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Systematics of the South American Hypochaeris sessiliflora Complex (Asteraceae, Cichorieae) 1

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El complejo Hypochaeris sessiliflora comprende nueve especies de Sudamérica (sección Achyrophorus Scop.) con cabezuelas sésiles o cortamente pedunculadas y con una roseta de hojas basales. Habitan entre los 1430–5100 m a lo largo de los Andes desde Venezuela hasta el centro de Chile y el centro oeste de Argentina. Dos especies, H. sessiliflora Kunth e H. meyeniana (Walp.) Benth. & Hook. f. ex Griseb., son extremadamente polimórficas (forma de los filarios, hojas e indumento) y presentan una amplia distribución, desde Venezuela al centro de Perú, y desde Perú al norte de Chile y noroeste de Argentina, respectivamente; principalmente crecen en ambientes secos y soleados, y florecen todo el año. Hypochaeris meyeniana se caracteriza por las hojas pinnatífidas y retrorsas (raro lobadas) y los aquenios ligeramente adelgazados en el ápice. Hypochaeris hohenackeri (Sch. Bip.) Domke e H. taraxacoides Ball son glabras, mientras que H. acaulis (J. Rémy) Britton tiene hojas híspidas; estas tres especies están asociadas a ambientes húmedos (e.g., vegas y mallines). Hypochaeris eriolaena (Sch. Bip.) Reiche e H. mucida Domke son níveo-tomentosas. Hypochaeris echegarayi Hieron. (corolas blancas) e H. eremophila Cabrera (corolas amarillas) son especies afines con pubescencia híspidas, comúnmente con cabezuelas pedunculadas y con considerable tolerancia ecológica, creciendo en ambientes secos o húmedos. La hipótesis del análisis cladístico morfológico y los recientes estudios moleculares muestran al complejo H. sessiliflora monofilético mientras que la hipótesis basada en datos moleculares excluye a H. acaulis del complejo H. sessiliflora, donde el hábito acaule es considerado una adaptación paralela como respuesta a la sobrevivencia en la alta montaña.
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SYSTEMATICS OF THE SOUTH
AMERICAN HYPOCHAERIS
SESSILIFLORA COMPLEX
(ASTERACEAE, CICHORIEAE)
1
Estrella Urtubey,
2
Tod F. Stuessy,
3
and
Karin Tremetsberger
4
ABSTRACT
The Hypochaeris sessiliflora complex (Asteraceae, Cichorieae) consists of nine species of the genus from South America (all
in section Achyrophorus Scop.) that have sessile or nearly sessile flowering heads surrounded by a rosette of leaves. They occur
at 1430–5100 m in elevation along the Andean chain from Venezuela to Chile and Argentina. Two species, H. sessiliflora
Kunth and H. meyeniana (Walp.) Benth. & Hook. f. ex Griseb., are extremely polymorphic, and they vary conspicuously in the
shape of the external phyllaries and presence or absence of different types of trichomes. They have the widest distributions
(Venezuela to central Peru, and Peru to northern Chile and northwestern Argentina, respectively), they flower throughout the
year, and they also are primarily associated with dry and sunny habitats. Hypochaeris meyeniana is characterized by retrorsely
pinnatifid leaves (rarely lobate) and slightly narrower cypselar apices. Hypochaeris hohenackeri (Sch. Bip.) Domke and H.
taraxacoides Ball are glabrous, whereas H. acaulis (J. Re
´my) Britton has scattered shaggy trichomes on the leaves; all three
occur in humid places, such as seeps or bogs. Hypochaeris eriolaena (Sch. Bip.) Reiche and H. mucida Domke are pubescent,
with long whiplike trichomes on leaves and phyllaries, giving a niveous-tomentose appearance. Hypochaeris echegarayi
Hieron. (white corollas) and H. eremophila Cabrera (yellow corollas) are two related species with shaggy trichomes on the
abaxial surfaces of the phyllaries, both with considerable ecological tolerance, that grow in dry as well as humid sites.
Morphological cladistic analyses suggest a hypothesis of relationships within the complex. Surprisingly, H. acaulis from Chile
and Argentina, although fitting morphologically within the H. sessiliflora complex, based on amplified fragment length
polymorphism (AFLP) data, clearly does not seem to belong to this group. Instead, the species ties to H. palustris (Phil.) De
Wild. and H. tenuifolia (Hook. & Arn.) Griseb., also from the southern Andes. The acaulescent habit of H. acaulis seems best
interpreted as a parallel adaptation to survival at high elevations.
RESUMEN
El complejo Hypochaeris sessiliflora comprende nueve especies de Sudame
´rica (seccio
´nAchyrophorus Scop.) con cabezuelas
se
´siles o cortamente pedunculadas y con una roseta de hojas basales. Habitan entre los 1430–5100 m a lo largo de los Andes
desde Venezuela hasta el centro de Chile y el centro oeste de Argentina. Dos especies, H. sessiliflora Kunth e H. meyeniana
(Walp.) Benth. & Hook. f. ex Griseb., son extremadamente polimo
´rficas (forma de los filarios, hojas e indumento) y presentan
una amplia distribucio
´n, desde Venezuela al centro de Peru
´, y desde Peru
´al norte de Chile y noroeste de Argentina,
respectivamente; principalmente crecen en ambientes secos y soleados, y florecen todo el an
˜o. Hypochaeris meyeniana se
caracteriza por las hojas pinnatı
´fidas y retrorsas (raro lobadas) y los aquenios ligeramente adelgazados en el a
´pice. Hypochaeris
hohenackeri (Sch. Bip.) Domke e H. taraxacoides Ball son glabras, mientras que H. acaulis (J. Re
´my) Britton tiene hojas
´spidas; estas tres especies esta
´n asociadas a ambientes hu
´medos (e.g., vegas y mallines). Hypochaeris eriolaena (Sch. Bip.)
Reiche e H. mucida Domke son nı
´veo-tomentosas. Hypochaeris echegarayi Hieron. (corolas blancas) e H. eremophila Cabrera
(corolas amarillas) son especies afines con pubescencia hı
´spidas, comu
´nmente con cabezuelas pedunculadas y con
considerable tolerancia ecolo
´gica, creciendo en ambientes secos o hu
´medos. La hipo
´tesis del ana
´lisis cladı
´stico morfolo
´gico y
los recientes estudios moleculares muestran al complejo H. sessiliflora monofile
´tico mientras que la hipo
´tesis basada en datos
moleculares excluye a H. acaulis del complejo H. sessiliflora, donde el ha
´bito acaule es considerado una adaptacio
´n paralela
como respuesta a la sobrevivencia en la alta montan
˜a.
Key words: Achyrophorus, Asteraceae, Cichorieae, Hypochaeris, South America.
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1
We express thanks to John McNeill and Werner Greuter for help with the nomenclature of Hypochaeris taraxacoides;A.
Migoya and M. Theiller for illustrations of the species and H. Calvetti for help with maps; the curators of BM, CONC, K, LP,
LPB, M, MCNS, MO, NY, OS, P, SGO, SI, UC, US, W, and WU for loans or permission to consult herbarium material; M.
Mun
˜oz (SGO) and L. Willemse (L) for digital images of the types of Distoecha taraxacoides and H. ornata and H. parvifolia,
respectively; F. Felbe and F. Jacquemoud for information on types at NEU and G, respectively; V. Noble, N. Hind, and S.
Barrier for their assistance during visits to BM, K, and P; S. E. Freire and D. Giuliano for valuable comments on the
manuscript; A. Luck for editorial help with various drafts of the manuscript; and financial support from the Consejo Nacional
de Investigaciones Cientı
´ficas y Te
´cnicas (CONICET, PEI and PIP6510), Myndel Botanica Foundation, a small grant from the
University of Vienna, and the Austrian National Science Foundation (FWF, grant numbers P15225-BIO and P18446-B03).
2
Instituto de Bota
´nica Darwinion, Labarde
´n 200, San Isidro, B1642HYD, Casilla de Correo 22, Buenos Aires, Argentina.
eurtubey@darwin.edu.ar.
3
Department of Systematic and Evolutionary Botany, Faculty Center Biodiversity, University of Vienna, Rennweg 14, A-
1030 Vienna, Austria. tod.stuessy@univie.ac.at.
4
Institute of Botany, Department of Integrative Biology and Biodiversity Research, University of Natural Resources and
Applied Life Sciences, Vienna, Gregor-Mendel-Straße 33, A-1180 Vienna, Austria. karin.tremetsberger@boku.ac.at.
doi: 10.3417/2006136
ANN.MISSOURI BOT.GARD. 96: 685–714. PUBLISHED ON 00 MONTH 2009.
Hypochaeris L. consists of about 60 species of
annual or perennial herbs in Asteraceae, tribe
Cichorieae Lam. & DC., subtribe Hypochaeridinae
Less., which are characterized by paleae on recepta-
cles, a plumose pappus, and style branches papillose
to well below the bifurcation (Fig. 1A–C). The genus
has a disjunct distribution, with ca. 12 species
scattered in the Mediterranean region, two in Central
Europe, one in Asia, and the remainder (some 40 to 50
species) in South America. Hoffmann (1893) reduced
the 10 sections of Bentham and Hooker (1873) to five,
based on the number of rows of pappus bristles,
shapes of heads, arrangements of phyllaries, and
shapes of the cypselae. One of these sections,
Achyrophorus Scop., is characterized primarily by
having a uniseriate pappus; all species from South
America fall into this group.
Previous evolutionary studies have been completed
on species of Hypochaeris from South America.
Chromosome counts from many species and popula-
tions have revealed most members of Hypochaeris to
be diploid, with some infraspecific polyploidy, and
with some species uniformly tetraploid (Baeza et al.,
2000; Weiss et al., 2003; Weiss-Schneeweiss et al.,
2007, 2008). Although all diploids uniformly have 2n
58, small karyotypic differences allow hypotheses
regarding modes of cytogenetic evolution (Weiss-
Schneeweiss et al., 2003). Sequences of nuclear
ribosomal ITS (nrITS) and chloroplast DNA have
revealed that the South American species are
monophyletic and that they are closest to H.
angustifolia Maire from Morocco (Samuel et al.,
2003; Tremetsberger et al., 2005). Sequence variation
within species from South America is minimal, and no
clades can be recognized with high levels of
confidence. Amplified fragment length polymorphism
(AFLP) studies have helped reveal phyletic assem-
blages within the South American taxa (Tremetsberger
et al., 2006), as well as indicate phylogeographic
trends at the populational level in H. acaulis (J. Re
´my)
Britton, H. palustris (Phil.) De Willd., and H.
tenuifolia (Hook. & Arn.) Griseb. (Tremetsberger et
al., 2003a, b; Muellner et al., 2005).
Among the South American taxa, a morphologically
distinctive group is formed by species that have single
sessile or nearly sessile capitula nestled among a
rosette of basal leaves, the so-called Hypochaeris
sessiliflora complex. These taxa occur from Venezuela
south into Chile and Argentina and occur at 1430–
5100 m, usually in dry habitats or in seeps within
these high Andean regions. This paper focuses on this
complex as a beginning for understanding detailed
morphological relationships within the South Ameri-
can species of Hypochaeris. A revision of all taxa is
included plus a morphological cladistic analysis.
CLADISTIC ANALYSIS
MATERIALS AND METHODS
Morphological characters and states were obtained
primarily from study of herbarium material from BM,
CONC, K, LP, LPB, M, MCNS, MO, NY, OS, P, SGO,
SI, UC, US, W, and WU, plus our additional
collections and field observations. Trichomes and
stomatal types and terminology are used according to
Metcalfe and Chalk (1950, 1979), Uphof (1962), and
Harris and Harris (1994). For study of vascularization
of corollas and cypselae, the clearing and staining
method of Fuchs (1963) was used. Information
regarding color, habitat, and phenology of plants was
taken from specimen labels, the literature, and our
field observations. Chromosome numbers were ob-
tained from the reviews of Weiss et al. (2003) and
Weiss-Schneeweiss et al. (2007, 2008).
The ingroup consisted of all nine South American
species of the Hypochaeris sessiliflora complex: H.
acaulis,H. echegarayi Hieron., H. eremophila Cab-
rera, H. eriolaena (Sch. Bip.) Reiche, H. hohenackeri
(Sch. Bip.) Domke, H. meyeniana (Walp.) Benth. &
Hook. f. ex Griseb., H. mucida Domke, H. sessiliflora
Kunth, and H. taraxacoides Ball. We included five
additional South American species of Hypochaeris to
serve as outgroup. These species were selected to
represent morphological diversity within Hypochaeris
on the continent, two of which (H. hookeri Phil. and H.
caespitosa Cabrera) are believed to be close relatives
of the H. sessiliflora complex. Hypochaeris hookeri is
closely related morphologically to the complex
(especially in its acaulescent habit). Its leaves are
long, linear, and erect, the peduncle is shorter than
the leaves, and the paleae have trichomes on the
adaxial surface; it is distributed in central-western
Argentina and central-eastern Chile. Hypochaeris
caespitosa is also an acaulescent herb, the peduncle
is longer than the leaves, and it inhabits the mountain
ranges of central Argentina. We also examined three
caulescent species, H. argentina Cabrera of central
Argentina, and H. chillensis (Kunth) Hieron. and H.
elata (Wedd.) Benth. & Hook. f. ex Griseb. from Peru,
south into Argentina.
To provide a context for selection of characters and
states in the cladistic analysis, and to better clarify
the character data used in the descriptions of each
taxon, a sketch of the morphology of the complex
follows here (see also Fig. 1). All nine species of the
Hypochaeris sessiliflora complex are dwarf perennial
herbs, from only 1 cm (H. mucida) to 13 cm (in H.
sessiliflora) tall. They are acaulescent plants with
thick rhizomes and rosette leaves. The capitula are
always solitary, and they can be sessile or shortly
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Figure 1. Selected morphological features of taxonomic efficacy in the Hypochaeris sessiliflora complex. —A. Palea. —B.
Plumose pappus bristles. —C. Papillose style. D–H. Trichomes. —D. Papillate. —E. Conical (many-celled). —F. Whip. —G.
Shaggy. —H. Glandular. I, J. Corolla vascularization. —I. Type 1. —J. Type 2. K–M. Anther tails. —K. Type 1. —L. Type 2.
—M. Type 3. N–P. Basal appendages. —N. Type 1. —O. Type 2. —P. Type 3.
Volume 96, Number 4 687
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Urtubey et al.
The Hypochaeris sessiliflora Complex (Asteraceae)
pedunculate (never longer than the leaves). The leaves
are entire or pinnatisect, the leaf base is attenuate in a
long petiole and tomentose, and the blades are
commonly lanceolate, oblanceolate, or linear to
elliptic. The leaf margin is entire or dentate. The
epidermal cells are lightly undulate, and the stomata
are anomocytic.
The involucre is commonly campanulate, campan-
ulate-cylindrical (Hypochaeris eremophila), or cylin-
dric (H. taraxacoides), and sometimes hemispheric (H.
acaulis,H. meyeniana, and H. sessiliflora). The
phyllaries are in three to six series. The outer ones
are lanceolate to broadly ovate, often becoming
gradually lighter toward the hyaline margin. The
maximum expression of this character state occurs in
some populations of H. sessiliflora in which the apical
margin is broadly expanded, called ‘‘cucullate’’
(hooded), and it can be entire or divided. The
phyllaries are either glabrous or ciliate, sometimes
lanuginous (whip trichomes) and/or setaceous (shaggy
trichomes). The adaxial side of the paleae is glabrous.
The corolla is of typical Lactuceae-type, i.e.,
ligulate and 5-lobed, with epidermal cells lightly
undulate and with epicuticular plated waxes (very
conspicuous in Hypochaeris mucida), and it has short
trichomes on the middle third. The vascularization of
corollas is characterized by the adjacent marginal
bundles united and fused at the apex of their lobes,
and there are two different types of vascularization.
The most common pattern has five marginal bundles
united at the apex of the lobes (type 1, Fig. 1I); the
other type comprises the same number of bundles, but
these are repeatedly divided in the throat and ligule of
the corolla (type 2, Fig. 1J). The color of the corolla
varies: white in H. echegarayi and H. taraxacoides;
yellow in H. acaulis,H. eremophila, and H. hohenack-
eri; or white, yellow, or orange in H. eriolaena and H.
sessiliflora. Some plants with pink corollas in H.
hohenackeri and H. sessiliflora have been collected.
We have no information about the corolla color of H.
mucida, because the available specimens are too
faded, none of the herbarium labels mention the color,
and we have not collected the taxon.
The five stamens have short anther tails, with basal
appendages with pollen sacs extending almost to the
end (1/2 to 3/4, Fig. 1K), into the middle or less (1/4
to 1/2, Fig 1L), or basal appendages with only sterile
cells (Fig. 1M). The antheropodium consists of
epidermal cells that are axially elongated with
thickened and lignified walls, and it can be shorter
than, the same length as, or longer than the basal
appendages (Fig. 1N–P).
The cypselae are commonly erostrate or narrowed
toward the apex; in Hypochaeris acaulis they are
rostrate. The cypselae are usually 5-ribbed (some-
times inconspicuous), and the surface is smooth or
scaly and striate.
