striking character of this species.
Discussion. is 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 in-
cluded in the synonymy were also described within the genus Senecio.
Senecio melanandrus is a highly variable species that has been variously interpreted.
e poor condition of the type material probably helped to maintain the uncertainty
Discoid caespitose Andean Senecio 115
Figure 1. Senecio melanandrus. Habit (drawn from Buchtien 1589). Illustration by Alice Tangerini.
surrounding the application of this name. Weddell (1856) described the leaves as “in-
tegerrimis vel nonnullis dente triangulari, […] glabriusculis vel inconspicue ciliolatis”
[entire or with a few triangular teeth, rather glabrous or inconspicuously ciliate]. Sev-
eral years later Rockhausen (1939), who published the rst comprehensive taxonomic
revision of the genus Werneria, stated that the leaves have “marginibus obsolete glandu-
loso-ciliolata” [margins scarcely glandular-ciliate]. On the basis of our studies, this spe-
cies displays an unusually wide variability in leaf margin and indumentum of leaves and
involucre, which is reected in the number of names included in the synonymy. e
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). e glabrous forms also ap-
Joel Calvo et al. / PhytoKeys 132: 111–130 (2019)
116
Figure 2. Senecio melanandrus A–C variability of leaves D capitulum E, F supplementary bracts G invo-
lucre H style I achene and oret J stamen. All details drawn from Weberbauer 5446 except for A (drawn
from Calvo & Zárate 7872), C (drawn from Montesinos 3103), and G (drawn from Buchtien 1589). Il-
lustration by Alice Tangerini.
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)]. ese glabrous plants usually have
dentate leaves, but forms exist with rather subentire leaves (Gonzáles 3568, USM). e
Discoid caespitose Andean Senecio 117
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, Potosí, Kari Kari; Calvo &
Zárate 7872) C habit (Chile, Tarapacá, Colchane; Moreira-Muñoz 2876) D leaves (Chile, Arica-Parina-
cota, Las Cuevas; Moreira-Muñoz & Luebert 2470) E habit F leaves (Chile, Antofagasta, Pacana; Calvo
7926). Picture A by Jim Farfán B, E, F by Joel Calvo C, D by Andrés Moreira-Muñoz.
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 Mon-
tesinos. is form was also collected near the Bolivian locality of Ulla Ulla (Menhofer
1901, US). is puzzling distribution pattern and a continuum of intermediates sug-
Joel Calvo et al. / PhytoKeys 132: 111–130 (2019)
118
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.
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. e 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 char-
acter, 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.
e name Senecio vegetus var. lobatus Cabrera, here synonymized with S. melanan-
drus, 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]) con-
tains mixed material and some plants certainly correspond to S. melanandrus (the indi-
vidual on the left hand and likely the fragment at the right hand below). e syntype P
Discoid caespitose Andean Senecio 119
Figure 5. Distribution map of Senecio melanandrus (black circle), S. pygmophyllus (red circle), and
S.digitatus (blue circle).
[P01816917], not containing admixtures, shows plainly glabrous plants with the leaves
entire and obtuse at the apex. erefore, and in disagreement with Cabrera, we consider
that S. vegetus and S. melanandrus correspond to distinct taxonomic entities. e 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 posi-
tion 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 eorts 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 cor-
respond to the other varieties described by Weddell. For that reason, we selected as the
Joel Calvo et al. / PhytoKeys 132: 111–130 (2019)
120
neotype a Mandon collection that perfectly matches the diagnosis provided by Wed-
dell. Moreover, it was identied as S. humillimus var. melanolepis by Schultz Bipontinus
[see Mandon (1865)], which supports our interpretation of this taxon. e specimen
P03730757 is excluded because it contains mixed material.
e holotype of the name S. canoi should be housed at USM (Montesinos-Tubée
et al. 2015); however, it was not located. e 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 An-
derberg 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 decient for a prop-
er study of the diagnostic characters, we consider it appropriate to designate an epitype.
