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Morphology and Taxonomic Revision of Calycera

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DOI: 10.1600/036364414X683877

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Systematic Botany (2014), 39(4)
© Copyright 2014 by the American Society of Plant Taxonomists
DOI 10.1600/036364414X683877
Date of publication September 3, 2014

Morphology and Taxonomic Revision of Calycera

Silvia S. Denham, Lucio Zavala Gallo, and Raúl E. Pozner1
Instituto de Botánica Darwinion (ANCEFN-CONICET), Labardén 200, Postal Code 22, B1642HYD San Isidro,
Buenos Aires, Argentina.
1
Author for correspondence (rpozner@darwin.edu.ar)

Communicating Editor: Min Feng

Abstract—This taxonomic treatment is the first in the genus Calycera based on a detailed study of morphology and a critical analysis of
species boundaries. In this revision, nine species and four varieties are recognized, one new combination (C. crassifolia var. spinulosa),
nine new synonymies (Anomocarpus axillaris, Boopis integrifolia, Calycera boopidea, C. crenata, C. foliosa, C. intermedia, C. pulvinata f. cauligera,
C. squarrosa, and Gymnocaulus viridiflorus), six new lectotypifications (for Anomocarpus, Boopis gracilis, Calycera crenata, C. pulvinata f.
cauligera, C. sessiliflora, and C. sympaganthera), and one neotypification (for Calycera involucrata) are established. Updated morphological
descriptions (including an emended description of C. herbacea var. sinuata) and geographical distributions are included for each species.
Analyses of their life forms and inflorescence, flower, and fruit morphology are presented.

Keywords—Calyceraceae, flower morphology, fruit polymorphism, life forms, taxonomy.

Calycera Cav. is the largest genus of Calyceraceae, with
a total of 14 species, four varieties, and two forms sensu
Chiapella (1999a) and Zanotti and Pozner (2008). Species
are distributed in southern South America, from south-
ern Peru to northern Patagonia in Argentina and Chile
(Hellwig 2007).
Despite a lack of sharp boundaries among Calyceraceae
genera (cf. Hellwig 2007; Zavala et al. 2010), Calycera is
considered a monophyletic group based on shared mor-
phological characters (Hellwig 2007). Calycera is easily dis-
tinguished from other Calyceraceae genera by its dimorphic
(armed and unarmed) cypselas borne on the same receptacle
(Hellwig 2007). Additionally, as noted by botanists of the
nineteenth century, species of Calycera also show floral poly-
morphism, particularly in the calyx of flowers within
each inflorescence (Rémy 1847[1848]; Philippi 1856; Weddel
1857[1858]; Miers 1860–1869; Bentham and Hooker 1873).
Calycera has been partially studied in regional floras
(Reiche 1902; Hauman-Merk 1918; Hicken 1919; Muñoz-
Pizarro 1959; Pontiroli 1963, 1993; Cabrera and Zardini 1978;
Navas-Bustamante 1979; Roig 1981; Lamberto et al. 1997;
Hoffmann et al. 1998; Chiapella 1999b), but a taxonomic
overview of all species is needed to assess species bound-
aries and suggest reliable terminals for phylogenetic studies.
As a first contribution toward a complete and comprehensive
taxonomic revision of Calyceraceae, we present here the
first taxonomic revision of Calycera, recognizing nine spe-
cies and four varieties, and including descriptions and a
complete list of synonymies for each species. This revi-
sionary effort includes one new combination, nine new
synonyms, six new lectotypifications, one neotypification,
an emended variety, a key to the species, updated geo-
graphic distributions, references to iconographies, and the
first illustration of C. calcitrapa Griseb.
The species of Calycera grow in diverse environments,
from the humid Atlantic seashore to high-altitude arid habi-
tats, including the high mountain deserts of the Puna,
open grassland, and meadows (Pontiroli 1963; Chiapella
1999b; Hellwig 2007). Calycera comprises hemicryptophytic,
chamaephytic, cryptophytic, and therophytic (cf. Raunkiaer
1934) species. We included an analysis of their life form
related to geographical distribution and the diversity of
environments where these species grow.
We also performed here an evaluation of fruit and flower
morphology to test if calyx polymorphism occurring in the
flowering stage is correlated to calyx polymorphism in the
fruiting stage.
Materials and Methods
We studied collections from BCRU, CONC, LIL, LP, and SI, as well
as the original diagnoses and type material housed in BA, BAF, BM,
CONC, GOET, K, LIL, LP, LPS, M, MA, S, SGO, and SI (some directly
as herbarium vouchers, and others as digital images from JSTOR or
herbarium web pages). Herbarium material was rehydrated following
Venning (1953). Vegetative, floral, and fruit structures were observed
with a Wild (Heerbrugg) M5–85943 scope. Specimens were also studied
during fieldwork in Argentina and Chile.
In the taxonomic treatment, descriptions of phytogeographic distribu-
tion were based on the biogeographic scheme by Cabrera and Willink
(1980). When describing the inflorescence structure, we followed Pozner
et al. (2012).
Results
Life Forms in Calycera—Calycera species grow according
to four life forms: hemicryptophytes, chamaephytes, crypto-
phytes, and therophytes (cf. Raunkiaer 1934). All of these
are related to semiarid habitats. Figure 1 summarizes the
relationship among life form and geographical distribution
of the species of Calycera.
Therophytic species of Calycera [C. eryngioides J. Rémy,
C. leucanthema (Poepp. ex Less) Kuntze, and C. sessiliflora
Phil.] are caulescent herbs endemic to central Chile, a
geographic area with argillic soils (Alfisols, Vertisols and
Inceptisols; Ibáñez 2008a) and strongly contrasting season-
ality with humid winters and dry summers (Ibáñez 2008b)
that expose the soil to alternating cycles of expansion and
contraction resulting in poor ventilation conditions for roots.
The only cryptophytic species, C. crassifolia (Miers) Hicken,
is a rhizomatous herb that grows in sandy soils (Entisols and
Aridisols) from two different habitats: foothills [C. crassifolia
var. spinulosa (Gillies ex Miers) Zavala, S. Denham and
Pozner] and the Atlantic seashore (C. crassifolia var. crassifolia).
Both varieties are similar in vegetative, floral, and fruit
morphology. Although they grow in different, disjunct
habitats in central Argentina, there are some records of
C. crassifolia var. spinulosa from northern Patagonia (Cabrera
 SYSTEMATIC BOTANY [Volume 39

Fig. 1. Distribution and life forms of the species of Calycera.