Five types of trichomes are present, three being
uniseriate: (1) papilla-like (Fig. 1D), a small unicel-
lular epidermal cell thickened at the apex, being
present on lobes of the corolla; (2) conical (Fig. 1E),
2- to many-celled, e.g., on the margins of the palea or
pappus bristles (2-celled), or comprising the cilia of
the phyllaries (2- to many-celled); and (3) whip
(Fig. 1F), many-celled, with a very long apical cell,
present on the surfaces of leaves and phyllaries of
Hypochaeris eriolaena and H. mucida. Other types of
trichomes are multiseriate: (1) shaggy (Fig. 1G),
many-celled, ending in one to few cells, forming a
setaceous indumentum (these trichomes are often
brown or red, i.e., on phyllaries of H. echegarayi,H.
eremophila, and H. sessiliflora); and (2) biseriate-
glandular (Fig. 1H), present on the leaf surface of
some populations of H. sessiliflora. We have used the
term glabrous when trichomes are relatively few or
totally absent.
From this spectrum of morphological variation, 16
morphological characters (five vegetative and 11
floral) were selected for assessing evolutionary
relationships among the taxa. These are listed in
Appendix 1, and selected features are highlighted in
Figure 1. Autapomorphic characters were not includ-
ed in the analysis. The data matrix of the 14 species
and 16 characters is given in Appendix 2; vouchers
are listed in Appendix 3. A complete listing of
specimens cited in this study is presented in
Appendix 4. Polymorphisms represent nearly 20%
of the data. Character 2 was uninformative within the
complex, but it was included because it is important
for circumscribing all the caulescent species of
Hypochaeris.
Phylogenetic analyses were performed with Pee-
Wee 3.0 (Goloboff, 1998). Initial PeeWee runs were
conducted using sequence commands holding 10000:
rseed 0, hold/40, poly5, amb-; mult*500; fit* 5fit
rescaled. All characters were weighted equally and
character states were treated as unordered. Jackknife
support values for nodes were calculated with 1000
replicates, two search replicates (mult*2), and two
starting trees per replication (hold/2). Character
distributions were studied and strict consensus trees
were calculated using Winclada (Nixon, 1999).
RESULTS
The analyses resulted in one most parsimonious
tree of 40 steps, with fit 5110 and fit* 557%;
synapomorphies and homoplasies are mapped onto the
phylogeny (Fig. 2). All species except Hypochaeris
argentina are supported by the presence of type 1
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corolla vascularization (character 11, state 1). The
pair H. chillensis–H. elata is supported by corollas
with the same length as the involucre (character 10,
state 1) and a jackknife value of 64. Hypochaeris
caespitosa and the rest of the species are supported by
possessing linear-lanceolate leaf laminas (character 3,
state 4), whip trichomes on the abaxial surface of the
phyllaries (character 9, state 1), and smooth cypselar
walls (character 15, state 1). Hypochaeris hookeri is
the sister species of the ingroup, a clade supported by
acaulescent plants or plants with shortly pedunculate
heads (character 1, state 1) and jackknife support of 56.
The H. sessiliflora complex is supported as a monophy-
letic clade by glabrous phyllaries and paleae (character
9, state 3 and character 16, state 1) and with jackknife
support of 59, and H. sessiliflora is the basal species.
The clade of the remaining species is supported by
oblong leaves (character 3, state 1). Hypochaeris
hohenackeri through H. acaulis are supported by
lanceolate phyllaries (character 8, state 1). The clade
of H. mucida and the rest of the species is supported by
sessile heads (character 7, state 1). The remaining three
groups are supported primarily by homoplasies, the first
homoplasybeingareversal(scalycypselarwall,
character 15, state 0), followed by a parallelism (divided
leaves, character 4, state 2). The unresolved complex
consisting of H. meyeniana,H. echegarayi,andH.
eremophila issupported by a reversal (lanceolate leaves,
character 3, state 0) and one synapomorphy (shaggy
phyllaries, character 9, state 2).
DISCUSSION
Hypochaeris hookeri appears as the sister species of
the ingroup. Future studies, including the rest of the
South American species, must evaluate whether H.
hookeri should properly belong to the H. sessiliflora
complex or not. The results tentatively support
monophyly of the H. sessiliflora complex, but for a
critical morphological assessment of this question, a
cladistic analysis of all South American taxa must be
completed.
The basic structure of relationships (Fig. 2) makes
considerable sense. Hypochaeris sessiliflora is a large,
acaulescent herb that contains conspicuous morpho-
logical variation (Fig. 3) and has a broad range in the
northern Andes (Fig. 4A). Hypochaeris taraxacoides
and H. hohenackeri are very close relatives morpho-
logically (cf. Figs. 5, 6) and are found in similar seeps
above 2600 m in the central Andes (Fig. 4B, squares
and circles). Their evolutionary history must be
closely entwined.
Hypochaeris mucida and H. eriolaena are both very
tomentose with sessile heads (Figs. 7, 8). They are
easily distinguished from each other in features of the
involucre, corolla vascularization, and cypselar wall.
Hypochaeris mucida is a dwarf herb, up to only 1.5 cm
tall, densely pubescent, with shaggy and whip
trichomes on the backs of the phyllaries and leaves,
and growing above 4500 m in grassy steppes
associated with Pycnophyllum J. Re
´my (Caryophylla-
ceae Juss.) in southeastern Peru and northwestern
Bolivia on the eastern side of Lake Titicaca (Fig. 4B,
triangles). Hypochaeris eriolaena has whip trichomes
on the backs of the phyllaries and leaves (or glabrous),
and it inhabits dry places (between 2200 and 5100 m)
from Peru to Bolivia (Fig. 4C, squares).
The other rosette taxa with divided leaves,
Hypochaeris meyeniana,H. echegarayi, and H.
eremophila, are closely related. Morphologically, the
last two are similar (Figs. 9, 10), both with relatively
short peduncles, very divided leaves, phyllaries with
long trichomes on the midribs (Figs. 10C, D; 11C–E),
and plumose pappus bristles. They have different and
consistent floral colors: white versus yellow, respec-
tively. They are also both found in the junction
between southern Peru, southwestern Bolivia, and
northwestern Argentina. It is very likely that these two
taxa share a common origin. Hypochaeris meyeniana is
similar to the other two, but its heads are subsessile,
leaves less divided, phyllaries nearly glabrous, and
pappus bristles with very short side trichomes
(Fig. 11).
Hypochaeris acaulis, which occurs only in the
southern Andes (Fig. 13), is geographically far
removed from the other taxa of the H. sessiliflora
complex. Although morphologically H. acaulis fits
comfortably within this complex (it can be differen-
tiated from the other species of this complex by its
broader phyllaries and rostrate cypselae; Fig. 12; see
also key), recent AFLP data do not support this
inclusion. Based on a broad sampling of taxa of
Hypochaeris within South America (but not including
H. mucida due to lack of material), Tremetsberger et
al. (2006) have shown that H. acaulis does not at all
relate to the H. sessiliflora complex, but instead to H.
palustris and H. tenuifolia, which are also found at
high elevations along volcanoes of the southern Andes
in Chile and Argentina (Tremetsberger et al., 2003a,
b; Muellner et al., 2005). This makes sense geo-
graphically. Hypochaeris acaulis appears to represent
a case of extreme morphological parallelism as
adaptation for survival against the harsh environment
(especially low temperatures and wind). Hypochaeris
acaulis is included here for convenience; it will be
separated in a future comprehensive treatment of the
entire South American complex of the genus.
Considering the classical biogeographic division of
the Andes into three portions (Morrain, 1984; Taylor,
1991), the northern Andes (10uNto3uS), central
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The Hypochaeris sessiliflora Complex (Asteraceae)
Andes (3uSto18uS), and southern Andes (18uSto
54uS), only H. sessiliflora is present in the northern
Andes (Fig. 4A). Hypochaeris acaulis is the only taxon
in the southern Andes; as discussed above, it appears
to have had an independent evolutionary history. Most
of the species of the complex occur in the central
Andes; the major evolutionary and biogeographic
differentiation must have occurred in this general
region. Exactly what the mechanisms of speciation
and biogeography have been in this subgroup remain
to be determined; they likely have involved dispersal
throughout the zone, isolation due to diverse topog-
raphy, adaptation to local ecologic conditions, and
eventual local speciation. Suspected hybridization
between H. echegarayi and H. meyeniana (pers. field
obs.) in western Bolivia emphasizes the evolutionary
closeness of this group and perhaps also its recent
evolutionary origins.
TAXONOMIC TREATMENT
Hypochaeris L. Sp. Pl. 2: 810. 1753. TYPE:
Hypochaeris glabra L.
The genus Hypochaeris is characterized by the
presence of latex, ligulate corollas, receptacular
paleae, and a plumose pappus. It comprises approx-
imately 60 species with an important center of
diversification in South America (where more than
2/3 of the species occur). The rest of the taxa inhabit
the Old World and Asia. All South American species
fall under section Achyrophorus, characterized by the
presence of a uniseriate pappus.
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Figure 2. Most parsimonious tree of 40 steps (fit 5110; fit* 557%), showing all characters (numbers above line) and
states (numbers below line; see Appendix 1) for clades. Closed circles indicate synapomorphies; open circles indicate
homoplasies (parallelisms or reversals). The numbers in italics are jackknife support values greater than 50%taken from a
strict consensus tree.
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Figure 3. Hypochaeris sessiliflora. —A. Habit. —B–D. Variation in capitula and phyllaries. —E. Setaceous bract. —F–J.
Phyllaries. K–M. Trichomes occurring on some phyllaries. —K. Glandular trichomes. —L. Shaggy trichomes. —M. Conical
trichomes. —N. Corolla with anthers and style. —O. Palea, cypselae, and pappus. —P. Smooth achenial wall. —Q, R. Scaly
cypselar wall. A, C, F–I, N–P from Cuatrecasas et al. 25599 (US); B from Stuessy et al. 18538 (LP); and D, E, J–M, Q, R from
Plowman 1955 (US).
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The Hypochaeris sessiliflora Complex (Asteraceae)
The Hypochaeris sessiliflora complex comprises
nine South American species of dwarf (1–13 cm tall),
perennial, acaulescent herbs, with solitary capitula
that are sessile or pedunculate (shorter than the
leaves), which inhabit dry and humid places along the
Andean mountains, from southern Venezuela to
central-eastern Chile and northwestern Argentina,
between 1430 and 5100 m.
KEY TO SPECIES OF THE HYPOCHAERIS SESSILIFLORA COMPLEX
1a. Outer phyllaries broadly ovate (as wide as long); cypselae rostrate, proximally scaly . ............. 9.H. acaulis
1b. Outer phyllaries usually longer than wide; cypselae erostrate or narrower near apices, scaly or smooth.
2a. Involucre cylindric; corollas white . . ....................................... 2.H. taraxacoides
2b. Involucre campanulate-cylindric, campanulate to hemispheric, corollas yellow (white in H. echegarayi,
sometimes white in H. sessiliflora).
3a. Plants to 1.5 cm tall; phyllaries with both whip and shaggy trichomes ................. 4.H. mucida
3b. Plants 3–13 cm tall; phyllaries glabrous or with only one type of trichome.
4a. Leaves undivided (margins entire or dentate).
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Figure 4. —A. Distribution of Hypochaeris sessiliflora in the northern and central Andes. —B. Distribution in the central
and northern Andes of H. hohenackeri (&), H. mucida (m), and H. taraxacoides (N). —C. Distribution in the central Andes of
H. echegarayi (m), H. eremophila (w), and H. eriolaena (&). —D. Distribution of H. meyeniana in the central Andes.
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5a. Outer phyllaries lanuginous (whip trichomes) toward the apex ............... 5.H. eriolaena
5b. Outer phyllaries glabrous or setulose (shaggy trichomes).
6a. Leaves linear-lanceolate or elliptic-lanceolate . . . ................... 1.H. sessiliflora
6b. Leaves oblong .......................................... 3.H. hohenackeri
4b. Leaves pinnatipartite to pinnatisect.
7a. Corollas white ............................................... 7.H. echegarayi
7b. Corollas yellow.
8a. Heads usually sessile; involucre campanulate . . . ................... 6.H. meyeniana
8b. Heads usually pedunculate; involucre cylindric-campanulate . . .........8.H. eremophila
1. Hypochaeris sessiliflora Kunth, Nov. Gen. Sp.
[HBK], folio ed. 4: 2. 1818, quarto ed. 4: 2. 1820.
Oreophila sessiliflora (Kunth) D. Don, Trans.
Linn. Soc. London 16: 178. 1830. Achyrophorus
sessiliflorus (Kunth) DC., Prodr. (DC.) 7: 95.
1838. TYPE: Ecuador. Prov. Pichincha: ‘‘crescit
in alta convalli Quitensi, juxta montem ignivo-
mum Pichincha, alt. 1500 hex.’’ [ca. 3000 m], 14
Apr. 1802 [Sandwith, 1926], F. W. H. A. von
Humboldt & A. J. G. Bonpland s.n. (holotype, P!,
P photo MO!). Figure 3.
Hypochaeris sonchoides Kunth, Nov. Gen. Sp. [HBK], quarto
ed. 4: 2. 1818, quarto ed. 4: 2. 1820. Achyrophorus
sonchoides (Kunth) DC., Prodr. (DC.) 7: 95. 1838.
Achyrophorus quitensis Sch. Bip. var. sonchoides
(Kunth) Wedd., Chlor. And. 1: 219. 1857. TYPE:
Ecuador. Prov. Pichincha: ‘‘crescit cum praecedente,’’
14 Apr. 1802 [Sandwith, 1926], F. W. H. A. von
Humboldt & A. J. G. Bonpland s.n. (holotype, P!, P
photos LP!, MO!).
Achyrophorus barbatus Sch. Bip., Jahresber. Pollichia 16–17:
48. 1859. Achyrophorus sessiliflorus (Kunth) DC. var.
barbatus (Sch. Bip.) A. Gray, Proc. Amer. Acad. Arts 5:
146. 1861. Hypochaeris barbata (Sch. Bip.) Reiche,
Anales Univ. Chile 116: 589. 1905. TYPE: Colombia.
Dept. Norte de Santander: ‘‘Nova Granada, prov.
Pamplona in Paramo de San Urban,’’ 11,000 ft., Dec.
1842, J. J. Linden 746 (lectotype, designated here, P!;
isotypes, BM [2]!, K [2]!, P!, W!, W photo MO!).
Hypochaeris stuebelii Hieron., Bot. Jahrb. Syst. 21: 373.
1896. TYPE: Ecuador. Prov. Napo: ‘‘crescit in Pa
´ramo
montis Antisana,’’ Oct. 1871, M. A. Stu
¨bel 184b
(holotype, B [presumably destroyed], B photos LP!,
MO!).
Herbs to 13 cm tall. Leaves linear-lanceolate or
elliptic-lanceolate, 20–130 35–25 mm, attenuate in
a broad petiole with conical trichomes, margins
dentate to slightly dentate, both surfaces glabrous or
with some glandular trichomes. Capitula sessile or
pedunculate (to 10 cm); bracts on peduncle linear,
with long shaggy trichomes to 6 mm or glabrous;
involucres campanulate to hemispheric, 13–25 310–
25 mm; phyllaries 3- to 4-seriate; outer and middle
phyllaries oblong-lanceolate to lanceolate, 7–15 32–
4 mm, with shaggy trichomes on midribs to near apex
or glabrous, at margins ciliate; inner phyllaries linear-
lanceolate to linear, 11–25 32–4 mm, glabrous or
sparsely ciliate at margins, often divided at the apex;
paleae 12–23 mm; florets ca. 25. Corollas usually
yellow or white, rarely orange or purple, 12–27 mm;
tube 5.5–13 mm; ligule 6.5–14 mm; vascularization
type 1; stamens 7–19 mm; anthers 2.5–9 mm; basal
appendages types 2 and 3, 0.5–1 mm; filaments 5–
14 mm; antheropodium types 1 and 2. Styles 9–
22 mm with branches 1–2.5 mm. Cypselae 5-ribbed,
semirostrate, rostrate, or erostrate, 2–7 mm; walls
scaly or smooth; pappus 7–18 mm. Chromosome
numbers 2n58, 16 (Jansen & Stuessy, 1980; Olsen,
1980; Weiss et al., 2003; Weiss-Schneeweiss et al.,
2007, 2008).
Habitat and distribution. Hypochaeris sessiliflora
has been found from Venezuela to Bolivia, in drier
places, between 2000 and 4500 m (Fig. 4A).
Phenology. Hypochaeris sessiliflora flowers
throughout the year.
Common names. ‘‘Achicoria’’ (Ecuador, Pen
˜afiel
et al. 382); ‘‘cachu-cachu’’ (Peru, Macbride &
Featherstone 1791); ‘‘chicorea’’ (Colombia, Soejarto
494); ‘‘chicoria’’ (Venezuela, Jahn 146); ‘‘chicoria
blanca’’ (Venezuela, Aristeguieta 2438). The white
latex is reported as being used for toothache (Ecuador,
Ellemann 91672).
Morphological characters. Hypochaeris sessiliflora
is distinguished from other species of the complex by
having leaves that are linear-oblanceolate (or less
often elliptic-oblanceolate), glabrous, and toothed or
slightly toothed at the margin, and cypselae that are
cylindrical.