e 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 Ven-
tilla, 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. Mén-
dez 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 Apa-
cheta 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 Po-
tosí, 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 Oka-
ria (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. Solo-
mon & 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); Sa-
jama, nevado de Sajama, sur del cerro Jasasuni [Asa-asuni], 18°11'S, 68°55'W, 18
Discoid caespitose Andean Senecio 121
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 Medra-
no, 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 Po-
racota, 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 Soraco-
cha, 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); cordil-
lera 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. Del-
gado 19680 (USM); Huachocolpa, alrededores de la unidad minera Caudalosa,
13°4'S, 75°0'W, 23–31 Mar 2015, P. Gonzáles 3568 (USM); Castrovirreyna, cor-
dillera 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, Ubi-
nas, 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).
2. Senecio pygmophyllus (S.F. Blake) J.Calvo, A.Granda & V.A.Funk, comb. nov.
urn:lsid:ipni.org:names:60479387-2
Figs 3C, D, 4C, D, 6
Werneria pygmophylla S.F. Blake, J. Washington Acad. Sci. 18: 491. 1928. Type: Peru.
Moquegua: cordillera East of Carumas, 4500–4600 m, 7–8 Mar 1925, A. We-
Joel Calvo et al. / PhytoKeys 132: 111–130 (2019)
122
berbauer 7358 (holotype: F [F-552587]; isotypes: CONC [CONC-28864], G
[G00356025], US [US00622822]).
Senecio laucanus Ricardi & Martic., Gayana, Bot. 11: 17. 1964. Type: Chile. Arica-
Parinacota: camino de Putre a Chucuyo, km 17, 4250 m, 12 Feb 1964, C. Mar-
ticorena, O. Matthei & M. Quezada 208 (holotype: CONC [CONC-29864];
isotype: CONC), syn. nov.
Description. Caespitose perennial herb. Leaves long pseudopetiolate; leaf lamina 2.5–
5.5 mm long, 2.4–5.5 mm wide, ovate to suborbiculate, obtuse at the apex, rounded
to truncate at the base, typically crenate-lobate with 3–9 rounded lobes, revolute, usu-
ally strongly conduplicate downwards, pilose on both surfaces, somewhat eshy, glau-
cous; pseudopetiole 5–25 mm long, at, slightly broadened at the base, marginally
ciliate. Capitulum discoid, solitary, terminal, sessile or subsessile; involucre 6–8 mm
long, 7–10 mm wide. Involucral bracts 16–21, oblong-lanceolate, 2.5–4 mm long,
0.7–1.7mm wide, partially fused at the base, smooth, with trichomes on the abaxial
surface 0.5–0.8 mm long, dark purple- or blackish-tipped. Supplementary bracts ca.
3, linear, 6–7.5 mm long, 0.5–0.8 mm wide, smooth, three-quarters to as long as the
involucral bracts, with trichomes on the margins, dark purple- or blackish-tipped. Disc
orets 50–82, 3.5–5.1 mm long, 0.6–1 mm wide, 5-lobed, conspicuously dierenti-
ated in a distinct tube and campanulate limb, whitish. Anther bases auriculate, clearly
acute, dark purple to blackish; lament collar balusterform. Style branches truncate
with a crown of sweeping hairs, dark purple to blackish. Achenes 1.7–1.8(–2.5) mm
long, ca. 0.5 mm wide, brownish, covered by dense indumentum of obtuse whitish
myxogenic twin trichomes ca. 0.2 mm long; pappus 3–4.5 mm long, barbellate, whit-
ish. Chromosome number: unknown.
Additional iconography. Blake (1928: pg. 496, g. 1F, G, sub W. pygmophylla);
Ricardi and Marticorena (1964: pg. 19, g. 6, sub S. laucanus).
Distribution and habitat. Chile (Arica-Parinacota, N Tarapacá) and Peru (Mo-
quegua) (Fig. 5). e species is also expected in the Peruvian department of Tacna and
in the Bolivian region bordering northern Chile, although no collections have been
studied from there. It grows in exposed places on sandy soils, between elevations of
4100–4700 m.
Phenology. Collected in bloom from January to June, although full bloom prob-
ably takes place between March and April.
Etymology. e epithet refers to the resemblance of the leaves to a st.
