Content uploaded by Paola De Lima Ferreira
Author content
All content in this area was uploaded by Paola De Lima Ferreira on Oct 15, 2021
Content may be subject to copyright.
Phytotaxa 518 (2): 109–142
https://www.mapress.com/j/pt/
Copyright © 2021 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Alexander Sennikov: 29 May 2021; published: 3 Sept. 2021
https://doi.org/10.11646/phytotaxa.518.2.3
109
Touch me carefully: a step towards understanding morphological diversity in the
South American spiny sunflowers (Compositae, Barnadesioideae)
PAOLA DE LIMA FERREIRA1,2,3,4,7*, ALEXANDRE ANTONELLI2,3,5,6,8 & MILTON GROPPO1,9
1 Departamento de Biologia, Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto, Universidade de São Paulo, Avenida
Bandeirantes 3900, 14051-901, Ribeirão Preto, São Paulo, Brazil
2 Gothenburg Global Biodiversity Centre, Box 461, 405 30, Gothenburg, Sweden
3 Department of Biological and Environmental Sciences, University of Gothenburg, Box 461, 405 30, Gothenburg, Sweden
4 Institute of Entomology, Biology Centre of the Czech Academy of Sciences, 37005, České Budějovice, Czech Republic
5 Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, United Kingdom
6 Department of Plant Sciences, University of Oxford, South Parks Road, Oxford, OX1 3RB, United Kingdom
7
�
paolaferreira@alumni.usp.br; https://orcid.org/0000-0002-6957-4243
8
�
a.antonelli@kew.org; https://orcid.org/0000-0003-1842-9297
9
�
groppo@ffclrp.usp.br; https://orcid.org/0000-0003-2932-7798
* Author for correspondence
Abstract
The subfamily Barnadesioideae (Compositae) is endemic to South America, comprising 10 genera and 80 species of mostly
spiny herbs, subshrubs, shrubs, trees, or woody vines distributed from Venezuela to Argentina. Three genera, Dasyphyllum
(27 species), Chuquiraga (22 spp.) and Barnadesia (19 spp.) contain 85% of the species, while the other seven genera
(Archidasyphyllum, Arnaldoa, Doniophyton, Duseniella, Fulcaldea, Huarpea, and Schlechtendalia) are represented by up
to three species each. Most species are found in xeric areas in the Andean and Patagonian regions—as in the Páramos, Puna
and Patagonian steppe vegetation—with a secondary center of diversity in eastern South America. Previous phylogenetic
hypotheses have clarified the relationships within the subfamily, showing that there are many non-monophyletic groups
in different taxonomic ranks. As a result, taxonomic changes have been proposed over recent decades in order to reflect
classifications comprising only monophyletic groups. In the present study, we provide a generic synopsis of the subfamily
Barnadesioideae based on the most recent generic circumscriptions, including a key, expanded morphological descriptions,
information on geographical distribution and habitat, photographs and taxonomic notes for all genera.
Keywords: Asteraceae, distribution, Neotropical flora, morphology, taxonomy
Introduction
The Compositae subfamily Barnadesioideae (D.Don) K.Bremer & R.K.Jansen (Bremer & Jansen 1992: 415) comprises
ten genera and 80 South American species of perennial or annual herbs, subshrubs, shrubs, trees, or woody vines
distributed from Venezuela to Argentina (Fig. 1). Most taxa are found in xeric areas in the Andean and Patagonian
regions (Bremer 1994, Stuessy et al. 2009), but also in the Nothofagus forest (broadleaf and mixed beech forest) of
central Chile and adjacent central-western areas of Argentina. A secondary center of diversity spans in Brazil, Bolivia,
Paraguay, Uruguay and northeastern Argentina. The subfamily is characterized by having axillary spines, rarely solitary,
in pairs or fascicles, straight or curved, convergent (when the spines follow the same direction) or divergent (when the
spines follow different directions), and by the “barnadesioid trichomes”—a pubescence of unbranched three-celled
hairs on the corollas, cypselae and pappus (Cabrera 1959, Urtubey 1999, Erbar & Leins 2000, Stuessy et al. 2009).
Barnadesioideae was first recognized as a group by Don (1830), who described the tribe Barnadesieae (Don
1830: 273), classifying 12 species in two genera, Barnadesia and Chuquiraga. Later, the genera of Barnadesiodieae
were recognized in different taxonomic ranks (Lessing 1832, Candolle 1838, Bentham 1873, Hoffmann 1893, Cabrera
1961). With the advent of DNA technologies, phylogenetic studies recovered the subtribe Barnadesiinae as sister
group to the rest of the family (Jansen & Palmer 1987, 1988, Jansen et al. 1992), and the subtribe was elevated to the
subfamilial rank, as Barnadesioideae (Bremer & Jansen 1992).
FERREIRA ET AL.
110 • Phytotaxa 518 (2) © 2021 Magnolia Press
FIGURE 1. Distribution of the South American subfamily Barnadesioideae (Compositae).
Although Barnadesioideae comprises less than 1% of the species currently accepted in Compositae, the subfamily
presents a great diversity in many aspects, such as morphology (Ezcurra 1985, Urtubey 1999, Padin et al. 2015a, Stuessy
& Urtubey 2006, Stuessy et al. 2009, Svoma et al. 2019), chemical compounds (Bohm & Stuessy 1995, Mediondo et
al. 1997, 2000, Bohm & Stuessy 2001, Ccana-Ccapatina et al. 2018), pollen (Skvarla et al. 1977, Hasen 1991, Urtubey
1997, Urtubey & Telléria 1998, DeVore et al. 2000, Zhao et al. 2000, Suessy & Urtubey 2007, Telléria et al. 2015),
chromosome counts (Diers 1961, Heiser 1963, Olsen 1980, Ciadella & López 1981, Wulff 1984, Cristóbal 1986,
Wulff 1990, Stuessy & Sagástegui 1993, Strother & Panero 1994), and habitats, (Stuessy et al. 2009) which makes the
subfamily an interesting group for the study of evolutionary trends in the Neotropics. Moreover, since Barnadesioideae
is proposed as the sister group to the rest of the Compositae (Jansen & Palmer 1987, 1988), evolutionary studies allied
to morphology, taxonomy, nomenclature and historical biogeography can shed light into the early evolution of the
family.
Despite previous studies on the evolution of Barnadesioideae, understanding the relationships within the subfamily
remains a challenge. Today, there is no doubt that the subfamily is monophyletic (Gustafsson et al. 2001, Gruenstaeudl
et al. 2009, Padin et al. 2015b, Ferreira et al. 2019, Mandel et al. 2019). However, previous phylogenetic analyses have
provided different hypotheses regarding the monophyly of the subfamily, the relationships between genera, infrageneric
classification, and also species delimitation (Bremer 1994, Stuessy et al. 1996, Gustafsson et al. 2001, Urtubey &
Stuessy 2001, Gruenstaeudl et al. 2009, Padin et al. 2015b, Ferreira et al. 2019). Previous phylogenetic results could
have been affected by the taxa selected, choice of the molecular regions, and the phylogenetic reconstruction method
MORPHOLOGICAL DIVERSITY IN SPINY SUNFLOWERS Phytotaxa 518 (2) © 2021 Magnolia Press • 111
(Nabhan & Sarkar 2012). With the advance of high-throughput sequencing, phylogenies including thousands of
molecular markers and species, using new statistical models and coalescence methods, could provide better insights
into this group.
This work aims to increase the understanding of this diverse subfamily, providing updated generic circumscriptions
in addition to a key, morphological descriptions, photographs, geographical distribution, maps, habitat, and notes for
all genera currently circumscribed in Barnadesioideae.
Material and methods
This study is based on a bibliographic review of Barnadesioideae (Cabrera 1959, Ezcurra 1985, Hansen 1991, Harling
1991, Sagástegui & Sánchez 1991, Stuessy & Sagástegui 1993, Ferreyra 1995, Bremer 1994, Granda 1997, Katinas &
Stuessy 1997, Urtubey 1999, Hind 2001, Ulloa et al. 2002, Urtubey & Stuessy 2001, Stuessy & Urtubey 2006, 2007,
Stuessy et al. 2009, Funk & Roque 2011, Saavedra 2011, Saavedra et al. 2014, 2018, Ferreira et al. 2019, Svoma et al.
2019), examination of ca. 3,500 voucher materials from 36 herbaria: ALCB, B, BHCB, CEN, CESJ, COL, E, EAC,
ESA, F, FURB, GB, HJ, HRCB, HUEFS, HUFU, IBGE, ICN, K, LP, MBM, MO, NY, NYBG, P, QCA, QCNE, RB,
RFA, SI, SJRP, SPF, SPFR, UEC, US, WU (acronyms according to Thiers 2019), and observations of living plants
during fieldwork conducted in Brazil and Ecuador between 2012 and 2018. Information on pollen and chromosome
numbers present in the taxonomic treatment were directly extracted from the literature (Diers 1961, Heiser 1963, Olsen
1980, Cialdella & López 1981, Wulff 1990, Stuessy & Sagástegui 1993, Strother & Panero 1994, Stuessy et al. 1996,
Suessy & Urtubey, 2007, Watanabe et al. 2007).
We present a taxonomic key that reflects the recent modifications on generic circumscriptions and morphological
diversity within Barnadesioideae. Morphological terms follow Radford et al. (1974) for indument and shapes,
whereas corolla types follow Stuessy & Urtubey (2006), except for the genus Chuquiraga, which follows Ezcurra
(1985). Geographical distribution information was obtained from herbarium specimens and a review of the literature
cited above. For the maps, when the coordinates were not available, we added georeferences using the municipality
coordinates provided by NASA (https://mynasadata.larc.nasa.gov/, last accessed December 2018); the records were
plotted using QGIS version 3.2.1 (QGIS Development Team 2019).
Results
Taxonomic treatment
Barnadesioideae (D.Don) K.Bremer & R.K.Jansen (1992: 415). Type: Barnadesia Mutis ex L.f.
Annual, or perennial herbs, subshrubs, shrubs, trees, or woody vines, up to 30 m tall. Stems erect, decumbent or
scandent, single- to much-branched, with or without lenticels, cylindrical or rarely flat, with or without scales at
base, glabrous or with diverse types of trichomes, unarmed or armed with axillary spines—rarely solitary, in pairs or
fascicles, straight or curved, divergent or convergent (Fig. 2), glabrous, pubescent, villous or tomentose at the base
and becoming glabrous towards the apex. Leaves simple, alternate, opposite, or fasciculate, rarely rosulate, sessile to
petiolate, persistent or deciduous, blade linear, oval, elliptic, obovate or orbicular, chartaceous, coriaceous or succulent,
pale or lustrous, glabrous or with diverse types of trichomes, leaf base acute or obtuse, sometimes amplexicaul, margin
entire, flat, revolute or involute, rarely plicate, glabrous or commonly ciliate apex unarmed, mucronate or spiny;
nervation brochidodromous, eucamptodromous, actinodromous (with 3–5 basal or supra-basal veins), acrodromous,
parallelodromous or hyphodromous, venation immersed in the mesophyll and barely visible in succulent leaves.
Capitulescence (the secondary arrangement of capitula) terminal or axillary, monocephalous, cymose, corymbiform,
paniculiform, racemiform, spiciform or umbelliform. Capitula 1–135-flowered, homogamous or heterogamous,
discoid, disciform or radiate, sessile to pedunculate, involucre cylindrical to campanulate, 3–14-seriate, phyllaries
imbricate, scarious to coriaceous, commonly colorful, erect to reflexed, glabrous to densely villous, triangular-ovate to
linear, apex unarmed, mucronate or spiny, margin flat or reflexed, glabrous or commonly ciliate. Receptacle flat, rarely
convex, pilose, rarely glabrous. Flowers isomorphic or dimorphic, bisexual or pistillate by the androecium atrophy or
staminate by the suppression of the style, corolla bilabilate (3+2), subbilabiate (4+1, 3+1), tubular (5+0, 4+0, 3+0), or
FERREIRA ET AL.
112 • Phytotaxa 518 (2) © 2021 Magnolia Press
ligulate (5+0), 3–5-lobed, cream, white, yellow, orange, red, pink to purple, glabrous to villous on both surfaces. Anthers
5, rarely 3 or 4, apical appendage acute, bilobed, apiculate, emarginated, obtuse, lanceolate, basal appendage ecaudate
or caudate, ecalcarate or calcarate, inserted from the base to the throat of corolla, filaments free, glabrous, rarely
partial connate or connate into a staminal tube, villous; staminodes shorter than the stamens or indistinct from these.
Style cylindrical or rarely swollen bellow the branching point, cream, white, yellow, red, purple or orange; pistillodes
usually shorter than the fertile pistils. Cypsela cylindrical, fusiform or turbinate, villous or rarely glabrous. Pappus
barbellate, plumose, scaly, setaceous, rarely absent, 1-seriate, connate at base, shorter or equal to the corolla length,
glabrous or with bristles pink or red. Pollen with or without intercolpal depressions, psilate, lophate, microechinate,
sparsely microechinate, scabrate-microechinate (Stuessy & Urtubey 2007). Chromosome number (haploid numbers)
= 8, 12, 24, 25, 26, 27, 31, 48, 50, 54 (Diers 1961, Heiser 1963, Olsen 1980, Cialdella & López 1981, Wulff 1990,
Stuessy & Sagástegui 1993, Strother & Panero 1994, Stuessy et al. 1996, Watanabe et al. 2007).
FIGURE 2. Synapomorphies of Barnadesioideae. A–C: Diversity of spines. D: Barnadesioid trichomes. A–B: Dasyphyllum vagans. A:
Spines in pairs, curved, and convergent. B: Spines in pairs, straight, and convergent. C: Barnadesia parviflora. Spines in fascicles, straight,
and divergent. D: SEM photograph of Barnadesioid trichomes.
Barnadesioideae comprises 10 genera and 80 species endemic to South America. Three genera, Dasyphyllum (27
species), Chuquiraga (22 spp.) and Barnadesia (19 spp.), contain 85% of the species, while the other seven genera
(Archidasyphyllum, Arnaldoa, Doniophyton, Duseniella, Fulcaldea, Huarpea, and Schlechtendalia) are represented
by up to three species each. Most species are found in the Andes, especially in montane, xeric vegetations in the
Páramos (alpine meadows), the Puna (highland and lowland steppe), and in the Patagonian steppe and semidesert, but
also in the Andean tropical and subtropical mixed (deciduous-evergreen) forests (especially species of Barnadesia). A
secondary center of diversity spans in Brazil, Bolivia, Paraguay, Uruguay and northeastern Argentina, with species of
MORPHOLOGICAL DIVERSITY IN SPINY SUNFLOWERS Phytotaxa 518 (2) © 2021 Magnolia Press • 113
Dasyphyllum occurring in many types of vegetation, Schlechtendalia occurring in the Pampas (grassland in south Brazil
and adjacent areas in Uruguay and Argentina), and Archidasyphyllum restricted to the Nothofagus forest (broadleaf and
mixed beech forest) of central Chile and adjacent central-western areas of Argentina (Fig. 1).
Species of Barnadesioideae are mainly perennial or annual herbs, subshrubs, or shrubs, but there are also trees
(some up to 30 m high, as in Archidasyphyllum) or woody vines. Representatives of the subfamily are characterized by
morphological synapomorphies that include the presence of axillary spines (in pairs or fascicles, rarely solitary), and
by the “barnadesioid trichomes”—a pubescence of unbranched three-celled hairs on the corollas, cypselae and pappus
(Cabrera 1959, Urtubey 1999, Erbar & Leins 2000, Stuessy et al. 2009, see Fig. 2).
Key to genera of Barnadesioideae
1. Herbs or subshrubs .............................................................................................................................................................................2
- Shrubs, trees or woody vines ..............................................................................................................................................................5
2. Axillary spines present ....................................................................................................................................................Doniophyton
- Axillary spines absent .........................................................................................................................................................................3
3. Capitula discoid; leaves with parallelodromous nervation .........................................................................................Schlechtendalia
- Capitula disciform or radiate; leaves with hypodromous or actinodromous nervation; nervation immersed in the mesophyll and
barely visible in succulent leaves .......................................................................................................................................................4
4. Leaves succulent, capitula with marginal flowers female and corolla tubular 10–40, disc flowers hermaphroditic and corolla
tubular 30–95, anthers caudate ........................................................................................................................................... Duseniella
- Leaves not succulent; capitula with marginal flowers female and corolla subbilabiate 5, disc flower hermaphroditic and corolla
tubular 1, anthers ecaudate .................................................................................................................................................... Huarpea
5. Capitula uniflowered; style swollen below the branching point ......................................................................................... Fulcaldea
- Capitula multiflowered; style not swollen below the branching point ...............................................................................................6
6. Capitula radiate ..................................................................................................................................................................Barnadesia
- Capitula discoid ..................................................................................................................................................................................7
7. Leaves with brochidodromous or eucamptodromous venation; anthers with apical appendage emarginated ........Archidasyphyllum
- Leaves with actinodromous or hypodromous venation; anthers with apical appendage acute, bilobed, lanceolate or obtuse .........8
8. Corolla orange, red, purple, yellow, or rarely white (A. argentea); anthers with apical appendage acute, lanceolate or obtuse .......9
- Corolla white or yellow; apical appendage bilobed ........................................................................................................Dasyphyllum
9. Corolla subbilabiate; anthers with filaments inserted at the corolla throat ...........................................................................Arnaldoa
- Corolla tubular; anthers with filaments inserted at the base of the corolla ......................................................................Chuquiraga
1. Archidasyphyllum (Cabrera) P.L.Ferreira, Saavedra & Groppo (2019: 13)
Type:—Archidasyphyllum diacanthoides (Less.) P.L.Ferreira, Saavedra & Groppo
Trees, up to 30 m tall, multi-stemmed. Stems erect or decumbent, much-branched, lenticelate, cylindrical, scales
imbricate at base, glabrous, strigose, villous, or velutinous, unarmed or with axillary spines in pairs of fascicles,
straight, convergent or rarely divergent, glabrous or rarely sparse pubescent at base and becoming glabrous towards
the apex. Leaves alternate, spiral, sessile to petiolate, persistent or deciduous, blade narrow elliptic, elliptic, obovate
or orbiculate, coriaceous, pale or lustrous, glabrous or pubescent in both surfaces, base acute or obtuse, margin flat,
ciliate, apex mucronate or spiny; venation brochidodromous or eucamptodromous. Capitulescence terminal or axillary,
monocephalous or spiciform. Capitula 20–35-flowered, homogamous, discoid, sessile to pedunculate, involucre
campanulate, 3–4-seriate, phyllaries coriaceous, green or brown, erect or rarely slightly reflexed, glabrous or densely
villous, ovate-triangulate grading to lanceolate, apex unarmed or mucronate, margin flat, ciliate. Receptacle flat, pilose.