Hypochaeris sessiliflora is the most polymorphic
with respect to vegetative and reproductive characters
(especially shape and pubescence of leaves and
phyllaries, and variation in floral color) of all members
of the H. sessiliflora complex (Fig. 3B–D). The heads
can be pedunculate (to 10 cm) or sessile, the outer
phyllaries can have the margin hyaline or opaque, and
in some populations (described as H. stuebelii by
Hieronymus, 1896) the phyllaries are inflated with the
apex either divided or entire. Another variable
character is the color of corollas, with white, yellow,
or rarely purple often encountered in the same
population. Hypochaeris sessiliflora var. barbata
(described as Achyrophorus barbatus, Schultz Biponti-
nus, 1859) was characterized by having linear
peduncular bracts with long shaggy trichomes to
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Figure 5. Hypochaeris taraxacoides. —A. Habit. —B. Leaf. —C–E. Phyllaries. —F. Trichomes (in cilia) on phyllaries. —
G. Corolla with anthers and style. —H. Palea, cypselae, and pappus. —I. Smooth cypselar wall. From Stuessy et al. 18074 (LP).
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Figure 6. Hypochaeris hohenackeri. —A. Habit. —B–E. Phyllaries. —F. Cilia on phyllaries. —G. Corolla with anthers
and style. —H. Palea, achene, and pappus. —I. Smooth cypselar wall. From Lewis 35168 (LP).
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Figure 7. Hypochaeris mucida. —A. Habit. —B–E. Phyllaries. —F. Whip trichomes on phyllaries. —G. Shaggy
trichomes on phyllaries. —H. Trichomes (in cilia) on phyllaries. —I. Corolla with anthers and style. —J. Palea, cypselae, and
pappus. —K. Smooth cypselar wall. From Ceballos et al. 614 (SI).
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Figure 8. Hypochaeris eriolaena. —A. Habit. —B. Leaf. —C–F. Phyllaries. —G. Trichomes (in cilia) on phyllaries. —H.
Whip trichomes on phyllaries. —I. Corolla with anthers and style. —J. Palea, cypselae, and pappus. —K. Smooth cypselar
wall. From Smith & Buddensiek 11133 (LPB).
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Figure 9. Hypochaeris echegarayi. —A. Habit. —B–D. Phyllaries. —E. Shaggy trichomes on phyllaries. —F. Trichomes
(in cilia) on phyllaries. —G. Corolla with anthers and style. —H. Palea, achene, and pappus. —I. Smooth cypselar wall. —J.
Scaly cypselar wall. A–H, J from Valenzuela 998 (LPB); I from Hunziker & Caso 6039 (LP).
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Figure 10. Hypochaeris eremophila. —A. Habit. —B. Leaf. —C–E. Phyllaries. —F. Shaggy trichomes on phyllaries. —G.
Trichomes (in cilia) on phyllaries. —H. Corolla with anthers and style. —I. Palea, cypselae, and pappus. —J. Scaly cypselar
wall. A, C–J from Stuessy et al. 18064 (LP); B from Fabris 1369 (LP).
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The Hypochaeris sessiliflora Complex (Asteraceae)
6 mm, setaceous phyllaries with yellow-green and
purple trichomes, and outer and middle phyllaries
that are divided at the apex (Fig. 3D). Despite these
conspicuous morphological features, this form like-
wise seems to have no geographic integrity; we deem it
not worthy of formal recognition. We have seen
variation in capitula as represented in Figure 3B–D
within a single population on Volca
´n Pichincha
(Stuessy et al. 18539) in Ecuador. Schultz Bipontinus
(1845: 120) also struggled with the morphological
variation within this species (as A. quitensis, nom.
illeg.). He recognized a set of unnamed ‘‘Formae’’ in
the following manner: I, entire plant glabrous; IIa,
heads large; IIb, heads half as small; IIIa, plants
subacaulescent; IIIb, plants caulescent. This is a
variable species. More populational sampling within
its broad geographic range using morphometric and/or
molecular analyses would be worthwhile.
Observations. Hypochaeris sessiliflora and H.
sonchoides were published by Kunth (1818) in the
same volume (even on the same page), and either
could be selected for use (McNeill et al., 2006: Art.
11.5). The former is the much more widely used name,
however, and therefore it has been retained.
Schultz Bipontinus (1845: 120) published the
names Achyrophorus quitensis Sch. Bip., Scorzonera
quitensis Humb., and S. sessilis Humb. under
Hypochaeris sessiliflora. Because they were not
accompanied by descriptions or specimens, they
remain prosynonyms. Hypochaeris sessiliflora f. cau-
lescens Hieron. (Bot. Jahrb. Syst. 28: 658. 1901) is
another invalid name because it was published
without description.
Schulz Bipontinus (1859: 52) also published
Achyrophorus albiflorus Sch. Bip. and A. humboldtii
Sch. Bip., but both are nom. illeg., being superfluous
names of A. quitensis Sch. Bip., which itself is
illegitimate, having been cited in synonymy of
Hypochaeris sessiliflora. Bortiri (1997) lectotypified
both A. albiflorus and A. humboldtii, but being
illegitimate, this is unnecessary. Hypochaeris sessili-
flora var. albiflora Hieron. (Bot. Jahrb. Syst. 28: 658.
1901), therefore, is also an illegitimate name.
Reiche (1905) may have had a confused concept of
Hypochaeris barbata, as he cited it as occurring in
northern Chile (Prov. [5Region] Tarapaca
´), where H.
sessiliflora is not known to occur (Fig. 4A); he
probably had material of H. eremophila, as he cited
Distoecha taraxacoides Phil. in synonymy.
Representative specimens. BOLIVIA. Cochabamba: Tu-
naria [probably Cerro Tunari], Mu
¨sch 151 (p.p., SI).
COLOMBIA. Boyaca
´:entre Gachoque y Toca, Zuloaga &
Landono 4136 (SI). Caldas: Nevado del Cocuy, alto valle de
Las Lagunillas, Cuatrecasas 1484 (US). Cauca: Cordillera
Central, E slopes of pa
´ramo del Purace, around la laguna de
San Rafael, Cuatrecasas & Willard 26299 (US). Cundina-
marca: Pa
´ramo de Chisaca
´,Soderstrom 1254 (NY).
Magdalena: Sierra Nevada de Santa Marta, valley descend-
ing SW from Pico Reina and Ojeda, Cuatrecasas & Romero
Castaneda 24560 (US). Narin
˜o: Pasto, volca
´n de El Galeras
above Pasto, Ewan 16324 (US); Pasto. Volca
´n Galeras,
Plowman 1955 (US); University of Narin
˜o property, vic.
Granja Botana, 3.5 km down rd. from entrance, ca. 7 km S of
Pasto, Leiteyn, Dumont & Buritica 4995 (NY). Santander:
Pa
´ramo de Mogotocoro, near Vetas, Killip & Smith 17641
(US); Molina & Barkley 18 S.388 (US). ECUADOR. Azuay:
Cuenca, Parroquia Ban
˜os, hacienda de Yanasacha, Boecke &
Jaramillo 2493 (US). Azuay-Loja: Nudo de Cordillera
Occidental y Cordillera Oriental entre On
˜a y Rancho
Ovejero, Barclay & Juajibioy 8472 (US). Carchi: Julio
Anrade-playo
´n de San Francisco rd., La Palestina, Boelke &
Jaramillo 2348 (NY). Chimborazo: along the rd. to
Riobamba, ca. 20 km SW of Riobamba, King & Garvey
6968 (US). Cotopaxi: 5.5 km E of Pujilı
´,Stuessy et al.
18549 (LP, WU). Imbabura: Cotacachi Canton, Res. Ecol.
Cotacachi-Cayapas, laguna de Cuicocha, Pen
˜afiel et al. 382
(US). Loja: 12 km NW of Saraguro on loma Paredones,
Ellemann 91672 (US). Napo: Tena Canto
´n, Parque Nac.
Llaganates, vı
´a Salcedo-Tena, de laguna Chaloa Cocha
desvı
´o a Rayo Filo, Vargas, Narva
´ez & Orellana 2619 (NY,
US). Napo-Pastaza: alrededores de Los Llanganati, entre
Ainchilibı
´yRı
´o Portrero, E de Romo Pa
´ramo, Barclay &
Juajibioy 9210 (US). Pichincha: 23 km E of Pifo, Stuessy et
al. 18540 (WU). Tungurahua: Patate Canto
´n, Parque Nac.
Llanganates, laguna de Aucacocha-cerro Pan de Azu
´car,
Vargas, Ronquillo & Granda 2887 (US). PERU. Amazonas:
Chachapoyas, Quebrada Molino, 5 km below Chachapoyas,
Wurdack 761 (US). Ancash: Huaylas, distr. Pamparnas, path
Karka to laguna Negra Huacanam, around the laguna itself,
Weigend 2000/616 (NY). Celendı
´n: 17–24 km from Cel-
endı
´n, Smith & Cabanillas 7328 (US). Cuzco: Cuzco,
alrededores de las ruinas de Sacsahuama
´n, Beck 8356 (LPB).
Junı
´n: Huancayo, Killip & Smith 23369 (US). Lima:
´o
Blanco, Killip & Smith 21680 (US). Puno: Cavasaya, hda.
Sojela, Vargas 21784 (LP). VENEZUELA. Apure: a lo largo
del Rı
´o Talco (Oira
´) y sus afluentes, en pa
´ramos entre Alto
de Cruces y Tierra Negra, Steyermark & G. C. K. & E.
Dunsterville 101105 (NY). Me
´rida: Me
´rida, pa
´ramo at
laguna de Mucubaji, near Apartaderos and San Rafael,
35 km NE of Me
´rida, Maguire 39405 (NY).
2. Hypochaeris taraxacoides Ball, J. Linn. Soc.,
Bot. 22: 48. 1885. Oreophila taraxacifolia Meyen
& Walp., Nov. Actorum Acad. Caes. Leop.-Carol.
Nat. Cur. 19(suppl. 1): 291. 1843, non Oreophila
taraxacifolia Loisel., 1827. TYPE: ‘‘Peruvia: in
planitie circa Tacoram,’’ 14,000–17,000 ft.,
1831. F. J. F. Meyen s.n. (holotype, B [presumed
destroyed, Stafleu & Cowan, 1981]). Figure 5.
Hypochaeris stenocephala (A. Gray ex Wedd.) Kuntze var.
integrifolia Kuntze, Revis. Gen. Pl. 3(2): 160. 1898.
Hypochaeris stenocephala (A. Gray ex Wedd.) Kuntze
var. taraxacoides (Meyen & Walp.) Kuntze, Revis. Gen.
Pl. 3(2): 160. 1898. TYPE: Bolivia. Dept. Oruro: Oruro,
1892, C. E. O. Kuntze s.n. (holotype, NY!).
Herbs to 7 cm tall. Leaves lanceolate or oblong,
1.5–11 30.3–1.2 cm, pinnatifid to lobate, glabrous,
at base attenuate. Capitula pedunculate to 40 mm;
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involucre cylindrical, 14–23 35–10 mm; phyllaries
3- to 4-seriate, lanceolate, apex rounded, margin
ciliate; outer phyllaries 8–11 32–3 mm; inner
phyllaries 15–19 33–3.5 mm; paleae 12–18 mm;
flowers ca. 30 per capitula. Corollas white (upper
surface) to dark blue-black at tips of ligules
underneath, 15–21 mm; tube 7–9 mm; ligule 8–
12 mm; vascularization type 2; stamens 10–17 mm;
anthers 4–5 mm; basal appendages type 3, ca. 1 mm;
filaments 6–12 mm; antheropodium type 1; style 12–
22 mm, with style branches 1.8–2 mm. Cypselae
erostrate, 1.8–2 mm; wall smooth; pappus 12–
17 mm. Chromosome number 2n58, 16 (Parker,
1971; Weiss et al., 2003 [as Hypochaeris stenoce-
phala]; Weiss-Schneeweiss et al., 2007, 2008).
Habitat and distribution. Hypochaeris taraxa-
coides inhabits bogs and seeps from Colombia to
northern Chile and northwestern Argentina, 2640–
5000 m (Fig. 4B, circle).
Phenology. Hypochaeris taraxacoides flowers
throughout the year.
Common names. ‘‘Achicoria’’ (Argentina, Haber
142); ‘‘chicoria’’ (Peru, Mexia 4191); ‘‘diente de leo
´n’’
(Bolivia, Schulte 9); ‘‘flor de cie
´naga’’ (Argentina,
Cabrera et al. 22484); ‘‘lechero’’ (Argentina, Haber 5);
‘‘mula-siki’’ (Bolivia, Wolstenholme 6); ‘‘pilly’’ (Peru,
Mexia 4191).
Morphological characters. Hypochaeris taraxa-
coides is readily distinguishable from other taxa by
the combination of peduncular (rarely sessile) cylin-
drical heads, glabrous phyllaries (ciliate on margins)
and leaves, and corollas white on the upper surface to
dark blue-black at the tips of ligules underneath
(abaxial surface), and being longer than the involucre.
Observations. Although no isotype has been
located, the detailed description and locality combine
to leave no doubt as to the biologic affinity of this
name. The locality of plains near ‘‘Tacoram’’ presum-
ably refers either to the town of Tacora and/or Volca
´n
Tacora, both of which are now within the boundaries
of Chile, Region I, Parinacota Province (just across
the border from Peru), Tacna Department. At the time
of collection and for another half century, this region
was within Peru (e.g., Enock, 1908: cf. map).
Ratzeburg (1843: xviii) tells us that Meyen went up
from Arica, Chile, toward Lake Titicaca, and Tacora is
on this route. This species is known from this general
region (Fig. 4B, circle).
In publishing Achyrophorus taraxacoides Walp.
(Repert. Bot. Syst. 6: 336. 1846, nom. illeg.) [non A.
taraxacoides (D. Don) Steud. (Steudel, 1841: 226],
Walpers (1846) purported to make a new combination
based on Oreophila taraxacoides Meyen & Walp., but
gave reference to the place of publication of O.
taraxacifolia Meyen & Walp. Whether or not this
incorrect epithet used in the combination should be
treated as an error to be corrected to ‘‘A. taraxacifolia’’
or considered deliberate in view of the existence of A.
taraxacifolia Moench (Moench, 1802) is unimportant
because, in either case, the name is illegitimate, with
Steudel having validly published the name A.
taraxacoides five years earlier. Hypochaeris taraxa-
coides (Walp.) Benth. & Hook. f. (Gen. Pl. 2: 519.
1873) has also been used in some reports (e.g., Bortiri,
1999, as synonym) and in much herbarium material,
but this also uses the incorrect epithet in combination.
More importantly, Bentham and Hooker (1873: 519)
did not actually make the combination. They only
listed ‘‘. . . in Achyrophoro taraxacoide, Walp.’’ under
Hypochaeris (as ‘‘Hypochoeris’’), a particular technique
in this work that is specifically disallowed as a new
combination in Article 33.1 of the International Code
of Botanical Nomenclature (and exemplified in
McNeill et al., 2006: Art. 33, Ex. 2). The first
publication of a legitimate name for the species in
Hypochaeris followed shortly thereafter: H. taraxa-
coides Ball (J. Linn. Soc., Bot. 22: 48. 1885). It is
curious that Walpers, one of the two original authors,
cited the original epithet taraxacifolia incorrectly
when transferring it into Achyrophorus, and in this was
followed by Bentham and Hooker and Ball under
Hypochaeris. They may have been aware of the earlier
competing name, H. taraxacifolia Moench (Suppl.
Meth. 224. 1802), now referable to Crepis albida Vill.
(Babcock, 1947: 310) or to the earlier O. taraxacoides
D. Don, which does not belong to the H. sessiliflora
complex, but there is no evidence of this. It is
fortunate for maintaining stability of usage that H.
taraxacoides Ball can be treated as a new name in
1885, for a new combination based on the epithet that
was actually used would have resulted in the
illegitimate H. taraxacifolia (Meyen & Walp.) Ball,
a later homonym of H. taraxacifolia Moench (Moench,
1802). An earlier H. taraxacoides Pourr. ex Steud.
(Nomencl. Bot. [Steudel], ed. 1, 422, 618. 1821) also
exists, but appears as a synonym of Picridium albidum
DC. (5Crepis albida Vill., Babcock, 1947: 310); the
name is not validly published and so creates no
competing difficulty.
Achyrophorus stenocephalus A. Gray ex Wedd. is an
invalid name because it was listed by Weddell (1857)
as synomyn under Achyrophorus taraxacoides (Meyen
& Walp.) Walp. Hypochaeris stenocephala (A. Gray ex
Wedd.) Kuntze [Kuntze, 1898]), therefore, is an
illegitimate combination.
The characters used by Kuntze (1898: 160) to
differentiate Hypochaeris stenocephala var. integrifolia
from variety taraxacoides are leaf margins entire or
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denticulate versus runcinate, respectively, which were
essentially validating the listed, but not named,
varieties in Achyrophorus taraxacoides (Meyen &
Walp.) Walp. by Weddell (1857). This variable
feature (leaf margin) is insufficient for varietal
recognition.
The name Hypochaeris taraxacoides (Meyen &
Walp.) Ball var. lanuginosa Herzog is a nomen
nudum because it was only mentioned in a list of
Bolivian plants published by Herzog (1923: 228).