Discussion. Blake (1928) placed his new species within Werneria arguing that
the involucral bracts were connate half way. Otherwise, he assumed a close similarity
between it and a Senecio species collected by Pennell here identied as S. moqueguensis
Montesinos (see protologue of W. pygmophylla). It is certain that the involucral bracts
of S. pygmophyllus are usually partially fused at the base; however, this character alone
cannot be used as diagnostic to place a species in one or another genus. Rather, we
prefer to base a decision on a set of characters, i.e., presence or absence of genuine
supplementary bracts, involucral bracts free or fused at the base, achene trichomes
Discoid caespitose Andean Senecio 123
myxogenic or not, and rosettiform or caespitose habit. Accordingly, we consider that
this species should be placed within Senecio on the basis of the following characters:
presence of supplementary bracts, myxogenic achene trichomes, and caespitose habit.
is decision is also supported by the fact that it was inaccurately considered a syno-
nym of S. digitatus for a long time.
Rockhausen (1939) was the rst author who treated W. pygmophylla as a heterotypic
synonym of S. digitatus. Since then, most authors followed his treatment (e.g., Cabrera
1949; Freire et al. 2014). Ricardi and Marticorena (1966), in disagreement, concluded
that they correspond to two distinct taxonomic entities. We agree with Ricardi and
Marticorena’s treatment after studying the respective type materials, further collections
from southern Peru and northern Chile, and living plants. e two species can be dif-
ferentiated by their leaf shape and indumentum type. Senecio pygmophyllus has a lamina
clearly dierentiated from the pseudopetiole (petioliform base); usually the lamina is
remarkably reduced when compared with the pseudopetiole length (at least in the more
basal leaves). e lamina we observed were ovate to suborbicular, typically crenate-
lobate with 3–9 rounded lobes and revolute margins (Fig. 4C). In contrast, S. digitatus
Figure 6. Senecio pygmophyllus A habit (drawn from Funk et al. 13153) B leaf (drawn from Weberbauer
7358). Illustration by Alice Tangerini.
Joel Calvo et al. / PhytoKeys 132: 111–130 (2019)
124
has linear to slightly spatulate leaves narrowed at the base (Fig. 3E, F). is latter spe-
cies is extremely variable with regard to the leaf margin, which can be dentate, pinnati-
partite or distantly pinnatisect, with clearly acute teeth; however, specimens with entire
leaves and even individuals displaying both entire and dentate leaves were occasionally
observed. e leaf apex is acute and usually shows a whitish callus-like tip, whereas in S.
pygmophyllus the apex is always plainly obtuse and unadorned (Fig. 3C, 4D). Both spe-
cies usually have abundant indumentum on the leaves, involucre, and supplementary
bracts but the type of trichomes diers and is useful to separate them from one other.
e indumentum of S. pygmophyllus is pilose whereas in S. digitatus the trichomes are
clearly arachnoid, longer, and intermingled. Moreover, the indumentum of S. digitatus
is essentially concentrated on the adaxial surface, whereas in S. pygmophyllus the leaf
lamina has trichomes on both surfaces. eir distribution areas do not overlap (Fig. 5).
Senecio pygmophyllus might be confused with those forms of S. melanandrus dis-
playing pilose, dentate leaves. A useful character to discriminate them from each other
is the leaf shape, although some overlap has been detected in a few specimens. In S.
pygmophyllus the leaves are clearly pseudopetiolate and the ratio lamina/pseudopetiole
length usually is very low in the more basal leaves (Fig. 4D). In contrast, S. melanan-
drus displays a lamina progressively narrowed at the base (Fig. 3B). e distinctive
pseudopetiole length of S. pygmophyllus might be an adaptation to the sandy soils
where this species thrives because the plants usually appear to be partially sunken. Ad-
ditionally, the number of disc orets tends to be higher in S. pygmophyllus (50–82 vs.