Flowers isomorphic, bisexual, or pistillate by the gynoecium atrophy, corolla tubular (5+0), ligulate (5+0), subbilabiate
(4+1) or bilabiate (3+2), 5-lobed, white to cream, externally sericeous at lobes apex. Anthers 5, apical appendage
emarginated, basal appendage lanceolate, acute to rounded, caudate, calcarate or ecalcarate, inserted at the throat of
the corolla, filaments free, glabrous. Style cylindrical, cream to yellow. Cypsela fusiform, densely villous. Pappus
plumose, slightly shorter or equal to corolla length, bristles whitish. Pollen with intercolpal depressions (Suessy &
Urtubey 2007). Chromosome number unknown.
Fig. 3a–b; 10e.
Distribution and habitat:—Archidasyphyllum comprise two species restricted to the Nothofagus (southern
beech) forest (broadleaf and mixed beech forest) of central Chile and adjacent central-western areas of Argentina from
above an altitude of 1,200 m a.s.l. (Cabrera 1959, Ferreira et al. 2019).
FERREIRA ET AL.
114 • Phytotaxa 518 (2) © 2021 Magnolia Press
Notes:—Archidasyphyllum can be distinguished from the rest of Barnadesioideae by a set of characters:
monoecious or gynodioecious trees reaching 30 m tall, discoid capitula, and anthers with emarginated apical appendages.
Cabrera (1959), in the taxonomic revision of Dasyphyllum, considered Archidasyphyllum as a subgenus due to the
high morphological similarity, because both genera have discoid capitula, a monoecious or gynodioecious breeding
system, pollen with intercolpar depressions, and similar corolla shape and color. Moreover, Cabrera (1959) believed
that Dasyphyllum, especially the subgenus Archidasyphyllum (Archi—from the ancient Greek ἀρχι- chief, early +
Dasyphyllum—a genus of Barnadesioideae) was the most primitive group and therefore the ancestor of the other genera
within the subfamily. However, molecular phylogenies recovered Dasyphyllum subgenus Archidasyphyllum as sister to
the Arnaldoa and Fulcaldea clade (Gustafsson et al. 2001, Gruenstaeudl et al. 2009, Ferreira et al. 2019). Therefore,
Ferreira et al. (2019) elevated the subgenus Archidasyphyllum to the rank of genus based on molecular phylogenetic,
morphology, and biogeographical evidence. Although all phylogenetic analyses based on molecular data recover
Archidasyphyllum as sister to Fulcaldea and Arnaldoa, this clade is morphologically diverse and synapomorphies that
support the relationships are still unknown (Funk & Roque 2011, Ferreira et al. 2019).
Iconography:—Poeppig & Endlicher (1835–1836), Cabrera (1959).
Accepted species:—1.1. Archidasyphyllum diacanthoides (Less.) P.L.Ferreira, Saavedra & Groppo; 1.2.
Archidasyphyllum excelsum (D. Don) P.L.Ferreira, Saavedra & Groppo.
Selected specimens examined:—ARGENTINA. Río Negro: Bariloche, near Blest, 800 m, 41°02’S, 71°49’W, 03
March 1979, K. Rahn 4643 (MO; 1.1). Parque Nacional Nahuel Huapi, January 1968, Esk 150 (MBM; 1.1). CHILE.
Araucanía: Cautín, 3 km E of Melipeuco, 600 m, 38°51’S, 71°42’W, 03 February 1990, M.F. Gardner 4704 (E;
1.1). Malleco, Camino entre Curacautín y Malalcahuello, 3 km antes de Manzanar, 700 m, 38°28’S, 71°47’W, 07
January 1977, C. Marticorena 1225 (B; 1.1). Caraucautín, Estrada para Parque Nacional Tolhuaca, próximo à Terma
de Tolhuaca, na beira da estrada, no lado esquerdo, 1065 m, 38°14’43’’S, 71°44’44.2’’W, 25 September 2013, C.L.
Silva-Luz 200 (SPFR, SPF; 1.1). Bío-bío: Recinto, N from Hotel Lleuques, toward small hill, 750 m, 14 January 1993,
T.F. Stuessy 12703 (WU; 1.1). Los Lagos: Llanquihue, Lago de todos los santos, 41°13’S, 72°16’W, K.W. Wolffhügel
s.n. (B; 1.1). Osorno, Parque Nacional Puyehue, Termas de Puyehue, 560 m, 40°40’S, 72°20’W, 01 February 1988,
M.F. Gardner 4045 (E; 1.1). Los Rios: Valdivia, Cordillera de la Costa, road to Curiñanco on the way to Parque Oncol.,
379 m, 30°46’16.9’’S, 73°19’W, 18 January 2003, M.F. Gardner 8 (E; 1.1). Valparaíso: Quillota, Olmué, Cordillera de
la Costa, Parque Nacional La Campana (sector Granizo), 700 m, 32°57’S, 71°05’W, 01 February 1998, P. Baxter 2 (E;
1.2). Quillota, Parque Nacional Cerro Campana, 17 January 1993, T.F. Stuessy 12749 (WU; 1.2).
2. Arnaldoa Cabrera (1962: 39)
Type:—Arnaldoa weberbaueri (Muschl.) Ferreyra
Arching shrubs, up to 4 m tall, multistemmed. Stems erect or decumbent, much-branched, lenticelate, cylindrical,
glabrous, velutinous or densely tomentose, axillary spines in pairs, straight, convergent or divergent, glabrous or
tomentose at base and becoming glabrous towards the apex. Leaves alternate, petiolate, persistent or deciduous, blade
elliptic, ovate or oblate, coriaceous, pale or lustrous, adaxially glabrous to tomentose, abaxially glabrous to densely
tomentose, villous, floccose, lanose, apex acute or obtuse, margin flat or slightly revolute, glabrous or ciliate, apex
mucronate or spiny; venation actinodromous with 3 basal suprabasal nerves. Capitulescence terminal, monocephalous.
Capitula 30–95-flowered, homogamous, discoid, sessile, involucre campanulate, 8–15 series, phyllaries coriaceous,
brown or black, erect or reflexed, glabrous to densely tomentose, ovate—triangulate grading to lanceolate, apex
mucronate or spiny, margin flat, ciliate. Receptacle flat, pilose. Flowers isomorphic, bisexual, corolla subbilabiate
(4+1), 5-lobed, red, purple, orange or white, villous. Anthers 5, apical appendage lanceolate or obtuse, basal appendage
truncate, caudate, calcarate, inserted at the throat of the corolla, filaments free, glabrous or villous. Style cylindrical, red,
purple, orange, white. Cypselae turbinate or cylindrical, densely villous. Pappus plumose, slightly shorter or equal to
MORPHOLOGICAL DIVERSITY IN SPINY SUNFLOWERS Phytotaxa 518 (2) © 2021 Magnolia Press • 115
FIGURE 3. Archidasyphyllum and Arnaldoa. A–B: Archidasyphyllum diacanthoides. A: Habit. B: Multi-stemmed tree with fasciculate
spines. C–D: Arnaldoa weberbaueri. C. Habit. D. Capitulum. Photos by Marcelo Monge (A–B), Gari Ccana-Ccapatinta (C), and Daniel
Montesinos (D).
FERREIRA ET AL.
116 • Phytotaxa 518 (2) © 2021 Magnolia Press
corolla length, bristles whitish. Pollen with intercolpal depression, microechinate (Suessy & Urtubey 2007). Chromosome
number = 24–27 (Stuessy & Sagástegui 1993).
Fig. 3c–d; 10f.
Distribution and habitat:—Arnaldoa comprise three shrubby species found in Monte Bajo Puna vegetation
(shrubby dry slopes) above an altitude of 1,500 m in northern Peru and southern Ecuador (Stuessy & Sagástegui-Alva
1993, Ulloa et al. 2002).
Notes:—Arnaldoa is morphologically close to Chuquiraga, sharing the shrubby habit, discoid capitula, colorful
corollas and stigma, long caudate anthers, and microechinate pollen. However, Arnaldoa is distinguished by the
subbilate corolla (vs. tubular in Chuquiraga sensu Ezcurra 1985) and stamens inserted on the throat of corolla tube (vs.
base of corolla in Chuquiraga). Phylogenetic hypotheses based on molecular data recovered Arnaldoa as closely related
to Fulcadea (Gustafsson et al. 2001, Gruenstaeudl et al. 2009, Funk & Roque 2011, Ferreira et al. 2019). Although the
morphological differences between Arnaldoa and Fulcaldea are remarkable, Arnaldoa by having solitary capitula with
30–95-flowered, subbilabiate corollas, caudate anthers, microechinate pollen, cylindrical style vs capitula arranged
in synflorescence with 1-flowered, tubular corollas, ecaudate anthers, spinulose pollen, and swollen style below the
branching point in Fulcaldea, they share the arching shrubby habit, red corollas, and geographical distribution in
southern Ecuador and Northern Peru.
Iconography:—Stuessy & Sagástegui-Alva (1993), Ferreyra (1995), Ulloa et al. (2002).
Accepted species:—2.1 Arnaldoa argentea C. Ulloa, P. Jørg. & M.O. Dillon; 2.2. Arnaldoa macbrideana Ferreyra;
2.3. Arnaldoa weberbaueri (Muschl.) Ferreyra.
Selected specimens examined:—ECUADOR. Loja: Amaluza, road to the antennas above town, at the top, 2400
m, 4°33’58”S, 79°26’26’’W, 05 November 2000, P.M. Jørgensen 2234 (QCNE, MO, WU; 2.1). Bellavista, Vicinity of
Bellavista, at “Las Antenas” (7 km from Amaluza), 2400 m, 04°33’S, 79°26’W, 14 August 2001, J.E. Madsen 8341
(MO; 2.1). Vicinity of Bellavista, at “Las Antenas” (6–7 km from Amaluza), 2200–2300 m, 04°33’S, 79°27’W, 31 may
2001, J.E. Madsen 8175 (MO; 2.1). PERU. Amazonas: Chachapoyas, western base of the Cerros Calla Calla, 22km
east of Balsas on the road to Leymebamba, 1940 m, 30 May 1964, P.C. Wright 5454 (K, MO; 2.3). Cordillera de Calla
Calla, Balsas-Leymebamba road near km 359, 6400 ft. (c. 1951m) 9 July 1977, T. Ducan 2602 (MO; 2.3). Ancash:
Chingas, north of the village of Llamellin, on the west flanks of the Rio Marañón, 2669 m, 09°04’08’’S, 76°59’39’’W,
27 May 2011, E. Gagnon 3157 (K; 2.3). Huari, Falda de los Cerros de los rios Mosna y Puchca, 2400–2800 m, 27 May
1962, M.Z. Agnes 41 (MO; 2.3). Cajamarca: Celendín, Rio Marañon valley, Celedín-Balsas road, 25 km, and onward
from Celendín, 1000–2500 m, 06°51’S, 78°05’W, 16 July 1983, D.N. Smith 4323 (MO; 2.3). Celendín, Had. El Limón
(Celendín-Balsas), 2150 m, 5 May 1970, A. Sagastégui A. 7410 (US; 2.3). Celendín, Hacienda Limon, 33 kms E of
Celendín on road to Rio Marañon, 2050 m, 10 July 1992, T.F. Stuessy 12524 (WU; 2.3). Lambayeque: Lambayeque,
Alredores del Cuello Porculla, 1800 m, 24 July 1991, J. Mostacero L. 2296 (MO; 2.2). Lambayabeque, Road from
Olmos to Abra Porculla, 1500 m, 22 February 1976, T. Plowman 5516 (US; 2.2). Lambayabeque, west side Abra
Porculla Km. 35 east of Olmos. Mesones-Muro Highway between Olmos y Río Maranon, 1650 m, 26 September
1957, C. Hutchison 1379 (US; 2.2). Libertad: Pataz, road Buldibuyo to Tayabamba, dryish inner andean valley, mostly
cultivated, but steeper slopes with scrub forest, 3141 m, 08°12.571’S, 77°19.665’W, 24 April 2004, M. Weigend 8031
(B; 2.3). Pataz, Huancaspata-Puente Mamahuaje, 3000 m, 26 June 1974, A. López M. 8218 (MO; 2.3). Bolivar, District
La Longotea, section Pampa Colorada, 1914 m, 13 May 2011, 07°02’31’’S, 77°55’08’’W, R.W. Bussman 16871 (MO;
2.3). Piura: Abra de Porculla, entre Olmos y Jaen, 2100–2200 m, 28 June 1959, R. Ferreyra 13731 (MO; 2.2). Bajando
Abra de Porculla y Jaen, cerca del Abra, 1800–1900 m, 26 June 1959, R. Ferreyra 13628 (MO; 2.2).
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
MORPHOLOGICAL DIVERSITY IN SPINY SUNFLOWERS Phytotaxa 518 (2) © 2021 Magnolia Press • 117
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
FERREIRA ET AL.
118 • Phytotaxa 518 (2) © 2021 Magnolia Press
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 west-
southern 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. Jørgensen 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
MORPHOLOGICAL DIVERSITY IN SPINY SUNFLOWERS Phytotaxa 518 (2) © 2021 Magnolia Press • 119
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).
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,
FERREIRA ET AL.
120 • Phytotaxa 518 (2) © 2021 Magnolia Press
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).
4. Chuquiraga Jussieu (1789: 178)
Type:—Chuquiraga jussieui J.F. Gmel.
Shrubs or dwarf cushion-forming shrubs, up to 2 m tall. Stems erect, much-branched, without lenticels, cylindrical,
rarely scales imbricate at base, glabrous, tomentose, strigose, villous, sericeous, unarmed or with axillary spines in
pairs or fascicles, straight, convergent or divergent, glabrous, villous or pubescent at base and becoming glabrous
towards the apex. Leaves alternate or opposite, sessile or subsessile, persistent or deciduous, blade acicular, linear, or
ovate, coriaceous, pale or commonly lustrous, glabrous to densely sericeous on both surfaces, base acute or obtuse,
margin flat or involute, glabrous or ciliate, apex mucronate or spiny; venation hyphodromous or actinodromous with 3
MORPHOLOGICAL DIVERSITY IN SPINY SUNFLOWERS Phytotaxa 518 (2) © 2021 Magnolia Press • 121
basal supra-basal nerves. Capitulescence terminal, monocephalous. Capitula 5–100-flowered, homogamous, discoid,
sessile, involucre cylindrical, turbinate or campanulate, 4–14-seriate, phyllaries scarious or commonly coriaceous,
green, brownish, yellow to orange, erect or reflexed, glabrous to densely sericeous, triangular-ovate to lanceolate, apex
mucronate or spiny, margin flat or reflexed, glabrous or ciliate. Receptacle flat, pilose. Flowers isomorphic, bisexual,
tubular (5+0), 5-lobed often unequal and deep incisions between lobes, yellow, orange, villous. Anthers 5, apical
appendage lanceolate, obtuse or acute, basal appendage rounded or acute, caudate, calcarate or ecalcarate, inserted at
the base of the corolla, filaments free, glabrous. Style cylindrical, yellow to orange. Cypselae turbinate, densely villous.
Pappus plumose, shorter than or equal to corolla length, bristles whitish. Pollen without intercolpar depressions,
microechinate (Suessy & Urtubey 2007). Chromosome number = 24–27 (Diers 1961, Wulff 1990).
Fig. 5; 10b.
FIGURE 5. Chuquiraga. A–B: Chuquiraga aurea. A: “Cushion” Habit. B: Capitulum. C–D: Chuquiraga longiflora. D: Capitulum. D:
Tubular corolla. E: Chuquiraga jussieui. Capitulum. F: Chuquiraga weberbaueri. Shrub Habit. Photos by Darwinion Institute (A–D),
Paola Ferreira (E), Gari Ccana-Ccapatinta (F).
FERREIRA ET AL.
122 • Phytotaxa 518 (2) © 2021 Magnolia Press
Distribution and habitat:—Chuquiraga is the second largest genus of Barnadesioideae, comprising 22 species
(Ezcurra 1985, Harling 1991, Sagástegui & Sánchez 1991, Granda 1997). The species are found in the Andes and
Patagonia, from Colombia to Argentina, in the Páramos (alpine meadows), in the Puna (Highlands and steppe), and
open woodlands. Some species can be found at 4,200 m. a.s.l, but there are also records in lowland Puna near sea
level.
Notes:—Species of Chuquiraga contain several medicinal properties commonly used into the phytopharmaceutical
industry and commercialized by European countries. In addition, Chuquiraga is largely commercialized in some
medicinal marketplaces in Peru and Ecuador and it is indicated for the treatment of several conditions (Ccana-
Ccapatinta et al. 2018).
The species of Chuquiraga are classified into two sections (following Ezcurra 1985): Chuquiraga sect.
Acanthophylla, comprising nine species distributed from northwest Bolivia to Chile and Argentina (Fig. 5a, b) and
characterized by leaves with the margin involute, naviculate or cymbiform (Fig. 5b), abaxial surface without the
prominent midvein, and absence of axillary spines; and Chuquiraga sect. Chuquiraga, the most species-rich in the
genus, comprising 13 species distributed from Colombia to Argentina (Fig. 5 c–e) and characterized by flat leaves,
the abaxial surface with prominent midvein and by the presence of axillary spines, rarely reduced or absent. Within
the Chuquiraga section Chuquiraga, two series are recognized: Chuquiraga and Parviflorae, distinguished by the
length of capitula (>3cm vs. 0–3cm), corolla (>17mm vs. <16mm) and anthers (15–20mm vs. 8–12mm), and by the
geographical distribution (from Colombia to northwest Argentina vs. western Bolivia to Argentinean Patagonia).