et al. 32822, Bacigalupo & Nicora s. n., Durango s. n.) that Chamaephytic species of Calycera grow below 2,000 m ele-
extend its distribution towards the Atlantic shore along the vation [C. calcitrapa and C. sympaganthera (Ruiz and Pav.)
Rı́o Negro basin. This basin could probably be a biological Kuntze]. Calycera sympaganthera is the only species grow-
corridor connecting the distribution of both varieties. ing in Ultisoles of humid environments in a restricted area
2014] DENHAM ET AL.: MORPHOLOGY AND TAXONOMIC REVISION OF CALYCERA
from the Chilean Subantarctic Province (Cordillera de
Nahuelbuta, IX Región de la Araucanı́a) with cold, rainy,
seashore weather (Ibáñez 2008b). Calycera calcitrapa grows
in Entisols and Aridisols from valleys and mountain
bases (Moscatelli 1990) in the north of Monte Province
and its border with the Prepuneña Province (Cabrera and
Willink 1980).
Hemicryptophytic species of Calycera (C. herbacea Cav.,
C. horrida Hicken, and C. pulvinata J. Rémy) grow in higher
altitudes. Calycera horrida and C. pulvinata grow in Andean
arid environments (between 2,000 and 2,500 m, and above
4,000 m in elevation, respectively). In C. herbacea, the length
and development of scapiform peduncles and branches
show continuous variation from long peduncles and branches
at lower altitudes, to progressively shorter ones at higher
altitudes. In C. herbacea var. sinuata (Miers) Pontiroli, with
a wider geographic distribution, this altitudinal variation is
better recorded. Between 2,500 and 3,000 m, rosulate forms
with long peduncles (12 –15 cm) or long, erect stems that
are basally branched are present (Sanzin 345, Burkart 9311,
Covas 6980, Kiesling et al. 6856). Between 3,000 and 3,500 m,
individuals produce creeping stems, branched or not, and
shorter peduncles (5–8 cm) (Cabrera et al. 32470, Kiesling
3283, Kiesling et al. 9430). Between 3,500 and 4,000 m, indi-
viduals are strongly rosulate, unbranched, and possess the
shortest peduncles (1–3 cm) (Nicora et al. 8324, Zuloaga
et al. 9306b, Donadı́o et al. 121). Between 4,300 and 4,800 m,
individuals are strongly rosulate and stemless (Arroyo et al.
97034, Martin 458).
Inflorescence, Flower and Fruit Morphology—Variation
in calyx morphology and floral structure among Calycera
species are illustrated in Figs. 2 and 3, respectively. The
nine species and four varieties of the genus are illustrated
individually in Figs. 4–14, accompanying the descriptions
of the taxa.
The inflorescence structure of Calycera species is a cephalioid
(cf. Pozner et al. 2012). The terminal/central flower and soli-
tary (distal) flowers are present in every species of Calycera
(Fig. 2). Most species (C. calcitrapa, C. crassifolia, C. eryngioides,
C. herbacea, C. horrida, C. leucanthema, and C. sympaganthera)
develop peripheral (basal) cymose groups of up to seven
flowers each, a flat or dish-shaped involucre with a reduced
or more or less developed fused base, and free lobes
well differentiated from the normal leaves (Figs. 2, 6, 7).
Cephalioids of these species are not closely associated in
secondary inflorescences. However, the two remaining
species (Calycera sessiliflora and C. pulvinata) develop dif-
ferent inflorescences. Calycera sessiliflora does not pro-
duce cymose groups, but a terminal flower surrounded
by 3–5 solitary flowers (Fig. 2); the involucre has a
deep campanulate fused base with short lobes (Fig. 13);
cephalioids are usually nutant and associated in a sec-
ondary cymose inflorescence. In contrast, C. pulvinata pro-
duces peripheral, 2–5-flowered cymose groups, but the
involucre remains undifferentiated; normal leaves pro-
gressively reduce their petioles and blades, becoming
shorter, sessile, and less lobed; they are eventually expressed
as bracts of every lateral cephalioid, then they become bracts
of every cymose group, and finally bracts of every soli-
tary flower. Cephalioids are condensed in the center of the
rosulate plant, with a central, terminal cephalioid and sev-
eral surrounding lateral cephalioids. The transition between
the outermost cymose groups of the central cephalioid and
the first lateral cephalioids is so progressive that it is hard
to discern a sharp limit. Both C. sessiliflora and C. pulvinata
could be described as having secondary inflorescences.
With respect to the flower, diversity exists in flower
size, calyx morphology, hypanthium length, corolla tube
length, staminal tube length, and position of nectar glands
(Figs. 2 and 3).
Flower size ranges from ca. 3 mm in C. leucanthema and
C. sessiliflora, up to 9.5 mm in C. crassifolia and C. horrida.
Two types of calyx are present in the same cephalioid:
armed and unarmed, depending on the presence or absence
of spiniform sepals; intermediate forms may be present
(polymorphic calyx) or not (dimorphic calyx), and the
position of the two types within the inflorescence also
varies. Calycera crassifolia, C. pulvinata, C. sessiliflor, and
C. sympaganthera show a sharp dimorphism, with flowers
with spiniform and non-spiniform sepals. The remaining
species produce intermediate forms between the armed
and the unarmed types, and in these cases it is best to
describe this variation as calyx polymorphism. Flowers
with an armed calyx can appear at any position within
the inflorescence (Fig. 2). The central, terminal flower has
an armed calyx except for C. eryngioides and C. crassifolia
(the terminal flower of C. horrida has intermediate spiniform
sepals). Solitary flowers usually have non-spiniform sepals,
except for those of C. horrida and C. leucanthema that pro-
duce intermediate spiniform sepals. Except for C. sessiliflora,
spiniform sepals are always present in the flowers of
cymose groups. In Calycera pulvinata, C. sympaganthera,
and C. crassifolia, with calyx dimorphism, the armed calyx
of cymose groups are restricted to the first order flower. In
species with calyx polymorphism, the largest spiniform
sepals appear in the first order flower of the outermost
cymose groups, intermediate armed forms may appear in
the first order flower of inner groups (Calycera calcitrapa
and C. eryngioides), or in the second and third order flower
within each cymose group (Calycera herbacea, C. horrida, and
C. leucanthema). Flowers with armed calyces vary in three
aspects: 1) sepal morphology, 2) sepal number and relative
size, 3) sepal consistency at anthesis. Spiniform sepals can
be foliose (orbicular with flat section) with a pungent
apex (C. crassifolia, Fig. 5 B), long triangular with a narrow
naviculate transverse section (C. sympaganthera, Fig. 14 C),
acicular with a circular to triangular transverse section
(C. calcitrapa, C. eryngioides, C. herbacea, C. horrida, and
C. pulvinata; Figs. 4 H, 7 D–E, 8 B, 10 B, 12 B–C),
or basally navicular and distally acicular (C. sessiliflora;
Fig. 13 E). Usually, species with acicular spiniform sepals
produce one to three spiniform sepals per flower, unequal
in length, while those species with foliose, wide or narrow
pungent sepals (C. sympaganthera and C. crassifolia) keep
four to five spiniform sepals per flower, more or less equal
in length. Acicular spiniform sepals are hard and lignified
in cypselas, but they can be herbaceous (C. leucanthema) or
partially lignified at anthesis (i.e. C. calcitrapa, C. eryngioides,
C. herbacea, and C. horrida). In the latter case, spiniform
sepals become hardened from the distal end to base,
allowing basal growth during cypsela ripening. Foliose
or navicular spiniform sepals are usually herbaceous at
anthesis, becoming lignified at cypsela ripening.
The relative lengths of the hypanthium, corolla tube, and
corolla lobes (Fig. 3) produce perianth types according to
the species: long hypanthium and corolla tube, short lobes
 SYSTEMATIC BOTANY [Volume 39

Fig. 2. Calyx polymorphism according to flower position in Calycera species. Schematic drawings of half longitudinal sections of cephalioids.
Print version: terminal central flower (dark), solitary flowers (white), and flowers of the cymose groups (grey). Online version: terminal central
flower (red), solitary flowers (grey), and flowers of the cymose groups (green). Receptacle and bracts were represented in grey without outline;
dotted lines within the receptacle are artificial marks to assist cymose group interpretation. Cephalioid structure was based on Pozner et al. (2012).
All the cephalioids and flowers are at the same scale.
2014] DENHAM ET AL.: MORPHOLOGY AND TAXONOMIC REVISION OF CALYCERA

Fig. 3. Flower structure diagrams (style removed) of Calycera species. A. Calycera calcitrapa. B. Calycera crassifolia. C. Calycera eryngioides.
D. Calycera herbacea. E. Calycera horrida. F. Calycera leucanthema. G. Calycera pulvinata. H. Calycera sessiliflora. I. Calycera sympaganthera. Flower structure
follows Erbar (1993), where the tube between the sepals and the insertion of the staminal tube is recognized as the hypanthium. Ovary (o), sepals (s),
hypanthium (h), corolla tube (ct), and corolla lobes (cl) as a grey silhouette; staminal tube (st), filaments and anthers (a), in white. Nectarial glands (n)
are represented as black dots.

(C. horrida), long hypanthium and corolla lobes, short corolla
tube (C. sympaganthera, C. crassifolia), with corolla lobes
becoming progressively shorter in C. herbacea, C. calcitrapa,
C. eryngioides, and C. pulvinata. Calycera leucanthema and
C. sessiliflora have the smallest flowers in the genus, with
a very reduced hypanthium.
Calycera species also vary in development of the staminal
tube and filaments, anther size, and position of nectar
glands (Fig. 3). The staminal tube is usually short, between
0.2–0.5 mm long, but it is almost absent in C. sessiliflora,
and reaches 2 mm in C. horrida. Staminal filaments are
usually 0.3–0.5 mm long, reaching 0.7 mm in C. eryngioides,
and being almost absent in C. leucanthema. Anthers are
usually 1–1.5 mm long, reaching 2 mm in C. horrida, or
reduced to 0.6 mm in C. sessiliflora. Concerning anther
morphology, Calycera species may have anthers obtuse or
sagittate at base. In obtuse anthers, the base of each theca
is roundish and fused to the connective and filament. In
sagittate anthers, the base of each theca is acute, separate
and divergent from the connective and filament as an
arrowhead base. In both types, a short, solid, conical pro-
jection may grow at the very end of the base of each
theca, and called a “caudicle” in this revision. In some
specimens of C. eryngioides, C. horrida, and C. sessiliflora,
the connective is slightly apiculate.
There are always five nectar glands alternating with the
staminal filaments. Nectar glands are similar in morphology
to those described by Erbar (1993) for Acicarpha. They can
be seen as ovoid swellings, between 0.1–0.6 mm, differen-
tiated just at the junction of the corolla tube, hypanthium
and staminal tube, so that the upper half of the gland is
within the base of the staminal tube, and the lower half
of the gland is within the hypanthium (Fig. 3).
Finally, calyx morphology in the cypsela is congruent
with calyx morphology in the flower and with flower
position within the inflorescence. Calycera species with
dimorphic calyces produce dimorphic cypselas (armed and
unarmed), and those species with polymorphic calyces
produce polymorphic cypselas with several intermediate
armed forms. Usually, armed cypselas have a smooth
pericarp surface, while unarmed cypselas have irregu-
lar, transverse wrinkles on the pericarp. Intermediate
armed cypselas of species with fruit polymorphism also
develop a partially wrinkled pericarp at the cypsela base
(Fig. 9 C–G).
Taxonomic Treatment
CALYCERA Cav., Icon. [Cavanilles] 4: 34, 1797.—TYPE: Calycera
herbacea Cav.
 SYSTEMATIC BOTANY [Volume 39