20–45), as well as the number of involucral bracts (16–21 vs. 11–16). Since the men-
tioned morphology coincides with geographical separation, we consider it appropriate
to recognize it as a distinct species.
e name S. laucanus Ricardi & Martic. was described from northern Chile (Arica-
Parinacota) and it was hitherto considered endemic to this country (Moreira-Muñoz
et al. 2016). It is included in the synonymy of S. pygmophyllus since we failed to iden-
tify any diagnostic character to dierentiate them. In some specimens from Chile the
more basal leaves are not so long pseudopetiolate as in the typical forms (e.g., Moreira-
Muñoz & Luebert 2470), but it is considered as part of the variability encompassed by
this species; indeed, this morphology probably responds to the fact that these plants
grow on less sandy soils.
Specimens examined. Senecio digitatus. ARGENTINA. Salta: Los Andes, Huaiti-
quina, 23°44'S, 67°12'W, 27 Feb 1972, Cabrera et al. 22559 (LP). BOLIVIA. Potosí: Sud
Lípez, a 1 km al W de salar Chalviri, 22°30'S, 67°38'W, 7 May 1999, N. Massi & C. Salles
726 (LPB) [rst record for Bolivia]. CHILE. Antofagasta: El Loa, camino entre Ascotán
y San Pedro de Conchi, 21°58'S, 68°26'W, 4 Apr 1985, M. Arroyo 85-606 (CONC); El
Loa, cerro Losloyo, ladera SE, 23°9'S, 67°15'W, 9 Apr 1997, M. Arroyo, L. Cavieres & A.
Humaña 97331 (CONC); El Loa, cerro Nevados de Poquis, ladera SO, 23°4'S, 67°5'W,
9 Apr 1997, M. Arroyo, L. Cavieres & A. Humaña 97343 (CONC); El Loa, pampa La-
guna Helada, 23°6'S, 67°5'W, 9 Apr 1997, M. Arroyo, L. Cavieres & A. Humaña 97403
(CONC); El Loa, pampa Loyoques, 23°11'S, 67°12'W, 9 Apr 1997, M. Arroyo, L. Ca-
vieres & A. Humaña 97408 (CONC); El Loa, cordón cerro de la Pacana, cuesta entre salar
de Aguas Calientes y quebrada Quepiaco, 23°3'S, 67°29'W, 11 Apr 1997, M. Arroyo, L.
Discoid caespitose Andean Senecio 125
Cavieres & A. Humaña 97477 (CONC); El Loa, cordón cerro de la Pacana, cuesta entre
salar de Aguas Calientes y quebrada Quepiaco, 23°4'S, 67°30'W, 11 Apr 1997, M. Arroyo,
L. Cavieres & A. Humaña 97498 (CONC); El Loa, Toconao, camino a Tara, monjes de
La Pacana, 23°3'S, 67°29'W, 6 Mar 2019, J. Calvo 7926 (SGO); cruce camino internac-
ional Paso Jama con camino a salar de Tara, 23°3'S, 67°29'W, 19 Dec 1996, A. Moreira-
Muñoz 317 (SGO); Machuca-Copacoya, 22°28'S, 68°2'W, 18 Feb 1885, F. Philippi s.n.
(LP, SGO); laguna de Llaillai, 21°55'S, 68°12'W, 23 Feb 1885, F. Philippi s.n. (CONC,
LP, SGO, SI); El Loa, Ascotán, 21°27'S, 68°21'W, 23 Jan 1943, E. Pisano & J. Venturelli
1753 (SGO); El Loa, entre Machuca y Tatio, 15 Feb 1943, E. Pisano & J. Venturelli 1866
(CONC, SGO); Tarapacá: [without locality], Feb 1885, F. Philippi s.n. (K); Iquique,
Collaguasi, San Carlos, 20°58'S, 68°41'W, 22 Jan 1994, S. Teillier 3286A (CONC).