The great morphological variation in habit, leaf shape, venation, trichomes, spines, involucre shape, size, bract
colors, and others, makes Chuquiraga the most taxonomically complex group of Barnadesioideae. After analyzing
hundreds of specimens and other material from this genus, we decided to follow the corolla classification sensu Ezcurra
(1985), and do not use the classification sensu Suessy & Urtubey (2006). Although the study by Stuessy & Urtubey
(2006) represents a significant step towards understanding corolla evolution in Barnadesioideae, in Chuquiraga it is
difficult to apply a corolla classification, since this genus has different lengths of corolla incisions splitting the lobes.
Therefore, we prefer to classify the corolla as tubular with different incision lengths (see Sagástegui & Sánchez 1991,
Figure 1.I; for the corolla line drawing).
The most comprehensive phylogeny at species-level for Chuquiraga was performed based on molecular data by
Padin et al. (2015b). The results were incongruent regarding the monophyly of the genus in relation to the position
of Doniophyton or Duseniella. Within the genus, the sections were found to be monophyletic, but the results do not
corroborate the series of Chuquiraga sect. Chuquiraga, a finding that was also corroborated by phylogenomic studies
(Ferreira et al. in prep.). Further taxonomical and morphological studies are therefore needed in order to propose a new
classification and species circumscription.
Iconography:—Ezcurra (1985), Harling (1991), Sagástegui & Sánchez (1991), Ferreyra (1995), Granda (1997).
Accepted species:—Chuquiraga section Acantophylla C. Ezcurra: 4.1 Chuquiraga acanthophylla Wedd.;
4.2 Chuquiraga atacamensis Kuntze; 4.3 Chuquiraga aurea Skottsb.; 4.4 Chuquiraga echegarayi Hieron.; 4.5.1
Chuquiraga erinacea subsp. erinacea D. Don; 4.5.2 Chuquiraga erinacea subsp. hystrix (D.Don) C.Ezcurra; 4.6
Chuquiraga kuschelii Acevedo; 4.7 Chuquiraga rosulata Gaspar; 4.8 Chuquiraga ruscifolia D.Don; 4.9.1 Chuquiraga
ulicina subsp. acicularis (D.Don) C.Ezcurra.; 4.9.2 Chuquiraga ulicina subsp. ulicina Hook. Chuquiraga section
Chuquiraga series Chuquiraga: 4.10 Chuquiraga arcuata Harling; 4.11 Chuquiraga calchaquina Cabrera; 4.12
Chuquiraga jussieui J.F.Gmel.; 4.13 Chuquiraga longiflora (Griseb.) Hieron; 4.14 Chuquiraga oblongifolia Sagást.
& Sánchez Veja; 4.15 Chuquiraga raimondiana A.Granda; 4.16.1 Chuquiraga spinosa subsp. australis C.Ezcurra;
4.16.2 Chuquiraga spinosa subsp. huamanpinta C.Ezcurra; 4.16.3 Chuquiraga spinosa subsp. rotundifolia (Wedd.)
C.Ezcurra; 4.16.4 Chuquiraga spinosa subsp. spinosa D.Don; 4.17 Chuquiraga weberbaueri Tovar. Chuquiraga sect.
Chuquiraga series Parviflorae C.Ezcurra: 4.18 Chuquiraga avellanedae Lorentz; 4.19 Chuquiraga morenonis (Kuntze)
C.Ezcurra; 4.20 Chuquiraga oppositifolia D.Don; 4.21 Chuquiraga parviflora (Griseb.) Hieron; 4.22 Chuquiraga
straminea Sandwith.
Selected specimens examined:—ARGENTINA. Catamarca: Andalgalá, Las Niñas, 05 March 1916, P. Jörgensen
1671 (US; 4.5.1). Andalgalá, Rio Potrero sup., Cuenca Pelada, 2750 m, 27 February 1951, H. Sleumer 1959 (B; 4.13).
Chubut: 25 kms N of Comodoro Rivadavia on rte 3 (ca. 5 kms S of Estancia Begonia), 220 m, 16 February 1993, T.F.
Stuessy 12928 (MO; 4.3). Florentino Ameghino, Ruta 3, Sarmiento, A.L. Cabrera 33267 (SI; 3.5.2). Río Senguerr, Rio
Mayo, 17 December 1981, A.L. Cabrera 33187 (SI; 4.19). Sarmiento, Ruta 26, 50 km al E de Sarmiento, 17 December
1981, A.L. Cabrera 33245 (SI; 4.19). Sierra de San Bernardo, 15 December 1981, A.L. Cabrera 33225 (SI; 4.3). 25
kms N of Comodoro Rivadavia on rte 3, ocean can be seen in the distance, 200 m, 16 February 1993, T.F. Stuessy
12929 (WU; 4.18). 96 kms E of Sarmiento on rte 26 to Comodoro Rivadavia (ca. 56 kms W of Comodoro Rivadavia),
MORPHOLOGICAL DIVERSITY IN SPINY SUNFLOWERS Phytotaxa 518 (2) © 2021 Magnolia Press • 123
600 m, 17 February 1993, T.F. Stuessy 12940 (WU; 4.19). Jujuy: Abra de Pipes, 11 November 1973, A.L. Cabrera
24037 (SI; 4.16.1). Rinconada, desde la Cienaga a Lipan, 2800 m, 23°41’8.74’’S, 65°33’058’’W, 08 January 2001,
P.M. Simon 512 (MO; 4.5.1). Santa Catalina, Ruta provincial 5, 20 km de la Quiaca caminho a Santa Catalina, Cuesta
de Toquero, 3660 m, 22°07’S, 65°45’W, 16 February 1998, O. Morrone 2671 (MO; 4.1). Tumbaya, along rte 40 to El
Moreno ca. 7 km S of its intersection with rte 16, 3310 m, 23°42’01’’S, 65°50’22’’W, 15 March 1994, C.M. Taylor
11189 (MO; 4.2). Tumbaya, Ruta Nac. 9.4 km de Volcán caminho a S.S. de Jujuy, cantera sobre el Rio Grande, 2040
m, 23°56’S, 65°27’W, 10 December 1998, O. Morrone 3173 (MO; 4.21). Valle Grande, entre três morros y Cerro
Hermoso, Serranía Callileagua, 2700 m, 23°37’15.02’’S, 64°54’35.78’’W, 18 April 2016, C.M. Martín 736 (SPFR;
4.13). Yavi, La Quiaca, 2km antes de llegar al aeropuerto, 3400 m, 28 January 1999, A. Chapin 25951 (MO; 4.1). Yavi,
cerca de La Quiaca, 20 January 1942, A.L. Cabrera 7780 (K; 4.1). La Pampa: Monte em Laguna Amarga, 17 February
1948, A. Burkart 15951 (SI; 4.5.1). Zwischen Santa Isabel und 25 de Llago, 19 November 1965, H. Walter 456 (B;
4.7). La Rioja: Chilecito, caminho a mina el Oro (cerca de Chilecito), 2000 m, 29°05’7.81’’S, 67°41’8.76’’W, 23
January 2001, P.M. Simon 720 (MO; 4.21). General Lamadrid, Ruta providencial 76 de Punta de Agua a Laguna Brava,
3300 m, 28°30’23’’S, 68°47’51’’W, 24 January 2012, F.O. Zuloaga 13753 (MO; 4.5.2). Río Amarillo, sierra de
Famatina, 3900 m, 15 January 1947, J.H. Hunziker 1959 (MO; 4.8). Mendoza: Along route #143, ca 150 kms generally
NW of San Rafael, 2700 ft. (c. 822m), 15 January 1985, R.M. King 9436 (MO; 4.5.1). Caracoles de Villavicencio,
cerca del empalme a la Quebrada del Toro, 2700 m, 11 May 1980, O. Bottino 85 (WU; 4.8). High desert near Uspallata,
route to Mina Monya (talc) 18 km E of Uspallata, 2800 m, 32°37’S, 69°32’W, 12 January 1996, R.R. Brooks MS32
(MO; 4.5.2). Marlargüe, Valle de Río Salado, a 13 km de Los Molles hacia las Leñas, puente sobre el Arooyo el
Deshecho, 2000 m, 12 January 1988, B. Leunberger 3813 (B; 4.20). San Rafael, cerca de Cerro Bola, 17 January 1982,
A.L. Cabrera 33319 (SI; 4.7). Ruta 40 a 24 km S de Calmuco, 01 February 1963, O. Boelcke 10421 (SI; 4.18).
Neuquén: Añelo, Ruta Prov. 17, 6 km al E de Añelo em la meseta, 11 November 1981, J.J. Valla 27 (MO; 4.7). Chos
Malal: 27 km de Chos Malal, por ruta prov. 43, em dirección a Andacollo, em Cañadón, 1200 m, 37°15’37’’S,
70°29’32’’W, 27 February 1994, H. Fortunato 4322 (US; 4.20). Pehuenches, Laguna Auquinco, January 1984, A.
Wulff 359 (SI; 4.18). Along the gravel road to route #40, ca 68 kms from route #40, 4700 ft.(c. 1432m), 12 January
1985, R.M. King 9404 (US; 4.20). Along route #40. Ca. 26 kms from route #237, 1800 ft. (c. 548m), 09 January 1985,
R.M. King 9378 (MO; 4.22). Río Negro: Adolfo Alsina, road to San Antonio 100km west of Viedma, growing along
roadside in clay sand, 30 m, 19 December 1938, W.J. Eyerdam 23496 (MO; 4.18). 25 de Mayo, 9 km al W de Ing.
Jacobacci en dirección a Bariloche, 800 m, 41°19’59’’S, 69°40’09’’W, 04 March 1994, H. Fortunato 4373 (US; 4.22).
General Roca, 250–360 m, Sep 1914–February 1915, W. Fischer 11 (MO; 4.7). Manuel Choique, Winderosionsfäche
in Talgrund am westlichen Ortsausgang, 800 m, 41°45’S, 70°10’W, 13 November 1984, Hager 1008 (B; 4.3). Along
the route #22m ca 36 kms W of Rio Colorado, 200 ft.(c. 60.9m), 05 February 1985, R. M. King 9336 (MO; 4.5.1).
Salta: Chicoana, Quebrada de Escoipe, entre Escoipe y San Fernnando, 1900–2000 m, 15 December 1987, L. Novara
9200 (B; 4.21). San Antonio de Los Cobres, 21 October 1948, H. Humbert 21072 (P; 4.2). San Antonio de Los Cobres,
3800 m, 13 December 1946, A. Krapovickas 3131 (MO; 4.2). Chicoana x Cachi, Valle Encantado, 3000–3400 m, 24
June 1980, I. Novara 1109 (B; 4.11). Valle Encantado (Mirador), 3273 m, 25°10’55.3’’S, 65°51’11.9’’W, 29 February
2016, C.M. Martín 608 (SPFR; 4.13). San Juan: Calingasta, Reserva El Leoncito, subiendo el Cordón del Naranjo por
la Quebrada de Las Minas, 17 km al este de la Ruta Prov N. 412, paraje llamado Pampa de Los Pozos, 3325 m, 20
March 1999, F. Biurrun 5749 (SI; 4.4). Calingasta, de confluência a Alojo Las minitas, 2500–3000 m, 17 February
1988, R. Kiesling 6857 (SI; 4.4). Calingasta, Reserva El Leoncito, Rancho del Cura, 2802 m, 31°46’S, 69°13’W, 15
March 2005, R. Kiesling 10206 (SI; 4.4). Calingasta, limite com Mendoza, Yalguaraz, 21 January 1987, R. Kiesling
6514 (SI; 4.5.2). Calingasta, Qda. De Santa Clara, 1700 m, 32°06’49’’S, 68°57’44’’W, 17 February 2000, T.S. Tombesi
79 (SI; 4.7). Iglesia, Quebrada del Agua Negra, 3000 m, 23 February 1979, A.L. Cabrera 30131 (SI; 4.8). Zonda,
camino a estancia maradona, 1300 m, 27 January 1981, R. Kiesling 3310 (SI; 4.5.1). Zonda, Cno. a Estancia Maradona,
2100 m, 12 November 1982, R. Kiesling 4079 (SI; 4.7). Santa Cruz: Deseado, SW Cañadon Seco, 350 m, 03 January
1991, B. Leuenberger 4095 (B; 4.3). Deseado, em la estepa, 25 November 1979, R. Méndez 9868 (SI; 4.19). Corpen
Aike, Laguna Grande a Gob Gregores, 8 km, 49°34’S, 70°14’W, 24 January 1967, O. Boelke 12770 (SI; 4.3). Tucumán:
Tafí, Tafí del Valle, Cerro Pelado, 2250 m, 24 January 1950, H. Sleumer 132 (B; 4.11). Tafí, subida al Infiernillo, 07
November 1978, A.L. Cabrera 29699 (SI; 4.11). Tafí, quebrada del Río blanco, cerca del Valle Tafí, 2600 m, 26
January 1950, H. Sleumer 105 (B; 4.13). Tafí, 3000 m, 24 April 1926, S. Venturi 4180 (US; 4.21). BOLIVIA. Chuquisaca:
Cinti, 4000 m, January 1846, A. Weddel 3974 (P; 4.1). Oropeza, between Tipoyo y Cerro Obispo on Sucre—Quila
Quila road, 2900 m, 24 January 1999, J.R.I. Wood 14397 (SPF; 4.21). Cochabamba: Carrasco, El Churo-Sunchal, ca.
de 10 km. Hacia Pojo, lado del caminho carretero Comparapa-Cochabamba, 2780 m, 17°47’50.5’’S, 64°47’24.3’’W,
I.G. Vargas 7051 (B; 4.21). La Paz: Murillo 9.5 km al oeste de Ventilla por el caminho entre Palca y Calacoto (La Paz),
FERREIRA ET AL.
124 • Phytotaxa 518 (2) © 2021 Magnolia Press
4200 m, 16°32’S, 67°58’W, 15 March 1987, J.C. Solomon 16358 (MO; 4.12). La Paz, 10.000 ft. (c. 3048m), 1889,
A.M.B. Lectae 138 (K; 4.21). Potosí: Above Quechisla (Palcahuasí), 3600 m, November 1946, M. Cárdenas 3725 (US;
4.2). Daniel Campos, Llica, 45 km. Hacia el salar de Uyuni, 19°52’57’’S, 68°13’15’’W, 22 September 2006, S.G. Beck
32409 (US; 4.2). Miraflores, 3250 m, 19°27’4’’S, 65°15’W, 04 December 1986, S. Riera 668 (B; 4.21). Sud Chicas,
Mina Usca-Isca above the village of La Torre, north of Tupiza, 3875–3880 m, 02 December 1967, B. Vuilleumier (SI;
4.16.1). Santa Cruz: Vallegrande, entre Las Lajas y Abra Grande, 2741 m, 18°16’57’’S, 64°17’59’’W, 05 December
2011, G.A. Parada 4084 (MO; 4.21). Tarija: Cercado, ladera sur, 2260 m, 15 February 2006, F. Zenterro 3739 (US;
4.21). Tarija, de Tarija a Iscayachi, 2000–3000 m, 20 March 1982, R. Kiesling 3847 (US; 4.13). CHILE. Antofagasta:
Ascotan, 3900 m, 08 March 1992, O. Kuntze s.n. (US; 4.2). Guatin, San Pedro de Atacama, Provincia Loa, 3500 m,
22°43’9.20’’S, 68°00’7.95’’W, 05 February 2001, M. Ackermann 50 (B; 4.2). El Loa, NE of Calama, along the road
from near Chiu Chiu to Caspana, ca. 46 km from its turnoff from the Calama-Chiu Chiu road, 3500 m, 22°20’36’’S,
68°15’30’’W, 10 April 1994, C.M. Taylor 11548 (MO; 4.2). Coquimbo: Elqui, Río Turbío, road from Rivadavia to
Laguna Dam, ca 75 km, 2000 m, 05 November 1938, C.R. Worth 16404 (SI; 4.9.2). Illapel, Loma Negra in the Andes
between La Veja Escondida and Cuncumen, 2300 m, 27 February 1939, J.L. Morrison 17438 (MO; 4.9.1). Rivadavia,
800 m, November 1923, E. Werdermann 102 (MO; 4.9.1). Maule: Talca, Upper valley of Rio Maule, 1500 m, 35°55’S,
70°43’W, 26 January 1990, M.F. Gardner 4579 (E; 4.20). Metropolitana: La Parva, just 0.3 km up from village towards
ski slopes, 2800 m, 16 January 1993, T.F. Stuessy 12727 (WU; 4.20). 3.7 km W of Farellones, on rd back to Santiago,
2000 m, 16 January 1993, T.F. Stuessy 12726 (WU; 4.20). O’Higgins: Copada, 1750–2000 m, F.W. Pennel 12271 (US;
4.20). Tarapacá: Arica, 90 km E of the Panamericana on Ruta 11 towards Putre, just before the frontier to Prov.
Parinacota, 2950–3000 m, 18°24.27’S, 69°39.22’W, 17 September 1997, U. Eggli 2813 (B; 4.6). Arica, antes de
Zapahuira, 3000 m, 22 February 1978, H. Escobar A. X232 (SI; 4.6). Parinacota, Putre, Cordillera de Los Andes,
Ch11—road to Putre at kilometer 102, 3300 m, 18°19’49’’S, 69°34’41’’W, 15 December 1999, M.F. Gardner 6261 (E;
4.16.3). COLOMBIA. Nariño: Cumbal, Arriba de Cumbal, alto de Machines, 3200 m, 02 September 1962, L.E. Mora
2346 (COL; 4.12). Quíndio: Génova, vereda Pedregales, prédio Valle Lindo, 3500 m, March 2004, E. Méndez 3042
(COL; 4.12). Tolima: Cajamarca, Valle del rio Anaime, Páramo de los Valles, 3550–3800 m, 11 February 1991, J.I.