Representative specimens. ARGENTINA. Catamarca:
Antofagasta de la Sierra Hoyada de Antofagasta, Haber 5
(SI). Jujuy: Capital, refugio del nevado de Chan
˜i, Fabris,
Cano & Tello 4035 (LP); Cochinoca, Abra Pampa, cerro
Huancar, Cabrera et al. 15256 (LP); Humahuaca, 31.4 km W
of Huamahuaca on gravel rd. to El Aguilar, Stuessy, Urtubey
& Tremetsberger 18089 (LP, WU); Humahuaca, 0.8 km NW
of Chiapi Rodeo on dirt rd. to Iruya, Stuessy, Urtubey &
Tremetsberger 18074 (LP, WU); 4.7 km NW of Chiapi Rodeo
on dirt rd. to Iruya, Stuessy, Urtubey & Tremetsberger 18075
(LP, WU); Iturbe, Meyer, Cuezo & Legname 21040 (LP);
Rinconada, Mina Pirquitas, Schwabe, Ancibor & Vizinis 641,
818,819 (LP); Tilcara, lecho de Rı
´o Grande cerca de
Huacalera, Werner 820 (LP); Tumbaya, camino de El Angosto
al Chan
˜i, encrucijada Rı
´o Chan
˜i, Cabrera et al. 22484 (LP);
Valle Grande, Caspala
´,Burkart & Troncoso 11822 (SI); Yavi,
Quebrada de Toquero, A. L. & S. M. Cabrera, Malacalza &
von Schmiden 17656 (LP); Lecho, 1–2 km S RP 5, Km 13, 4–
5 km SW de Yavi, Tolaba, Acun
˜a, Arapa, Gutie
´rrez, Quiroga,
Ragno, Ramallo & Da Silva 1539 (MCNS). La Rioja:
Famatina, Sierra de Famatina, Rohweder T-12 (LP). Salta:
Los Andes, Vega de Tocomar, Cabrera et al. 22531 (LP);
Poma, Quebrada entre Mun
˜an
˜o y Tastil, Cabrera 8993 (LP);
San Antonio de Los Cobres, San Antonio de Los Cobres,
Krapovickas 3155 (K, SI); Santa Victoria, 43.7 km from Santa
Victoria on rd. to La Quiaca, Hawkes, Hjerting & Rahn 3900
(K, LP). Tucuma
´n: Tafı
´, Cerro Mu
´n
˜oz, La Banda, Fabris
1520 (LP). BOLIVIA. Cochabamba: Chapare, Quebrada de
Colomi, Balls B-6253 (K, UC); Quillacollo, 63 km de
Cochabamba en direccio
´n al poblado de Cami, alrededores
del cerro Pytuljata, Liberman 2343 (LPB, US); Tapacarı
´,
Qachi un
˜kata, cerca del Rı
´o Chankha, 3 km al E de Japo
K’asa, Pestalozzi 284 (LPB). La Paz: Aroma, Kanton
Callamarca, 6 km no
¨rdlich Calamarca an der Straße La
Paz-Oruro, Krach 7492 (SI); Camacho, Puerto Acosta, 10 km
hacia La Paz, al borde del Rı
´o Huaycho, cerca del Lago
Titicaca, Beck 7712 (SI); Fray Tamay, Pelechuco, Krach
9391 (SI); Ingavi, low ridge along the main rd. from La Paz to
Viacha, ca. 8 km NE of Viacha, Solomon & Nee 14233 (LPB,
NY); La Paz, Palca, zona basal del Illimani, Ceballos et al.
548 (SI); Pacajes, La Paz entre Corocoro y Topohoco,
Ceballos et al. 164 (SI); Los Andes, Kanton Pen
˜as, Straße zur
Mina Fabulosa, Km 6.5 von der Abzweigung von der Straße
La Paz-Huarina, Krach 8435 (SI); Larecaja, viciniis Sorata,
inter Ancohuma et Turilaque, Mandon 276 (BM, K, W);
Murillo, 4.6 km S jct. rd. to Chacaltaya on rd. from Milluni to
El Alto (La Paz), Solomon 13177 (LPB); Nor Yungas, Canton
Pocollo, La Cumbre, Krach 8672 (SI); Omasuyos, Canto
´n
Huarina, comunidad de Moco-Moco, Loza de la Cruz 72
(LPB); Tamayo, Ulla-Ulla, estribaciones de la cordillera de
Apolobamba, Menhofer 1036 (LPB); Saavedra, bei Curva
großes Moor oberhalb der Endnora
¨nen oberhalb Canizaya,
Freueler 4384 (SI); Tamayo, Ulla-Ulla, Okaria, Menhofer
1600 (LPB). Oruro: Sajama, Canto
´n Lagunas, puna y
vegetacio
´n alto andina, Loza de la Cruz 254 (LPB). Potosı
´:
Frı
´as, cerro Khare-Khare detra
´s de la ciudad, Schulte 9
(LPB). Tarija: Jose
´Marı
´a Avilez, Pampa Tajzara, Arenales,
Beck & Paniagua 27086 (LPB). COLOMBIA. Boyaca
´:
Cordillera Oriental Pa
´ramo de Huma, entre Bele
´n y Susaco
´n,
Barclay & Juajibioy 7642 (US). PERU. Ancash: along Peru
´
hwy. 105 to Chavin de Huantas, ca. 26 km E of Catac, King
& Collins 9064 (US). Arequipa: Arequipa, just SW of Puno
border, Paso del Co
´ndor, Iltis & Ugent 1489 (UC).
Ayacucho: ca. 10 km N of Mataral, on trail to Ayacucho,
West 3663 (UC). Cuzco: Urubamba, Zamalloa Dı
´az 78 (LP);
Chanchis, La Raya pass 1 km WNW of La Raya on Cuzco
rd., Iltis & Ugent 1256 (US); Chumbivilcas, 15 km de Sto.
Toma
´s a Yairi, Hoogte & Roessch 2534 (NY); Paucartambo,
Paucartambo, Accanaco, Balls B6711 (K). Huancavelica:
Huancavelica, Ojapampa, entre Laria y Tambopata, a 25 km
de Conaica, Tovar 850 (LP [2]); Tayacaja, quebradas
westward from Huancavelica, Stork & Horton 10838 (UC).
Hua
´ncayo: Huancayo, 30 km N of Huancayo, Blair 636 (K).
Huanuco: Huanuco, 15 mi. NE of Huanuco, Macbride &
Featherstone 2167 (US). Junı
´n: Concepcio
´n, Km 112, rd.
Concepcio
´n–Satipo, Saunders 1106 (K, UC); Junı
´n, Capilla-
cocha, a 20 km E of Carhuamayo, Tovar 390, 393 (LP);
Llanos de Junı
´n, 76u009W, 11u109S, Smith et al. 5652 (NY);
Tarma, lago Junı
´n, 13 km N of Junı
´n, Hutchison, Wright &
Straw 5892 (UC); Yauli, Jaranacu, cerca de Chacalpa, Ochoa
264 (US). La Libertad: Bolivar, laguna de Loa Ichus
Nevado, Cajamarguilla, Lo
´pez & Saga
´stegui 3223 (LP); Pataz,
Huancaspata-Tajabamba, Lo
´pez & Saga
´stegui 8245 (LP);
Santiago de Chuco, Quesquenda, Jalca Quiruvilca, Lo
´pez &
Saga
´stegui 2888 (LP). Lima: Canta, Carhuapampa (camino a
Canta–Lachaqui), Mezar 162 (LP); Yauyos, Huancracha
arriba de Tupe, Cerrate & Tovar 1174 (LP). Puno: Patanca,
Fiebrig 3188 (BM, K).
3. Hypochaeris hohenackeri (Sch. Bip.) Domke,
Notizbl. Bot. Gart. Berlin-Dahlem 13: 251. 1936.
Basionym: Achyrophorus hohenackeri Sch. Bip.,
Bonplandia 4: 54. 1856. TYPE: Peru. ‘‘Tobina in
Cordill. sum. jug. [summit ridge],’’ July 1854, W.
Lechler 2111a (holotype, P!). Figure 6.
Hypochaeris parvifolia J. Kost., Blumea 5(3): 661, fig. 3c.
1945. TYPE: Bolivia. Dept. La Paz: ‘‘Rosettenpolster
auf Alpenwiesen im Tcacota-Thal’’ [most probably
Teacota River valley, 16u439S, 67u259W, Division of
Geography, 1955; protologue gives ‘‘Tcacota-Thal,’’ but
label ‘‘Teacota-Thal’’], 4300 m, Oct. 1911, T. Herzog
2425c (holotype, L, digital image!).
Herbs to 4 cm tall. Leaves oblong, 20–30 32–
3 mm, base attenuate, margin entire or slightly
dentate, glabrous. Capitula pedunculate to 20 mm
(rarely sessile). Involucre campanulate, 11–15 36–
8 mm; phyllaries 4- to 6-seriate, lanceolate, apex
acute or slightly rounded, margin ciliate; outer
phyllaries 8–9 32.5–3 mm; inner phyllaries 13–
15 3ca. 2 mm; paleae 11–15 mm; florets ca. 15
per capitula. Corollas yellow, rarely pink (Smith
& Cabanillas 7212), 10–20 33–8 mm; ligule
7–12 mm; vascularization type 1; stamens 11.5–
20.5 mm; anthers 8–12 mm; basal appendages type
1 or 2, 0.50–0.75 mm; filaments 3.5–8.5 mm; anther-
opodium type 1 or 2; style 10–12 mm; style branches
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1.5–2.5 mm. Cypselae erostrate, 2–2.5 mm; wall
smooth; pappus 9.5–15 mm. Chromosome number
unknown.
Habitat and distribution. Hypochaeris hohenackeri
occurs from Peru to Bolivia, 3200–4500 m, in humid
grasslands, mountain pastures, riverbanks, and bogs
associated with Juncus L. and Stipa L. (Fig. 4B, squares).
Phenology. Hypochaeris hohenackeri flowers from
January to September.
Morphological characters. Hypochaeris hohenack-
eri is characterized by oblong, sometimes dentate,
leaves, a pedunculate head (rarely sessile), and dark,
glabrous phyllaries that are acute or rounded at the apex.
Observations. Achyrophorus hohenackeri [‘‘Hohe-
nakeri’’] was published earlier by Schultz Bipontinus
(1855: 236) as a nomen nudum. Although without
description, he did cite a specimen (W. Lechler
2111a), which was later taken by him (1856: 54) as
the original specimen to accompany his valid
description (1856: 54) in the same journal. Hypo-
chaeris hohenackeri Herzog (Herzog, 1923) is an
illegitimate name because no basionym was provided.
Although the type specimen of Hypochaeris parvi-
flora J. Kost. is small, there is no doubt of the affinity
of this name; the original description is also
accompanied by a clear illustration that shows the
diagnostic characters of this species.
Representative specimens. BOLIVIA. Cochabamba:
Arani, near the summit of the pass through the cordillera
de Cochabamba, Eyerdam 24839 (UC). La Paz: Inquisivi,
along the trail betw. Pongo Chico and laguna Naranjani, ca.
3 km W of Quime, Lewis 35168 (LPB, NY); W slope of cerro
Kharrata SSE, Camillaya, Mu
¨ller & Heinrichs 6428 (LPB);
1 km NE of Mina Argentina and 10 km S of Coquetanga,
Lewis 38124 (LPB); Saavedra, el camino de Charazani a
Hayrapata, Menhofer X-1832 (LPB, SI). PERU. Amazonas:
Chachapoyas, Chachapoyas–Celendı
´n rd., cerro Calla-Calla,
Smith & Cabanillas 7212 (US); cerro de Calla-Calla, betw.
Leimebamba–Balsas rd. pass and the camino de herradura,
Wurdack 1225 (LP). Huancavelica: Huancavelica, 4 km
from Conayca, Tovar 222 (LP).
4. Hypochaeris mucida Domke, Notizbl. Bot. Gart.
Berlin-Dahlem 13: 250. 1936. TYPE: Bolivia. La
Paz: ‘‘Mittlere Anden: Su
¨dlich des Titicaca-Sees
u
¨ber Ancoraime,’’ 13,500 ft., 12 Feb. 1903, A.
W. Hill 299 (lectotype, designated here, K!).
Figure 7.
Hypochaeris mucida Domke var. integrifolia Cuatrec., Proc.
Biol. Soc. Wash. 77: 156. 1964. TYPE: Peru. Puno:
WSW of Checayani, NE of Azangaro, puno, 4150 m, 29
Mar. 1957, H. Ellenberg 495 (holotype, U not seen;
isotype, US not seen, US photo LP!).
Herbs 1–1.5 cm tall. Leaves oblong, 8–20 31.5–
4 mm, slightly toothed or pinnatifid, adaxial sur-
face sericeous with whip and some shaggy tri-
chomes, abaxial surface with whip trichomes,
petiole sericeous. Capitula sessile. Involucre cam-
panulate, 8–12 39–10 mm; phyllaries 3- to 4-seriate;
outer phyllaries lanceolate, to 12 mm, lanuginous
with shaggy trichomes; inner phyllaries with the
same indument or glabrous, margin ciliate. Paleae
9–11 mm; florets ca. 12. Corolla 9.5–11.5 mm, color
unknown; tube 3.5–5 mm; ligule 6–6.5 mm; vascu-
larization type 1; stamens 9–10.2 mm, anthers 4–
4.2 mm; basal appendages type 1, 0.8–1 mm; fila-
ments 4.5–6 mm; antheropodium type 1; style 8–
10.5 mm, with branches 2–3 mm. Cypselae 5-ribbed,
unbeaked, 2–3 mm; wall smooth; pappus ca. 8 mm.
Chromosome number unknown.
Habitat and distribution. Hypochaeris mucida is
known from Peru (Puno) and Bolivia (La Paz). It
inhabits the Andean steppe at 4150–4700 m, associ-
ated with Pycnophyllum (Caryophyllaceae) (Fig. 4B,
triangles).
Phenology. Hypochaeris mucida flowers from
February to April.
Morphological characters. Hypochaeris mucida is
the smallest species of Hypochaeris, growing to only
1.5 cm tall. The adaxial surfaces of the leaves and
phyllaries are tomentose (with whip and some shaggy
trichomes), giving it a distinctive appearance.
Observations. Cuatrecasas (1964) considered Hy-
pochaeris mucida var. integrifolia to be unique in
having leaves entire to sinuate, but this admittedly
extreme leaf shape is sporadic and shows no
geographic coherence. Hypochaeris mucida var. in-
tegrifolia is not recognized.
The holotype deposited at the Berlin herbarium was
destroyed during World War II; the isotype specimen
at K is here designated as the lectotype.
Specimens examined. BOLIVIA. La Paz: Camacho,
Carabuco, Mina Matilde, Ceballos et al. 614 (SI); Tamayo,
Ulla-Ulla, estribaciones de la cordillera de Apolobamba,
Menhofer X-2197 (LPB). PERU. Puno: ENE of Checayani,
Ellenberg 638 (US, photo LP).
5. Hypochaeris eriolaena (Sch. Bip.) Reiche,
Anales Univ. Chile 116: 589. 1905. Basionym:
Achyrophorus eriolaenus Sch. Bip., Bonplandia 4:
54. 1856. TYPE: Peru. Puno: ‘‘Cordiller. pascuis
sterilibus pr. Azangaro,’’ June 1854, W. Lechler
1754 (lectotype, designated by Bortiri [1997:
228], P!; isotypes, K!, P!). Figure 8.
Achyrophorus cryptocephalus Sch. Bip., Bonplandia 4: 54.
1856. Hypochaeris cryptocephala (Sch. Bip.) Domke,
Notizbl. Bot. Gart. Berlin-Dahlem 13: 251. 1936, as
cryptocephalus. TYPE: Peru. Puno: ‘‘provincia Cara-
baya in cacumine Cordill. Agapata [Ayapata],’’ June
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1854, W. Lechler 1963 (lectotype, designated here, P!;
isotypes, P!, W!).
Hypochaeris spinneri Beauverd, Bull. Soc. Bot. Gene
`ve, ser.
2, 14: 177. f. XIII. 1922 [1923]. TYPE: Peru.
Huancavelica: Huancavelica, above Huancavelica ‘‘in
pascuis petrosis,’’ 4700 m, 1915, E. Godet 38 (holo-
type, presumably NEU, not located).
Herbs to 5 cm tall. Leaves lanceolate or oblongate,
25–70 35–16 mm, base attenuate in a broad petiole,
margin toothed or entire, glabrous to sericeous (whip
trichomes, to 35 mm) or with shaggy trichomes.
Capitula sessile. Involucre campanulate or hemi-
spheric, 10–30 315–25 mm; phyllaries 4- to 5-
seriate; outer phyllaries ovate or lanceolate, lanugi-
nous (with whip trichomes), 5–12 32–3 mm; inner
phyllaries lanceolate to linear, 12–21 31.5–2.5 mm,
lanuginous to glabrous, ciliate; paleae 15–28 mm;
florets numerous. Corollas white or yellow to golden-
yellow, 18–38 mm; tube 8–18 mm; ligule 10–20 mm;
vascularization type 2; stamens 11–22 mm; anthers 3–
7 mm; basal appendages type 1 or 2, 0.7–1 mm;
filaments 8–15 mm; antheropodium type 1 or 2; style
15–26 mm, with branches 1.5–3 mm. Cypselae 5-
ribbed, transversely wrinkled, erostrate or slightly
narrower at the apex, 1.5–4 mm; wall scaly; pappus
11–29 mm. Chromosome number unknown.