Senecio pygmophyllus. CHILE. Arica-Parinacota: cerca de laguna de Cotacotani,
camino a Guane Guane, 18°10'S, 69°14'W, 9 Mar 1984, M. Arroyo 84-724 (CONC);
portezuelo entre cerro Guane Guane y cerro Larancagua, 18°9'S, 69°19'W, 22 Apr
1984, M. Arroyo 84-935 (CONC); Las Cuevas, antes del Chaku, 18°11'S, 69°25'W,
20 Mar 2015, A. Moreira-Muñoz & F. Luebert 2470 (SGO); camino de Putre a Por-
tezuelo de Chapiquiña, 18°20'S, 69°30'W, 28 Mar 1961, M. Ricardi, C. Marticorena
& O. Matthei 277 (CONC); Tarapacá: Colchane, géiser Puchultiza, 100 m antes
del géiser, 19°24'S, 68°57'W, 16 Jun 2018, A. Moreira-Muñoz 2876 (SGO). PERU.
Moquegua: minera Quellaveco, 17°6'S, 70°36'W, 8 Apr 1999, ESCO 7238 (US); area
between the carretera-binacional and the interoceanica sur, on unpaved road that con-
nects the two main roads and borders a large bofedal, 16°51'S, 70°32'W, 12 Mar 2014,
V.A. Funk, M. Diazgranados & E. Cochachin 13153 (US, USM); Mariscal Nieto,
Carumas, Ancolacaya, 16°38'S, 70°19'W, Mar–Apr 2018, V. Morales 140 (USM); 5
km East of lago Suche, 16°55'S, 70°19'W, 19 Jan 1952, O.P. Pearson 5 (CONC, UC).
New synonyms
1. Senecio casapaltensis Ball, J. Linn. Soc., Bot. 22: 47. 1885.
Senecio sanmarcosensis H. Beltrán, Arnaldoa 15: 211. 2009. Type: Peru. Ancash: Huari, San
Marcos, Ccolla Chica, 09°40'28"S, 77°03'10"W, 5600 m, 4 May 2008, H. Beltrán
6476 (holotype: USM [USM-00277272]; isotypes: CUZ n.v., HUT n.v.), syn. nov.
Senecio repens var. taraxacifolius A. Gray [“taraxicifolius”], nom. nud. in sched. (Tur-
land et al. 2018, ICN Art. 38.1) (US [US00829056]).
Type. Peru. Lima: supra Casapalta, 4265–4360 m, 22 Apr 1882, J. Ball s.n. (lectotype,
designated here: K [K000497782]; isolectotype: E [E00417028]).
Discussion. Senecio casapaltensis Ball was described from central Lima near the
border between Lima-Junín departments, whereas the type material of S. sanmarco-
sensis H. Beltrán comes from southeastern Ancash Department. After studying several
specimens from both regions, we can conclude that the dierences concerning the
shape and size of the leaves are not signicant. e populations from Ancash tend to
Joel Calvo et al. / PhytoKeys 132: 111–130 (2019)
126
have a denser indumentum composed of capitate trichomes, whereas those specimens
from Lima are glabrescent or the indumentum is rather deciduous and composed of
shorter glandular trichomes. Nonetheless, the existence of intermediate specimens
makes their recognition as distinct species inadvisable and, therefore, S. sanmarcosensis
is here synonymized with S. casapaltensis.
Among the located original material of S. casapaltensis, the specimen at K is desig-
nated as the lectotype due to it being more complete than the duplicate at E.
2. Senecio expansus Wedd., Chlor. Andina 1: 107. 1856.
Senecio macrorrhizus Wedd., Chlor. Andina 1: 108. 1856. Type: Peru. Cusco: dept.
de Cuzco, Oct 1839–Feb 1840, C. Gay 1870 (lectotype, designated here: P
[P01816797]), syn. nov.
Type. Bolivia. Potosí: montagnes des lagunas de Potosí, [without date], A. d’Orbigny
1418 (lectotype, designated by Cabrera (1966) as “typus”, pg. 21: P [P01816805];
isolectotypes: BR [BR00000552801], G [G00356020], K [K000497783]).
Discussion. Senecio macrorrhizus Wedd. was described from Cusco (Peru) and
distinguished from S. expansus Wedd. mainly by having a thicker, longer, and more
sinuous rhizome, larger capitulum, and rosettes less spread out (Weddell 1856). e
mentioned dierences fall within the variability encompassed by S. expansus, and here
we synonymize them.