Barrera 07 (COL; 4.12). Valle del Cauca: Cordillera Central, Cerro de Pan de Azúcar, páramo, 3700 m, 26 February
1969, J. Cuatrecasas 27561 (COL; 4.12). ECUADOR. Azuay: Road Cuenca–Loja road, km 31 from Cuenca outskirts,
3150 m, 03°08’S, 79°02’W, 30 November 1996, G.P. Lewis 2869 (US; 4.12). Bolívar: Cordillera Occidental, high
country at headwaters of Río Gradas, 15 km s. of Chimborazo Volcano, 4000 m, 01°35’S, 78°51’W, 10 January 1945,
F.R. Fosberg 22484 (US; 4.12). Cañar: between Biblián and Cañar, 3250 m, 21 September 1955, E. Asplund 17679 (S;
410). Vicinity of Cañar, near paramo, 15 September 1918, J.N. Rose 1918 (US; 4.12). Carchi: Mira Cantón, 30 km al
oeste de San Isidro, 3000 m, 00°35’S, 78°05’W, 22 July 1990, E. Gudiño 420 (MO; 4.12). Chimborazo: ca 7 kms up
road toward Chimborazo (Mt.) from main road N of Riobamba (ca. 20 kms NW of Riobamba), 3700 m, T.F. Stuessy
12432 (WU; 4.12). Cotopaxi: 25 km W of Pujilí on road to Quevedo, 3850 m, 20 August 1991, T.F. Stuessy 12411
(WU; 4.12). Imbabura: side of Cayambo mountain, 13.500 ft. (c. 4,115m), 12 June 1961, P.C.D. Cazalet 5700 (B;
4.12). Loja: Road to Jimbura to Zumba, páramo de Cordillera de Las Lagunitas km 19, 3380 m, 04°43’34”S,
79°26’00”W, 06 November 2000, P.M. Jørgensen 2251 (WU; 4.12). Napo: Rd from Quito to Baeza, just at the pass.,
3000 m, 22 February 1992, V. Funk 11026 (US; 4.12). Pichincha: Cerro Pichincha, up from Quito, on road to microondas
towers above city, 3700 m, T.F. Stuessy 12417 (WU; 4.12). Tungurahua: Near Calamaca, app. 20 km W of Ambato
along old road Amabato-Guaranda, 3400–3900 m, 01°16’S, 78°48’W, 22 June 1985, S. Laegaard 54562 (MO; 4.12).
PERU. Amazonas: along road Jumbilla—Molinopampa until ca 1 ½ hours by car from Jumbilla, 2500–3000 m,
06°04’44’’ S, 77°39’47’’W, 07 November 2012, H. van der Werff 25304 (MO; 4.17). Ancash: collected near road Km.
25 towards Carpa, 4260 m, 24 August 1977, K.A. de Mayolo 332 (US; 4.16.2). Arequipa: Arequipa, South slopes of
Nevado Chachani, on the road to Sumbay, ca 10 km north of Arequipa, 3500 m, 30 November 1964, P.C. Hutchison
7234 (K; 4.16.3). Ayacucho: amostra comercial comprada no mercado “Central de Huamanga”, January 2015, G.
Ccapatinta 24 (SPFR; 4.16.2). Huanta, north of Huanta, near Huaillay, 4900–4000 m, 13–14 March 1926, A. Weberbauer
7593 (US; 4.16.4). Cajamarca: Celendín, Jalca de Jumulca (Ruta a Celendín), 3800 m, 19 August 1984, A. Sagástegui
A. 12268 (MO; 4.17). San Miguel, sobre el desvio a Tongod, em el lugar llamado Toropampa, Antes de Quilcata, 3300
m, 13 September 1991, I. Sánchez Veja 5761 (MO; 4.14). San Miguel, sobre el desvio a Tongod, a 10 km de la Carretera
Cajamarca–Hualgayoc, 3300 m, 06 November 1991, I. Sánchez Vega 6076 (MO; 4.14). San Miguel, 76 kms NNW of
Cajamarca on road toward Hualgayoc, detour to Tongod, 3320 m, 15 July 1992, T.F. Stuessy 12625 (WU; 4.14). Cuzco:
Combepala, 26 January 1969, Soukup 6342/6132 (US; 4.16.2). Junín: entre Carhuamayo y Junín, 4100–4200 m, 09
January 1949, R. Ferreyra 5237 (K; 4.16.2). Tarma, Tarma, Carretera La Oroya, 28 January 2015, G. Ccapatinta 51
(SPFR; 4.16.2). Huancayo, amostra comercial comprada no mercado “Modelo”, January 2015, G. Ccapatinta 22
MORPHOLOGICAL DIVERSITY IN SPINY SUNFLOWERS Phytotaxa 518 (2) © 2021 Magnolia Press • 125
(SPFR; 4.16.3). Huancayo, road from Huancayo to Huancavelica, 3543 m, 12°23’38’’ S, 74°54’15’’W, 16 September
2001, M. Weigend 5807 (B; 4.16.4).
Huacanvelica: Ccaullapa, a 5 km. O. de Comaica, 4100–4150 m, 18 March 1951, O. Tovar 230 (US; 4.16.2).
Huacanvelica, Amostra comercial comparada no mercado de “Abastos”, January 2015, G. Ccapatinta 32 (SPFR;
4.16.4). Tayacaja, quebrada, 3 km north of Salcabamba Village, 3500 m, 09 January 1939, H.E. Stork 10329 (K; 4.17).
Huánuco: Ambo, Tomayquichua, camino de Estanco, aos arredores de lagoa, 27 January 2016, G. Ccapatinta 48
(SPFR; 4.15). Huanuco, amostra comercial comprada no mercado “Modelo”, January 2015, G. Ccapatinta 17 (SPFR;
4.16.2). Quina, October 1927, M. Sawada P84 (US; 4.16.4). Huaraz: Recuay, Cátac, a caminho de Pastoruri, 4206 m,
09°53’02.3’’ S, 77°17’36.1’’W, 25 January 2015, G. Ccapatinta 46 (SPFR; 4.16.2). La Libertad: Sanchez Carrión,
señal Huaylliyas, NW-facing, 4200 m, 07°53’S, 78°02’W, 21 August 1982, D.N. Smith 2269 (MO; 4.17). Uuncha, fila
de Andosa, 3802 m, 07°06’52’’ S, 77°49’30’’W, 12 November 2013, C.V. Ocaña 45 (MO; 4.17). Lima: Lima, amostra
comercial comprada no mercado “La Parada”, January 2015, G. Ccapatinta 15 (SPFR; 4.16.2). Pasco: Cerro de Pasco,
amostra comercial comprada no mercado “Central”, G. Ccapatinta 26 (SPFR; 4.16.2). Piura: Huacabamba, trilha para
as lagoas de Las Huaringas, January 2015, G. Ccapatinta 10 (SPFR; 4.12). Puno: Azángaro, Watasani, 4100 m, 04
April 1959, C. Vargas 812539 (US; 4.12).
5. Dasyphyllum Kunth (1820: 17)
Type:—Dasyphyllum argenteum Kunth
Shrubs, trees or woody vines, up to 15 m tall, commonly mutistemmed. Stems erect, decumbent or scandent, much-
branched, lenticelate, cylindrical, with or without scales imbricate at base, glabrous, villous, sericeous, tomentose or
velutinous, unarmed or with axillary spines in pairs or fascicles, straight or curved, convergent or divergent, glabrous
or sparse pubescent at base and becoming glabrous towards the apex. Leaves alternate, spiral, rarely in fascicles,
subsessile or petiolate, persistent or deciduous, blade narrow elliptical, obovate or orbiculate, chartaceous or coriaceous,
pale or lustrous, glabrous or pubescent, villous, tomentose, lanose in both surfaces, base acute or obtuse, margin flat
or revolute, glabrous or cilate, apex unarmed, mucronate or spiny; venation actinodromous with 3 or more basal or
suprabasal main nerves. Capitulescence terminal or axillary, monocephalous, racemose, paniculiform, corymbiform
or umbelliform. Capitula 5–60-flowered, homogamous, discoid, sessile to pedunculate, involucre cylindrical to
campanulate, 6–14-seriate, phyllaries scarious or commonly coriaceous, green, yellow-golden, brown or black, erect
or reflexed, glabrous or densely villous, ovate—triangulate grading to lanceolate, apex unarmed, mucronate or spiny,
margin flat, ciliate. Receptacle flat, pilose. Flowers isomorphic, bisexual or pistilate by the gynoecium atrophy, corolla
tubular (5+0), ligulate (5+0), subbilabiate (4+1), or bilabiate (3+2), 5-lobed, white to cream, externally glabrous or
sericeous, internally sericeous. Anthers 5, apical appendage bilobed, basal appendage lanceolate, acute to rounded,
caudate or ecaudate, calcarate, inserted at the base or between the base and the throat, filaments free, glabrous. Style
cylindrical, cream to yellow. Cypsela cylindrical or fusiform, densely villous or rarely glabrous. Pappus plumose,
shorter than or equal to corolla length, bristles whitish. Pollen with or rarely lacking intercolpal depressions, sparsely
microechinate (Suessy & Urtubey 2007). Chromosome number = 27 (Watanabe et al. 2007).
Fig. 6; 10a.
Distribution and habitat:—Dasyphyllum is the largest genus of Barnadesioideae, comprising 27 species. The
last revision of the genus was by Saavedra (2011), who recognized 33 species—two of them were later transferred to
Archidasyphyllum (Ferreira et al. 2019, see above), plus four probably new species remain undescribed (not counted
here). Species of the genus are distributed throughout South America in two main centers of diversity: the first, in the
Andes, from Venezuela to northern-western Argentina, occurring in environments such as the Puna (highlands and
steppe, till 3,700 m high) or in dry to tropical to subtropical mixed (deciduous-evergreen) montane forests; and the
second, in Brazil, Bolivia, and Paraguay, in savannas, such as the Brazilian Cerrado, in dry to tropical to subtropical
mixed (deciduous-evergreen) forests, such as the Atlantic Forest, in the Caatinga (Brazilian thorn forest and shrubs), in
the Chaco (dry forests and open woodland), in the Brazilian Campos Rupestres (open vegetation in rocky outcrops), in
gallery forests (Cabrera 1959, Sagástegui 1980, Sagástegui & Dillon 1985, Saavedra et al. 2014, Ferreira et al. 2019,
Saavedra 2020)
Notes:—Dasyphyllum can be distinguished from the other genera by the following characters: arching shrubs,
trees, or woody vines, discoid capitula, cream to white corolla, bisexual or pistilate by the gynoecium atrophy and
bilobed apical appendage.
FERREIRA ET AL.
126 • Phytotaxa 518 (2) © 2021 Magnolia Press
FIGURE 6. Dasyphyllum. A: Dasyphyllum lanceolatum. Capitulescence. B: Dasyphyllum reticulatum. Capitulum. C: Dasyphyllum
sprengelianum. Capitulum. D: Dasyphyllum vagans. Capitulescence. E: Dasyphyllum leiocephalum. Habit. Photos by Carolina Ferreira
(A and D), Mauricio Mercadante (B), Paola Ferreira (C), and Gari Ccana-Ccapatinta (E).
The circumscription of Dasyphylllum presented here correspond to the species of Dasyphyllum subgenus
Dasyphyllum (Dasyphyllum “sensu stricto”) since the other subgenus—Dasyphyllum subgenus Archidasyphyllum—
was erected to the generic rank after Ferreira et al. (2019). Differences between these two groups are discussed above
under Archidasyphyllum.
MORPHOLOGICAL DIVERSITY IN SPINY SUNFLOWERS Phytotaxa 518 (2) © 2021 Magnolia Press • 127
Cabrera (1959) recognized two sections in his Dasyphyllum subgenus Dasyphyllum (Dasyphyllum “senso stricto”):
sections Microcephala (23 species) and Macrocephala (11 species). Saavedra et al. (2018) recognized only six species
in section Macrocephala. The two sections are distinguished by their involucre size and capitula arrangement, with
section Macrocephala having involucre longer than 20 mm in length and arranged in a solitary or small group of
heads and section Microcephala having heads arranged in synflorescence (corymbiform cymes) smaller than 18 mm
in length. Ferreira et al. (2019) investigated the phylogenetic relationships of Dasyphyllum based on molecular data. In
this study, both sections were rejected and further studies are necessary to propose a new infrageneric classification.
Iconography:—Cabrera (1959), Harling (1991), Ferreyra (1995), Saavedra et al. (2014, 2018).
Accepted species (Saavedra 2011):—5.1 Dasyphyllum argenteum Kunth; 5.2 Dasyphyllum armatum (J.Kost.)
Cabrera; 5.3 Dasyphyllum brasiliense (Spreng.) Cabrera; 5.4 Dasyphyllum brevispinum Sagást. & Dillon; 5.5
Dasyphyllum colombianum (Cuatr.) Cabrera; 5.6 Dasyphyllum cryptocephalum (Baker) Cabrera; 5.7 Dasyphyllum
diamantinense Saavedra & M.Monge; 5.8 Dasyphyllum donianum (Gardner) Cabrera; 5.9 Dasyphyllum ferox (Wedd.)
Cabrera; 5.10 Dasyphyllum flagellare (Casar.) Cabrera; 5.11 Dasyphyllum floribundum (Gardner) Cabrera; 5.12
Dasyphyllum fodinarum (Gardner) Cabrera; 5.13 Dasyphyllum hystrix (Wedd.) Cabrera; 5.14 Dasyphyllum inerme
(Rusby) Cabrera; 5.15 Dasyphyllum lanceolatum (Less.) Cabrera; 5.16 Dasyphyllum leiocephalum (Wedd.) Cabrera;
5.17 Dasyphyllum leptacanthum (Gardner) Cabrera; 5.18 Dasyphyllum maria-lianae Zardini & Soria; 5.19 Dasyphyllum
popayanense (Hieron.) Cabrera; 5.20 Dasyphyllum reticulatum (DC.) Cabrera; 5.21 Dasyphyllum retinens (S.Moore)
Cabrera; 5.22 Dasyphyllum spinescens (Less.) Cabrera; 5.23 Dasyphyllum sprengelianum (Gardner) Cabrera; 5.24
Dasyphyllum trichophyllum (Baker) Cabrera; 5.25 Dasyphyllum vagans (Gardner) Cabrera; 5.26 Dasyphyllum varians
(Gardner) Cabrera; 5.27 Dasyphyllum vepreculatum (D.Don) Cabrera.
Selected specimens examined:—ARGENTINA. Misiones: Cainguás, ruta 14, 5 km E de Aristóbulo del Valle, 29
July 1987, R. Vanni 836 (K; 5.3). BRASIL. Bahia: Caraíbas, próximo à cancela, 13°05’S, 41°22’W, 15 August 2005,
E.C. Oliveira 111 (HUEFS; 5.23). Coribe, estrada vicinal partindo da estrada São Félix do Coribe-Coribe, ca. 28km de
São Félix do Coribe, cerca de 4km após o distrito de Colônia do Formoso, entrar a direta por 2km até as margens do
Rio Formoso, 448 m, 13°34’53’’S, 44°19’36’’W, 20 July 2009, M.M. Saavedra 997 (RB; 5.11). Entre Rios, Fazenda
do Rio do Negro, área antropizada, próximo a plantio de eucalipto, 143 m, 12°01’04’’S, 38°02’56’’W, 04 September
2009, M.M. Saavedra 934 (RB; 5.15). Érico Cardoso, localidade da porteira, estrada para Capão, ca. 15 km de Érico
Cardoso, 851 m, 13’18’5’’S, 42’08’31’W, 23 July 2009, M.M. Saavedra 1000 (RB, SPFR; 5.3). Jacaraci, 18 September
2010, J.G. Carvalho-Sobrinho 2864 (RB; 5.8). Mucugê, Capão do Correia, estrada vicinal saindo de Caraíba a 17.5
Km da BA-142, base de campo rupestre com solo pedregoso, 1221 m, 13°6’38’S, 41°22’39’W, 11 July 2009, M.M.
Saavedra 968 (RB; 5.7). Palmeiras, Morro do Pai Inácio, ao redor da trilha iniciada para a torre de transmissão, 1103
m, 12°27’21.6’’S, 41°28’23’’W, 17 May 2013, P.L. Ferreira 09 (SPFR; 5.7). Piatã, Fazenda Dragão Verde, 1318 m,
13°05’45’’S, 41°51’12’’W, 07 July 2017, N. Roque 5012 (ALCB; 5.23). Rio dos Pires, beira do riacho da Forquilha,
1500 m, 13°54’S, 42°29’W, 24 July 1993, W. Ganev 1945 (HUEFS, K; 5.7). São Desidério, assentamento Thainá, ca.
40 km da BA-467, 732 m, 12°35’46’’S, 44°50’28’’W, 16 July 2009, M.M. Saavedra 993 (RB, SPFR; 5.8). Ceará:
Crato, estrada para casa do guarda, Belmonte, 1010 m, 07°16’27’’S, 39°27’03’’W, 26 October 2010, M. Mata 2241
(HUEFS; 5.23). Crato, Estrada Crato-Exu, Carrasco, 964 m, 07.1457 S, 39.2940 W, 13 October 2011, M.I.B. Loiola
1372 (EAC; 5.23). Distrito Federal: Áreas da FERCAL, A 6,7 km da fábrica de cimento da CIPLAN, na DF-250 Leste,
850 m, 26 July1990, L.B. Bianchetti 891 (HUFU; 5.25). Brasília, Bacia do Rio São Bartolomeu, 06 May 1981, E.P.
Heringer 6908 (RB; 5.15). Recanto das EMAS, Núcleo Rural Monjolo, 11 August 2009, A.E. Ramos 1903 (RB; 5.8).