Habitat and distribution. Hypochaeris eriolaena
occurs in Peru and Bolivia in dry, stony Andean
ridges, with scattered grass and limestone outcrops,
2200–5100 m (Fig. 4C, squares).
Phenology. Hypochaeris eriolaena flowers from
May to November.
Common names. ‘‘Cebollana’’ (Peru, Angulo &
Lo
´pez 1358); ‘‘qachi tika’’ (Bolivia, Pestalozzi 606).
Morphological characters. Hypochaeris eriolaea is
clearly distinguishable from other members of the H.
sessiliflora complex by long sericeous trichomes on
broad petioles plus lanuginous phyllaries. This
species often has no developed leaves when the
capitulum opens.
Observations. Schultz Bipontius (1859) in his
revision of Hypochaeris (as Achyrophorus) distin-
guished H. cryptocephala from H. eriolaena by
abaxially sericeous phyllaries in the former and
niveous-tomentose phyllaries in the latter. Indument
on phyllaries is a variable character in H. eriolaena,
which can have phyllaries with both lanuginous (with
long whip trichomes) and matted indument; further-
more, sometimes phyllaries in the inner series are
glabrous. Because of this range of variation, we treat
H. cryptocephala as conspecific with H. eriolaena.
Despite no type having been located of Hypochaeris
spinneri, the figure associated with the protologue
leaves little doubt of the biological placement of this
name.
Hypochaeris eriolaena var. hispida Herzog (Herzog,
1923) is a nomen nudum appearing only in a list of
plants from Bolivia.
Representative specimens. BOLIVIA. Cochabamba: Ar-
que, cerca camino a Tacopaya, Ibisch & Rojas 439 (LPB);
Tapacarı
´, Jacha Rancho, comunidad Japo (Km 125 de la
carretera Cbba.–Oruro), Pestalozzi 606 (LPB). La Paz:
Achachicala (cumbre), subiendo 3 km de la mina Kaluyo
hacia la cumbre, Beck 11914 (LPB, SI). CHILE. Tarapaca
´:
Cosapilla, Limani, Wevi 129 (LP). PERU. Ancash: Huara
´z-
Marcara
´,Velarde Nu
´n
˜ez 3225 (LP); Yungay, Yungay-
Llanganuco, Lo
´pez, Saga
´stegui & Aldave 7421 (LP). Caja-
marca: Cajamarca, Majada Pampa, Jalca Alta, Becker &
Terrones 1433 (US). Contumaza
´:Shano
´n (arriba de
Contumaza
´), Saga
´stegui, Alvitez & Mostacero 9006 (SI).
Cuzco: Canchis, en direccio
´n al cementerio (parte alta),
Vargas 11355 (US); Espinar, alrededores de Yauri, Vargas
11350 (p.p.) (US). Huancavelica: Huancavelica, Huayta-
nayoc-Tansiri, Conayca, Tovar 2512 (LP). Junı
´n: Pampa de
Junı
´n, cerro de Pasco and Junı
´n, Canne & Schunke 232 (US).
La Libertad: Otuzco, carretera a Shoreyo, ladera rocosa,
Angulo & Lo
´pez 1358 (LP); Sa
´nchez Carrio
´n, summit above
Aricapampa on rd. to Huamachuco, Hutchison, Wright &
Straw 6267 (K, US); Santiago de Chuco, Huillillas, N of
Cachicadan, Stork & Horton 10000 (UC). Lima: Canta, La
Viuda (cerca de Cullhuay), Meza 207 (LP). Puno: Puno, San
Antonio de Esquilante, Stafford 865 (K); Juliaca, Hillside
Capachica, Penı
´nsula Lake Titicaca, Tutin 1209,1210 (BM).
6. Hypochaeris meyeniana (Walp.) Benth. &
Hook. f. ex Griseb., Abh. Ko
¨nigl. Ges. Wiss.
Go
¨ttingen 19: 199. 1874. Basionym: Oreophila
meyeniana Walp., Nov. Actorum Acad. Caes.
Leop.-Carol. Nat. Cur. 19(suppl. 1): 292. 1843.
Achyrophorus meyenianus (Walp.) Walp., Repert.
Bot. Syst. 6: 336. 1846. TYPE: ‘‘Peruvia: in
planitie circa Tacoram,’’ 14,000–17,000 ft.,
1831 [one label on type, ‘‘4/31,’’ presumably
indicating Apr. 1831, but another label in
another hand gives ‘‘1833’’], F. J. F. Meyen 22
(holotype, B [presumably destroyed], B photos
LP!, MO!). Figure 11.
Achyrophorus meyenianus (Walp.) Walp. var. ciliatus Wedd.,
Chlor. And. 1: 220. 1857. Hypochaeris meyeniana
(Walp.) Benth. & Hook. f. ex Griseb. var. ciliata
(Wedd.) Perkins, Bot. Jahrb. Syst. 49: 232. 1913.
TYPE: Bolivia. Tarija: ‘‘en monte Calama,’’ June 1846,
H. A. Weddell 4017 (lectotype, designated by Bortiri
[1997: 227], P!; isotype, P!).
Hypochaeris meyeniana (Walp.) Benth. & Hook. f. ex Griseb.
var. eriolaenoides Cabrera, Revista Invest. Agric. 11:
410. 1957. TYPE: Argentina. Jujuy: Sierra de Zenta,
4500 m, Feb. 1931, E. Budin s.n. (holotype, LP!;
isotype, LP!).
Hypochaeris meyeniana (Walp.) Benth. & Hook. f. ex Griseb.
var. brachylepis Cabrera, Fl. Prov. Jujuy 10: 677. 1978.
TYPE: Argentina. Jujuy: Yavi, ‘‘alrededores de La
Quiaca,’’ T. Meyer, A. R. Cuezzo & V. Legname 21270
(holotype, LP!; isotype, LP!).
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Figure 11. Hypochaeris meyeniana. —A. Habit. —B–F. Phyllaries. —G. Trichomes (in cilia) on phyllaries. —H. Corolla
with anthers and style. —I. Palea, cypselae, and pappus. —J. Scaly cypselar wall. A–D, G–J from Stuessy et al. 18062 (LP); E
from Meyer et al. 21306 (LP); F from Meyer et al. 21107 (LP).
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Urtubey et al.
The Hypochaeris sessiliflora Complex (Asteraceae)
Herbs to 8 cm tall. Leaves lanceolate-elliptic or
obovate, 15–150 35–40 mm, pinnatisect (rarely
pinnatifid), runcinate, base attenuate, margin toothed
or entire, glabrous or hispid on the margin (or also on
the midrib). Capitula sessile or rarely with a very short
peduncule. Involucre campanulate to hemispheric, 8–
30 38–25 mm; phyllaries 3- to 4-seriate; outer
phyllaries lanceolate to ovate, 7–11 33–7 mm,
attenuate to rounded or acute, glabrous, hirsute
(with shaggy trichomes) or lanuginous (with whip
trichomes), margin ciliate; middle phyllaries lanceo-
late to ovate-lanceolate; inner phyllaries lanceolate,
13–22 32–2.5 mm, glabrous, slightly ciliate; paleae
13–22 mm; florets ca. 25. Corollas yellow, yellow-
orange, or golden-yellow, 13–24 mm; tube 7–11 mm;
ligule 6–13 mm; vascularization types 1 and 2;
stamens 10–22 mm; anthers 4–6 mm; basal append-
ages types 2 and 3, 0.3–1 mm; filaments 6–16 mm;
antheropodium type 1; style 11–22.5 mm, with
branches 2–4.5 mm. Cypselae slightly constricted at
apex, 2–10 mm; wall scaly or smooth; pappus 7–
17 mm. Chromosome numbers 2n58, 16 (Diers,
1961; Weiss et al., 2003; Weiss-Schneeweiss et al.,
2007).
Habitat and distribution. Hypochaeris meyeniana
occurs from Peru to northwestern Argentina, on rocky
and/or sandy slopes, in dry and humid places, 1700–
4800 m (Fig. 4D). Hypochaeris meyeniana grows
sympatrically with H. taraxacoides, but they occupy
different microhabitats, dry places versus bogs,
respectively.
Phenology. Hypochaeris meyeniana flowers
throughout the year.
Common names. ‘‘Achicoria’’ (Argentina, Cabezas
25,Cabrera 7723,12121); ‘‘chambi’’ (Bolivia, Pesta-
lozzi 206); ‘‘kawi kawi’’ (Bolivia, Pestalozzi 939).
Morphological characters. Hypochaeris meyeniana
is polymorphic, with phyllaries ranging from lanceo-
late to ovate, generally glabrous, sometimes with
shaggy trichomes. As a result of these and other
morphological variations, several varieties have been
described within the species. Hypochaeris meyeniana
var. brachylepis was described by Cabrera (1978) to
have phyllaries wider (more than 5 mm) than those
normally encountered in this species. Likewise,
Hypochaeris (as Achyrophorus)meyeniana var. ciliata
was regarded as distinct by Weddell (1857) for plants
with pubescent phyllaries. Variety eriolaenoides was
described by Cabrera (1957) for plants characterized
by deeply runcinate leaves and tomentose phyllaries.
None of these features shows a consistent geographic
pattern to warrant varietal status, and hence we treat
H. meyeniana as a taxon in a broad sense.
Conservation status. Our field studies reveal that
the conservation status of Hypochaeris meyeniana is
very good. We found many and large populations, in
both dry and humid rocky places in puna and prepuna
(with Trichocereus pasacana (F. A. C. Weber) Britton
& Rose, ‘‘cardon,’’ Cactaceae Juss.) in Jujuy Province,
Argentina, and in La Paz Department, Bolivia. In
Cochabamba Department, Bolivia, it occurs in Poly-
lepis tomentella Wedd. (‘‘tabaquillo,’’ Rosaceae Juss.)
woods.
Observation. The type of Achyrophorus meyenia-
nus was probably collected in Chile. See comments on
this same locality under Hypochaeris taraxacoides.
Bortiri (1997) placed Hypochaeris meyeniana
var. ciliata into synonymy of H. sessiliflora,a
referral with which we do not agree. Description of
‘‘feulles roncinees’’ suggests placement here in H.
meyeniana.
Representative specimens. ARGENTINA. Catamarca:
Andalgala
´, subida al cerro Yutuyaco desde Capillitas,
Sleumer 2730 (LP); Bele
´n, faldeos al N de Portezuelo del
´o Blanco, arriba de Granadilla, Sleumer & Vervoorst 2589
(LP, UC). Jujuy: Cachi, camino a Mina Don Otto, 4–5 km al
S de la ruta a Cachi, Novara 10644 (MCNS); Cochinoca,
Abra Pampa, Vignati 445 (LP [2], NY); Humahuaca, 20.7 km
N of Humahuaca on rte. 9, Stuessy,Urtubey & Tremetsberger
18068 (LP [2], WU); Rinconada, Mina Pirquitas, Schwabe,
Ancibor & Vizinis 1098 (LP); Mina Aguilar, II-1968,
Ferna
´ndez 2029 (LP); entre Cachi y San Carlos, camino a
Isonza, del cruce de la R 33 a la bifurcacio
´n del Santuario,
De la Fuente & Dı
´az 10905 (MCNS); San Carlos, Cuesta de
Isonza, 20 km S de Piedra del Molino, Novara & Neumann
9781 (MCNS); Susques, planicie al pie del cerro Tuzgle,
Werner 89, 91, 103, 131 (LP); Tilcara, abra de Yala, Fabris,
Crisci & Petriella 6514 (LP); Tumbaya, Stuessy,Urtubey &
Tremetsberger 18062 (LP [3], WU); Valle Grande, Santa Ana,
Cerro Ronque, Kiesling et al. 581 (LP); Yavi, Quebrada de
Cajas, Cabrera 7826 (LP); Chalguamayoc, 15–20 km SE de
Yavi, 15 km S RP 5, Km 35–38, Tolaba, Acun
˜a, Arapa,
Gutie
´rrez, Quiroga, Ragno, Ramallo & da Silva 1677
(MCNS); Abra Colorada, costados de RP 5, Km 40–45
(camino a Santa Victoria), pasando quebrada de Cajas,
Tolaba, Acun
˜a, Arapa, Gutie
´rrez, Quiroga, Ragno, Ramallo &
Da Silva 1652 (MCNS). La Rioja: Famatina, entre Los
Corrales y Cueva de Pe
´rez, Cabrera et al. 27174 (SI). Salta:
Iruya, Loma
´n, alrededores del pueblo de Delfina Dı
´az, 2 km
al W de San Isidro, 14 km al NW de Iruya, Tolaba, Ragno &
Quiroga 1235 (MCNS); Pantipampa, alrededores del puesto
de Concepcio
´n Bustamante, filos del cerro al N de San Isidro,
12 km NW de Iruya, Tolaba, Ragno & Quiroga 1198
(MCNS); Ora
´n, Caucillar, Hilgert & Lamas 1654 (MCNS);
Poma, Abra Mun
˜an
˜o, Cabrera 8982 (LP); Santa Victoria, RP
7, Km 12, Deginani, Cialdella & Bortiri 806 (SI); Nazareno,
200 m al S del pueblo, Adler 2 (MCNS); San Carlos, Alturas
de Amblayo, valle Isonza, 10 km S de valle Encantado,
Novara 6356 (MCNS). BOLIVIA. Chuquisaca: Colomi, near
Cochabamba, Balls 6232 (K). Cochabamba: Apopaya,
cuenca Rı
´o Tambillo, estancia Pajchanti, Baar 386 (LPB);
Arque, proximidades a la comunidad de Kutimarca y
Sumuruni, camino hacia Arque, Mercado 2181 (K); Cha-
parre, Cerro Guakanquı
´,Steinbach 9696 (K); Cochabamba,
Vacas, Ca
´rdenas 4347 (US); Quillacollo, camino Sipe-Sipe-
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Lipichi, Hensen 789 (LPB); Mizque, Sacha Ioma, Ramı
´rez
203 (LPB). La Paz: Chaparre, about 50 km SE of
Cochabamba, on rd. to Chimore, Eyerdam 25059 (K, UC);
Ingavi, Tihuanaco, Balls B6361 (K, UC); Inquisivi, Cruz
Sayaquira, 10 km SW of Quime, Lewis 37146 (US); Larecaja,
ca. 23 km N of Sorata, along rd. toward Consata, Luteyn &
Dorr 13780 (US); Los Andes, valle de Hichu Kkota, Ostria
132 (LPB); Murillo, La Paz, Calacoto, 60 km hacia Nevado
Illimani, Panguyo, Beck 9060 (SI); Omasuyos, el camino
principal a Pen
˜as, 18 km via mina Fabulosa, Hichukkota,
Beck 2861 (LPB, US); La Paz, Challapampa, Asplund 4886
(US); Pacajes, 39 km from La Paz along rd. to Guaqui, King
& Bishop 7504 (US). Oruro: Saavedra, Kanton Amarete
Felsen enmittelbar no
¨rdlich des Ortes, Krach 8417 (SI).
Potosı
´:above presa Incachaca, 10 km from La Paz pa
´ramo,
Stuessy, Tremetsberger & Ho
¨ssinger 18504 (LP); Chayanta,
Cruz Kasa, Zamora 156 (LPB); Frı
´as, Potosı
´, ‘‘Las
Lecherı
´as,’’ Schulte 129 (LPB). Tarija: Avile
´z, lagunas de
Tajsara, Meyer, Cuezzo & Legname 21517 (LP); Me
´ndez,
cerca Pasajes, Bastia
´n 334 (US); Tarija, de Tarija a Iscayada,
Kiesling et al. 3801,3826 (SI). CHILE. Regio
´nI:Arica,
camino de Arica a Portezuelo de Chapiquin
˜a, Ricardi,
Marticorena & Matthei 343 (CONC); Parinacota, camino
entre Putre y Socoroma, Km 17, Matthei & Rodriguez 290
(CONC). Regio
´n III: Huasco, Vallenar, Rı
´o Laguna Grande,
Werdermann 247 (LP [the label of this collection bears the
herbarium name, Hypochoeris glacialis Weder., which was
never published]). PERU. Ancash: Bolognesi, Ocros, Cerrate
6025 (SI); Huaylas, Distr. Pamparomas, Path Karka to laguna
Negra Huacanam, Weigend 2000/615 (NY). Arequipa:
Arequipa, Paso del Co
´ndor, i.e., pass on continental divide
on Puno–Arequipa Rd., Iltis & Ugent 1495 (US). Ayacucho:
betw. Huanta and hacienda Pargora, Killip & Smith 22191
(US). Cajamarca: Cajamarca, entre la Encan
˜ada y Jalca de
Kumulca, Lo
´pez Vega 1607 (SI); Contumaza
´, Cascabamba
(arriba de Contumaza
´), Saga
´stegui et al. 10006 (US).