Leucocera Turcz., Bull. Soc. Imp. Naturalistes Moscou 21 (1): sis. Armed calyx usually in the terminal central flower and
583, 1848.—TYPE: Leucocera annua Turcz. [= Calycera the first order flower of the outer cymose groups. Hypan-
leucanthema (Poepp. ex Less.) Kuntze]. thium tubulose, long to short. Corolla with campanulate or
cylindrical tube and free lobes variable in length, with dif-
Gymnocaulus Phil., Linnaea 28: 705, 1856.—TYPE: Gymnocaulus
ferent proportions according to the species. Staminal tube
viridiflorus Phil. [= Calycera herbacea Cav. var. herbacea].
and filaments reduced to well developed, with different
Anomocarpus Miers, Ann. Mag. Nat. Hist. ser. 3, 6(35): 351, proportions according to the species. Anthers oblong, thecae
1860. [Nov 1860].—TYPE: designated here, Anomocarpus basally obtuse or sagittate, caudicles present or absent.
axillaris Miers [= Calycera sessiliflora Phil.]. Nectar glands 5, alternating the stamens just at the junc-
tion of the corolla tube, hypanthium and staminal tube.
Chamaephytic subshrubs, cryptophytics, hemicryptophytic
Cypselas prismatic, narrow at base, with 3–5 ridges, crowned
rosulate, or therophytic caulescent herbs, glabrous or pilose.
by short, reduced triangular to rounded sepals, or wide/
Leaves sessile or petiolate, blade lanceolate-oblong, entire to
squarrose-slashed, margin usually toothed, teeth mucronate. long triangular to conic-spiniform, lignified sepals, accord-
Inflorescence a cephalioid, solitary or arranged in secondary ing to the flower position and sepal dimorphism or poly-
inflorescences. Peduncles indistinct from the uppermost morphism described above. Usually, ridges of unarmed
internodes of the stem or very much distinct and scapiform. cypselas have transverse wrinkles, while armed cypselas
Inflorescence bracts clearly differenciated from distal upper- have smooth ridges. Intermediate cypselas may have a
most leaves. Involucre differentiated or not; in the first few wrinkles at base.
case, the outermost bracts are clearly differentiated in Distribution—The nine species and four varieties included
shape and size from the inner bracts (usually spathulate in the genus Calycera are South American endemics, dis-
to linear), having a fused base (and involucral bracts as tributed across Argentina, Bolivia, Chile, and Peru. The
lobes) or not (and involucral bracts free); in the second phytogeographic distribution of Calycera corresponds with
case, there is a progressive reduction in shape and size the “Andino-Patagónico” (Andean-Patagonian) Domain
of the outermost bracts, towards the central, innermost (Altoandina, Puneña, del Desierto, Chilena Central, and
bracts of the cephalioid. Receptacle reduced, punctiform. northern Patagonian Provinces), with the “Chaqueño” Domain
Flowers pentamerous, rarely tetramerous. Calyx basically (Prepueña, del Monte, and Pampeana Provinces), and with
of two types in the same cephalioid: armed or unarmed, the Subantarctic Domain (northen Subantarctic Province)
without intermediate forms (dimorphic calyx) or with inter- (Cabrera and Willink 1980).
mediate forms (polymorphic calyx). Armed calyx usually Notes—The species Anomocarpus axillaris is selected as
with 2–5 unequal, needle shape or flat sepals, lignified or the type of the genus Anomocarpus because the author did
not at anthesis. Unarmed calyx with 5 reduced, triangular not designate a type species in the original publication.
to round, herbaceous sepals. Intermediate calyx usually The features of A. axillaris match exactly the protologue of
with 5 unequal, long triangular sepals, herbaceous at anthe- the genus.

Key to the Species and Varieties of CALYCERA

1. Involucre indistinct. Althougth floral bracts are clearly different from the normal leaves, the outermost are not arranged
into a distinct involucre. All floral bracts are wide lanceolate, free, the outermost with apex entire or 3-toothed,
the inner ones progressively smaller with apex entire. Rosulate herbs with a central cephaloid surrounded by
several smaller ones crowded at the center of the plant. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calycera pulvinata
1. Involucre distinct. Floral bracts clearly different from the normal leaves, variable in shape, the outermost arranged
into a distinct involucre (sometimes with a fused base), the inner ones narrow and very much reduced or absent . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Fused base of the involucre absent or reduced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Flowers tetramerous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calycera leucanthema
3. Flowers pentamerous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Spiniform sepals not hardened at anthesis. Anthers sagittate at base. Cypselas armed and unarmed,
without intermediate forms. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calycera sympaganthera
4. Spiniform sepals already hardened at anthesis. Anthers obtuse at base. Cypselas armed and unarmed,
with progressive intermediate forms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5. Suffrutex. Leaves sessile, blade lanceolate-oblong, entire. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calycera calcitrapa
5. Rosulate herbs. Leaves petiolate, blade oblong, squarrose-slashed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calycera herbacea var. herbacea
2. Involucre with a fused campanulate, dish- or wheel-shaped base, involucral bracts as lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6. Involucre base campanulate. Cephaliods on short peduncles (shorter than the involucre),
with few (about 10) flowers; cymose groups absent. Total length of hypanthium and corolla, 2.5 mm. . . . . . . . . . . . . . . Calycera sessiliflora
6. Involucre base dish- or wheel-shaped. Cephaliods on long peduncles (longer than the involucre), with
many (more than 20) flowers; cymose groups few to many. Total length of hypanthium and corolla, 4 mm or longer . . . . . . . . . . . . . . . . 7
7. Plants woolly on stems, leaves, peduncles and bracts. Corolla tube long (4 mm). Filaments short (0.1–0.3 mm).
Anthers sagittate at base. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calycera horrida
7. Plant glabrous (sometimes only pilose on peduncles). Corolla tube short (1 mm). Filaments long (0.3–0.6 mm).
Anthers obtuse at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Spiniform sepals 5, wide triangular, flat, herbaceous at anthesis, hardened and pungent in the cypsela.
Plants with deep rhizomes and emergent, aerial, erect shoots. Upper leaves sessile . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9. Aerial erect shoots 20–75 cm tall. Lower blades oblong, margin toothed-apiculate.
Involucre with lobes triangular. Staminal tube 1/2 the filament length, filaments 1/3 the
anther length. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calycera crassifolia var. crassifolia
9. Aerial erect shoots 15–30 cm tall. Lower blades oval, margin toothed-aristate. Involucre
with lobes triangular to rectangular. Staminal tube as long as the filament, filaments
1/5 the anthers length. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calycera crassifolia var. spinulosa
2014] DENHAM ET AL.: MORPHOLOGY AND TAXONOMIC REVISION OF CALYCERA
8. Spiniform sepals usually less than 5, needle-shaped or long lanceolate, middle transverse
section triangular or circular, usually hardened at anthesis. Plants without rhizomes,
rosulate perennials or caulescent annuals. Upper leaves petiolate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10. Terminal central flower with spiniform sepals. Peduncles glabrous. Rosulate to
caulescent herbs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calycera herbacea var. sinuata
10. Terminal central flower without spiniform sepals. Peduncles woolly. Caulescent herbs. . . . . . . . . . . . . . . . . Calycera eryngioides
CALYCERA CALCITRAPA Griseb., Abh. Königl. Ges. Wiss.
Göttingen 19: 163. 1874.—TYPE: ARGENTINA. Catamarca:
“in convallibus arenosis inter Nacimiento et Laguna
Blanca”, Jan 1872, P. G. Lorentz 432 (holotype: CORD
00006342!; isotypes: BA 42208!, K 000588753 not seen,
photo of K at SI!).
Chamaephytic, glabrous subshrubs, with erect stems about
50–60 cm tall. Leaves sessile, blade lanceolate-oblong, entire,
margin toothed, teeth mucronate, apex obtuse, apiculate or
aristate. Peduncles indistinct from the uppermost inter-
nodes of the stem. Involucre bracts clearly differentiated
from distal uppermost leaves, with a reduced, wheel-shaped
fused base and triangular lobes, the lobes with margin
entire or with mucronate teeth, apex mucronate. Receptacle
punctiform, with spathulate to linear bracts towards the
centre. Flowers pentamerous. Calyx polymorphic: 2–3 unequal
sepals, up to 5 mm, acicular, with triangular section, ligni-
fied at anthesis (terminal central flower, and terminal flower
of the outer cymose groups); 5 unequal sepals, shorter, up
to 3 mm, flat-triangular, with plane section, non-lignified
at anthesis (terminal flower of the inner cymose groups);
5 equal sepals, up to 0.5 mm, lanceolate, non-lignified at
anthesis (second and third order flowers of cymose groups
and solitary flowers). Hypanthium 3.5–4 mm, green, tubulose.
Corolla white, deeply lobed; tube reduced, 0.6–0.7 mm;
lobes 1–1.2 mm, oblong, flat, recurved, apex obtuse, non
uncinate. Staminal tube 0.15 mm. Filaments 0.3 mm. Anthers
1.3 – 1.4 mm, oblong, thecae basally obtuse. Nectaries
0.3 mm long. Cypselas prismatic, armed and unarmed,
narrow at base, with 3–5 smooth ridges, crowned by short
lanceolate -or triangular- to conic-spiniform sepals accord-
ing to the flower position and sepal polymorphism described
above. Figure 4.
Distribution—Endemic to northwestern Argentina, from
San Juan, La Rioja, Catamarca, to southern Salta and north-
western Tucumán, between 400 and 1,900 m in elevation.
Habitats of Calycera calitrapa include valleys and foothills
with Entisols and Aridisols from “El Monte” phytogeo-
graphic Province and its limit with the “Prepuneña” phyto-
geographic Province.
Notes—Calycera calcitrapa is a subshrub with sessile leaves.
Upper leaves are sessile also in C. crassifolia, but the latter
is a rhizomatous cryptophyte. The terminal flower of
C. calcitrapa may eventually have an intermediate armed
calyx, like second- and third-order flowers of cymose groups,
or unarmed at all.
Representative Specimens Examined—ARGENTINA. Catamarca:
Andalgalá, Campo del Arenal, 4 Dec 1917, Jörgensen 1845 (SI); Belén,
30 Jan 1973, Ulibarri 303 (SI); Santa Marı́a, El Cajón, Saladillo, 13 Jan
1914, Castillón 3339 (LIL, SI); Tinogasta, Ruta Nac. N
45, entre
Alpasinche y Tinogasta, a 35 km de la primera, 29 Nov 1997, Biurrun
et al. 4938 (SI). La Rioja: Arauco, Aimogasta, 1 Mar 1944, A. T.
Hunziker 5001 (SI); Gral. F. Varela, Talampaya, al S de Villa Unión,
20 Nov 1982, Kiesling and Saenz 4270 (SI); Independencia, Guayapa
(15 km al SW de Patquı́a), 16 Sep 1947, Lourteig 1195 (SI). Salta:
Cafayate, 16 Jan 1976, Cabrera et al. 27277 (SI); Guachipas, Quebrada
de Guachipas, Dec 1896, C. Spegazzini s. n. (SI 12667); San Carlos,
Amblayo, Jan 1897, C. Spegazzini s. n. (SI 12666). San Juan: Caucete,
al Este de Caucete, médanos, 20 Mar 2005, Kiesling 10227 (SI); Ullún,
entre la Estación y el Dique de Ullún, 11 Feb 1984, Kiesling 4366
(SI); San Agustı́n del Valle Fértil, 23 Nov 1986, Haene 475 (SI); Valle
Fértil, camino a Usno, without date, Kiesling 3075 (SI). Tucumán:
Tafı́ del Valle, Amaicha al Bañado, 1,840 m, 12 Feb 1913, Castillón
2636 (SI).
CALYCERA CRASSIFOLIA (Miers) Hicken, Physis (Buenos Aires)
2: 117. 1916. Acicarpha crassifolia Miers, Ann. Mag. Nat.
Hist. ser. 3, 6(36): 402. 1860 [Dec 1860]. Boopis crassifolia
(Miers) A. Gray, Proc. Amer. Acad. Arts. 5: 321. 1861.—
TYPE: URUGUAY. Maldonado: date not indicated,
J. Tweedie 1068 (holotype: K 000588754 not seen, photo
of K at SI!; isotype: BM 000947754 not seen, photo of
BM at SI!).
Cryptophytic herbs with deep, white rhizomes and
emergent, aerial, erect shoots: erect shoots about 15–75 cm
tall above ground, glabrous. Lower leaves with the base
attenuate into a rectangular short petiole, blade oval or
oblong, margin toothed-apiculate or toothed-aristate, apex
obtuse, apiculate or aristate. Upper leaves sessile, equal in
shape as the lower ones. Peduncles indistinct from the
uppermost internodes of the stem. Involucre bracts clearly
differentiated from distal uppermost leaves with a fused
dish-shaped base and lobes triangular and rectangular,
the lobes with margin entire, toothed-apiculate or toothed-
mucronate, and apex mucronate or aristate. Receptacle
punctiform, bracts lanceolate to spathulate towards the
centre. Flowers pentamerous. Calyx dimorphic: 5 equal
sepals, 0.7 mm, orbicular, with flat section, non-lignified
at anthesis (terminal central flowers, and all remaining
flowers except the terminal flower of the outer cymose
groups); 5 equal sepals, 1.5 mm wide, triangular, with
flat-naviculate section, and pungent to spinulose apex,
non-lignified at anthesis (the terminal flower of the outer
cymose groups). Hypanthium 4 mm, green, tubulose.
Corolla white, deeply lobed; tube 1 mm, reduced; lobes
3 mm, oblong, flat, apex obtuse, non-uncinate. Staminal
tube 0.25 mm. Filaments 0.5 mm. Anthers 1.5 mm, oblong,
thecae basally obtuse. Nectaries 0.5 mm long. Cypselas
prismatic, very narrow at base, with 4–5 smooth ridges,
crowned by suborbicular (unarmed cypselas) or triangular
spiniform sepals (armed cypselas), according to the flower
position and sepal dimorphism described above. Figures 5
and 6.
Distribution—Calycera crassifolia typically grows in sandy
soils (Entisols and Aridisols) with a wide geographical
range from the foothills in Mendoza and Neuquén to the
Atlantic shore of Buenos Aires and Uruguay.
Notes—Calycera crassifolia is similar to C. calcitrapa in
the shape of the leaves; the latter species differs by being
a subshrub.
CALYCERA CRASSIFOLIA (Miers) Hicken var. CRASSIFOLIA.
Boopis crassifolia (Miers) A. Gray var. spinuligera Speg.,
Anales Soc. Ci. Argent. 48: 174. 1899. Calycera crassifolia
(Miers) Hicken var. spinuligera (Speg.) Hicken, Reun. Nac.
Soc. Arg. Cienc. Nat. 1: 244. 1919.—TYPE: ARGENTINA.
 SYSTEMATIC BOTANY [Volume 39