Gay’s specimen P01816797 perfectly matches the protologue information, and
therefore, it is designated as the lectotype of the name S. macrorrhizus.
Key to the discoid caespitose Senecio species from Bolivia and Peru
e dwarf shrubs developing erect stems are excluded (e.g., S. apolobambensis Cabrera,
S. puchei Phil., S. trifurcifolius Hieron.). Senecio aquilaris Cabrera was cited for Bolivia
(Beck and Ibáñez 2014) and Peru (Gonzáles et al. 2016); it is not included in the key
because its identication is doubtful and further studies are required. e rosettiform
species S. expansus and S. hyoseridis (Benth.) L. Salomón & S.E. Freire were placed
in S. ser. Culcitium (Bonpland) Cabrera (Freire et al. 2014; Salomón et al. 2018) but
they are included in the key because they t well within the discoid caespitose species
group. e color of the anthers, style branches, and corollas has a relevant taxonomic
value within the group and it is readily noticeable on living plants. However, on dried
specimens a careful study is required in order to avoid misidentications.
1 Plants in rosette form .................................................................................. 2
– Plants developing prostrate or decumbent stems ......................................... 6
Discoid caespitose Andean Senecio 127
2 Leaves pinnatilobate to lyrate-pinnatisect ............................. S. casapaltensis
– Leaves subentire to pinnatipartite ...............................................................3
3 Capitula sessile, solitary or several; leaf lamina longer than or similar to the
pseudopetiole .............................................................................................. 4
– Capitulum shortly pedunculate, solitary; leaf lamina clearly shorter than the
pseudopetiole .............................................................................................. 5
4 Leaves densely white tomentose on both faces, concolorous ....... S. expansus
– Leaves only densely white tomentose beneath, discolorous ............................
....................................................S. hyoseridis s.l. (further research needed)
5 Leaves ovate-deltate, crenate-dentate, puberulous on both faces ... S. genisianus
– Leaves elliptic-suborbicular, subentire, with scattered long hispid trichomes
above and nearly glabrous beneath .......................................... S. sagasteguii
6 Anthers and style branches yellowish; corolla yellowish ...............................7
– Anthers and style branches blackish; corolla whitish .................................16
7 Leaves and involucre covered by whitish lanate indumentum......................8
– Leaves and involucre glabrous or covered by arachnoid or pilose indumen-
tum .............................................................................................................9
8 Leaves arranged along true stems; leaves oblanceolate to oblong ...................
..................................................................................... S. carhuanishoensis
– Leaves arranged in rosettiform clusters arising directly from rhizome-like
stems; leaves obovate-spatulate ...................................................S. evacoides
9 Involucre 4–5 mm long; involucral bracts 8(–9) .................... S. humillimus
– Involucre 6–12 mm long; involucral bracts (9–)12–15(–20) .....................10
10 Achenes with indumentum .......................................................................11
– Achenes glabrous ......................................................................................13
11 Leaves glabrous, entire or subentire ................................................S. woodii
– Leaves covered with trichomes, dentate .....................................................12
12 Involucral bracts 9–12; leaves sparsely covered with trichomes ......................
...........................................................................................S. moqueguensis
– Involucral bracts 15–20; leaves densely covered with trichomes ....................
......................................................... S. ticsanicus (further research needed)
13 Leaves dentate or lobate, rarely only shallowly crenate ..............................14
– Leaves entire .............................................................................................15
14 Leaves dentate, rarely only shallowly crenate ........................... S. menesesiae
– Leaves pinnatilobate..........................................................S. pinnatilobatus
15 Leaves (15–)20–50 mm long, arranged along the stems .................. S. algens
– Leaves 5–10 mm long, arranged in rosettiform clusters ............. S. gamolepis
16 Achenes papillose, with visible ribs; leaves linear ............. S. scorzonerifolius
– Achenes silky-pubescent, usually with invisible ribs; leaves linear, linear-
oblong or spatulate ...................................................................................17
17 Leaf lamina glabrous .................................................................................18
– Leaf lamina with indumentum .................................................................19
Joel Calvo et al. / PhytoKeys 132: 111–130 (2019)
128
18 Supplementary bracts 4.1–5.7 mm long, a third to a half as long as the invo-
lucral bracts, glabrous; leaves linear-oblong, at, acute at the apex ................