Huancavelica: Castrovirreina, near Co
´rdova, Metcalf
30280 (US); Huancavelica, Occopampa, entre Laria y
Tambopata, a 25 km de Conaica, Tovar 842 (LP). Junı
´n:
Jauli, Anticona bajo, Guille
´n & Riccio 10784 (US). Lima:
Canta, Ijadero, a 20 km NE de Canta, Meza 149 (LP);
Huarochiri, Distr. San Mateo, Rı
´o Rimae, Saunders 822 (K).
Puno: Huarochiri, Distr. Matucana, Km 85 on hwy.,
Saunders 379 (BM). Tacna: 20 km above Candasave on
rd. from Mazo Cruz just S of Volca
´n Tutupaca, Weigend &
Fo
¨rther 97-790 (K).
7. Hypochaeris echegarayi Hieron., Bol. Acad.
Nac. Ci. 4(1): 51. 1881. TYPE: Argentina. San
Juan: Barriales de Leoncito, Dec. 1875–Jan.
1876, D. S. Echegaray s.n. (holotype, CORD, not
located; isotype, LP!). Figure 9.
Achyrophorus setosus Wedd., Chlor. Andina 1: 220. 1857.
Hypochaeris setosa (Wedd.) Rusby, Bull. New York Bot.
Gard. 4: 402. 1907, non Hypochaeris setosa Forma
´nek,
1897. TYPE: Bolivia. Potosı
´: ‘‘environs de Potosi,’’ 15–
28 Mar. 1833 [Papavero, 1971], A. C. V. D. d’Orbigny
1425 (lectotype, designated by Cabrera [1978: 674], P!;
isotypes, MO!, P!, W!).
Hypochaeris ornata J. Kost., Blumea 5(3): 660, fig. 3a, b.
1945. TYPE: Bolivia. Oruro: ‘‘am cerro de Oruro auf
steinigem trockenem Boden,’’ 3800 m, Nov. 1911, T. C.
J. Herzog 2522c (holotype, L, digital image!).
Hypochaeris meyeniana (Walp.) Benth. & Hook. f. ex Griseb.
var. leucantha Cabrera, Revista Invest. Agric. 11(4):
410. 1957. TYPE: Argentina. Jujuy: Yavi, Quebrada de
Cajas, 4000 m, 1 Feb. 1943, A. L. Cabrera 7837
(holotype, LP!).
Herbs to 5 cm tall. Leaves oblanceolate or oblong,
20–70 33–10 mm, apex acute, base attenuate in a
broad petiole, pinnatipartite to pinnasect, runcinate,
margin entire or toothed, hirsute to glabrous. Capitula
pedunculate (to 2 cm), rarely sessile. Involucre
campanulate, 10–25 38–20 mm; phyllaries 3- to 4-
seriate; outer phyllaries lanceolate, 7–11 32–2.5 mm,
apex acute or semiobtuse, hirsute on middle nerves
(many to few shaggy trichomes), margin hyaline; inner
phyllaries oblongate or lanceolate, 12–14 3ca. 2 mm,
hirsute (shaggy trichomes) to glabrous; paleae 7–
17 mm; florets 10 to 20. Corollas white, 13.5–20 mm;
tube 7–10 mm; ligule 6.5–10 mm; vascularization type
1; stamens 14–17 mm; anthers ca. 5 mm; basal
appendages type 2, 0.6–0.8 mm; filaments 9–12 mm;
antheropodium type 1; style 12–15 mm, branches 2–
3 mm. Cypselae 5-ribbed, unbeaked or slightly
narrower at the apex, 2–6 mm; wall scaly or smooth;
pappus 12–14 mm. Chromosome number unknown (a
possible hybrid with Hypochaeris meyeniana, however,
has been reported as 2n516; Weiss-Schneeweiss et
al., 2007).
Habitat and distribution. Hypochaeris echegarayi
occurs in the mountains of Peru to northwestern-
western Argentina (to San Juan Province), in bogs or
dry places, 3100–4950 m (Fig. 4C, triangles).
Phenology. Hypochaeris echegarayi flowers from
December to March.
Common name. ‘‘Q’ausilla’’ (Bolivia, Pestalozzi
194).
Morphological characters. Hypochaeris echegarayi
has shaggy trichomes on the leaves, phyllaries, and
corollas. This species is closely related to H.
eremophila, which has yellow corollas.
Observations. Achyrophorus sessiliflorus (Kunth)
DC. var. subruncinatus A. Gray (1861: 146) is an
illegitimate name that included both A. setosus Wedd.
and A. eriolaenus Sch. Bip. as synonyms. The
diagnosis by Gray (1861) corresponds more closely
to the former than to the latter.
During 1874 and 1883, Hieronymus worked in the
herbarium CORD of Argentina (Stafleu & Cowan,
1979), and his main collection is deposited there. In a
thorough search of that herbarium, we have been
unable to locate holotype material.
Cabrera (1957) originally described Hypochaeris
meyeniana var. leucantha as new based on its white
flowers. In 1978, he relegated this name to synonymy
under H. echegarayi Hieron.
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The Hypochaeris sessiliflora Complex (Asteraceae)
The small specimen of Hypochaeris ornata, plus the
clear figure that accompanies the protologue, showing
diagnostic features of campanulate involucre, setulose
phyllaries, and runcinate leaves, allows referral to H.
echegarayi. The protologue (Koster, 1945) does not
mention the color of the corollas, and the colors of the
specimen are faded.
Representative specimens. ARGENTINA. Jujuy: Cochi-
noca, Mina Aguilar, Meyer, Cuezzo & Legname 21305 (LP);
Humahuaca, Mina Aguilar, Cabrera et al. 19002 (LP);
Rinconada, Mina Pirquitas, Quebrada Cortadera, Schwabe,
Ancibor & Vizinis 909 (LP); Santa Catalina, cie
´naga cerca de
Cienaguillas, Cabrera et al. 15393 (LP, NY); Tumbaya, Abra
Grande de Volca
´n, Sleumer 3507 (SI); Yavi, Quebrada de
Cajas, Cabrera et al. 21505 (LP); alrededor de La Quica,
Hunziker & Caso 6039 (LP). San Juan: Iglesia, Reserva San
Guillermo, Los Caserones, Cazal, Pujalte & Reca 17 (SI).
Tucuma
´n: Tafı
´, Cumbre de Chaquivil, Olea 241 (UC).
BOLIVIA. Cochabamba: Tapacarı
´, 2 km al E de Japo K’asa
(Km 125 Cochabamba–Oruro), cerro Kampiyani, Pestalozzi
194 (LPB); 42 km from Caracollo, on the rd. to Cochabamba,
King & Bishop 7530 (US). La Paz: Aroma, Fisel U-123
(LPB); Juan Bautista Saavedra, Mergel-Kalksandstein-Horn-
stein-Lager, ca. 1 km S Kreuzung an der Straße nach
Escoma, Krach 7672 (LP, SI); Inquisivi, Quime, 25 km
hacia Caxata, Beck 17244 (LPB, US); Pedro Domingo
Murillo, vic. of lago Zongo at the head of the Zongo valley,
Solomon 13153 (LPB, NY); Presa Incachaca, 10 km NE from
La Paz, Stuessy, Tremetsberger & Ho
¨ssinger 18503 (LP);
13.4 km N of Huilacala, Stuessy, Tremetsberger & Ho
¨ssinger
18515 (LP); laguna Can
˜uma, N.P., Stuessy, Tremetsberger &
Ho
¨ssinger 18516 (LP); 3 km E from Kalullo (ca. 38 km NNE
from La Paz), valle de Achachicala, Stuessy, Tremetsberger &
Ho
¨ssinger 18525 (LP); Franz Tamayo, Puyo-Puyo (Ulla-Ulla),
Menhofer X-1880 (LPB); Pacajes, entre Corocoro y Topohoco,
Ceballos et al. 180 (SI). Oruro: Sajama, Canto
´n Lagunas,
Loza de la Cruz 253 (LPB). Potosı
´:N de Potosı
´, entre Bolivar
y Oruro, Ferna
´ndez Casas 8025 (NY). PERU. Lima:
Huarichori, Casapalca, Marcapomacocha rd. to Ratontay,
Sanderman 853 (K). Puno: Puno, San Antonio de Esqui-
lante, minas de San Antonio, Sanderman 3915 (K).
8. Hypochaeris eremophila Cabrera, Notas Mus.
La Plata 13: 22. 1948. Replacement name for
Distoecha taraxacoides Phil., Anales Mus. Nac.
Santiago de Chile. 8: 37, tab. 2, fig. 2. 1891, non
Distoecha taraxacoides Ball, 1885. TYPE: Chile.
Regio
´n III: Copiapo
´, Colorados [26u599S,
68u569W; Mun
˜oz Pizarro, 1960: 171], 3600 m,
15 Jan. 1885, A. Philippi s.n. (lectotype,
designated here, SGO 65208!, SGO photo LP!,
SGO digital image!). Figure 10.
Herbs to 7 cm tall. Leaves lanceolate or obovate,
20–70 35–12 mm, generally deeply runcinate,
glabrous or hispid principally on the midrib, base
attenuate in a broad petiole, margin toothed or entire.
Capitula pedunculate to 30 mm, rarely sessile.
Involucre campanulate-cylindrical, 13–18 310–
18 mm; phyllaries 3-seriate, lanceolate, apex acute,
margin with shaggy trichomes to 7 mm long; outer
phyllaries 10–14 32.5–4 mm; inner phyllaries acute
or slightly obtuse, glabrous or only slight trichomes,
14–17 32–3 mm; paleae 11–14 mm; florets 15 to 25.
Corollas yellow to yellow-orange, 12–20 mm; tube 5–
10 mm; ligule 7–10 mm; vascularization type 1;
stamens 11–18 mm; anthers 5–6 mm; basal append-
ages type 2 or 3, 0.8–1 mm; filaments 6–12 mm;
antheropodium type 1; style 10–14 mm, with branch-
es ca. 3 mm. Cypselae 5-ribbed, transversely wrin-
kled, erostrate, narrowed at the apex, 2.3–4 mm; wall
scaly; pappus 7–11 mm. Chromosome number 2n58
(Weiss-Schneeweiss et al., 2007).
Habitat and distribution. Hypochaeris eremophila
occurs from southern Peru to northwestern Argentina,
inhabiting dry places, bogs, and ‘‘mallines,’’ 2800–
4700 m (Fig. 4C, stars).
Phenology. Hypochaeris eremophila flowers from
November to April.
Common names. ‘‘Achicoria’’ (Argentina; Cabrera
8399); ‘‘co
´ndor siki’’ (Bolivia; Meneses P602-6).
Morphological characters. Hypochaeris eremophila
can be distinguished from the other acaulescent
species of Hypochaeris by its peduncules with
cylindric-campanulate involucres, phyllaries with
shaggy trichomes on the abaxial surface, and yellow
(sometimes yellow-orange) corollas.
Observation. Cabrera (1948: 22) proposed the name
Hypochaeris [Hypochoeris]eremophila when he moved
Distoecha taraxacoides into Hypochaeris, because H.
taraxacoides Ball was already occupied. See more detailed
comments under H. taraxacoides in this treatment.
Mun
˜oz Pizarro (1960) listed two original specimens
of Distoecha taraxacoides from the type locality in
Philippi’s herbarium at SGO. We have selected
SGO65208 as lectotype because it is the better of
the two specimens.
Representative specimens. ARGENTINA. Catamarca:
Bele
´n, camino a Antofagasta, Pasto Ventura, Cabrera, Botta
& Deginani 32555 (SI). Jujuy: Susques, Quebrada de
Tocomar, Cabrera 8278 (LP); al pie del Cerro Tuzgle, Cabrera
8399 (LP); Tumbaya, 33.3 km W of Purmamarca, Stuessy,
Urtubey & Tremetsberger 18064 (LP [2]); Valle Grande, subida
a cerro Amarillo desde Alto de Calilegua, Kiesling, Ulibarri &
Lo
´pez 1720 (SI). Salta: Cachi, Cerro de Cachi, Spegazzini s.n.
(LP); Guachipas, Pampa Grande, Spegazzini s.n. (LP); Pastos
Grandes, Abra del Gallo, Cabrera 9051 (LP). Tucuma
´n: Tafı
´,
Tafı
´, cumbres Calchaquı
´es, Co. Bayo alrededores del refugio,
Go
´mez Sosa & Mu
´lgura 157 (SI [2]). BOLIVIA. Cocha-
bamba: at Challa, on the rd. to Cochabamba, King & Bishop
7538 (US). CHILE. Regio
´nII:Loa, Collahuasi, quebrada
Capella, Pisano & Venturelli 1726 (LP). PERU. Puno: Grau,
Chuquibambilla, Pennell 13362 (US).
9. Hypochaeris acaulis (J. Re
´my) Britton, Bull.
Torrey Bot. Club 19: 371. 1892. Basionym:
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Figure 12. Hypochaeris acaulis. —A. Habit. —B–F. Phyllaries. —G. Trichomes (in cilia) on phyllaries. —H. Corolla with
anthers and style. —I. Palea, cypselae, and pappus. —J. Scaly cypselar wall. From Lo
´pez 1825 (LP).
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Urtubey et al.
The Hypochaeris sessiliflora Complex (Asteraceae)
Achyrophorus acaulis J. Re
´my, in Gay, Fl. Chil.
3: 448. 1847 [late 1848 or early 1849; Stafleu &
Cowan, 1976]. TYPE: Chile. Region VI: Cacha-
poal, ‘‘Se cria en los prados pantanosos de las
altas cordilleras de Talcaregue, provincia de
Colchagua,’’ 5–7 Feb. 1831 [Mun
˜oz Pizarro,
1944], C. Gay s.n. (holotype, P not seen, P photo
SI!; isotype, K!). Figure 12.
Herb to 5 cm high. Leaves oblongate, 32–60 312–
20 mm, pinnatisect, with divisions antrose, base
attenuate in a broad petiole, margin entire or toothed,
with some shaggy trichomes (also on nerves) or
glabrous. Capitula sessile. Involucre campanulate or
hemispheric, 13–23 310–25 mm; phyllaries 4- to 6-
seriate; outer phyllaries broadly ovate, 8–10 36–
8 mm; inner phyllaries lanceolate, 8–15 33–5 mm,
margin ciliate; paleae 13–16 mm; florets ca. 40.
Corollas yellow, 10–14 mm; tube 5–6.5 mm; ligule 5–
7.5 mm; vascularization type 1; stamens 7–9 mm;
anthers ca. 2 mm; filaments 5–7 mm; basal append-
ages type 1, ca. 0.3 mm; antheropodium type 3; style
7–10 mm, with branches ca. 1.5 mm. Cypselae 5-
ribbed, transversely wrinkled, 3.5–8 mm, rostrate;
wall scaly; pappus 10–15 mm. Chromosome number
2n58 (Wulff, 1998; Weiss et al., 2003).
Habitat and distribution. Hypochaeris acaulis
grows only in the southern Andean Cordillera across
a range about 600 3150 km, from the regions of
Santiago Metropolitan to Araucanı
´a of Chile and
provinces of Mendoza to Neuque
´n of Argentina
(Fig. 13). It occurs between 1430 and 3000 m in
bogs, marshy places, humid meadows, stream banks,
and Araucaria araucana (Molina) C. Koch woods. It
also can be found occasionally in disturbed areas near
roads and buildings.
Phenology. Hypochaeris acaulis flowers from De-
cember to March.
Morphological characters. Hypochaeris acaulis can
be clearly distinguished from the other acaulescent
species of the genus by its widely ovate outer
phyllaries, yellow corollas slightly longer than the
involucre, and the long rostrate cypselae. Tremetsber-
ger et al. (2003a) have completed a molecular (AFLP)
study of seven populations of H. acaulis from Chile and
Argentina. All data point to restricted gene flow among
populations, probably due to limited dispersal capa-
bility. There is also the expected positive correlation
between genetic variation and population size. The
species is postulated to be inbreeding (autogamous),
based on pollen:ovule ratios, which would be consistent
with the observed pattern of genetic variation. Based on
other AFLP data (Tremetsberger et al., 2006), the
evolutionary relationships of this taxon appear to reside
with H. palustris and H. tenuifolia and not with the
other acaulescent members of the South American
species of the genus. This appears to be a rather
dramatic case of parallelism of morphological features
for survival in the high Andes.
Representative specimens. ARGENTINA. Mendoza: Ma-
largu
¨e, Paso Pehuenche, sobre la frontera con Chile, Boelcke,
Bacigalupo & Correa 10379 (LP, SI). Neuque
´n: Minas,
extremo S laguna Varvarco Campos, cajo
´n del Varvarco,
Annals of the Missouri Botanical Garden mobt-96-04-06.3d 26/10/09 13:50:32 710 Cust #2006136
Figure 13. Distribution of Hypochaeris acaulis in the southern Andes (Chile and Argentina).