Fig. 4. Calycera calcitrapa. A. Habit. B. Inflorescence. C. Solitary flower. D. Terminal flower of the outer cymose group. E. Diagrams of flowers
illustrating proportions and shape of hypanthium and corolla. F–I. Cypsela polymorphism. From Kiesling 3075 (SI).

Buenos Aires: Médanos de Punta Rubia, Bahı́a de Both varieties can be distinguished by the characters in
San Blas, 3 Feb 1898, C. Spegazzini s. n. (holotype: LPS the key.
11029!; isotype: CORD 00005205, not seen, photo of Distribution—This variety is known from Argentina and
CORD at SI!). Uruguay; it grows in Rı́o de la Plata river dunes (Uruguayan
2014] DENHAM ET AL.: MORPHOLOGY AND TAXONOMIC REVISION OF CALYCERA

Fig. 5. Calycera crassifolia var. crassifolia. A. Habit. B–C. Cypsela dimorphism. D. Diagrams of flowers illustrating proportions and shape of
hypanthium and corolla. A. From Cabrera 32314 (SI). B–C. From Zanotti 144 (SI).

shore) and in dunes of the Atlantic seacoast. The northern
limit of its distribution is Cabo Polonio (Department of
Rocha) in Uruguay, and its southern limit at Department
of Biedma (Chubut Provincie) in Argentina.
Representative Specimens Examined—ARGENTINA. Buenos Aires:
Partido Cnel. Rosales, Pehuén Co, 14 Oct 1960, Verettoni 1912 (SI); Partido
Cnel. Dorrego, Monte Hermoso, 12 Jan 2007, Deginani 2022 (SI); Partido
Gral. Alvarado, Miramar, Jan 1949, Burkart 17824 (SI); Partido Gral.
Pueyrredón, Mar del Plata, 2 Feb 1937, Nicora 1335 (SI); Partido de la
Costa, San Clemente del Tuyú, 30 Jan 1939, Cabrera 4923 (SI); Partido
Mar Chiquita, Mar Chiquita del Sud, Dec 1905, Fablet s. n. (SI); Partido
Necochea, playa, 9 Dec 1964, Boelcke et al. 11989 (SI); Partido Tres Arroyos,
San Francisco de Bellocq, 14 Nov 1942, Mollura 1008 (SI); Partido Villa
Gessel, Mar Azul, a 15 km al S de Villa Gesell, 15 Jan 2008, Zanotti and
Donadı́o 144 (SI). Chubut: Biedma, Puerto Madryn, en las dunas
playeras enfrente del CENPAT, 6 Nov 2006, Zanotti 61 (SI); Biedma,
Penı́sula Valdés, Jan 1971, Cabrera 21202 (LP). Rı́o Negro: Adolfo
Alsina, Embouchure der Rı́o Negro, Feb 1912, Hauman-Merck 259 (SI).
URUGUAY. Maldonado: Piriápolis, Jan 1911, Hicken 181 (SI). Rocha:
Isla Gorriti y Cabo Polonio, Jan 1922, Felippone 4941 (SI); La Pedrera,
Jan 1981, Cabrera 32314 (SI). San José: Estancia Pascual [Playa Pascual],
28 Feb 1931, Herter 86707 (SI).
Calycera crassifolia (Miers) Hicken var. spinulosa (Gillies
ex Miers) Zavala, S. Denham and Pozner, comb. nov.
Calycera spinulosa Gillies ex Miers, Ann. Mag. Nat. Hist.
ser. 3, 6(36): 399. 1860 [Dec 1860].—TYPE: ARGENTINA.
Mendoza: on sandy soil, near El Totoral and Los Arbolitos,
4 Nov 1824, J. Gillies s. n. (holotype: K 000588765 not
seen, photo of K at SI!; isotype: BM 000947751 not
seen, photo of BM at SI!, E 00392101 not seen).
Calycera spinulosa Gillies ex Miers var. serratifolia Miers, Ann.
Mag. Nat. Hist. ser. 3, 6(36): 399. 1860 [Dec 1860].—
TYPE: ARGENTINA. Mendoza: near La Casa Pintada,
date not indicated, J. Gillies s. n. (holotype: K 000677638
not seen, photo of K at SI!).
Calycera foliosa Phil. ex Reiche, Anales Univ. Chile 106: 1046.
1900, syn. nov.—TYPE: ARGENTINA. Mendoza: Valle
del Tunuyán, 1871, F. Leybold s. n. (holotype: SGO not
seen; isotype: SI 12652!).
Calycera neuquenensis Suess., Repert. Spec. Nov Regni Veg.
51: 197. 1942.—TYPE: ARGENTINA. Neuquén: Cobunco,
700 m, date not indicated, E. Ammann 12 (holotype: M
not found).
 SYSTEMATIC BOTANY [Volume 39

Fig. 6. Calycera crassifolia var. spinulosa. A. Habit. B–C. Cypsela dimorphism. D. Diagrams of flowers illustrating proportions and shape of
hypanthium and corolla. A. From Zuloaga et al.12250 (SI). B–C. From Cabrera 32822 (SI).