Brasília, condomínio Mini Chácaras, próximo à barragem do Paranoá, 1100 m, 15º48’20’’S, 41º47’32’’W, 05 October
2001, G. Pereira da Silva 5507 (CEN; 5.23). Brasília, Brasilândia, Fazenda do Sr. Pereira, 1200 m, 15º36’19’’S,
48º06’02’’W, 08 October 2015, B.T.P.M. Goes 87 (IBGE; 5.23). Brasília, bacia do Rio São Bartolomeu, 05 August
1981, E.P. Heringer 7323 (MO; 5.3). Espírito Santo: Afonso Cláudio, três pontões, 1018 m, 20º31’28’’S, 41º07’14’’W,
09 September 2007, M.M. Saavedra 520 (RB, SPFR; 5.3). Castelo, Distrito de Forno Grande, localidade Rio Manso,
1480 m, 20º31’28’’S, 41º07’14’’W, 20 September 2008, M.M. Saavedra 825 (RB; 5.22). Castelo, Forno Grande,
localidade Balança, 1250–1600 m, 20º31’37’’S, 41º06’06’’W, 18 July 2008, M.M. Saavedra 796 (RB; 5.10). Várzea
Alegre, mata do Fausto, próximo ao colégio, 27 September 2000, V. Demuner 1305 (RB; 5.15). Goiás: Caiaponia, Rio
Borá, 24 July 1977, G. Hatschbach 40069 (MO; 5.3). Corumbaíba, margem esquerda do rio Corumba, 300 m montante
da alternativa 4, 24 April 1993, S.P. Cordovil 289 (UEC; 5.25). Doverlândia, Conglomerado, GO-561, subunidade 2,
subparcela 1, individuo 11, 12 June 2018, G. Allegretti IFN 522 (CEN; 5.23). Niquelândia, Macedo, ca 18 km de
Niquelândia, 24 August 1994, T.S. Filgueiras (RB; 5.8). Piranhas, Serra do São João, estrada para a cachoeira São
Domingos a partir da zona urbana de Piranhas, 35.6 km da zona urbana, 748 m, 16º35’44.5’S, 51º55’57.7’W, 06
November 2012, B. Loeuille 770 (SPF; 5.23). Pirenópolis, Parque Estadual dos Pirineus, 15 km da base da Serra.
FERREIRA ET AL.
128 • Phytotaxa 518 (2) © 2021 Magnolia Press
Cerrado com campo rupestre, afloramentos de quartzito, 1273 m, 15º48’8’S, 48º51’11’W, 19 July 2007, M.M. Saavedra
425 (RB; 5.8). Mato Grosso: Chapada dos Guimarães: road to radarstation, 11 July 1994, B. Dubs 1502 (K; 5.21).
Nobres, cerca de 30 km NE, em linha reta, BR-242, ca. 2 km do entroncamento com a linha BR 364/163, crista da serra
do Caixa Furada, 14°42’ S, 56°11’W, 19 May 1997, V.C. Souza 16463 (ESA; 5.23). Mato Grosso do Sul: Três Lagoas,
Km 85 da estrada para o alto Sucuriu, Fazenda Barreirinho, 24 July 1983, F. Barros 858 (UEC; 5.25). Minas Gerais:
Alto do Caparaó, Parque Nacional campo de altitude ao lado da trilha, 2200 m, L.S. Leoni 3970 (RB; 5.10). Araporã,
Reserva vegetal da usina Alvorada, 07 June 2001, A.A. Arantes s.n. (RB; 5.25). Baependi, Toca dos Urubus, face
sudeste da serra de Sta. Maria, 08 October 2006, F.M. Ferreira 1167 (RB; 5.15). Botumirim, no barranco à beira da
estrada de retorno da Serra do Botumirim, próximo a cidade, 1021 m, 16°51’59’’S, 43°00’52’’W, 10 July 2007, M.M.
Saavedra 578 (RB, SPFR; 5.24). Botumirim, trilha para o Cruzeiro, na entrada da cidade, pelo acesso a partir de
Cristália, 16°51’26’’S, 43°00’46’’W, 07 July 2001, P. Fiaschi 851 (SPFR; 5.24). Brazilândia de Minas-MG, 17°04’24’’S,
45°54’16’’W, 11 July 2000, J.A. Lombardi 3963 (SPF; 5.3). Cata altas, RPPN Santuário do Caraça, trilha para o tanque
grande, 1166 m, 20°06’10’’S, 43°29’38’’W, 26 April 2010, L.L. Giacomin 1139 (RB; 5.12). Cristália, Morro da Antena
de Televisão, 1200 m, 16°43’28’’S, 42°55’42’’W, V.C. Souza 25816 (ESA; 5.24). Diamantina, Estrada Diamantina-
Datas, próximo ao km 602 da estrada, 1367 m, 18°18’26’’S, 43°41’04’’W, 26 May 2009, M.M. Saavedra 888 (RB;
5.20). Felício dos Santos, APA Felício dos Santos, Interior de Mata semidecidua, 27 August 2008, N.F.O. Mota 1414
(RB; 5.12). Jaboticatubas, PARNA Serra do Cipó, cachoeira da farofa, portaria do retiro, coletas realizadas ao longo
da encosta após a bifurcação para a cachoeira da farofa até o riacho, 883 m, 19°20’56’’S, 43°34’38’’W, 16 October
2013, M. Verdi 6717 (SPFR; 5.20). Lima Duarte, Distrito de Conceição do Ibitipoca, Serra do Ibitipoca, estrada para o
Parque Estadual do Ibitipoca, próximo a Pousada dos Manacas, 1370 m, 21°42’32’S, 43°54’12’W, 09 May 2002, R.
Marquete 3258 (RB; 5.10). Mato Verde, estrada Mato verde—Rio Pardo de Minas, ca de 8 km de entrocamento Rio
Pardo/Mato verde/Mentezuma em direção a Rio Pardo, 1030 m, 15°26’08’’S, 42°39’36’’W, 20 July 2005, R.C. Forzza
4110 (RB, SPFR; 5.8). Ouro Preto, Serra de Ouro Preto, Fazenda brigida, 1538 m, 20°21’51’’S, 43°31’02’’W, 17 May
2011, R.C. Forzza 6343 (RB; 5.12). Santa Bárbara, Serra do Caraça, estrada para a Capelinha, 23 May 1983, J.R.
Pirani 716 (SPF; 5.24). Santana do Pirapama, Serra do Cipó, acesso pela fazenda Inhame, afloramento rochoso e mata
seca ao norte da fazenda Inhame, 750 m, 18°55’54’’S, 43°48’33’’W, 14 March 2009, D.C. Zappi 2124 (RB; 5.25).
Santana do Riacho, estrada para Conceição do Mato Dentro (MG 10), lado esquedo do morrote, 940 m, 19°18’31.8’’S,
43°36’23’’W, 23 August 2006, R. Marquete 3806 (RB; 5.20). Santana do Riacho, morro de calcário na base da Serra
do Cipó, 24 May 2009, M.M. Saavedra 869 (RB; 5.25). Santana do Riacho, Serra do Cipó, Morro da Pedreira, 3° grupo
de calcário, 935 m, 19°18’18.4’’S, 43°36’47.9’’W, 17 June 2007, M. Groppo 1452 (SPFR; 5.23). São Roque de Minas,
Parque Nacional da Serra da Canastra, 1307 m, 20°15’13.7’’S, 46°24’17.2’’W, 29 October 2016, P.L. Ferreira 57
(SPFR; 5.23). Paraná: Arapoti, Rio das Perdizes, 06 April 1970, G. Hatschbach 24104 (RFA; 5.10). Bituruna, Faxinal,
12 February 1966, G. Hatschbach 13853 (MBM; 5.22). Campina Grande do Sul, Serra do Capivari, 14 July 1986, J.M.
Silva 121 (MBM; 5.22). Curitiba, Jardim das Américas, próximo à fábrica da Coca-Cola, esquina das ruas S. Saza
Lattes e Dr. Paulo Assumpção, M.M. Saavedra 521 (RB, SPFR; 5.3). Morretes, Zoador, 20–50 m, 21 July 1987, G.
Hatschbach 51270 (WU; 5.3). Pernambuco: Moreilândia, Mata Agreste, 07 August 1999, F.S. Cavalcante 528 (EAC;
5.23). Moreilândia, Serra do Catolé, refúgio das abelhas Professor Paulo Nogueira Neto, 880 m, 7.2751 S, 39.28132
W, V.M. Mascena 54C (EAC; 5.23). Rio de Janeiro: Itatiaia, Sítio Vista linda, 750 m, 22°15’00’’S, 44°34’00’’W, 13
September 1994, M.P.M. de Lima 278 (RB; 5.17). Itatiaia, Parque Nacional do Itatiaia, Três picos, área perturbada,
1200 m, 22°15’00’’S, 44°34’00’’W / 22°28’00’’S, 44°45’00’’W, 37 March 1995, R.R. Guedes 2533 (NY; 5.22). Nova
Friburgo, subida para o Pico da Caledônia, trilha de acesso ao Pico, 2190 m, 22°21’11.56’’S, 42°35’4.83’’W, 20 April
2009, L.L. Giacomin 951 (RB; 5.17). Nova Friburgo, Reserva Ecológica Municipal de Macaé de Cima, estrada para
Sitio Sophronites, 1000 m, 22º00’S, 42º03’W, 03 June 1990 S.V.A. Pessoa 522 (NY; 5.22). Paraty, pico do Cairuçu,
acesso pela Praia Negra, 25 November 1990, C. Farney 2558 (RB; 5.10). Petrópolis, Vale das Videiras, Morro do
Cuca, 1500–1750m, 10 October 1979, G. Martinelli 6148 (RB; 5.6). Petrópolis, Araras, Pico da Maria Comprida,
Vertente NE, 1900 m, 22°24’22’’S, 43°12’62’’W, G. Martinelli 10696 (RB, SPFR; 5.6). Petrópolis, Vale das Videiras,
Morro do Cuca, esqueda da trilha próximo ao topo do morro, 1636 m, 22°24’57.9’’S, 43°16’09.6’’W, 03 May 2013,
P.L. Ferreira 01 (SPFR; 5.17). Petrópolis, entre Araras e Vale das Videiras, morro do cuca, 1050–1200 m, 02 June
1984, G. Martinelli 9884 (RB; 5.10). Reserva Biológica de Araras, Vale das Videiras, Pedra Comprida, 22°25’25’’S,
43°15’58’’W, 28 March 2018, T.M. Portugal 139 (RB; 5.6). Rio de Janeiro, 1874, A.F.M. Glaziou 7697 (NY; 5.25).
Teresópolis, Parque Nacional da Serra dos Órgãos, trilha para a Pedra do Sino, 2118 m, 22°27’54’’S, 43°01’56’’W, 19
November 2006, E.J. Lucas 556 (RB, K; 5.17). Rio Grande do Sul: Bom Jesus, Fazenda do Cilho, 1050 m, 28º31’45’’S,
50º26’37’’W, 25 February 1999, M. Rossato 24 (NY; 5.22). Encruzilhada do Sul, fazenda Xafri, ca. 20 km distante do
centro da cidade, 17 April 2004, C. Giongo 353 (ICN; 5.22). Jaquirana, beira da mata, 800 m, 01 July 1989, R. Wasum
MORPHOLOGICAL DIVERSITY IN SPINY SUNFLOWERS Phytotaxa 518 (2) © 2021 Magnolia Press • 129
s.n. (MO; 5.3). Santa Catarina: Anita Garibaldi, Portão de entrada da UHE Barra Grande, sentido Anita Garibaldi, 1006
m, 27°46’21’’S, 51°10’36’’W, 02 February 2006, E.S.G. Guarino 972 (CEN; 5.22). Lages, 10 April 2008, M. Verdi
22695 (FURB; 5.22). São Paulo: Altinópolis, Morro do Forno, 762 m, 21°05’47.3’’ S, 47°20’01.0’’ W, 18 August 2014,
P.L. Ferreira 34 (SPFR; 5.23). Anhembi, sítio Ribeirão Bonito, 22°48’43.5’’ S, 48°12’51.6’’ W, 22 March 1994, K.D.
Barreto 2194 (SPF; 5.25). Campinas, Santa Genebra Forest Reserve, Barão Geraldo near Campinas, 22º52’S, 46º05’W,
550 m, 26 August 1987, A. Gentry 58680 (MO; 5.3). Campos do Jordão, Horto Florestal, 22º42’31’’S, 45º27’58’’W,
22 September 1993, K.D. Barreto 1240 (ESA; 5.22). Cunha, Parque Estadual da Serra do Mar, picada do rio bonito,
18 August 1994, G.A.D.C. Franco 1242 (HRCB; 5.12). Cruzeiro, estrada da fazenda Boa Vista, 22º29’02’’S,
45º02’00’’W, 05 April 1995, J.L.A. Moreira 32 (UEC; 5.22). Jundiaí, Reserva Biológica Municipal da Serra do Japi,
trilha Base/Mirante/DAE/Rua Bauru/Cidinho/Base, 850–1178 m, 23º13’49’’S, 46º56’8’’W, 24 June 2008, J.A.
Lombardi 7443 (RB; 5.10). Ilha Solteira, Fazenda São José, 05 August 1995, M.R. Pereira-Noronha 1379 (HRCB;
5.26). Itaporanga, Patrimônio do Mosteiro Santa Cruz, 19 July 1949, A.B. Joly s.n. (SPF; 5.25). Ribeirão Preto, Estação
Ecológica de Santa Tereza, 582m, 21°13’30.5’’S, 47°50’57.7’’W, 13 September 2016, P.L. Ferreira 67 (SPFR; 5.15).
São Paulo, cidade jardim, August 1941, W. Hoehne s.n. (SPF; 5.15). Tocantins: Dianópolis, Serra do Ouro, fazenda
Cachoeira do Sr. Martins, 15 July 1963, A.L. Costa s.n. (RB; 5.21). BOLIVIA. Chuquisaca: Padilla, ca. 10 km west of
Padilla, 9 April 1994, J.R.I. Wood 8219 (K; 5.2). Zudañez, on ascent c. 20 km SE of Icla on road to Cordillera de los
Sombreros and Azurduy, 3000 m, 13 March 1999, J.R.I. Wood 14663 (SPF, K; 5.13). Cochabamba: Ayopaya, ca. 1 km
above Independencia in wooded valley above ford and footbridge, 2500 m, 11 March 2000, J.R.I. Wood 16012 (K;
5.2); Pasorapa, ca. 36 km de Pasorapa, y unos 4 km antes de Buena Vista, 2426 m, 18º31’S, 64º33’W, 24 March 2003,
J.R.I. Wood 19461 (K; 5.2). La Paz: Calamarca, on Antiplano, 55 km South of La Paz, 4200 m, 16º59’S, 68º07’W, 28
September 1947, F.R. Fosberg 28698 (MO; 5.9). Larecaja, along road Caranavi-Guanay, outskirts of Guanay, 600m,
05 August 1998, D.C. Wasshausen 2204 (K; 5.21). Trail from Cota Cota to Muela del Diablo, 3500–3800 m, 16º32’S,
68º03’W, 13 April 1982, J.C. Solomon 7473 (MO; 5.9). Potosi: Charcas, Toro Toro, Sendero bajando al Cañon del
Vergel, 2558m, 18º07’S, 65º46’W, 26 February 2003, J.R.I. Wood 19231 (K; 5.16). Cornélio Saavedra, Betanzos area,
in the valley NW of Ticoya, 3500–3700 m, 28 December 1994, J.R.I. Wood 9046 (SPF, K; 5.13). Santa Cruz: Andres
Ibanez, along road from Santa Cruz to Samaipata, 2–4 km SW of Angostura, 700 m, 18º10’S, 63º32’W, 12 July 1987,
M. Nee 35138 (GB; 5.3). Caballero, Comarapa, ca. 2 km E de Comunidad Candelaria, subiendo unos 200m hacia cerro
Empinao, 2275 m, 17º54’S, 64º34’W, 27 March 2004, M. Mendoza 936 (K, NY). Ñuflo de Chávez, ca. 3–4 km SW of
San Javier on road to San Ramon, 500 m, 24 July 2000, J.R.I. Wood 16507 (K; 5.21). Tarija: Tarija, 3000–3500 m, 04
February 1904, K. Fiebrig 3038 (K; 5.13). COLOMBIA. Boyacá: Valle del Cocuy, toward southeast, 2880–3000 m, J.
Cuatrecasas 1735 (P; 5.5). Norte de Santander: Between Mutiscua and Pamplona, 2700–3400 m, 23 February 1927,
E.P. Killip 19690 (US; 5.5). Valle del Cauca: Cordillera Central, vertiente occidental, hoya del río Bugalagrande, Loma
de Barragám, Hacienda de San José, 2580–2600 ft. (c. 786–792m), 15 March 1946, J. Cuatrecasas 20005 (F; 5.5).
Cordillera Central, vertiente occidental, márgenes del río Bugalagrande, entre El Puente y Salinas, 2000 m, 27 March
1946, J. Cuatrecasas 20468 (F; 5.5). ECUADOR. Azuay: 7 kms N of Nabon on road toward Cuenca, 2720 m, 26
August 1991, T.F. Stuessy 12464 (US; 5.1). Km 86 out of Cuenca en route to Loja, 3000 m, 25 September 1959, B.
Maguire 44308 (GB; 5.1). Between Oña and Cuenca, 2700–3300 m, 9–10 September 1923, A.S. Hitchcock 21589 (US;
5.1). Vicinity of Nabón, 26 September 1918, A. Pachano 23055 (US; 5.1). Carchi: El Angel, Carretera entre San Isidro
y Chulte, 3200 m, 00°36’S, 78°02’W, 08 April 1991, W. Palacios 7016 (MO; 5.19). Imbabura: Selva Alegre-Otavalo,
W side of pass, 2800–3000 m, 00°15’S, 78°25’W, 04 December 1988, P.M. Jørgensen 65606 (QCA; 5.19). Loja: on
road from Loja to Cuenca, 2900 m, 01 October 1961, C.H. Dodson 858 (MO; 5.1). Loja-Gonzanama, 8 km SW of
Gonzanama, 1 km along track from sign to PREDESUR, 1750 m, 04°14’9.09’’S, 79°28’5.91’’W, 13 May 1997, G.P.
Lewis 3277 (QCA; 5.19). Pichincha: Camino al Hospital entre el Mauca Quito y el Sincholagua, 2400–2750 m,
00°05’S, 78°30’W, 21 May 1988, C.E. Cerón 3769 (MO; 5.19). Ca 2 kms W of Calacali on dirt rd to Nebli, Just outside
Reserva Pululahua, 3020 m, 14 August 1991, T.F. Stuessy 12308 (WU; 5.19). Piura: Ayabaca, 3.6 kms W of Arreipite
Norte, 16.6 kms W of Ayabaca, 21 July 1992, T.F. Stuessy 12700 (WU; 5.19). PARAGUAY. Amambay: Parque Nacional
Cerro Corá, cerro Trébol, table mountain, 22°37’10’’S, 56°02’36’’W, 13 June 1996, E.M. Zardini 45013 (MO; 5.23).