710 Annals of the
Missouri Botanical Garden
Boelcke et al. 14111 (LP, SI); N
˜orquı
´n, Copahue, Cabrera
1440 (LP); Termas de Copahue, Cabrera 6154 (LP [2]);
Picunches, ca. 1 km down from Pino Hachado, small laguna,
Stuessy & Baeza 15593 (CONC, WU); Zapala, Comber 1257
(K). CHILE. Regio
´n VI: Cachapoal, Rancagua, Cordillera
de Codegua, Barros 3790 (LP). Regio
´n VII: Curico
´,
Cordillera Volca
´n Peteroa, Wedermann 610 (BM, K, LP [2],
SI, UC); Talca, laguna de Maule, Stuessy & Baeza 15571
(CONC, WU); Alto de Vilches, camino a laguna El Alto,
Finot & Lo
´pez 1825 (CONC). Regio
´n VIII: N
˜uble, valle de
Las Nieblas (near Termas de Chilla
´n), Stuessy & Baeza
15565 (CONC, WU); Bio-Bio, Los Barros, ca. 1 km N of
reten de carabineros, Stuessy & Baeza 15703 (CONC, WU).
Regio
´n IX: Malleco, below volca
´n Tolhuaca, Stuessy &
Baeza 15817 (CONC, WU); Cautı
´n, Cordillera Las Raı
´ces,
Lonquimay, Burkart 9528 (SI). Regio
´n Metropolitana:
Santiago, Valle Largo, (Mapochotal) bei Las Condes
Hochkordillera, C. & G. Grandjot s.n. (LP).
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APPENDIX 1. Morphological characters and character states
used for cladistic analysis of the Hypochaeris sessiliflora
complex. See also Figure 1.
1. Habit: caulescent or peduncle longer than leaves (0);
acaulescent or peduncle never longer than the leaves
(1).
2. Stem: simple or bifurcating (0); ramified (1).
3. Leaf shape: lanceolate (0); oblong (1); obovate (2);
elliptic (3); linear-lanceolate (4); linear-filiform (5).
4. Leaf margin: dentate (0); entire (1); divided (2).
5. Leaf vestiture: shaggy (0); whip (1); shaggy and whip (2);
glabrous (3).
6. Involucre: campanulate (0); hemispheric (1); cylindric
(2); cylindric-campanulate (3).
7. Capitulum: pedunculate (0); sessile (1).
8. External phyllary shape: linear-lanceolate (0); lance-
olate (1); ovate (2).
9. Phyllary vestiture (abaxial): whip and shaggy tri-
chomes (0); whip trichomes (1); shaggy trichomes (2);
absent (3).
10. Length of corolla: longer than the involucre (0); as
long as involucre (1).
11. Corolla vascularization: type 2 (0); type 1 (1).
12. Basal appendages: type 2 (0); type 1 (1); type 3 (2).
13. Antheropodium: type 1 (0); type 3 (1); type 2 (2).
14. Cypselar shape: rostrate (0); semirostrate (1); erostrate
(2).
15. Cypselar wall: scaly on all surfaces (0); smooth (1);
scaly on main portion only (2).
16. Palea surface: pubescent (0); glabrous (1).
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712 Annals of the
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APPENDIX 3. Vouchers for the cladistic analysis.
Outgroup
H. argentina:Burkart et al. 10486 (LP); Krapovickas 7806
(LP); Urtubey & Tremetsberger 146 (LP, WU)
H. caespitosa:Castellanos s.n. (LP); Hunziker 8649 (LP);
Urtubey & Tremetsberger 145 (LP, WU)
H. chillensis:Cabrera 5688, 8595 (LP); Urtubey &
Tremetsberger 114 (LP)
H. elata:Cabrera et al. 16971, 17657 (LP); Stuessy et al.
18080, 18083 (LP, WU)
H. hookeri:Stuessy et al. 18019, 18044 (LP)
Ingroup
H. acaulis:Boelcke et al. 10379 (LP); Cabrera 6154 (LP);
Lo
´pez 1825 (LP)
H. echegarayi:Hunziker & Caso 6039 (LP); Cabrera et al.
15393 (LP); Meyer, Cuezzo & Legname 21305 (LP); Stuessy et
al. 18025 (LP)
H. eremophila:Burkart 5234 (LP); Cabrera 8278, 8399
(LP); Stuessy et al. 18064 (LP)
H. eriolaena:Angulo & Lo
´pez 1358 (LP); Hutchison, Wright
& Straw 6267 (US); Macbride & Featherstone 2487 (LP, US);
Velarde Nu
´n
˜ez 3225 (LP)
H. hohenackeri:Lewis 35168 (LPB); Menhofer X-1832 (SI,
LPB); Smith & Cabanillas 7212 (US).
H. meyeniana:Bundin s.n. (LP); Fabris 6514 (LP);
Ferna
´ndez 2029 (LP); Meyer 21270 (LP); Stuessy et al.
18062 (LP)
H. mucida:Ceballos et al. 614 (SI); Menhofer X-2197 (LPB)
H. sessiliflora:Cuatrecasas et al. 25599 (LP); Plowman
1955 (US); Molina & Barclay 18S388 (US); Stuessy et al.
18538 (WU)
H. taraxacoides:Meyer et al. 21040 (LP); Stuessy et al.
18074, 18075 (LP)
APPENDIX 4. Index to numbered collections cited. Numbers
in parentheses correspond to the species number in the List
of Species.
1. Hypochaeris sessiliflora Kunth
2. Hypochaeris taraxacoides Ball
3. Hypochaeris hohenackeri (Sch. Bip.) Domke
4. Hypochaeris mucida Domke
5. Hypochaeris eriolaena (Sch. Bip.) Reiche
6. Hypochaeris meyeniana (Walp.) Benth. & Hook. f. ex
Griseb.
7. Hypochaeris echegarayi Hieron.
8. Hypochaeris eremophila Cabrera
9. Hypochaeris acaulis (J. Re
´my) Britton
Adler 2 (6); Angulo & Lo
´pez 1358 (5); Aristeguieta 2438 (1);
Asplund 4886 (6). Baar 386 (6); Balls 6232 (6), B-6253 (2),
B-6361 (6), B-6711 (2); Barclay & Juajibioy 7642 (2), 8472,
9210 (1); Barros 3790 (9); Bastia
´n334(6); Beck 2861 (6),
7712 (2), 8356 (1), 9060 (6), 11914 (5), 17244 (7); Beck &
Paniagua 27086 (2); Becker et Terrones 1433 (5); Blair 636
(2); Boecke & Jaramillo 2348,2493 (1); Boelcke et al. 10379,
14111 (9); Bundin s.n. (6); Burkart 5234 (8), 9528 (9);
Burkart & Troncoso 11822 (2). Cabezas 25 (6); Cabrera 1440,
6154 (9); 7723,7826 (6), 8278,8399 (8), 8982 (6), 8993 (2),
9051 (8), 12121 (6); Cabrera et al. 15256 (2), 15393 (7),
17656 (2), 19002 (7), 21505 (7), 22484,22531 (2), 27174 (6),
32555 (8); Canne & Schunke 232 (5); Ca
´rdenas 4347 (6);
Cazal et al. 17 (7); Ceballos et al. 164 (2), 180 (7), 548 (2),
614 (4); Cerrate 6025 (6); Cerrate & Tovar 1174 (2); Comber
1257 (9); Cuatrecasas 1484 (1); Cuatrecasas & Romero
Castaneda 24560 (1); Cuatrecasas & Willard 26299 (1).
Deginani et al. 806 (6); de la Fuente & Dı
´az 10905 (6).
Ellemann 91672 (1); Ellenberg 638 (4); Eyerdam 24839 (3),
25059 (6); Ewan 16324 (1). Fabris 1369 (8), 1520 (2); Fabris
et al. 4035 (2), 6514 (6); Ferna
´ndez 2029 (6); Ferna
´ndez
Casas 8025 (7); Fiebrig 3188 (2); Finot & Lo
´pez 1825 (9);
Fisel U-123 (7); Freueler 4384 (2). Go
´mez Sosa & Mu
´lgura
157 (8); Grandjot, C. & Grandjot, G. s.n. (9); Guille
´n & Riccio
10784 (6). Haber 5,142 (2); Hawkes et al. 3900 (2); Hensen
789 (6); Hilgert & Lamas 1654 (6); Hoogle & Roessch 2534
(2); J. H. Hunziker & Caso 6039 (7); Hutchison et al. 5892 (2),
6267 (5). Ibish & Rojas 439 (5); Iltis & Ugent 1256,1489 (2),
1495 (6). Jahn 146 (1). Kiesling et al. 581 (6), 1720 (8), 3801,
3826 (6); Killip & Smith 17641,21680 (1), 22191 (6), 23369
(1); King & Bishop 7504 (6), 7530 (7), 7538 (8); King &
Collins 9064 (2); King & Garvey 6968 (1); Krach 7492 (2),
Annals of the Missouri Botanical Garden mobt-96-04-06.3d 26/10/09 13:50:37 713 Cust #2006136
APPENDIX 2. Data matrix used in the cladistic analysis of the Hypochaeris sessiliflora complex. Dashes indicate missing
data or inapplicable states.
12345 6 78910111213141516
Outgroup
H. argentina 00 00 30 0 000000000
H. caespitosa 0-- 401 3 0 0 0 10 1 0 1 1 10
H. chillensis 01 002133 0 00131111000
H. elata 001 0 0 3 0 0 0 01 1 0 1 0 00
H. hookeri 1 -- 5 01 13 3 0 0 1 0 1 1 0 2 1 0
Ingroup
H. acaulis 1-- 1 2 0301 1 2 30 1 1 1 0 21
H. echegarayi 1 -- 01 2 03 0 01 1 20 1 0 01201 1
H. eremophila 1 -- 02 02 03 3 01 1 2 0 1 02 0 02 0 1
H. eriolaena 1 -- 01 01 013 01 1 12 1 0 0 01 02 12 0 1
H. hohenackeri 1-- 101 3 0 0 1 30 10102 2 11
H. meyeniana 1 -- 023 2 03 01 01 12 123 0 01 02 0 1 01 1
H. mucida 1-- 102 2 0 1 1 00 1 1 0 2 11
H. sessiliflora 1 -- 34 0 3 01 01 012 23 0 1 02 02 12 01 1
H. taraxacoides 1-- 0102 3 2 0 0 30 0 2 0 2 11
Volume 96, Number 4 713
2009
Urtubey et al.
The Hypochaeris sessiliflora Complex (Asteraceae)
7672 (7), 8417 (6), 8435,8672 (2), 9391 (2); Krapovickas
3155 (2). Leiteyn et al. 4995 (1); Lewis 35168 (3), 37146 (6),
38124 (3); Liberman 2343 (2); Lo
´pez & Saga
´stegui 2888,
3223,8245 (2); Lo
´pez et al. 7421 (5); Lo
´pez Vega 1607 (6);
Loza de la Cruz 72 (2), 253 (7), 254 (2); Luteyn & Dorr 13780
(6). Macbride & Featherstone 1791 (1), 2167 (2), 2487 (5);
Maguire 39405 (1); Mandon 276 (2); Marticorena & Matthei
343 (6); Mathei & Rodrı
´guez 290 (6); Meneses P6O2-6 (8);
Menhofer 1036 (2), 1600 (2), X-1880 (7), X-1832 (3), X-2197
(4); Mercado 2181 (6); Metcalf 30280 (6); Meyer 21270 (6);
Meyer et al. 21040 (2), 21305 (7), 21517 (6); Meza 149 (6),
207,209 (5); Mezar 162 (2); Molina & Barkley 18 S.338 (1);
Mu
¨ller & Heinrichs 6428 (3); Mu
¨sch 151 (p.p.) (1). Novara
6356,10644 (6); Novara & Neumann 9781 (6). Ochoa 264
(2); Olea 241 (7); Ostria 132 (6). Pennell 13362 (8); Pen
˜afiel
et al. 382 (1); Pestalozzi 194 (7), 206 (6), 284 (2), 292 (6), 606
(5), 939 (6); Pisano & Venturelli 1726 (8); Plowman 1955 (1).
Ramı
´rez 203 (6); Ricardi et al. 343 (6); Rohweder T-12 (2).
Saga
´stegui et al. 9006 (5), 10006 (6); Sanderman 853,3915
(7); Saunders 379,822 (6), 1106 (2); Schulte 9 (2), 129 (6);
Schwabe et al. 641 (2), 818,819 (2), 909 (7), 1098 (6);
Sleumer 2730 (6), 3507 (7); Sleumer & Vervoorst 2589 (6);
Smith & Buddensiek 11133 (5); Smith & Cabanillas 7212
(3), 7328 (1); Smith et al. 5652 (2); Soejarto 494 (1);
Soderstrom 1254 (1); Solomon 13153 (7), 13177 (2);
Solomon & Nee 14233 (2); Spegazzini s.n.,s.n. (8); Stafford
865 (5); Steinbach 9696 (6); Steyermark et al. 101105 (1);
Stork & Horton 10000 (5), 10838 (2); Stuessy & Baeza
15565,15571,15593,15703,15817 (9); Stuessy et al.
18062 (6), 18064 (8), 18068 (6), 18074,18075 (2), 18089
(2), 18503 (7), 18504 (6), 18515,18516,18525 (7), 18538,
18540,18549 (1). Tolaba et al. 1198,1235 (6), 1539 (2),
1652,1677 (6); Tovar 222 (3), 390,393 (2), 842 (6), 850 (2),
2512 (5); Tutin 1209,1210 (5). Vargas 11355,11350 (p.p.)
(5), 21784 (1); Vargas et al. 2619,2887 (1); Velarde Nu
´n
˜ez
3225 (5); Vignati 445 (6). Wedermann 247 (6), 610 (9);
Weigend 2000/615 (6), 2000/616 (1); Weigend & Forther
97-790 (6); Werner 89,91,103,131 (6), 820 (2); West 3663
(2); Wevi 129 (5); Wurdack 761 (1), 1225 (3). Zamalloa Dı
´az
78 (2); Zamora 156 (6); Zuloaga & Landono 4136 (1).
Annals of the Missouri Botanical Garden mobt-96-04-06.3d 26/10/09 13:50:41 714 Cust #2006136
714 Annals of the
Missouri Botanical Garden
... Description: Readily distinguishable from other taxa by the combination of peduncular (rarely sessile) cylindrical heads, glabrous phyllaries (ciliate on margins) and leaves, and corollas longer than the involucre, white on the upper surface to dark blue-black at the tips of ligules underneath (Urtubey et al., 2009) Nesom (1990). ...
... This species is closely related to H. eremophila, which has yellow corollas. Details:Urtubey, Stuessy & Tremetsberger (2009).Description: Distinguishable from other members of the H. sessiliflora complex by long sericeous trichomes on broad petioles plus lanate phyllaries. This species often has no developed leaves when the capitulum opens. ...
... Corolla white or yellow. Details:Urtubey, Stuessy & Tremetsberger (2009).Ecology: Andean III: 4000-> 4500 m. Dry, stony Andean ridges, with scattered grass and limestone outcrops. ...
Book
The catalog describes 1362 species of ferns and flowering plants for the department of Arequipa, more than 200 species more as reported in the last systematic inventory by Quipuscoa, Dillon, & Ortíz in 2006. Ninety-five species are mentioned for the first time for Arequipa and 26 species mentioned in the literature for the department were excluded. In addition to a brief description of the species, the catalog includes information on their ecology, distribution, and human use. In addition to information on systematics and phylogeny, the most important synonyms are listed in the appendix.
... Dispersion followed by geographic isolation and selection within different ecological zones enabled the development of high levels of morphological divergence, indicating a pattern of speciation by adaptive radiation . Population genetic studies based on molecular markers support the assumption that isolation by distance and founder events shaped the geographic distribution patterns of the South American complex of Hypochaeris species (Muellner et al. 2005;Tremetserger et al. 2009;Urtubey et al. 2009;Rodrigues et al. 2016). Furthermore, biogeographic events and climate changes also played important roles in the distribution and isolation of populations. ...
... Several Hypochaeris species in South America have either endemic or widespread distribution (Bortiri 1999;Cabrera et al. 2000;Azevêdo-Gonçalves and Matzenbacher 2007). Some species are capable of colonizing a wide variety of habitats from sea level to over 5000 m (Tremetsberger et al. 2006;Urtubey et al. 2009), while others are restricted to coastal or high-elevation zones or are capable of growing in grasslands or modified environments (Reck et al. 2011;Rodrigues et al. 2016). In Brazil, the Hypochaeris genus is represented by approximately 12 species, most of which occur in Rio Grande do Sul State (Ruas et al. 2005;Azevêdo-Gonçalves and Matzenbacher 2007;Rodrigues et al. 2016). ...
Article
Full-text available
The genus Hypochaeris was likely introduced into South America by a unique long-distance dispersal event from an ancestral Northwest African species, with subsequent radiation and diversification throughout the temperate zones of the continent. Hypochaeris catharinensis is an endemic species of Southern Brazil growing mostly in high-elevation grasslands with dry, rocky, and shallow soils. Here, we used nine microsatellite loci and a total sample of 441 individuals to assess genetic variability patterns within and among 13 H. catharinensis populations. The results showed a total of 215 alleles in the nine loci tested, the expected and observed heterozygosities average were 0.31 and 0.43, respectively, and allelic richness varied from 4.09 to 7.23. The levels of allelic richness showed that genetic variability is maintained by gene flow, thus ensuring that these populations are not genetically isolated. We found no correlation between geographic and genetic distances, with high levels of genetic variability within, rather than among, populations. The low levels of genetic structure in the species are consistent with recent evolutionary processes of the Hypochaeris genus in South America; after adaptive radiation and species differentiation, dispersal to similar habitats led to the establishment of new populations. Along with initial founder effects, the existence of gene flow explains the current genetic variation across the H. catharinensis populations. In addition, historic environmental changes reported for Southern Brazil may explain the current clustered distribution of the species.