Distribution—Endemic to Argentina, growing at the
Andean mountain bases of Mendoza and northern Neuquén,
and extending to western La Pampa and northern Rı́o
Negro, from 170 to about 2,600 m in elevation.
Notes—When comparing Calycera spinulosa and C. crassifolia,
taxa formerly treated as distinct species, we observed that
morphological differences between them are polarized at
the extremes of a geographical distribution of what is better
considered a single species: the Andean piedmont and
the Atlantic coast. These taxa are likely connected along
the basins of Colorado and Negro rivers. We thus con-
sider C. spinulosa a variety of C. crassifolia.
Calycera foliosa Phil. ex Reiche had been placed under the
synonymy of Calycera crassifolia (Miers) Hicken (Hicken
1919). However, the specimens cited in the protologue were
collected in the Cordillera de Santiago and in Tunuyán,
while C. crassifolia var. crassifolia grows in the Atlantic
sea-shore dunes from Argentina and Uruguay. Although
the type material we examined does not possess flowers
or cypselas, vegetative features and the original description
2014] DENHAM ET AL.: MORPHOLOGY AND TAXONOMIC REVISION OF CALYCERA
by Reiche (1900) make clear that C. foliosa is conspecific
with C. crassifolia var. spinulosa.
Representative Specimens Examined—ARGENTINA. La Pampa: Chical-
Co, about 5 km S of Cerro Centinela, 1 Jan 1982, Pedersen 13329 (LP);
Puelén, Mar 1932, Durango s. n. (SI 107617). Mendoza: Malargüe, Sa.
del Nevado, portezuelo de los Cos. de Mondaca al SE del Co. Negro,
1,760 m, 12 Dec 1973, Boelcke et al. 15857 (SI); San Carlos, Cerro
Guanacos, 2,600 m, Jan 1921, Carette 393 (SI); San Rafael, Ruta Provin-
cial 180 entre cruce de la Ruta Provincial 184 y El Nihuil, 34
570 0500 S,
68
400 1500 W, 1,300 m, 20 Nov 2010, Zuloaga et al. 12271 (SI); San Rafael,
Ruta Nacional 184, de Los Toldos a Ruta Nacional 180, 20 Nov 2010,
Zuloaga et al. 12250 (SI); Tupungato, Los Álamos, 2,300 m, 27 Dec
1949, Paci and Melis 49 (LIL). Neuquén: Confluencia, Plaza Huincul,
3 Dec 1943, Platnick 54 (LP); Pehuenches, along route 40 generally
1–2 km N of Barrancas, 3,200 ft, 14 Jan 1985, King and Heinz 9421 (SI);
Añelo, Ruta Provincial 17, entre Aguada Pichana y Añelo, 38
260 0500 S,
68
570 2000 W, 420 m, 1 Dec 2010, Zuloaga et al. 12671 (SI); Collón Curá,
Ruta Nacional 237, entrada Dique Piedra del Águila, 3,6 km S Piedra
del Águila, 40
040 2700 S, 70
070 1800 W, 544 m, 16 Dec 2008, Nicola 115 (SI).
Rı́o Negro: Avellaneda, Chelforó, 6 Dec 1981, Cabrera et al. 32822 (SI);
Gral. Roca, Paso Córdoba, en médano al pie de las bardas, 6 Nov
1972, Bacigalupo and Nicora s. n. (BAA 11619, SI).
CALYCERA ERYNGIOIDES J. Rémy, Fl. Chil. [Gay] 3(3): 254, 1847
[1848]. Anomocarpus eryngioides (J. Rémy) Miers, Ann.
Mag. Nat. Hist. ser. 3, 6(35): 353. 1860 [Nov 1860].—
TYPE: CHILE. Prov. Santiago, date not indicated, C. Gay
1066 (holotype: P not seen; isotypes: BM 000947744
not seen, K 000588755 not seen, photo of K at SI!, K
000588756 not seen, photo of K at SI!, LIL 281133!).
Therophytic, caulescent herbs, about 15 cm tall. Leaves
spathulate, base attenuate into a narrow petiole, blade lan-
ceolate, margin entire, slightly sinuate or crenate-mucronate,
apex obtuse, mucronate. Peduncles woolly, indistinct from
the uppermost internodes of the stem. Involucre bracts
clearly differentiated from distal uppermost leaves with a
wheel-shaped fused base and triangular, accrescent lobes,
the lobes with margin entire, apex mucronate. Receptacle
punctiform, bracts spathulate to linear towards the centre.
Flowers pentamerous. Calyx polymorphic: 5 equal, up to
1 mm, suborbiculate, emarginate, margin crenate, non-
lignified sepals at anthesis (terminal central flower, soli-
tary flowers, and flowers of the inner cymose groups);
2–5 unequal, up to 2 mm, narrowly triangular, non-lignified
at anthesis sepals (the terminal flower of the intermediate
cymose groups); 2–5 unequal, up to 3 mm, acicular with
triangular section, lignified at anthesis sepals (terminal
flower of the outermost cymose groups). Hypanthium green,
tubulose. Corolla white, deeply lobed; tube reduced; lobes
oblong, flat, slightly recurved, apex obtuse, straight. Staminal
tube 0.2–0.3 mm. Filaments 0.7 mm, ½ the anthers length.
Anthers 1.5 mm, lanceolate, thecae basally obtuse. Nec-
taries 0.4 mm long. Cypselas prismatic, with 3–5 wrinkled
(unarmed cypselas) or flat ridges (armed cypselas), crowned
by roundish-folded, triangular or spiniform sepals sur-
rounding a central apiculum, according to the flower posi-
tion and sepal polymorphism described above. Figure 7.
Distribution—Although this species has been recorded
for Argentina (Hicken 1919; Pontiroli 1963) there is no
specimen cited to substantiate that it grows outside of Chile.
The specimen Philippi s. n., from Argentina, cited in Zanotti
and Pozner (2008) is a misidentification. We conclude that
Calycera eryngioides is endemic to “Chilena Central” Prov-
ince, from IV Región de Coquimbo to VI Región del
Libertador Gral. Bernardo O’Higgins, growing on argillic
soils (Alfisols and Vertisols).
Notes—Calycera eryngioides might be confused with
C. calcitrapa by the inflorescence and the erect, caulescent
habit (Pontiroli 1963). However, C. calcitrapa is a completely
glabrous subshrub with sessile leaves that is endemic to
northwestern Argentina.
Representative Specimens Examined—CHILE. IV Región de Coquimbo:
Limarı́, Cordillera Ovalle, La Hualtata, 2,300 m, 29 Oct 1949, Jiles-P.
1558 (LIL). VI Región del Libertador Gral. Bernardo O’Higgins:
Cachapoal, Cordillera de la Compañı́a, Nov 1853, R. A. Philippi s. n.
(SI 12662). Región Metropolitana: Santiago, Piuquencillos, Valle del Rı́o
Colorado, 2,000–3,500 m, 8–10 Dec 1942, Pisano-V. et al. 1646 (CONC).
CALYCERA HERBACEA Cav., Icon. [Cavanilles] 4: 34. 1797.—
TYPE: CHILE. Ex Cordillera del Portillo in Regno
Chilensi, date not indicated, L. Née s. n. (holotype: MA
475481 not seen, photo of MA at SI!).
Hemicryptophytic, rosulate or caulescent herbs, with
erect or prostrate stems, glabrous, up to 30 cm tall, with
contractile root. Leaves spathulate, base attenuate into a
rectangular petiole, blade oblong, squarrose-slashed, margin
with a distinct narrow, semi-translucent rim and few to abun-
dant glandular trichomes, toothed or mucronate. Peduncles
scapiform, long, clearly distinct from the stem. Involucre
with or without a dish-shaped fused base, involucral bracts
clearly differentiated from distal uppermost leaves, rectangu-
lar or triangular, with margin entire or toothed-mucronate,
apex obtuse, mucronate. Receptacle punctiform, bracts
linear, spathulate and rectangular, irregularly arranged.
Flowers pentamerous. Calyx polymorphic: 2–5 unequal
sepals, up to 8 mm, acicular, triangular section, lignified
at anthesis (terminal central flower and terminal flower
of cymose groups); 2–5 subequal, long triangular, flat,
1.5–2 mm, non-lignified at anthesis (second and third
order flowers of cymose groups); 5 equal, triangular,
short, flat, 0.3–0.5 mm sepals, non-lignified at anthesis
(solitary flowers and second and third order flowers of
cymose groups). Hypanthium 3–4 mm, green, tubulose.
Corolla green or white, deeply lobed; tube reduced, 1mm;
lobes 1.8–2 mm, oblong, flat, slightly recurved, apex obtuse,
uncinate. Staminal tube 0.4–0.5 mm. Filaments 0.3–0.4 mm.
Anthers 1–1.6 mm, oblong, thecae basally obtuse, with or
without caudicles. Nectaries 0.5–0.6 mm. Cypselas pris-
matic, with 3–5 ridges, free (furrows visible) or connate
(furrows concealed); unarmed cypselas with ridges usually
wrinkled and vestigial or triangular sepals; armed cypselas
with smooth ridges and spiniform curved sepals, according
to the flower position and sepal polymorphism described
above. Figures 8 and 9.
Distribution—Calycera herbacea occurs in the southern
Andes, from northern Chile and northwestern Argentina
to Mendoza Province in Argentina.
Notes—Sometimes flowers of second and third order of
the outer cymose groups develop intermediate spiniform
sepals.
Typically, leaves of Calycera herbacea posses a distinct
narrow, semi-translucent rim all around the margin. That
foliar rim develops abundant and noticeable glandular
trichomes in individuals from northwestern Argentina
(C. herbacea var. sinuata), becoming gradually scarce and
unnoticed in individuals from the southern distribution
(C. herbacea var. herbacea).
Calycera herbacea might be misidentified as C. horrida
because of its hemicryptophytic life form and rosulate
habit, and scapiform peduncles. However, C. horrida is
 SYSTEMATIC BOTANY [Volume 39