Cordillera: in regione lacus Ypacaray, E. Hassler 11862 (MO; 5.26). Guairá: Mbocayaity-Melgarejo, cerro Naville,
25°43’S, 56°21’W, 06 July 1992, E.M. Zardini 32470 (MO; 5.18). Paraguarí: Pirajú, 25°30’S, 57°15’W, 15 May 1984,
W. Hahn 2572 (MO; 5.3). Ybicui, Parque Nacional Ybycu’i, alrededores de la administración, 15–20 m,15 August
1984, L. Pérez 257 (MO; 5.3). San Pedro: Laguna Blanca, 23°48’46’’S, 56°16’39’’W, 01 August 1996, E.M. Zardini
45480 (MO; 5.18). PERU. Ancash: Sihuas, road from Sihuas to Yanac, before Pomabamba turnoff, 3500–3800 m, 10
March 2001, M. Weigend 5078 (B; 5.9). Huaraz, 10 km by road from Cachabamba, 2870 m, 09°27’S, 77°51’W, 6–8
June 1985, D.N. Smith 10850 (MO; 5.9). Recuay, en ladera abierta, 3300 m, 25 May 1970, A. Lopez M. 7488 (MO;
FERREIRA ET AL.
130 • Phytotaxa 518 (2) © 2021 Magnolia Press
5.9). Apurimac: Andahuaylas: 2 km south of chincheros, 26 February1939, H.E. Stork 10737 (K; 5.16). Cajamarca:
Contumazá, 16 kms S of Contumazá, 2530 m, 17 July 1992, T.F. Stuessy 12689 (WU; 5.4). Desvío al Bosque de Cachil
(Cascas-Contumazá), 2200–2300 m, 25 June 1982, A. López M. 9136 (F; 5.4). Cajamarca, Namora, Ladera oeste que
converge a La Laguna San Nicolás, 2800 m, 05 March 2002, I. Sánchez Veja 11276 (US; 5.13). Cusco: Canchis,
Sicuani, beira da Estrada a caminho de Uyumiri, 06 February 2016, G. Ccapatinta 58 (SPFR; 5.9). Urubamba,
Urubamba, trilha para a lagoa Yanacocha, 3372 m, 13°18’04.4’’S, 72°03’08.4’’W, 10 January 2016, G. Ccapatinta 55
(SPFR; 5.16). km 34, road between Cusco and Urcos (near town of Huaro), along Río Urubamba, 3100 m, 25 May
1977, J.C. Solomon 3091 (MO; 5.9). Vilcabamba, Chuanquiri camino a resistencia, 1000 m, 12°50’30’’S, 73°09’10’’W,
19 October 2006, J. Farfán 1117 (MO; 5.3). Junín: Tarma, 3200 m, 27 January 1968, H.W. Schwabe s.n. (B; 5.9). La
Libertad: Otuzco, Abajo de José Balta, ruta hacia Agallpampa, 2540 m, 09 June 1994, S. Leiva G. 1133 (GB; 5.9).
VENEZUELA. Anzoátegui: forested rocky slopes along Río Querecual southwest of Bergantín, 500 m, 14 March
1945, J.A. Steyermark 61489 (US; 5.27). Bolívar: Pira, Isla en el Lago de Guri, Sector Danto Machado, 40 km al S de
la Presa R. Leoni, 270 m, 07°35’S, 62°58’W, 24 February 1992, G. Aymard 10166 (NYBG; 5.27). Distrito Capital:
lado norte de la Cordillera de La Costa, Osma, 7 km de distancia aerea al este de Los Caracas, a lo largo del río, 200
m, Mayo 1992, W. Meier 2107 (B; 5.27).
6. Doniophyton Weddell (1855: 7)
Type:—Doniophyton anomalum (D. Don) Kurtz
Subshrubs, up to 40 cm tall. Stems erect or decumbent, much-branched, lenticelate, cylindrical, scales imbricated at
base, tomentose, velutinous, axillary spines in fascicles, straight, divergent or convergent, glabrous or rarely pubescent
at base. Leaves alternate, sessile, persistent or deciduous, blade linear-lanceolate, chartaceous or coriaceous, pale or
lustrous, glabrous or tomentose on both surfaces, base truncate, leaf margin revolute or plicate, ciliate, abaxial with
prominent midvein, apex spiny; nervation hyphodromous. Capitulescence terminal, monocephalous. Capitula 40–135-
flowered, heterogamous, disciform, sessile or shortly pedunculate, involucre hemispherical or campanulate, 4–7-seriate,
phyllaries scarious, yellow or yellow and purple, erect or reflexed, hirsute or velutinous, lanceolate grading to linear,
apex spiny, margin flat, ciliate. Receptacle flat or convex, alveolate or tuberculate, pubescent. Flowers heteromorphic.
Ray flowers 10–40, pistillate, corolla narrowly tubular (5+0), 5-lobed, yellow, villous. Style cylindrical, yellow. Disc
flowers 30–95, bisexual, corolla tubular (5+0), 5-lobed, yellow, villous. Anthers 5, apical appendage acute or rarely
apiculate, basal appendage acute, ecalcarate, caudate, inserted at base of the corolla, filaments free, glabrous. Style
cylindrical, yellow. Cypsela turbinate, densely villous. Pappus plumose, shorter than or equal to corolla length, bristles
whitish. Pollen without intercolpal depression, scabrate-microechinate. (Suessy & Urtubey 2007). Chromosome
number = 24, 25 (Wulff 1990).
Fig. 7a–b; 10d.
Distribution and habitat:—Doniophyton comprises two species found in dry open areas from northern Chile to
Patagonian Argentina up to an altitude of 4,000 m (Katinas & Stuessy 1997). The species occur in the Puna vegetation
(highland and steppe) and in the Patagonian steppe and semidesert.
Notes:—Doniophyton is a xeromorphic genus that has always been proposed as being closely related to Chuquiraga
and Duseniella by sharing drier habitats, long caudate anthers, yellow corolla, and pollen without intercolpal depression.
However, this group has long been enigmatic, since previous phylogenetic studies have not resolved the relationships
among these genera, recovering Doniophyton as either nested in Chuquiraga (Gruenstaeudl et al. 2009, Padin et
al. 2015b), or as sister to Chuquiraga (Gustaffson et al. 2001). Morphologically, Doniophyton can be distinguished
from Chuquiraga by the subshrub habit (vs. shrub), chartaceous leaves (vs. coriaceous), heterogamous and disciform
capitula (vs. homogamous and discoid capitula), and female marginal flowers (vs. all flowers in the capitula are
hermaphroditic), and from Duseniella by its chartaceous or coriaceous leaves (vs succulent leaves in Duseniella), by
its perennial subshrubs habit, up to 40 cm tall (vs annual herbs up to 10 cm in Duseniella) and by the pappus scaly
overlapped (vs a plumose pappus in Duseniella).
Iconography:—Katinas & Stuessy (1997).
Accepted species:—6.1 Doniophyton anomalum (D. Don) Kurtz; 6.2 Doniophyton weddellii Katinas & Stuessy.
Selected specimens examined:—ARGENTINA. Chubut: Laguiñeo, 12 December 1981, A.L. Cabrera 33104
(SI; 6.1). 96 kms E of Sarmiento on rte 26 to Comodoro Rivadavia (ca. 56. kms W of Comodoro Rivadavia), 600 m,
T.F. Stuessy 12939 (WU; 6.1). Jujuy: Cochinoca, Abra pampa, 20 January 1976, A.L. Cabrera 27396 (MO, SI; 6.2).
La Pampa: Utracán, General Acha, 05 November 1953, A. Burkart 19219 (SI; 6.1). Mendoza: Malargue, camino a la
MORPHOLOGICAL DIVERSITY IN SPINY SUNFLOWERS Phytotaxa 518 (2) © 2021 Magnolia Press • 131
mina la Valenciana, 20 January 1982, A.L. Cabrera 33461 (SI; 6.1). San Rafael, hotel Termas del Sosneado, 2180 m,
34°46’12”S, 70°03’33’‘W, 22 November 2010, F.O. Zuloaga 12385 (SI; 6.1). Puente del Inca, 2700 m, 20 December
1946, B. Sparre 1515 (K). Neuquén: Collon Curá, RN 40 between bridge over Embalse Piedra de Aguila and Collon
Curá, 640 m, 40°21’27”S, 70°39’46’‘W, 13 January 2002, M. Weigend 5871 (B; 6.1). Zapala, 01 November 1925, H.F.
Comber 111 (K; 6.1). Río Negro: General Roca, Vicinity of General Roca, 250–360 m, September 1914–February
1915, W. Fischer 7 (MO; 6.1). Ñorquinco, entre Ruta Ruin y Manuel Chique, 11 December 1981, A.L. Cabrera 33045
(SI; 6.1). Santa Cruz: Deseado, Caleta Olivia, 250 m, 20 November 1929, A. Donat 127 (MO; 6.1). Ruta 40, 38 km
al ESSE de la intersección com ruta 37, 3 km al sur de Ruta 40, 47°58’23.3”S, 70°58’45.8’‘W, 18 January 2002, M.
Bonifacino 481 (LP; 6.1). San Juan: Iglesia, quebrada del Agua Negra, 3800 m, 10 January 1976, A.L. Cabrera 27070
(SI; 6.2). Pie de Palo, then 2 kms N toward the hills and sand dunes (Médanos), 600 m, T.F. Stuessy 12887 (WU; 6.1).
Calingasta, Manatiales, 27 March 1971, Zardini 175 (MO; 6.2). CHILE. Atacama: Copiapó, Rio Turbío, Co. Cadillal,
3.000 m, January 1926, E. Werdermann 953 (K; 6.2). Vallenar, Vicinity of Laguna Valeriano, 4.000 m, 29°03’S,
69°52’W, I.M. Johnston 6073 (K; 6.2). Coquimbo: Bãnos del Toro, 3500 m, December 1923, E. Werdermann 201 (K;
6.2). Elquí, Baños del Toro, 3200–3300 m, 05 February 1939, J.L. Morrison (K; 6.2). Cordillera Doña Ana, Cancha de
Sky, 3400 m, 29°51’S, 70°03’W, 06 January 1988, F.A. Squeo 88001 (MO; 6.2). 21.4 kms N of Junta del Toro (aduana
chilena) on road to Baños del Toro, 3100 m, 20 January 1993, T.F. Stuessy 12780 (WU; 6.2).
FIGURE 7. Doniophyton and Duseniella. A–B: Doniophyton. C–D: Duseniella. A: Doniophyton anomalum. B: Doniophyton weddellii.
C–D: Duseniella patagonica. Photos by Darwinion Institute.
FERREIRA ET AL.
132 • Phytotaxa 518 (2) © 2021 Magnolia Press
7. Duseniella Schumann (1902: 475)
Type:—Duseniella patagonica (O.Hoffm.) K.Schum.
Annual herbs, up to 10 cm tall, unarmed. Stems erect, much-branched, lenticelate, cylindrical or flat, scales imbricated at
base, sparsely sericeous. Leaves opposite at base, alternate in the upper parts, sessile, persistent, blade linear, succulent,
lustrous, adaxially glabrous or sparsely sericeous, adaxially sericeous, leaf margin flat, ciliate, apex mucronate;
venation actinodromous, with 3 basal nerves, not evident and immersed in the mesophyll. Capitulescence terminal,
monocephalous. Capitula heterogamous, disciform, 9–41-flowered, sessile, surrounded by subinvolucral bracts,
involucre campanulate, 4–5-seriate, phyllaries scarious, erect, glabrous, ovate–oblong grading to linear, apex spiny,
margin flat, ciliate. Receptacle convex, glabrous. Flowers heteromorphic. Ray flowers 4–16, pistillate, corolla tubular
(5+0), 5-lobed, yellow, villous at apex. Style cylindrical, yellow. Disc flowers 5–25, bisexual, corolla tubular (5+0),
5-lobed, externally sericeous at base of corolla and apical lobes, internally sericeous at tube margin petals. Anthers 5,
apical appendage lanceolate or obtuse, basal appendage acute, caudate, calcarate, inserted between the base and the
throat, filaments free, glabrous. Style cylindrical, yellow. Cypsela cylindrical to turbinate, densely villous. Pappus
scaly lanceolate, overlapped, shorter than the corolla tube, sericeous, ciliate. Pollen without intercolpal depression,
microechinate (Suessy & Urtubey 2007). Chromosome number 2n = 54 (Baeza et al. 2019).
Fig. 7c–d; 10e.
Distribution and habitat:—Duseniella is monotypic, represented only by D. patagonica, which is endemic to
xeric areas in Argentinean Patagonia and Monte vegetation up to an altitude of 1,500 m. It occurs in semi-desert and
steppe vegetation in Patagonia.
Notes:—Duseniella is a morphologically distinctive genus in the subfamily Barnadesioideae, being the only
genus of annual herbs with unarmed branches, scucculent leaves, and disciform capitula with pappus scales overlapped
and ciliate. Because of its distinctive morphology, Cabrera (1959, 1961) did not recognize the genus as a member of
the subtribe Barnadesiinae (tribe Mutiseae); instead, he placed Duseniella in the subtribe Gochnatiinae (Cabrera 1977).
Phylogenetically, Duseniella is recovered in a clade with Doniophyton and Chuquiraga, but its relationship to these
two genera is still unclear (see discussion above under Doniophyton). A phylogenomic study including a large number
of taxa from this clade could provide better insights into the evolution of this group.
Iconography:—Flora del Conosur available at http://www.darwin.edu.ar/Proyectos/FloraArgentina/
DetalleEspecie.asp?forma=&variedad=&subespecie=&especie=patagonica&genero=Duseniella&espcod=16513
Accepted species:—7.1 Duseniella patagonica (O. Hoffm.) K. Schum.
Selected specimens examined:—ARGENTINA. Chubut: Escalante, Pico Salamanca, 21 January 1965, E. De
Marco de Kreibohm 223 (SI). Escalante, Comodoro Rivadavia, punta Bargia, 08 February 1903, M.S. Pennington
128 (SI). Ruta 20, 20 km al este de Sarmiento, 45°38’ 33.4”S, 68°57’01.0’‘W, 14 January 2002, J.M. Bonifacino
0437 (US). Neuquén: Picún Leufú, 08 November 1972, E.M. Zardini 142 (MBM). Río Negro: Aguada Cecilio, 20
November 1980, Z.A. Manuel s.n. (SI). General Roca and vicinity, 250–360 m, 16 November 1914, W. Fischer 173
(MO). Camino de Cinco Saltos a Lago Pellegrini, E.M. Zardini 149 (MBM). General Roca, Labio S del Valle, frente
a General Roca, 15 km, 16 February 1944, E.G. Nicora 4061 (SI). Santa Cruz: La Angostura, 300m, 13 January 1930,
A. Donat 238 (MO). Corpen Aike, Laguna Grande hacia Gregores, 49°34’S, 70°14’W, 24 January 1967, O. Boelcke
12772 (SI). Deseado, 35 km de Jaramillo caminho al bosque petrificado, 10 January 1967, O. Boelcke 12169 (SI).
8. Fulcaldea Poiret (1817: 375)
Type:—Fulcaldea laurifolia (Bonpl.) Poir.
Arching shrubs or small trees, up to 10 m tall. Stems erect, much-branched, lenticelate, cylindrical or flat, scales
imbricate at base, glabrous, pubescent or rarely strigose, unarmed or with axillary spines in pairs, straight, convergent
or divergent, glabrous or rarely pubescent at base. Leaves alternate, spiral, subsessile or petiolate, persistent or
deciduous, blade elliptic or ovate, coriaceous, pale or rarely lustrous, glabrous, base acute or obtuse, margin flat,
glabrous, apex acute, mucronate; venation eucamptodromous or actinodromous with 3 basal nerves. Capitulescence
terminal or axillary, in corymbose or paniculate cymes. Capitula 1-flowered, homogamous, sessile or subsessile,
involucre narrow cylindrical to cylindrical, 5–13-seriate, phyllaries scarious, green or pale brown, apex purple, erect or
reflexed, glabrous or villous, ovate–triangulate grading to lanceolate, apex mucronate, margin flat, ciliate. Receptacle
convex, glabrous, or pilose. Flowers bisexual, corolla tubular (5+0), 5-lobed, white, red, purple, externally densely
MORPHOLOGICAL DIVERSITY IN SPINY SUNFLOWERS Phytotaxa 518 (2) © 2021 Magnolia Press • 133
villous. Anthers 5, apical appendage lanceolate, or obtuse, basal appendage decurrent, ecaudate, ecalcarate, inserted
between the base and the throat, filaments free, glabrous. Style cylindrical but with a swollen portion below the
branching point, white to cream or purple, apex purple. Cypsela cylindrical, densely sericeous. Pappus plumose,
longer than the corolla length, white, pink or red bristles. Pollen without intercolpal depressions, spinulose (Suessy &
Urtubey 2007). Chromosome number unknown.
Fig. 8a; 10f.
FIGURE 8. Fulcaldea and Huarpea. A: Fulcaldea. B–E: Huarpea. A: Fulcaldea stuessyi. Inflorescence with a white arrow showing
swollen style below the branching point. B: Huarpea andina. Habit. C: Capitulum showing five ray flowers and a disc flower. D: Lateral
view from the capitula. E: Dissected capitula showing three ray flowers (white) and a disc flower (yellow). Photos by Ivan Abreu (A) and
Darwinion Institute (B–E).
Distribution and habitat:—Fulcaldea comprises two species with a remarkable 4,000 km disjunct distribution
(Funk & Roque 2011). Fulcaldea laurifolia (Bonpl.) Poir. is restricted to the dry forest in intermontane regions of
southern Ecuador and northern Peru (Ferreyra 1995), and Fulcaldea stuessyi is restricted to seasonally deciduous
forest in northeastern Brazil in the Chapada Diamantina rocky chain (Funk & Roque 2011).