............................................................................................. S. madidiensis
– Supplementary bracts 4.2–7.5 mm long, almost as long as the involucral
bracts, usually pilose; leaves linear-oblong to spatulate, usually conduplicate
downwards, rather obtuse at the apex.................................. S. melanandrus
19 Lamina ovate to suborbicular, dierentiated from the pseudopetiole .............
........................................................................................... S. pygmophyllus
– Lamina linear, linear-oblong or narrowly spatulate, progressively narrowed at
the base ..................................................................................................... 20
20 Leaves dentate, pinnatipartite or distantly pinnatisect (rarely entire), arach-
noid, usually with a callus-like tip ...............................................S. digitatus
– Leaves entire, crenate or dentate, pilose, unadorned at the apex ....................
........................................................................................... S. melanandrus
Acknowledgements
We are grateful to the curators and sta of the herbaria mentioned in the text. We
extend special thanks to Florian Jabbour (P), Hamilton Beltrán (USM), Laura Ihar-
legui (LP), and Diego Gutiérrez (BA, LP) for facilitating some type material from the
respective herbaria. Alice Tangerini (US) created the illustrations of Senecio melanan-
drus and S. pygmophyllus. Mauricio Diazgranados (K), Andrés Moreira-Muñoz (Chile),
Rosa I. Meneses (LPB), and Jim Farfán (GLORIA project) kindly shared some pictures
of living plants. We are indebted to Pieter Pelser and Liliana Katinas for their insight-
ful comments. is work has been funded by FONDECYT from Chile by means of a
postdoctoral fellowship for the rst author (project N°3170270).
References
Beck SG, Ibáñez D (2014) Senecio L. In: Jørgensen PM, Nee MH, Beck SG (Eds) Catálogo
de las Plantas Vasculares de Bolivia. Monographs in Systematic Botany from the Missouri
Botanical Garden 127: 281–287.
Beltrán H (2008) Dos especies nuevas de Senecio (Asteraceae: Senecioneae) del Perú. Arnaldoa
15: 211–215.
Blake SF (1928) New South American species of Werneria. Journal of the Washington Academy
of Sciences 18: 485–498.
Cabrera AL (1949) El género “Senecio” en Chile. Lilloa 15: 27–501.
Cabrera AL (1955) Notas sobre los Senecio sudamericanos, VIII. Notas del Museo de La Plata.
Botánica 18(89): 191–240.
Cabrera AL (1966) El género “Senecio” en la República Argentina. I. La sección Brachypappus.
Revista del Museo de La Plata. Sección Botánica 10: 1–36.
Discoid caespitose Andean Senecio 129
Cabrera AL (1985) El género Senecio (Compositae) en Bolivia. Darwiniana 26: 79–217. http://
www.jstor.org/stable/23218127
Cabrera AL, Freire SE, Ariza Espinar L (1999) Senecio L. In: Hunziker AT (Ed.) Flora Faner-
ogámica Argentina, fasc. 62. CONICET-Proora, Córdoba, 12–158.
Calvo J, Fuentes AF (2018) ree new caespitose species of Senecio (Senecioneae, Compositae)
from Central Andes. Phytotaxa 375(1): 70–80. https://doi.org/10.11646/phytotaxa.375.1.3
Calvo J, Muñoz-Schick M, Moreira-Muñoz A (2018) Towards a better understanding of Xeno-
phyllum esquilachense (Senecioneae, Compositae), a poorly known Andean species. Phyto-
taxa 382(3): 288–292. https://doi.org/10.11646/phytotaxa.382.3.5
Freire SE, Ariza Espinar L, Salomón L, Hernández MP (2014) Senecio L. In: Zuloaga FO,
Belgrano MJ, Anton AM (Eds) Flora vascular de la República Argentina, vol. 7(3). Insti-
tuto de Botánica Darwinion, Consejo Nacional de Investigaciones Cientícas y Técnicas,
Argentina, 27–220.