Fig. 7. Calycera eryngioides. A. Habit. B–E. Cypsela polymorphism. F. Diagrams of flowers illustrating proportions and shape of hypanthium
and corolla. A. From Weddel (1857)[1858]. B–F. From Jiles-P 1558 (LIL).

an herb with gemiferous roots, pinnatisect leaves, woolly
indument, and a long corolla tube.
Richard (1820) reassigned the holotype of C. herbacea to
Calycera cavanillesi. Both names were used indistinctly in the
literature until Pontiroli (1963) placed Calycera cavanillesi
under Calycera herbacea. Calycera cavanillesi Rich., Mem.
Mus. Hist. Nat. 6: 34. 1820, is a nomen superfluum.
Calycera squarrosa Miers, Ann. Mag. Nat. Hist. ser. 3, 6(36):
CALYCERA HERBACEA Cav. var. HERBACEA.
Gymnocaulus viridiflorus Phil., Linnaea 28: 706. 1856, syn.
nov. Calycera viridiflora (Phil.) Miers, Ann. Mag. Nat.
Hist. ser. 3, 6(36): 399. 1860 [Dec 1860]. Calycera herbacea
Cav. var. viridiflora (Phil.) Pontiroli, Revista Mus.
La Plata, Secc. Bot. 9(41): 191. 1963.—TYPE: CHILE.
Cordillera de Maule, 1856, P. Germain s. n. (holotype:
SGO 057270 not seen; isotypes: K 000588766 not seen,
photo of K at SI!, SI 090136!, SI 090137!).
398. 1860 [Dec 1860], syn. nov. Calycera viridiflora (Phil.)
Miers f. squarrosa (Miers) Hicken, Reun. Nac. Soc.
Arg. Cienc. Nat. 1: 247. 1919.—TYPE: ARGENTINA.
On the Andes of Mendoza, date not indicated, J. Gillies
2014] DENHAM ET AL.: MORPHOLOGY AND TAXONOMIC REVISION OF CALYCERA

Fig. 8. Calycera herbacea var. herbacea. A. Habit. B–D. Cypsela polymorphism. E. Diagrams of flowers illustrating proportions and shape of hypan-
thium and corolla. A. From Chiapella (1999b, INTA, Flora Patagónica). B–D. From Pozner et al. 563 (SI).

s. n. (holotype: K 000588764 not seen, photo of K
at SI!; isotype: BM 000947747 not seen, photo of BM
at SI!).
Distribution—This variety is endemic to the southern
Andes; it grows between 2,200–3,000 m in elevation in the
Metropolitana Region and Cordillera de Maule (Chile), and
at lower altitude (1,650 m) in Mendoza Province (Argentina).
Notes—The two varieties can be distinguished by the
following characters: Calycera herbacea var. herbacea are
rosulate herbs possessing flowers with a green corolla,
anthers without caudicles, involucral bracts free, rectangular,
 SYSTEMATIC BOTANY [Volume 39

Fig. 9. Calycera herbacea var. sinuata. A. Habit. B. Leaf. C–G. Cypsela polymorphism. H. Diagrams of flowers illustrating proportions and shape
of hypanthium and corolla. A. From Kiesling et al. 9430 (SI). B. From Kiesling 3283 (SI). C–G. From Kiesling 6188b (SI).