Notes:—Fulcaldea is easily distinguished from the other Barnadesioideae genera by having a single-flower
capitulum, and by the swollen style below the branching point (Fig. 8a). The genus is part of a strongly supported clade
together with Archidasyphyllum and Arnaldoa, being sister (also with strong support) to the latter according to Ferreira
et al. (2019); however, this clade is morphologically diverse and synapomorphies that support the relationships among
the genera are still to be determined (Funk & Roque 2011, Ferreira et al. 2019). Funk & Roque (2011) hypothesized
that the remarkable disjunction distribution (4,000 km) may be the result of vicariance or long-distance dispersal, thus
being inconclusive on this subject.
Iconography:—Ferreyra (1995), Roque & Funk (2011).
Accepted species:—8.1 Fulcaldea laurifolia (Bonpl.) Poir.; 8.2 Fulcaldea stuessyi N. Roque & V.A. Funk.
Selected specimens examined:—BRASIL. Bahia, Chapada Diamantina, Rio de Contas, distrito de Arapiranga,
Fazenda Brandão, 5 August 2010, I.S. Abreu 123 (ALCB; 8.2). Rio das Contas, antiga estrada do Brandão, sentido do
sítio Marcela, 716 m, 13°28’11’’S, 41°46’32’’, July 2014, N. Roque 4386 (ALCB; 8.2). ECUADOR. Loja: Catacocha
FERREIRA ET AL.
134 • Phytotaxa 518 (2) © 2021 Magnolia Press
to Loma Quemada, km 8.5, 1600 m, 04°06’95”S, 79°36’312’‘W, 16 April 1996, G.P. Lewis 2247 (MO, K; 8.1).
Sozoranga, km 4 along track from Sozoranga-Macará road to Reserva natural El Tundo, propriedade de fundación
ARCOIRIS, 1850 m, 04°19’S, 79°49’W, 19 August 1997, G.P. Lewis 3497 (MO; 8.1). Sozoranga outskirts, 1 km along
track to Utuaña, 1700 m, 04°20’S, 70°47’W, 5 March 1977, G.P. Lewis 3038 (MO, K; 8.1). Km 2 road Sororanga-
Yaramine, 1750 m, 04°18’S, 79°48’W, 17 June 1997, B.B. Klitgaard 203 (QCA, K; 8.1). Manabí: Jama, 24 km south
of Pedernales (as the crow flies) northwest off coastal highway; 1.5 km east of Edualdo (Don Lalo; 8.1) Loor’s farm,
30–150 m, 00°04’31’’S, 80°09’22’‘W, 9 October 1999, T. Delinks 399 (MO; 8.1). Jama Cantón, Cerro del Matal,
3km northwest of Jama, 1 km inland from seashore, 100 m, 00°11’S, 80°18’W, D. Neill 11621 (MO; 8.1). Machalilla
National Park, 01°39”S, 80°41’W, C. Josse 1063 (GB, QCA; 8.1). Pedernales, Estación Biológica Lalo Loor, a 22
km al sur de Pedernales por la carretera costera, 2 km de la costa, 150 m, 00°05’07”S, 80°09’05’‘W, October 2005, J.
Cevallos 213 (MO; 8.1). San Vicente, thicket, 07 June 1955, E. Asplund 16595 (K; 1.1). PERU. Piura: Ayabaca, on road
to Ayabaca, 18 km above Puente Tandopa (Río Quiroz), 1700 m, P.C. Hutchison 6685 (MO, K; 8.1). Ayabaca, 12.7
kms up from Puente Tandopa over Rio Quiroz, 1.540 m, 20 July 1992, T.F. Stuessy 12693 (WU; 8.1). Huacabamba,
bajo de Cachaque, 1100–1200m, 02 May 1955, R. Ferreyra 10932 (MO; 8.1). Paimas, Ayabaca road, ca. 15 km W of
Ayabaca, 1900 m, 20 September 1991, A.H. Gentry 74930 (MO; 8.1).
9. Huarpea Cabrera (1951: 129)
Type:—Huarpea andina Cabrera
Rhizomatous subshrubs, up to 4.5 cm tall, unarmed; underground system formed by thickened rhizomatous stems,
occurring at ground level and below. Stems erect, single- to much-branched, lenticelate, cylindrical, scales imbricated
at base, tomentose. Leaves alternate, subrosulate, sessile, persistent, blade linear, coriaceous, lustrous, adaxial face
glabrous, abaxial lanate, base truncate, margin revolute, ciliate, apex spiny, venation hyphodromous. Capitulescence
terminal, monocephalous. Capitula 6-flowered, heterogamous, radiate, sessile, hidden by the leaves, involucre
cylindrical–campanulate, 5–7-seriate, phyllaries coriaceous, lanceolate, erect, lanate, apex spiny, margin flat, ciliate.
Receptacle flat, pilose. Flowers dimorphic. Ray flowers 5, bisexual, subbilabiate (4+1), 5-lobed, with the inner lobe
shorter than the outer lobes, externally hirsute-sericeous, white, internally glabrous, yellow. Disc flower 1, male,
tubular (5+0), externally sericeous-pilose, pistil not seen. Anthers 5, apical appendage lanceolate to obtuse, inserted at
the throat of the corolla (ray flowers) or inserted between the base and the throat (disc flowers), filaments free, glabrous.
Style cylindrical, yellow (ray flowers), not seen in disc flowers; Cypselae turbinate (ray flowers) or cylindrical (disc
flowers), densely villous. Pappus plumose or single villous bristle or absent, equal to the corolla length. Pollen lophate,
radially symmetrical, smooth (Suessy & Urtubey 2007). Chromosome number unknown.
Fig. 8b–e; 10c.
Distribution and habitat:—Huarpea is a monotypic genus restricted to the department of Iglesia in San Juan
province, Argentina (Cabrera 1951). The sole species, H. andina, is found in the dry Monte vegetation above an
altitude of 3,300 m (Stuessy et al. 1996).
Notes:—Huarpea can be distinguished from the other Barnadesioideae genera by being unarmed subshrubs up
to 4.5 cm tall, capitula hidden by the leaves, with 5 ray flowers, hermaphroditic and subbilabiate corollas and one-disc
flower with tubular corollas and atrophied gynoecium (Cabrera 1951, Stuessy et al. 2009). Phylogenetically, Huarpea
is always recovered as sister to Barnadesia (Bremer 1994, Stuessy et al. 1996, Gustafsson et al. 2001, Gruenstaeudl et
al. 2009, Ferreira et al. 2019). This clade is distinguished by the radiate and heterogamous capitula, slightly sagittate
or decurrent anther bases, lophate pollen, and atrophied gynoecium in the disc flowers (only in Barnadesia subgenus
Bacasia).
Iconography:—Cabrera (1951).
Accepted species:—9.1 Huarpea andina Cabrera
Selected specimens examined:—ARGENTINA. San Juan: Iglesia, Reserva de San Guillermo, Cordón del
Infiernillo, 17 January 1083, E. Nicora 8573 (MO). Iglesia, Reserva de San Guillermo, alrededores del refugio de San
Guillermo, al Sur, 11 November 1982, E.A. Ulibarri 1499 (US). Iglesia, Reserva de San Guillermo, Reserva de San
Guillermo, detras del refugio de Agua del Godo, 3000 m, R. Kiesling 4555 (MO).
MORPHOLOGICAL DIVERSITY IN SPINY SUNFLOWERS Phytotaxa 518 (2) © 2021 Magnolia Press • 135
10. Schlechtendalia Lessing (1830: 242–243)
Type:—Schlechtendalia luzulifolia Less.
Perennial herbs, up to 1 m tall, unarmed. Leaves opposite on stems, rosulate at base, amplexicaul, sessile, persistent,
blade linear, chartaceous, pale or lustrous, sericeous or lanate, margin flat or slightly revolute, ciliate, apex spiny;
nervation parallelodromous. Capitulescence terminal or axillary, monocephalous, cymose, corymbiform, racemose, or
umbellate. Capitula homogamous, discoid, 50–100 flowered, pedunculate, involucre turbinate or hemispherical, 5–7-
seriate, phyllaries chartaceous, green, brownish, erect or reflexed, sericeous, lanceolate, apex spiny, margin flat, ciliate.
Receptacle flat, pilose. Flowers isomorphic, bisexual, corolla subbilabiate (4+1), 5-lobed, yellow, externally and throat
villous. Anthers 5, apical appendage obtuse, basal appendage acute, short-sagittated, ecalcarate, ecaudate, inserted
between the base and the throat, filaments free, glabrous. Style cylindrical, yellow. Cypselae turbinate, densely villous.
Pappus scaly, lanceolate, shorter than the corolla, scarious, glabrous. Pollen with one depression per mesocolpus,
sparsely microechinate (Suessy & Urtubey 2007. Chromosome number = 8 (Cialdella & López 1981).
Fig. 9; 10d.
Distribution and habitat:—The genus Schlechtendalia is monotypic, represented only by S. luzulifolia, and
occurs in the Pampa vegetation (grassland) in southern Brazil, Uruguay, and adjacent areas of Argentina (Stuessy et al.
1996, 2009). In Brazil, Schlechtendalia luzulifolia is classified as an endangered species, since the Pampas biome has
lost almost 54% of its original vegetation (Nakajima et al. 2013).
Notes:—Schlechtendalia is clearly a member of Barnadesioideae since it displays the “barnadesioids trichomes”
and subbilabilate corollas (Gustafsson et al. 2001). However, it has a set of morphological features that diverge from
the rest of the subfamily, including the unarmed habit, and the opposite leaves on stems, basally rosulate leaves with
parallelodromous venation. Besides these traits, the pappus is glabrous and narrow-scaly (also found in Duseniella)
without the pappus bristles (Gustafsson et al. 2001). Moreover, it is the only genus totally absent in xeric areas (Stuessy
et al. 2009).
The phylogenetic position of Schlechtendalia within Barnadesioideae remains uncertain. It has been proposed
as the sister group to the rest of the subfamily (Stuessy et al. 1996, Urtubey & Stuessy, 2001, Gruenstaeudl et al.
2009, Ferreira et al. 2019, Ferreira et al. in prep.), and as sister to the Doniophyton and Duseniella clade (Bremer
1994); Chuquiraga and Doniophyton (Gustafsson et al. 2001); the Archidasyphyllum, Arnaldoa and Fulcaldea clade
(Gustaffson et al. 2001); or the Barnadesia and Huarpea clade (Gruenstauedl et al. 2009). New ongoing studies focusing
on Barnadesioideae phylogeny (Ferreira et al. in prep.) may clarify the phylogenetic position of Schlechtendalia.
Iconography:—Suessy & Urtubey (2007).
Accepted species:—Schlechtendalia luzulifolia Less.
Selected specimens examined:—ARGENTINA. Entre Ríos: Colón, paraje la Calera, márgenes del Río Uruguay,
desembocadura del arroyo Perucho Vema em Río Uruguay, 21–22 December 1998, P.M. Simon 88 (WU). Colón,
Parque Nacional El Palmar, Palmar de Butia Yatay, Sendero La Glorieta, 31°53’08’’, 58°16’22’’W, 14 December 2014,
M.S. Ferrucci 3196 (HUEFS). Confederación, Confederación, 18 December 1963, A. Burkart 24972 (SI). BRAZIL.
Rio Grande do Sul: Alegrete, Estância Cerro do Tigre, cerro da ponte, 141 m, 29°39’48’’S, 55°23’40’’W, 25 October
2012, G. Heiden 2008 (SPF). Itapoã, Chacara Weber, 18 December 1948, B. Rambo SJ 38950 (B). Jaguarão, BR 116,
entre Arroio Grande e Jaguarão, 44 m, 32°30’2’’S, 53°19’15’’W, 03 January 2011, G. Heiden 1455 (SPF). Livramento,
Cerro Palomas, D.F.M. Valls 1379 (RB). Porto Alegre, Morro da Policia, 18 January 1964, E. Pereira 8490 (RB). São
Lourenço do Sul, BR116, main road from Pelotas to Porto Alegre, between arroio S. Isabel & R. Camaqua (km 167),
02 December 1979, T.M. Pedersen 12612 (MBM). URUGUAY. Maldonado, Sierra Ballena, 23 December 1964, Del
Puerto 3867/4348 (US). Maldonado, Punta Ballena, Bosque Lussich, 09 February 1943, Rosengurtt B4507 (US).
Maldonado, Cerro San Antonio, just above Puerto Antonio, 0.9 km SE of Piriápolis (jct. rd to Punta del Este), 09
February 1993, T.F. Stuessy 12810 (WU). Maldonado, Punta Ballena (La Bellena), about ½ distance from Piriápolis
and Punta del Este (ca. 28 km E of Piriápolis), 09 February 1993, T.F. Stuessy 12820 (WU). San José, Balneario Kiyú
al E, 14 November 1991, D. Bayce s.n. (US).
FERREIRA ET AL.
136 • Phytotaxa 518 (2) © 2021 Magnolia Press
FIGURE 9. Schlechtendalia luzulifolia. A: Brazilian Pampas. B: Grassy-like habit. C: Capitulum with subbilabiate corollas. Photos by
Gustavo Heiden (A and C) and Mauricio Bonifacino (B).
MORPHOLOGICAL DIVERSITY IN SPINY SUNFLOWERS Phytotaxa 518 (2) © 2021 Magnolia Press • 137
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).
FERREIRA ET AL.
138 • Phytotaxa 518 (2) © 2021 Magnolia Press
Conclusions and Prospects
Taxonomy is one of the oldest branches of plant science, dating back to the Greek and Roman civilizations ca. 370–285
BC. Since then, taxonomic classifications have been radically improved, changing from descriptions mostly based on
morphology to integrated monographies using as many sources of information as possible (Grace et al. 2021). Such
integrative methods provide a better understanding of evolution and species delimitation.
Previous taxonomic studies in Barnadesioideae generally comprised short descriptions of genera and did not
highlight the great diversity in this group. Taxonomic revisions at generic level display a more detailed understanding
(Ezcurra 1985, Katinas & Stuessy 1997, Urtubey 1999, Saavedra 2011), but these are genus-specific and not comparable
across genera as they were not proposed using the same morphological characters. In this work, we assembled data
from previous studies and analyzed thousands of herbarium collections in order to present an integrated monography
of Barnadesioideae, providing extended descriptions including morphology, pollen, chromosome counts, taxonomic
and phylogenetic relationship comments, distribution maps, photos and a key for all ten genera circumscribed in this
subfamily.
Given the advances in molecular techniques this century, we now have a much better understanding of the
phylogenetic relationships, classification and evolution of morphological characters in the subfamily (Gustaffson et
al. 2001, Stuessy & Urtubey 2006, Gruenstauedl et al. 2009, Padin et al. 2015b, Ferreira et al. 2019, Svoma et
al. 2019). Although generic relationships are roughly stable (exceptions are found in the Chuquiraga/Doniophyton/
Duseniella clade and Schlechtendalia), infrageneric relationships and species delimitations often remain unclear, and
additional in-depth studies–for example using phylogenomics–will be required to shed further light on the evolution
of Barnadesioideae and the entire family.
Acknowledgments
This research was supported by FAPESP (grant #2016/06260-2) and CNPq to M.G.; the Swedish Foundation for
Strategic Research, the Swedish Research Council, the Knut and Alice Wallenberg Foundation and the Royal Botanic
Gardens, Kew to A.A.; a Doctoral Fellowship (CAPES, Finance Code 001) and a Fellowship for Internship abroad
from Coordination for the Improvement of Higher Education Personnel (CAPES, PDSE proc. 88881.132410/2016-
01), Missouri Botanical Garden Elisabeth E. Bascom scholarship, MEMOVA project (EU Operational Programme
Research, Development and Education No. CZ.02.2.69/0.0/0.0/18_053/0016982), and an International Association for
Plant Taxonomy Granted in 2017 to P.L.F. The authors thank all the herbarium curators and staff who have enabled the
development of this work. Special thanks to John Pruski for his invaluable support and enthusiasm during a period at
Missouri Botanical Garden. Thanks also to Gari V. Ccana-Ccapatinta, Cíntia Luz, Marcelo Monge, Mariana Saavedra,
Claudia Martín for sending herbarium samples that greatly contributed to this work; Laura Afonso for helping with the
distributional data and maps; Fernando Zuloaga for allowing us the use of the Darwinion Institute photos; Carolina
Siniscalchi, Danilo Marques, José Pirani, Marcelo Monge, and Vanessa Rivera for critically reading this manuscript;
and Rhian Smith for English editing and proofreading.
References
Bentham, G. (1873) Notes on the classification, history, and geographical distribution of Compositae. Journal of the Linnean Society,
Botany 13: 335–577.
https://doi.org/10.1111/j.1095-8339.1873.tb02575.x
Bohm, B.A. & Stuessy, T.F. (1995) Flavonoid chemistry of Barnadesioideae (Asteraceae). Systematic Botany 20: 22–27.
https://doi.org/10.2307/2419629
Bohm, B.A. & Stuessy, T.F. (2001) Flavonoids of the Sunflower Family (Asteraceae). Springer-Verlag Wien, Vienna, 831 pp.
https://doi.org/10.1007/978-3-7091-6181-4
Bremer, K. (1994) Asteraceae: cladistics and classification. Timber Press, Portland, 752 pp.
Bremer, K. & Jansen, R.K. (1992) A new subfamily of the Asteraceae. Annals of the Missouri Botanical Garden 79: 414–415.
https://doi.org/10.2307/2399777
MORPHOLOGICAL DIVERSITY IN SPINY SUNFLOWERS Phytotaxa 518 (2) © 2021 Magnolia Press • 139
Cabrera, A.L. (1951) Huarpea, nuevo genero de Compuestas. Boletín de la Sociedad Argentina de Botánica 4: 129–132.
Cabrera, A.L. (1959) Revisión del género Dasyphyllum (Compositae). Revista del Museo de La Plata, Sección botánica 38: 21–100.
Cabrera, A.L. (1961) Compuestas Argentinas. Clave para la determinación de los géneros. Revista del Museo Argentino de Ciencias
Naturales. Botánicas 2: 291–362.
Cabrera, A.L. (1962) Compuestas andinas nuevas. Boletín de la Sociedad Argentina de Botánica 10: 21–45.