Gonzáles P, Navarro E, Trinidad H, Cueva M, Cano A, Al–Shehbaz I, Ramírez DW (2016)
Doce nuevos registros de plantas vasculares para los Andes de Perú. Arnaldoa 23: 159–170.
Grisebach A (1874) Plantae Lorentzianae. Abhandlungen der Königlichen Gesellschaft der
Wissenschaften zu Göttingen 19: 49–279.
Kuntze CEO (1898) Revisio generum plantarum, pars 3(3). Arthur Felix, Leipzig; Dulau &
Co., London; U. Hoepli, Milano; Gust. E. Stechert, New York; Charles Klincksieck, Paris,
[1]–576.
Mandon MG (1865) Première liste des plantes des Andes boliviennes recueillies et distribuées.
Bulletin de la Société Botanique de France 12(2): 79–83. https://doi.org/10.1080/00378
941.1865.10827407
Montesinos-Tubée DB (2014) ree new caespitose species of Senecio (Asteraceae, Senecio-
neae) from South Peru. PhytoKeys 39: 1–17. https://doi.org/10.3897/phytokeys.39.7668
Montesinos-Tubée DB, Gonzáles P, Navarro E (2015) Senecio canoi (Compositae), una especie
nueva de los Andes de Perú. Anales del Jardín Botánico de Madrid 72(2): e26. https://doi.
org/10.3989/ajbm.2409
Moreira-Muñoz A, Muñoz-Schick M, Marticorena A, Morales V (2016) Catálogo de Aster-
aceae (Compositae) de la Región de Arica y Parinacota, Chile. Gayana. Botánica 73(2):
226–267. https://doi.org/10.4067/S0717-66432016000200226
Mukherjee SK, Nordenstam B (2012) Diversity of trichomes from mature cypselar surface of
some taxa from the basal tribes of Compositae. Compositae Newsletter 50: 78–125.
Nordenstam B, Pelser PB, Kadereit JW, Watson LE (2009) Senecioneae. In: Funk VA, Susanna
A, Stuessy TF, Bayer RJ (Eds) Systematics, Evolution, and Biogeography of Compositae.
International Association for Plant Taxonomy, Vienna, Austria, 503–525.
Ricardi M, Marticorena C (1964) Compuestas nuevas o interesantes para Chile. Gayana. Bo-
tánica 11: 3–28. https://biodiversitylibrary.org/page/28854300
Ricardi M, Marticorena C (1966) Plantas interesantes o nuevas para Chile. Gayana. Botánica
14: 3–29. https://biodiversitylibrary.org/page/28854740
Rockhausen M (1939) Verwandtschaft und Gliederung der Compositen-Gattung Werneria.
Botanische Jahrbücher für Systematik, Panzengeschichte und Panzengeographie 70:
273–342.
Joel Calvo et al. / PhytoKeys 132: 111–130 (2019)
130
Salomón L, Sklenář P, Freire SE (2018) Synopsis of Senecio series Culcitium (Asteraceae: Sene-
cioneae, Senecioninae) in the Andean region of South America. Phytotaxa 340(1): 1–47.
https://doi.org/10.11646/phytotaxa.340.1.1
iers B (2018) [continuously updated] Index Herbariorum: A global directory of public her-
baria and associated sta. New York Botanical Garden’s Virtual Herbarium. http://sweet-
gum.nybg.org/science/ih/ [accessed 23.09.2018]
Turland NJ, Wiersema JH, Barrie FR, Greuter W, Hawksworth DL, Herendeen PS, Knapp
S, Kusber W-H, Li D-Z, Marhold K, May TW, McNeill J, Monro AM, Prado J, Price
MJ, Smith GF (2018) International Code of Nomenclature for algae, fungi, and plants
(Shenzhen Code). Regnum Vegetabile 159. Koeltz Botanical Books, Glashütten. https://
doi.org/10.12705/Code.2018
Weddell HA (1856) Chloris andina, vol. 1, part 3. Chez P. Bertrand, Paris, 57–136.
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