small, reflexed and hidden by the flowers, with margin
entire, cypselas with 4 – 5 connate ridges concealing
the furrows, and armed cypselas with outward-curved
spiniform sepals. Calycera herbacea var. sinuata includes
rosulate or caulescent herbs, flowers with white corolla,
anthers usually with caudicles, involucre with a dish-
shaped fused base, lobes rectangular to triangular,
basally broad, spreading, not hidden by the flowers, with
margin entire or toothed-mucronate, cypselas with 3–5 free
ridges, visible furrows, armed cypselas with inward-curved
spiniform sepals.
Reproductive structures of C. herbacea var. viridiflora and
C. herbacea var. herbacea are identical, and diagnostic char-
acters described elsewhere for var. viridiflora (stem partially
branched, leaves ovate-oblong, irregularly pinnatifid) are
extremely variable within the species. Therefore, C. herbacea
var. viridiflora is reduced to the synonymy of the type variety.
Calycera nudicaulis Phil. ex Miers, Ann. Mag. Nat. Hist.
ser. 3, 6(36): 399. 1860 [Dec 1860], is a nomen nudum for
this taxon.
Representative Specimens Examined—ARGENTINA. Mendoza:
Malargüe, altos valles de El Sosneado, 2,200 m, 19 Feb 1942, Burkart
2014] DENHAM ET AL.: MORPHOLOGY AND TAXONOMIC REVISION OF CALYCERA
et al. s. n. (SI 14321); San Carlos, Reserva Laguna del Diamante,
Quebrada del Paso Cruz de Piedra, 34
130 5000 S, 69
250 1900 W,
2,646 m, 6 Dec 2009, Zavala 21 (SI); San Rafael, Hotel Termas del
Sosneado, 34
460 1200 S, 70
030 3300 W, 2,180 m, 22 Nov 2010, Zuloaga
et al. 12381 (SI); Sa. del Nevado, zanjón del Plateado frente a los
Cos. Morados, 35
400 S, 68
230 W, 1,650 m, 8 Dec 1973, Boelcke et al.
15732 (SI); Tupungato, Nov 1981, Wingenroth et al. 381 (SI).
CHILE. Región Metropolitana: Cordillera, Along the Embalse
El Yeso, near the upstream end, along the access road 6 km upstream
from the dam. Rock fields and alluvial fans, 33
350 4000 S, 70
100 1500 W,
2,520 m, 14 Jan 1993, Taylor and Gereau 10924 (CONC, SI); Cajón rı́o
Maipo, confluencia con rı́o Cruz de Piedra, 34
060 S, 70
030 W, 2,800 m,
20 Jan 2000, Teillier 4539 (CONC).
CALYCERA HERBACEA Cav. var. SINUATA (Miers) Pontiroli,
Revista Mus. La Plata, Secc. Bot. 9(41): 191, 1963, emend.
Calycera sinuata Miers, Ann. Mag. Nat. Hist. ser. 3, 6(36):
398. 1860 [Dec 1860]. Calycera viridiflora (Phil.) Miers f.
sinuata (Miers) Hicken, Reun. Nac. Soc. Arg. Cienc.
Nat. 1: 247. 1919.—TYPE: ARGENTINA. Mendoza:
“Puente del Inca, altit. 8000 pedes”, 1825, J. Miers s. n.
(holotype: BM 000947748 not seen, photo of BM at SI!).
Calycera intermedia Phil., Anales Univ. Chile 36: 173. 1870,
syn. nov.—TYPE: ARGENTINA. Mendoza, locality date
not indicated, R. A. Philippi s. n. (holotype: K 000588763
not seen, photos of K at SGO! and at SI!).
Calycera involucrata Phil., Anales Univ. Chile 36: 174.
1870.—TYPE: ARGENTINA. Mendoza: Dpto. Tunuyán,
El Guindo (valle del arroyo Manzano), 10 Mar 1945,
G. Covas 6980 (neotype designated here: SI!).
Calycera crenata R. E. Fr., Nova Acta Regiae Soc. Sci. Upsal.
ser. 4, 1(1): 98. 1905, syn. nov. Calycera pulvinata
J. Rémy f. crenata (R. E. Fr.) Hicken, Reun. Nac. Soc.
Arg. Cienc. Nat. 1: 253. 1919.—TYPE: ARGENTINA.
Jujuy: La Rinconada, 3,800 m, 6–8 Jan 1901, F. Kurtz
11365 (lectotype designated here: S 08–715 not seen,
photo of S at SI!; isolectotypes: BAF not seen, photo
of BAF at SI!, SI 353!).
Calycera pulvinata J. Rémy f. cauligera Hicken, Reun. Nac.
Soc. Arg. Cienc. Nat. 1: 253. 1919, syn. nov.—TYPE:
ARGENTINA. Tucumán: Lara, 3,200 m, 30 Jan 1912,
D. Rodrı́guez 295 (lectotype designated here: SI 358!;
isolectotypes: LIL 89439!, SI 104739!).
Distribution—This variety grows in northern Chile
(Region of Antofagasta) and northwestern Argentina, in
a wide area of the Andes and pre-Andean mountains
(Famatina, Aconquija, Calchaquı́es), from Salta and Jujuy
to northeastern Mendoza, along the Altoandina, Puneña,
and Prepuneña phytogeographic provinces, from 2,500–
4,900 m in elevation.
Notes—Calycera herbacea var. sinuata may be confused
with C. pulvinata because of the hemicryptophytic life
form with rosulate habit, and the cypselas. However,
C. pulvinata has vertical rhizomes, undifferentiated invo-
lucre, corolla half-lobed with campanulate tube and lobes
scimitar-shaped, and lanceolate anthers without caudicles.
Previous to Hicken’s (1919) nomenclatural review, Hauman-
Merk (1918) had suggested C. involucrata as a synonym
of C. sinuata, and discussed the possibility of reducing
C. sinuata, C. intermedia, and C. viridiflora to the synonymy
of C. herbacea. Although the diagnostic characters of
C. herbacea var. sinuata are not reliable (“se distingue de
la variedad herbacea por las ramificaciones que puede
presentar el tallo y por las hojas sinuoso-dentadas”,
Pontiroli 1963: 191), the analysis of reproductive structures,
particularly the involucre and cypsela, reveals new charac-
ters that support this variety as a valid taxon. Therefore,
we establish as new diagnostic characters of C. herbacea
var. sinuata (Art. 47, International Code of Nomenclature)
the white corolla limb (vs. green in var. herbacea), broad,
spreading involucral bracts, triangular to rectangular, with
margin toothed-mucronate to entire (vs. linear, small and
hidden in var. herbacea), anthers usually with caudicles
(without caudicles in var. herbacea), calyx lobes incurvate
in armed cypselas (recurvate in var. herbacea), and visible
furrows between the ridges of the cypsela (vs. connate
ridges concealing the furrows in var. herbacea).
Flower and cypsela morphology of Calycera intermedia
Phil. do not differ from the holotype and illustration of
Calycera herbacea var. sinuata. According to the holotype,
flowers of C. intermedia have white corollas. The remain-
ing diagnostic characters of C. intermedia can be also
found in C. herbacea var. sinuata. Also, both taxa share a
geographic distribution.
Calycera crenata R. E. Fr. has been considered as a syno-
nym of C. pulvinata J. Rémy. Nevertheless, its floral and
fruit characters match those of C. herbacea var. sinuata, and
vegetative characters agree with the morphological vari-
ability of var. sinuata. Thus, we consider C. crenata a syno-
nym of C. herbacea var. sinuata.
Hicken (1919) established Calycera pulvinata f. cauligera
based on habit and cypsela morphology. However, that
form clearly differs from the type form by the deeply lobed
corolla with reduced tube (vs. partially lobed with cam-
panulate tube in C. pulvinata f. pulvinata), flat corolla lobes
(vs. scimitar-shaped in C. pulvinata f. pulvinata), dish-shaped
involucre (vs. undifferentiated involucre in C. pulvinata
f. pulvinata), oblong anthers with caudicles (vs. lanceolate
anthers without caudicles in C. pulvinata f. pulvinata), and
shortened broad stems (vs. vertical rhizome in C. pulvinata
f. pulvinata). Furthermore, flower and fruit morphology, as
well as all diagnostic characters of C. pulvinata f. cauligera,
fits those of C. herbacea var. sinuata. The taxa differ only
by the armed calyx (shorter in cypselas of C. pulvinata
f. cauligera) and the position of stems and scapiform pedun-
cles (decumbent in C. pulvinata f. cauligera); these characters
are variable within the var. sinuata.
Since we did not find the Philippi’s typus in any of the
herbaria where his collection is housed (B, BAF, G, LE, P,
SGO, and W; Stafleu and Cowan 1988), or in CONC, we
propose the specimen G. Covas 6980 as neotype of Calycera
involucrata because it presents all the diagnostic characters
indicated in the protologue. In addition, Covas’s speci-
men was collected close to the original type locality.
There are two syntypes of C. crenata; one of them, Fries
861 (S 05–2021), is composed of four immature plants
which do not show diagnostic floral and fruit characters.
The second one, Kurtz 11365 (S 08–715), is selected as
lectotype since reproductive structures are well preserved
and it better matches the original description.
Among all syntypes of Calycera pulvinata f. cauligera
[Lillo 3432 (LIL 89436, SI 107634), Lillo 4894 (LIL 89437,
SI 107636), Jörgensen 1178 (LIL 89295, SI 107635), Holmberg
972 (SI 107637), Dinelli 512 (SI 108045), Gerling 91
(SI 108046, SI 108047), Rodriguez 295 (LIL 89439, SI 358,
SI 104739)], we choose Rodriguez 295 as lectotype. We choose
 SYSTEMATIC BOTANY
the voucher Rodriguez 295 (SI 358) as lectotype due to its
better preservation, greater number of flowers and cypselas,
and by carrying the collector’s label plus two labels with
Hicken’s handwriting, including notes and drawings.
Representative Specimens Examined—ARGENTINA. Catamarca:
Andalgalá, Cerros del Aconquija, 4,500 m, 20 Feb 1917, Jörgensen 1676
(SI); Belén: al pie de la cuesta de Randolfo, 3,200 m, 23 Feb 1981, Cabrera
et al. 32470 (SI); Santa Marı́a, La Hoyada: Quebrada del Zarzo, Sierra
Aconquija, 4,580 m, 20 Dec 1933, Peirano s. n. (LIL); Tinogasta, La
Crispita a Vallecito, 3,100 m, 6 Feb 1930, Schreiter 6254 (LIL). Jujuy:
Cochinoca, Miraflores, INTA, Jan 1975, Cabezas 48 (SI); Humahuaca,
Tres Cruces, 21 Jan 1976, Cabrera et al. 27452 (SI); Mina Aguilar, Cerro
Aguilar, arriba de la Mina, ca. Toma de Agua, 4,670–4,730 m, 4 Mar
1983, J. H. Hunziker et al. 10583 (SI); Rinconada, Mina Pirquitas, Cerro
Granadas, ladera Sud, Mar 1970, Fabris and Zuloaga 7739 (LP); Susques,
Cerro Tuzgle, 4,700–4,900 m, 10 Feb 1946, Cabrera 9101 (LP); Tilcara,
Maimará, Laguna Colorada, 4,000 m, 22 Jan 1906, Lillo 4894 (SI);
Valle Grande, subida a Cerro Amarillo, 3,000 m, 2 Jan 1978, Kiesling
et al. 1593 (SI); Caspalá, cumbres, 1 Mar 1940, Burkart and Troncoso
11872 (SI). La Rioja: Famatina, Cueva de Pérez, camino a la mina La
Mexicana, 28
590 5000 S, 67
430 5900 W, 3,818 m, 10 Jan 2009, Donadı́o et al.
121 (SI); Sierra de Famatina, La Mesada, 3,500 m, 29 Apr 1951, Sparre
8881 (LIL). Mendoza: Las Heras, Puente del Inca, 11 Jan 1914, Sanzin
345 (SI); Punta de Vacas, 4 Feb 1934, Burkart 9311 (SI); Tunuyán:
El Guindo (valle del arroyo Manzano), 10 Mar 1945, Covas 6980 (SI).
Salta: Cachi, Valle Encantado 5 km del desvı́o de la Cuesta del Obispo,
28 Apr 1977, Abbiatti and Garcı́a 4520 (LIL); Chicoana, Camino a Cachi,
Piedra del Molino, 3,500 m, 27 Mar 1979, Cabrera et al. 30739 (SI);
La Poma, Mina Esperanza, 12 Feb 1960, Hernández 18 (LP); Los Andes,
desvı́o de la ruta 51 hacia Viaducto La Polvorilla, 24
130 1500 S,
66
230 4300 W, 3,950 m, 13 Feb 2007, Zuloaga et al. 9306b (SI); San
Carlos, Camino a Mina Don Otto, 6 km de ruta prov. 33, 23 Feb
1987, Nicora et al. 9082 (SI); Santa Victoria, Ruta Prov. 145, de
Nazareno a Abra de Fundición, 22
270 4800 S, 65
080 0700 W, 4,300 m,
17 Feb 2009, Zuloaga et al. 10882 (SI). San Juan: Calingasta, de
confluencia a Alojo Las Minitas, 2,500–3,000 m, 17 Feb 1988, Kiesling
et al. 6856 (SI); Manantiales, 7 Dec 1994, Kiesling 8569 (SI); Iglesia,
Camino a Valle del Cura. Quebrada de la Vicuñita, 3,200 m, 23 Jan
1981, Kiesling 3283 (SI); Reserva de San Guillermo, vega Los Corrales,
borde de la vega, 23 Feb 1981, Nicora et al. 8324 (SI); Ullún, del refugio
de la Ea. “Don Carmelo” hacia el W, 30
550 0100 S, 69
080 1800 W, 3,455 m,
10 Feb 2000, Kiesling et al. 9430 (SI). Tucumán: Chicligasta, Estancia
Santa Rosa, 4,000 m, 13 Jan 1927, Venturi 4720 (SI); Lules, Cumbre
de Mala Mala, 3,400 m, 6 Apr 1904, Lillo 3432 (SI); Tafı́ del Valle,
Cuesta del Infiernillo, 3 Dec 1960, Burkart 22075 (SI); Trancas, Cumbres
Calchaquı́es, 27 Dec 1913, Castillón s. n. (SI).
CHILE. II Región de Antofagasta: El Loa, Tatio, 22
210 S, 68
020 W,
4,300 m, 9 Feb 1969, Martin 458 (SI); Camino a Portezuelo del Cajón,
Cerro Toco, ladera N, 22
550 S, 67
460 W, 4,700–4,850 m, 3 Apr 1997, Arroyo
et al. 97034 (CONC).
CALYCERA HORRIDA Hicken, Physis (Buenos Aires) 1: 129.
1912.—TYPE: ARGENTINA. Neuquén: Arroyo Huinganco,
Cordillera del Viento, 1,900 m, 21 Mar 1912, F. Pastore 76
(holotype: SI 355!; isotype: SI 28580!).
Hemicryptophytic, woolly, rosulate herbs, about 10–20 cm
tall, with gemmiferous roots. Leaves pinnatisect, base atten-
uate into a rectangular, woolly petiole, rachis woolly, blade
oblong, leaflets torn, lobes obtuse, mucronate. Peduncles
scapiform, long, clearly distinct from the shortened stem.
Involucre bracts clearly differentiated from distal upper-
most leaves, woolly, with a broad, fused wheel-shaped
base and rectangular lobes, the lobes wide rectangular,
margin entire or crenate-mucronate, apex obtuse, mucro-
nate. Receptacle punctiform, bracts spathulate, woolly.
Flowers pentamerous. Calyx polymorphic: 1 – 2 unequal
sepals, up to 12 mm, acicular, circular section, lignified
at anthesis (terminal flower of outer cymose groups);
5 unequal, lanceolate, flat sepals, up to 4 mm, non-lignified
at anthesis (terminal central flower, solitary flowers, and
terminal flower of inner cymose groups); 5 equal, minute,
[Volume 39
orbicular, sepals non-lignified at anthesis (second and third
order flowers of cymose groups). Hypanthium 3 mm,
green, tubulose. Corolla shortly lobed, tube 4 mm, funnel-
shaped, lobes 1 mm, rectangular, recurved, flat, apex
obtuse, straight. Staminal tube 1.8–2 mm. Filaments 0.3 mm,
incurved. Anthers 2 mm, lanceolate, thecae basally sagittate
with caudicles. Nectaries 0.5 mm. Cypselas prismatic, with
3–5 wrinkled (unarmed cypselas) or smooth (armed
cypselas) ridges, crowned by vestigial, roundish, lanceolate
or spiniform recurved sepals according to the flower posi-
tion and sepal polymorphism described above. Figure 10.
Distribution—This species is endemic to northern Neuquén
(Argentina); it occurs in Cordillera del Viento and Parque
Provincial Tromen, between 2,000 and 2,500 m.
Notes—Calycera horrida may be misidentified as C. herbacea
(see notes for the latter species).
In the protologue, Hicken cited a specimen from Franco
Pastore as the only reference material, without details of
the collector’s number. That specimen was mounted in
two sheets. One of them (SI 355) has original labels by
Pastore and Hicken, where “76”, “F. Pastore 76” and “N
98”,
can be read. The other sheet (SI 28580) only exhibits
one printed label with the reference to the original pub-
lication, and the name of the species and the number
“98” handwritten by Hicken. SI 28580 has been consid-
ered a paratype, assuming that “98” is the collector’s
number. However, Hicken (1912) did not cite any other
additional specimen, and “98” actually corresponds to
the record number of C. horrida in the Hicken0 s species
list. Therefore, SI 28580 is a duplicate of SI 355. We con-
sider SI 355 the holoype of C. horrida because this sheet
has the collector’s label, Hicken’s label, and notes pub-
lished in the protologue, together with a floral drawing
made by the author.
Representative Specimens Examined—ARGENTINA. Neuquén: Ñorquı́n,
Cerro Huaile, ladera NO, 37
040 2200 S, 70
070 1400 W, 2,550 m, 8 Jan 2001,
Biganzoli 1186 (SI); Pehuenches, inter Aguas Calientes et Laguna
del Tromen, 2,125 m, 24 Feb 1888, Kurtz 6142 (SI); Pque. Prov. Tromen,
a 45 km del cruce de ruta 40, 2,100 m, 28 Dec 1999, Ezcurra et al.
2592 (BCRU).
CALYCERA LEUCANTHEMA (Poepp. ex Less.) Kuntze, Revis. Gen.
Pl. 3[3]: 127, 1898 [28 Sep 1898]. Boopis leucanthema
Poepp. ex Less., Linnaea 6: 259. 1831. Anomocarpus
leucanthemus (Poepp. ex Less.) Miers, Ann. Mag. Nat.
Hist. ser. 3, 6(35): 355. 1860. [Nov 1860].—TYPE:
CHILE. Cordillera de Antuco: date not indicated,
E. F. Poeppig 774 (holotype: M not seen, photo 20567
Field Museum of Natural History in SI!; isotypes:
P 00852244 not seen, P 00852245 not seen, P 00852246
not seen).
Leucocera annua Turcz., Bull. Soc. Imp. Naturalistes Moscou
21(1): 583. 1848.—TYPE: CHILE. Sandy places on the
Andes, “Prope Colchagua reipubl. Chilensis”, date
not indicated, T. Bridges 1186 (holotype: K 000588758
not seen, photo of K at SI!).
Anomocarpus tenuis Miers, Ann. Mag. Nat. Hist. ser. 3, 6(35):
356. 1860. [Nov 1860].—TYPE: CHILE. Province de
Concepción: date not indicated, C. Gay 1495 (holotype:
P not seen; isotype: LP 907866!).
Anomocarpus tenuifolius Miers, Ann. Mag. Nat. Hist. ser. 3,
6(35): 356. 1860.—TYPE: CHILE. Cordillera de Chillán:
2014] DENHAM ET AL.: MORPHOLOGY AND TAXONOMIC REVISION OF CALYCERA