Cabrera, A.L. (1977) Mutisieae—systematic review. In: Heywood, V.H., Harborne, J.B. & Turner, B.L. (Eds.) The Biology and Chemistry
of the Compositae, vol. 2. Academic Press, London, pp. 1039–1066.
Candolle, A.P. de (1838) Prodromus Systematis Naturalis Regni Vegetabilis, vol. 7. Treuttel & Würtz, Paris, 801 pp.
Ccana-Ccapatinta, G.V., Monge, M., Ferreira, P.L. & Da Costa, F.B. (2018) Chemistry and medicinal uses of the subfamily Barnadesioideae
(Asteraceae). Phytochemistry Reviews 17: 471–489.
https://doi.org/10.1007/s11101-017-9544-y
Chung, I.C. (1965) Revision of Barnadesia. Published by the author, Chicago, Illinois.
Cialdella, A.M. & López de Kiesling, A.G. (1981) Cariología de Schlechtendalia luzulaefolia (Compositae). Darwiniana 23: 357–360.
Cristóbal, C.L. (1986) El número cromosómico de dos Compositae–Mutisieae. Boletín de la Sociedad Argentina de Botánica 24: 363–
380.
DeVore, M.L., Zhao, Z., Jansen, R.K. & Skvarla, J.J. (2000) Utility of trends in pollen morphology for phylogenetic analyses: an example
using subfamilies Barnadesioideae and Cichorioideae (Asteraceae). In: Harley, M.M., Morton, C.M. & Blackmore, S. (Eds.) Pollen
and Spores: morphology and biology. Kew Royal Botanic Gardens, London, pp. 399–412.
Diers, L. (1961) Der Anteil an Polyploiden in den Vegetationsgürteln der Westkordillera Perus. Zeitschrift für Botanik 49: 437–488.
Don, D. (1830) Descriptions of the new Genera and Species of the Class Compositae belonging to the Floras of Peru, Mexico, and Chile.
Transactions of the Linnean Society of London 16: 169–304.
https://doi.org/10.1111/j.1095-8339.1829.tb00136.x
Erbar, C. & Leins, P. (2000) Some interesting features in the capitulum and flower of Arnaldoa macbrideana Ferreyra (Asteraceae,
Barnadesioideae). Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 122: 517–537.
Ezcurra, C. (1985) Revisión del género Chuquiraga (Compositae–Mutisieae). Darwiniana 26: 219–284.
Ferreira, P.L., Saavedra, M.M. & Groppo, M. (2019) Phylogeny and circumscription of Dasyphyllum (Asteraceae: Barnadesioideae) based
on molecular data with the recognition of a new genus, Archidasyphyllum. PeerJ 7: e6475.
https://doi.org/10.7717/peerj.6475
Ferreyra, R. (1995) Family Asteraceae: Part IV, Tribe Mutiseae. In: Macbride, J.F. et al. (Eds.) Flora of Peru. Fieldiana Botany. N.S. 35:
1–101.
Funk, V.A. & Roque, N. (2011) The monotypic Andean genus Fulcaldea (Compositae, Barnadesioideae) gains a new species from
Northeastern Brazil. Taxon 60: 1095–1103.
https://doi.org/10.1002/tax.604012
Grace, O.M., Pérez-Escobar, O.A., Lucas, E.J., Vorontsova, M.S., Lewis, G.P., Walker, B.E., Lohmann, L.G., Knapp, S., Wilkie, P.,
Särkinen, T., Darbyshire, I., Lughadha, E.N., Monro, A., Woudstra, Y., Demissew, S., Muasya, A.M., Díaz, S., Baker, W.J. &
Antonelli, A. (2021) Botanical Monography in the Anthropocene. Trends in Plant Science 26: 433–441.
https://doi.org/10.1016/j.tplants.2020.12.018
Granda, P.A. (1997) Uma nueva espécie de Chuquiraga (Asteraceae-Mutiseae) del Perú. Kurtziana 25: 151–156.
Gruenstaeudl, M., Urtubey, E., Jansen, R.K., Samuel, R., Barfuss, M.H.J. & Stuessy, T.F. (2009) Phylogeny of Barnadesioideae (Asteraceae)
inferred from DNA sequence data and morphology. Molecular Phylogenetics and Evolution 51: 572–587.
https://doi.org/10.1016/j.ympev.2009.01.023
Gustafsson, M.H.G., Pepper, A.S.R., Albert, V.A. & Källersjö, M. (2001) Molecular phylogeny of the Barnadesioideae (Asteraceae).
Nordic Journal of Botany 21: 149–160.
https://doi.org/10.1111/j.1756-1051.2001.tb01352.x
Hansen, H.V. (1991) Phylogenetic studies in Compositae tribe Mutisieae. Opera Botanica 109: 1–50.
Harling, G. (1991) Compositae-Mutisieae. In: Harling, G. & Andersson, L. (Eds.) Flora of Ecuador. University of Gothenburg, Gothenburg,
pp. 1–105.
Heiser, C.B. (1963) Numeración cromosómica de plantas ecuatorianas. Ciencia y Naturaleza 6: 2–6.
Hind, D.J.N. (2001) A New Species of Barnadesia (Compositae: Barnadesieae) from Bolivia. Kew Bulletin 56: 705–710.
https://doi.org/10.2307/4117698
Hoffmann, O. (1893) Tubuliflorae–Mutisieae. In: Engler, A. & Prantl, K. (Eds.) Die natürlichen Pflanzenfamilien, vol. 4 (5). Engelmann,
Leipzig, pp. 333–350.
Jansen, R.K. & Palmer, J.D. (1987) Chloroplast DNA from lettuce and Barnadesia (Asteraceae): structure, gene localization, and
FERREIRA ET AL.
140 • Phytotaxa 518 (2) © 2021 Magnolia Press
characterization of a large inversion. Current Genetics 11: 553–564.
https://doi.org/10.1007/BF00384619
Jansen, R.K. & Palmer, J.D. (1988) Phylogenetic implications of Chloroplast DNA restriction site variation in the Mustiseae (Asteraceae).
American Journal of Botany 75: 753–766.
https://doi.org/10.1002/j.1537-2197.1988.tb13496.x
Jansen, R.K., Michaels, H.J., Wallace, R.S., Kim, K-J., Keeley, S.C., Watson, L.E. & Palmer, J.D. (1992) Chloroplast DNA Variation in
the Asteraceae: Phylogenetic and Evolutionary Implications. In: Soltis, P.S., Soltis, D.E. & Doyle, J.J. (Eds.) Molecular Systematics
of Plants. Springer, Boston, pp. 252–279.
https://doi.org/10.1007/978-1-4615-3276-7_11
Jussieu, A.L. (1789) Genera Plantarum Secundum Ordines Naturales Disposita. Herrisant & Barrois, Paris, 498 pp.
Katinas, L. & Stuessy, T.F. (1997) Revision of Doniophyton (Compositae, Barnadesioideae). Plant Systematics and Evolution 206: 33–
45.
https://doi.org/10.1007/BF00987939
Kunth, C.S. (1820) Compositae. In: Humboldt, F.W.H.A., Bonpland, A.J.A. & Kunth, K.S. (Eds.) Nova genera et species plantarum, vol.
4. Librariae Graeco-Latino-Germanico, Paris, 312 pp. + 113 tab.
Lessing, C.P. (1832) Synopsis generum compositarum. Duncker & Humblot, Berlin.
Mandel, J.R., Dikow, R.B., Siniscalchi, C.M., Thapa, R., Watson, L.E. & Funk, V.A. (2019) A fully resolved backbone phylogeny reveals
numerous dispersals and explosive diversifications throughout the history of Asteraceae. Proceedings of the National Academy of
Sciences 116: 14083 LP-14088.
https://doi.org/10.1073/pnas.1903871116
Mendiondo, M.E., Juárez, B.E. & Seeligmann, P. (1997) Flavonoid patterns of some Barnadesioideae (Asteraceae). Eventual
chemosystematic significance. Biochemical Systematics and Ecology 25: 673–674.
https://doi.org/10.1016/S0305-1978(97)00060-4
Mendiondo, M.E., Juárez, B.E. & Seeligmann, P. (2000) Flavonoid profiles of some Argentine species of Chuquiraga (Asteraceae).
Biochemical Systematics and Ecology 28: 283–285.
https://doi.org/10.1016/S0305-1978(99)00061-7
Mutis, J.C.B. (1781) Polyadelphia Pentandria. In: Linné, C. (Ed.) Supplementum Plantarum. Brunsvigae, pp. 55.
Nabhan, A.R. & Sarkar, I.N. (2012) The impact of taxon sampling on phylogenetic inference: a review of two decades of controversy.
Briefings in bioinformatics 13: 122–134.
https://doi.org/10.1093/bib/bbr014
Nakajima, J.N., Dematteis, M., Loeuille, B., Teles, A.M., Heiden, G., Schneider, A., Ritter, M., Oliveira, C.T., Hattori, E.K.O., Roque, N.,
Ferreira, S.C., Magenta, M., Bringel, J.B.A., Esteves, R., Almeida, G.S.S., Saavedra, M.M., Monge, M., Soares, N.P., Sancho, G.,
Mondin, C.A., Fernandes, A.C., Pereira, A.C.M., Kutschenko, D.C., Filho, L.A.F.S., Prieto, P.V., Borges, R.A.X., Penedo, T.S.A.,
Messina, T., Moraes, M.M.V., Moraes, M.A. & Coelho, M.A.C. (2013) Asteraceae. In: Martinelli, G. & Moraes, M.A. (Eds.) Livro
vermelho da Flora do Brasil. Centro Nacional de Conservação da Flora, Rio de Janeiro, pp. 203–286.
Olsen, J. (1980) Chromosome number reports LXVII. Taxon 29: 366–367.
https://doi.org/10.1002/j.1996-8175.1980.tb00607.x
Padin, A.L., Calviño, C.I. & Ezcurra, C. (2015a) Morfología y anatomía foliar comparada de Chuquiraga y géneros afines (Asteraceae).
Brittonia 67: 150–165.
https://doi.org/10.1007/s12228-015-9364-6
Padin, A.L., Calviño, C.I. & Ezcurra, C. (2015b) Molecular Phylogeny of Chuquiraga (Asteraceae-Barnadesioideae): Infrageneric
Classification and Generic Affinities. Systematic Botany 40: 316–326.
https://doi.org/10.1600/036364415X686602
Poeppig, E.F. & Endlicher, S.L. (1835–1836) Nova genera ac species plantarum, vol. 1. F. Hofmeister, Leipzig, 62 pp.
Poiret, J.L.M. (1817) Fulcaldea [under Turpinia]. In: Lamarck, M. & Poiret, J.L.M. (Eds.) Encyclopedie Methodique, Botanique, suppl.,
vol. 5. Paris, pp. 575.
QGIS Development Team (2019) QGIS Geographic Information System, version 3.4. Open Source Geospatial Foundation Project.
Available from: http://qgis.osgeo.org (accessed 20 February 2019)
Radford, A.E., Dickson, W.C., Massey, W.C. & Bell, C.R. (1974) Vascular Plant Systematics. Harper & Row, New York, 891 pp.
Saavedra, M.M. (2011) Sistemática de Dasyphyllum (Asteraceae). Phd Thesis. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro,
Rio de Janeiro, pp. 1–247.
Saavedra, M.M. (2020) Dasyphyllum in Flora do Brasil 2020. Jardim Botânico do Rio de Janeiro. Available from: http://floradobrasil.jbrj.
gov.br/reflora/floradobrasil/FB5291 (accessed 20 March 2021)
MORPHOLOGICAL DIVERSITY IN SPINY SUNFLOWERS Phytotaxa 518 (2) © 2021 Magnolia Press • 141
Saavedra, M.M., Guimarães, E.F., Loeuille, B. & Forzza, R.C. (2018) Taxonomic Revision of Dasyphyllum sect. Macrocephala (Asteraceae:
Barnadesioideae). Systematic Botany 43: 297–315.
https://doi.org/10.1600/036364418X696888
Saavedra, M.M., Monge, M., Guimarães, E.F. (2014) Dasyphyllum diamantinense (Asteraceae, Barnadesioideae): A new species from the
Chapada Diamantina, Bahia State, Brazil. Phytotaxa 174 (4): 231–236.
https://doi.org/10.11646/phytotaxa.174.4.4
Sagástegui, A.A. (1980) Compuestas andinas peruanas nuevas para la ciencia. Boletín de la Sociedad Argentina de Botánica 19 (1–2):
61–68.
Sagástegui, A.A. & Dillon, M. (1985) Four new species of Asteraceae from Peru. Brittonia 37 (1): 6–13.
https://doi.org/10.1007/BF02809659
Sagástegui, A.A. & Sánchez, V.I. (1991) Una nueva espécie de Chuquiraga (Asteraceae-Mutisieae) del Norte del Peru. Arnaldoa 2: 1–4.
Schumann, K.M. (1902) Neue Arten des Siphonogamen. Just’s botanischer Jahresbericht 28 (Abt. 1): 410–498.
Skvarla, J.J., Turner, B.L., Patel, V.C. & Tomb, A.S. (1977) Pollen morphology in the Compositae and in morphologically related families.
In: Heywood, V.H., Harborne, J.B. & Turner, B.L. (Eds.) The Biology and Chemistry of the Compositae. Academic Press, London,
pp. 141–248.
Strother, J.L. & Panero, J.L. (1994) Chromosome studies: Latin American Compositae. American Journal of Botany 81: 335–577.
https://doi.org/10.2307/2445657
Stuessy, T.F. & Sagástegui, A.A. (1993) Revision de Arnaldoa (Compositae, Barnadesioideae) género endémico del norte del Peru.
Arnaldoa 1: 9–21.
Stuessy, T.F. & Urtubey, E. (2006) Phylogenetic implications of corolla morphology in subfamily Barnadesioideae (Asteraceae). Flora:
Morphology, Distribution, Functional Ecology of Plants 201: 340–352.
https://doi.org/10.1016/j.flora.2005.07.009
Stuessy, T.F. & Urtubey, E. (2007) Barnadesieae. In: Kadereit, J.W. & Jeffrey, C. (Eds.) Kubtizki, K., The Families and genera of vascular
plants, vol. 8. Springer-Verlag, Berlin & Heidelberg, pp. 87–90.
Stuessy, T.F., Sang, T. & DeVore, M.L. (1996) Phylogeny and biogeography of the subfamily Barnadesioideae with implications for early
evolution of the Compositae. In: Hind, D.J.H. & Beentje, H.J. (Eds.) Compositae: Systematics. Proceedings of the International
Compositae Conference, Kew, 1994, vol. 1. Royal Botanical Garden, Kew, pp. 463–490.
Stuessy, T.F., Urtubey, E. & Gruenstaeudl, M. (2009) Barnadesieae (Barnadesioideae). In: Funk, V.A., Susanna, A., Stuessy, T. & Bayer,
R.J. (Eds.) Systematics, evolution, and biogeography of Compositae. IAPT, Vienna, pp. 215–228.
Svoma, E., Mayer, V., Stuessy, T.F. & Urtubey, E. (2019) Staminal features in Barnadesioideae (Asteraceae): description, evolution and
function. Botanical Journal of the Linnean Society 192: 474–497.
https://doi.org/10.1093/botlinnean/boz091
Tellería, MC., Palazzesi, L. & Barreda, V. (2015) Evolutionary significance of exine ultrastructure in the subfamily Barnadesioideae
(Asteraceae) in the light of molecular phylogenetics. Review of Palaeobotany and Palynology 221: 32–46.
https://doi.org/10.1016/j.revpalbo.2015.05.008
Thiers, B. (2019) Index Herbariorum: A Global Directory of Public Herbaria and Associated Staff. New York Botanical Garden’s Virtual
Herbarium. Available from: http://sweetgum.nybg.org/science/ih/ (accessed 20 February 2019)
Ulloa Ulloa, C., Jørgensen, P.M. & Dillon, M.O. (2002) Arnaldoa argentea (Barnadesioideae: Asteraceae), a new species and a new
generic record for Ecuador. Novon 12: 415–419.
https://doi.org/10.2307/3393091
Urtubey, E. & Stuessy, T.F. (2001) New hypotheses of phylogenetic relationships in Barnadesioideae (Asteraceae) based on morphology.
Taxon 50: 1043–1066.
https://doi.org/10.2307/1224720
Urtubey, E. (1997) Morfología del pólen de Barnadesia (Asteraceae, Barnadesioideae). Boletín de la Sociedad Argentina de Botánica 33:
69–75.
Urtubey, E. (1999) Revisión del género Barnadesia (Barnadesioideae, Asteraceae). Annals of the Missouri Botanical Garden 86: 57–
111.
https://doi.org/10.2307/2666218
Urtubey, E. & Tellería, M.C. (1998) Pollen morphology of the subfamily Barnadesioideae (Asteraceae) and its phylogenetic and taxonomic
significance. Review of Palaeobotany and Palynology 104: 19–37.
https://doi.org/10.1016/S0034-6667(98)00049-9
Watanabe, K., Yahara, T., Hashimoto, G., Nagatani, Y., Soejima, A., Kawahara, T. & Nakazawa, M. (2007) Chromosome numbers and
karyotypes in Asteraceae. Annals of the Missouri Botanical Garden 94: 643–654.
FERREIRA ET AL.
142 • Phytotaxa 518 (2) © 2021 Magnolia Press
https://doi.org/10.3417/0026-6493(2007)94[643:CNAKIA]2.0.CO;2
Weddell, H.A. (1855) Chloris andina: essai d’une flore de la región alpine des Cordillères de l’Amérique de Sud. Bertrand, Paris.
https://doi.org/10.5962/bhl.title.217
Wulff, A.F. (1984) Estudios cromosómicos en Compuestas de las floras patagónica y bonaerense. Darwiniana 25: 17–26.
Wulff, A.F. (1990) Estudios cromosómicos en Barnadesiinae (Mutisieae, Asteraceae). I. Chuquiraga y Doniophyton. Darwiniana 30:
185–193.
Zhao, Z., Skvarla, J.J., Jansen, R.K. & DeVore, M.L. (2000) Phylogenetic implications of pollen morphology and ultrastructure in the
Barnadesioideae (Asteraceae). Lundellia 3: 26–40.
https://doi.org/10.25224/1097-993X-3.1.26