... The geographic affinities of species of Hypochaeris in South America are within the South American Transition Zone (Urtubey et al. 2010; see individual distribution maps in this revision and in Urtubey et al. 2009 for detailed documentation). The broad patterns show ser. ...
... Among the South American species of Hypochaeris, this is the easiest series to recognize morphologically. For more information, including a morphological phylogenetic analysis, see Urtubey et al. (2009). The only confusing taxon morphologically is H. acaulis (here placed in ser. ...
Book
ABSTRACT. This revision treats the 41 South American species of Hypochaeris, plus the northwest African H. angustifolia, all now placed in a single monophyletic section Phanoderis. The New World species constitute subsection Phanoderis, and the single African species comprises subsection Africana. The species in South America represent a challenge in classification; molecular clock investigations suggest that the entire complex diverged approximately within the past 1 million years. From an ancestor in northwestern Africa, similar to the present-day H. angustifolia (the phylogenetic sister-group), have come 41 species in a relatively short geological time. These species are similar genetically; routine DNA sequencing with normal markers provides little differentiation. With Amplified Fragment Length Polymorphism (AFLP) analysis, distinct groups can be resolved, some with very strong statistical support and others less so. Species are morphologically reasonably well defined, but the differences are subtle, requiring field experience for accurate identification of taxa. Previous studies have focused on regional perspectives, and this has led to confusion in identification of herbarium specimens. New cytogenetic data have also revealed groups of phylogenetically related species, which to some extent correlate with the AFLP distinctions. Speciation has been almost totally at the diploid level (2n = 8). Eight species are known with tetraploid cytotypes, and one species (H. caespitosa) is uniformly tetraploid. Polyploidy has not been a major factor for diversification within the group. The most common type of speciation has been by eco-geographic divergence; dispersal to new regions and habitats has led to isolation, genetic modification, and eventual speciation. This process has obviously been rapid, representing a pattern of radiation within a continental region. This is particularly remarkable because the entire complex has developed after a single long-distance dispersal and founding event from northwestern Africa. Dispersal over long distances within the continent, followed by strong selection in different habitats, has led to the mosaic patterns of morphology now seen. Subsection Phanoderis is distributed throughout South America in temperate or high elevation regions from Colombia to southern Argentina and Chile. Species are found from sea level to over 5000 m. Species of series Oreophila occupy very high elevations in páramos and wind-swept grasslands in the high Andes primarily in Colombia, Ecuador, and Peru. Series Scorzonerae is confined to coastal areas of Chile, and series Apargioides is found concentrated in south-central Chile and adjacent Argentina. Series Caespitosa is centered in a small region of the mountain ranges of Cordoba, Argentina, and series Phanoderis and series Chondrilloides occur along both sides of the southern Andes. Series Pampasica, series Microcephala, and series Lutea have considerable areal overlap in southern Brazil, Uruguay, and the pampas region of Argentina. Several new taxa and combinations are presented in this monograph: Hypochaeris sect. Phanoderis; Hypochaeris subsect. Phanoderis and Hypochaeris subsect. Africana; Hypochaeris ser. Oreophila; and Hypochaeris laciniata. In addition, eight new series are described within Hypochaeris subsect. Phanoderis: Hypochaeris ser. Phanoderis, ser. Apargioides, ser. Caespitosa, ser. Chondrilloides, ser. Lutea, ser. Microcephala, ser. Pampasica, and ser. Scorzonerae.
... Los ejemplares colectados fueron prensados, secados y herborizados. La determinación taxonómica se realizó mediante la comparación con muestras de los herbarios de la Universidad Nacional Mayor de San Marcos (Herbario USM) y el herbario de la Facultad de Ciencias de la Universidad Nacional Agraria la Molina (Herbario MOL), con muestras digitalizadas en la base de datos Neotropical Herbarium del Field Museum, con la ayuda de taxónomos y el uso de claves dicotómicas y descripciones en literatura especializada (McBride, 1963(McBride, -1981Ballard, 1986;Luteyn, 1991;Funk y Zermoglio, 1999;Hofreiter y Rodríguez, 2006;Urtubey et al., 2009;Huamantupa, 2010;Gonzáles, 2012;León, 2012;Rodríguez y Sagástegui, 2014;Freire et al., 2016;Mione et al., 2016;Luteyn y Pedraza-Peñalosa, s. f.). La nomenclatura y estatus migratorio de cada especie fue corroborada en la base de datos Tropicos (MBG, 2019), y la escritura de los nombres quechua con el Diccionario de Quechua de Huánuco Rimaycuna (Weber et al. 2008). ...
Chapter
Full-text available
Los Andes constituyen uno de los centros de origen de plantas cultivadas donde la gran diversidad de condiciones climáticas y suelos dan origen a una gran variedad de recursos naturales utilizables, pero también a una constante incertidumbre ecoclimática. Por ello, los pobladores andinos han desarrollado, durante milenios, estrategias de manejo de riesgos. De acuerdo a varios cronistas, los peruanos pre-colombinos “comían todas las hierbas, dulces y amargas”, así como una amplia gama de animales. Sin embargo, con la colonización española gran parte de los alimentos andinos fueron violentamente reemplazados. A este proceso de erosión cultural se ha sumado en las últimas décadas el cambio climático, que limita la capacidad de adaptación de los pobladores. Antonio Brack reporta en el Perú 787 especies alimenticias: 107 domesticadas y exclusivamente cultivadas, 167 silvestres que también se cultivan, y el resto exclusivamente silvestres; el 70 % son de origen amazónico. Darwin desafió la dicotomía silvestre-domesticada en 1868 al proponer que la domesticación es un proceso continuo. En este sentido, De Wet & Harlan plantearon la existencia de una gama de estatus ecológicos de las plantas de acuerdo a su grado de dependencia del hombre, denominando silvestres a aquellas que solo crecen naturalmente fuera de los hábitats disturbados por el hombre, arvenses a aquellas que invaden hábitats disturbados (campos de cultivo, huertos, etc.), y ruderales a las que crecen alrededor de caminos y vías. Con ello se desafía también la dicotomía recolector-agricultor. Si bien actualmente ciertas variedades de unas pocas plantas domesticadas (trigo, arroz, maíz y papa) dominan los patrones alimentarios mundiales, las no domesticadas (silvestres, arvenses y ruderales) siguen siendo incorporadas en la dieta de sociedades tradicionales. Desde hace cerca de cuatro décadas estas plantas, y en especial aquellas nutracéuticas (que tienen usos medicinales además de alimenticios), han sido abordadas mediante estudios antropológicos, etnobotánicos y etnoecológicos sobretodo en Europa, Asia y África, así como en algunos países de América; en base a ello, la contribución nutricional y potencialmente medicinal de los vegetales de hojas verdes, en especial los no domesticados, ha sido reconocida. Así, difundir su consumo es de suma importancia en Perú, donde, en el 2017, la anemia afectó al 44,4% de los niños de 6 a 35 meses de edad, y al 52,6% en el área rural. Por ello, estas plantas pueden contribuir a mantener la seguridad alimentaria de las poblaciones que las usan. Además, al configurar la culinaria, cultura e identidad de estos pueblos estos alimentos participan de la soberanía alimentaria local. En el Perú, sin embargo, los estudios etnobotánicos, es decir, aquellos que investigan las interacciones personas-plantas, son mayormente etnomedicinales y orientados a producir inventarios. El objetivo general de este estudio fue contribuir al conocimiento acerca de los patrones alimentarios de dos comunidades campesinas altoandinas del distrito de Kichki desde una perspectiva etnobotánica, como aporte para posibles estrategias de adaptación local frente al cambio climático. Los objetivos específicos fueron (1) Elaborar un registro etnobotánico de especies de plantas silvestres, arvenses y ruderales alimenticias, sus formas de uso y manejo, (2) Identificar sus hábitats de obtención y los meses de consumo, y (3) Describir los patrones alimentarios de las comunidades desde una perspectiva etnobotánica, y la importancia de estas especies en la dieta.
... Los ejemplares colectados fueron prensados, secados y herborizados. La determinación taxonómica se realizó mediante la comparación con muestras de los herbarios de la Universidad Nacional Mayor de San Marcos (Herbario USM) y el herbario de la Facultad de Ciencias de la Universidad Nacional Agraria la Molina (Herbario MOL), con muestras digitalizadas en la base de datos Neotropical Herbarium del Field Museum, con la ayuda de taxónomos y el uso de claves dicotómicas y descripciones en literatura especializada (McBride, 1963(McBride, -1981Ballard, 1986;Luteyn, 1991;Funk y Zermoglio, 1999;Hofreiter y Rodríguez, 2006;Urtubey et al., 2009;Huamantupa, 2010;Gonzáles, 2012;León, 2012;Rodríguez y Sagástegui, 2014;Freire et al., 2016;Mione et al., 2016;Luteyn y Pedraza-Peñalosa, s. f.). La nomenclatura y estatus migratorio de cada especie fue corroborada en la base de datos Tropicos (MBG, 2019), y la escritura de los nombres quechua con el Diccionario de Quechua de Huánuco Rimaycuna (Weber et al. 2008). ...
Book
Full-text available
Referirnos a la diversidad biológica, los alimentos y el clima desde un antiguo país de montañas tropicales resulta ser muy pertinente, más aún en tiempos donde tanto el riesgo como el cambio climático se hallan globalizados. Los estudios que se presentan provienen de diferentes perspectivas (Estado, sociedad civil, comunidades), disciplinas (ciencias naturales y sociales) y escalas (regional, nacional e internacional). La investigación duró tres años (2016 – 2018) y en ella participaron cinco universidades, de las cuales tres fueron regionales: Universidad Nacional de San Antonio Abad del Cusco, Universidad Nacional Hermilio Valdizán – Huánuco y Universidad Nacional Micaela Bastidas de Apurímac; una fue de carácter nacional, la Universidad Nacional Agraria la Molina y, de México, la Universidad Nacional Autónoma de México. Desde la sociedad civil estuvieron las Organizaciones No Gubernamentales IDMA (Huánuco), CADEP JM (Cusco) y CCTA (Lima), así como siete comunidades campesinas: San Pedro de Cani, Santa Rosa de Monte Azul y Tres de Mayo de Huayllacayán (Huánuco), Llachua, Patahuasi, Pauchi Marcallac y Santiago de Queuñapampa (Apurímac). La publicación reúne 15 documentos entre revisiones, ensayos y trabajos de investigación (nueve), pertenecientes a 17 autores, referidos a temas relacionados con la agrobiodiversidad vegetal nativa alto andina de la sierra centro (Huánuco) y sur (Apurímac), priorizando a dos cultivos: la papa y el maíz, y su relación con el cambio climático a través de escenarios climáticos, su potencial de adaptación a condiciones ambientales cambiantes, y su aporte a crear mejores condiciones para garantizar la seguridad alimentaria considerada como la medida de adaptación más importante. El papel de los conocimientos tradicionales es abordado como un tema transversal a todas las investigaciones. El proyecto fue posible gracias al apoyo del Programa Nacional de Innovación Agraria (PNIA 027-2015-INIA-PNIA/UPMSI/IE). Finalmente es importante resaltar que el objetivo esencial de esta publicación es documentar y comunicar los hallazgos de este equipo conformado por investigadores andinos y mesoamericanos a los profesionales, estudiantes, técnicos, autoridades locales y a los miembros de la comunidad relacionados con los recursos naturales y el ambiente.
... El conocimiento sobre las especies y distribución de asteráceas en el Perú, cada vez se está extendiendo por el incremento de colecciones y estudios en áreas muy localizadas. Además, estudios a nivel molecular están fortaleciendo hipótesis justificadas en estructuras morfológicas y permitiendo establecer géneros nuevos y sus relaciones filogenéticas, los cuales derivan en cambios nomenclaturales (Freire et al., 2014, Funk, 1997a, 1997b,1997cHarling, 1992;Katinas, 1996Katinas, , 2012Linares et al., 2014;Pruski & Sancho, 2006;Pelser et al., 2007, Nordenstam & Pelser 2009, Robertson, 2010Schilling & Panero, 2011;Urtubey et al., 2009, Urtubey, 2011. ...
Article
Full-text available
A list of 343 species of Asteraceae for the department of Lima is presented, highlighting and comparing their presence in the hydrographic basins, in addition to its altitudinal range; 14 species are endemic to Lima. It is added 17 new records, one of them, Werneria microphylla, is a new species for the Peruvian flora.
... In this study, we used amplified fragment length polymorphism (AFLP; Vos et al., 1995) to investigate the diversification of the taxonomically difficult P. bipinnatifidus species complex across its geographic range. AFLP markers have been shown to provide signals for inferring phylogenetic relationships of closely related species or species complexes (e.g., Bussell et al., 2005;Triplett et al., 2010;Safer et al., 2011;Rebernig et al., 2012), especially when nuclear DNA/ribosomal DNA and chloroplast DNA failed to distinguish these taxa (Despres et al., 2003;Urtubey et al., 2009). Recent evolutionary radiation could be inferred from the incongruence among independent genes for the reconstruction of phylogenetic trees, because speciation events may have occurred in rapid succession during this process (Knowles & Chan, 2008). ...
Article
The Panax bipinnatifidus species complex (Panax bipinnatifidus and its close relatives) in the Sino-Himalayan region has been taxonomically difficult. Evolutionary analyses using amplified fragment length polymorphism (AFLP) markers were conducted on 125 individuals representing eleven populations of the P. bipinnatifidus species complex and one population of P. stipuleanatus as an outgroup. The populations from the eastern Himalayan region sampled from Nepal and eastern Tibet formed two main groups in the neighbor-joining (NJ) and split network analyses. The Pailong population (Tibet-PL) in eastern Tibet showed a highly distinct AFLP profile and was placed as the most basally branched group in the NJ tree. The remaining Himalayan populations showed three subgroups: the Nepal-HB & Nepal-HS subgroup, the Nepal HH subgroup, and the Tibet-BY subgroup. The three Himalayan subgroups had very limited gene flow among them and showed subtle morphological differences. The populations in eastern, central and western China showed clear geographic patterns and can be sorted into several geographical groups. Each major group in the species complex has strong bootstrap support, but relationships among them are poorly resolved, which is consistent with a pattern of evolutionary radiation. The strong geographic grouping, high Nei's population differentiation index, and limited gene flow among populations in different regions support the importance of geographic isolation in the diversification of the P. bipinnatifidus species complex in the Sino-Himalayan region.
Article
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The family of the Asteraceae is the most diverse in the Chilean flora; nevertheless, only some tribes have undergone a recent systematic treatment, while the knowledge about the geographical distribution of the taxa is still very incomplete. From the review of herbarium specimens and fieldwork, a survey of the Compositae flora of the Arica y Parinacota Region was performed. The study shows the presence of 144 taxa of the Asteraceae family in the region of which 82.6% is native (119) and 9% is endemic to Chile (13), while 8.3% are considered as exotic species (12). The genus Senecio is the one showing the highest diversity in the region, with the presence of 27 species, including six endemic ones. The revision of the botanical material reveals the presence of five new species for the Chilean flora: Achyrocline ramosissima Britton, Gamochaeta humilis Wedd., Mniodes kunthiana (DC.) S.E. Freire et al. (=Lucilia violacea Wedd.), Pseudogynoxys cordifolia (Cass.) Cabrera, and Senecio evacoides Sch. Bip. A new species has been recently described based on our recent field surveys: Pseudognaphalium munoziae N. Bayón, C. Monti & S.E. Freire. Additionally, the correct denomination of the endemic species Stevia philippiana has been recently undertaken. The greatest diversity of species is found in the province of Parinacota, above 3,000 m asl. Compositae stand out in the region not only for its diversity but also as a crucial element of the vegetation, being a fundamental component of the precordillera and altiplano vegetation belts, known as "tolar", in which species pertaining to genera like Parastrephia, Lophopappus, Baccharis, or Senecio tend to dominate. Indeed, most of the vegetation belts described in the Parinacota province show members of the Asteraceae as dominant species. In spite of a big amount of the regional surface under formal protection, several species occur outside the protection units.
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Se presenta un documento técnico de divulgación sobre las plantas de los páramos del DMQ, se realiza una síntesis de los ecosistemas y se describen varias especies representativas. El propósito es dar a conocer la riqueza, la estructura y composición florística, el funcionamiento y los servicios ambientales que ofrece el páramo para los habitantes de Quito y zonas aledañas, además de concientizar acerca de la importancia de conservar y hacer un manejo sostenible de este ecosistema, mediante la divulgación del conocimiento.
Article
Chromosome counts of Compositae are reported from Mexico, Central America, Ecuador, and Chile. First counts are reported for 20 species in Baccharis, Calea, Erigeron, Eupatorium, Heliopsis, Isocarpha, Liabum, Monactis, Pinaropappus, Senecio, Sigesbeckia, Simsia, Spilanthes, Verbesina, and Viguiera. Additional counts also are provided for 65 populations of taxa counted previously, of which 10 are new numbers. The systematic implications of certain of these counts are discussed.