Fig. 10. Calycera horrida. A. Habit. B–E. Cypsela polymorphism. F. Diagrams of flowers illustrating proportions and shape of hypanthium
and corolla. A. From Chiapella (1999b, INTA, Flora Patagónica). B–E. From Kurtz 6142 (SI).
 SYSTEMATIC BOTANY
1856–1857, P. Germain s. n. (holotype: K 000588759 not
seen, photo of K at SI!).
Boopis integrifolia Phil., Anales Univ. Chile 85: 814. 1894,
syn. nov. Calycera integrifolia (Phil.) Reiche, Anales
Univ. Chile 106: 1044. 1900.—TYPE: CHILE. Maule:
“Thermae Longavi”, Jan 1888, O. Schönemann s. n. (holo-
type: SGO 057269 not seen, photo of SGO at SI!;
isotype: SI 12653!).
Therophytic, woolly, caulescent herbs, about 10–15 cm tall.
Leaves woolly, base attenuate into a petiole, blade lanceolate,
pinnatisect (rarely entire), lobes linear, margin entire, apex
obtuse. Peduncles woolly, indistinct from the uppermost
internodes of the stem. Involucre bracts almost free, clearly
differentiated from distal uppermost leaves, fused base
strongly reduced, bracts lanceolate to linear-lanceolate, apex
mucronate. Receptacle punctiform, bracts spathulate. Flowers
tetramerous. Calyx polymorphic: 3–4 unequal sepals, up to
1.5 mm, acicular, triangular section, non-lignified at anthesis
(terminal central flower and terminal flower of every cymose
group); 4 unequal, very reduced (0.5 mm) up to triangular, flat
sepals, non-lignified at anthesis (solitary flowers, and second
and third order flowers of cymose groups). Hypanthium
0.75 mm, green, tubulose. Corolla white, deeply lobed; tube
reduced, 0.5 mm; lobes 1.25 mm, oblong, flat, slightly recurved,
apex obtuse, non uncinate. Staminal tube 0.3 mm. Filaments
0.1 mm. Anthers 0.7 mm, oblong, thecae basally base sagit-
tate. Nectaries 0.1 mm. Cypselas prismatic, often narrow at
base, with 4 smooth ridges, crowned by reduced (unarmed
cypselas) to triangular-pungent compressed sepals (armed
cypselas) according to the flower position and sepal poly-
morphism described above. Figure 11.
Distribution—This species is endemic to central Chile;
it inhabits the VII Región del Maule and northern VIII
Región del Bı́o-Bı́o, between 300 and 700 m in elevation.
Notes—Calycera leucanthema is the only species of Calycera
with tetramerous flowers. It could be confused with
C. sessiliflora due to the pterophytic life form, herba-
ceous caulescent habit, and cypsela morphology. How-
ever, C. sessiliflora is completely glabrous, with lanceolate
leaves, pentamerous flowers, and slightly accrescent, cam-
panulate involucres.
The name Calycera integrifolia (Phil.) Reiche must be
placed in synonymy with C. leucanthema because it differs
only by the entire leaf blades, being indistinguishable in all
remaining morphological characters. Also, the holotype is
the only specimen of C. integrifolia, which was collected
within the distribution area of C. leucanthema.
The synonymy of Anomocarpus tenuis, A. tenuifolius and
Leucocera annua under Calycera leucanthema was proposed by
Reiche (1900, 1902) and Zanotti and Pozner (2008). Bentham
and Hooker (1873) placed Leucocera annua and Anomocarpus
tenuifolius (both = Calycera leucanthema) under Calycera lanata.
This last name probably corresponds to Calycera leucanthema
but its succinct description eliminates a trustworthy place-
ment to any of the species accepted in this present work.
Richard (1820) mentioned that the holotype was probably
in P, but he could not study it.
Acicarpha lanata Lag. ex Pers., Syn. Pl. (Persoon) 2(2): 488.
1807 [Sep 1807], nom. dub. Calycera lanata (Lag. ex Pers.)
Benth. and Hook. f., Gen. Pl. [Bentham and Hooker f.] 2(1):
162. 1873 [7 Feb 1880]. Cryptocarpha lanata (Lag. ex Pers.)
[Volume 39
Cass., Dict. Sci. Nat. (ed. 2) ed. 2, 12: 86. 1819. TYPE: “N.
Hispania”, date not indicated, L. Née s. n. (holotype, probable
in P, not located). This is a nomen dubium, probably a syno-
nym of C. leucanthema.
Representative Specimens Examined—CHILE. VII Región del Maule:
Curicó, Hacienda Monte Grande, 700 m, Dec 1924, Werdermann 552 (SI);
Linares, East and a little south of Linares along the Rı́o Ancoa, along
the road to Melado and Medina 38.2 km upstream from the intersec-
tion with the road to Peñasco, 35
50–520 S, 71
10–200 W, 750–900 m,
21 Jan 1993, Taylor and Gereau 10981 (CONC, MO); Talca, Cordillera de
Talca, El Picazo, 28 Dec 1936, Barros 2982 (SI); Roadside 10 km south
of Vilches Alto and 1 km south of village of Vilches, 35
340 S, 71
110 W,
700 m, 19 Jan 1991, DeVore 1486 (CONC). VIII Región del Bı́o-Bı́o:
Ñuble, Cordillera de Chillán, Dec 1855, R. A. Philippi s. n. (SI 12656);
Off Route N59 between Yungay and Pemuco, east side of road about
2 km south of Chillán-Yungay bridge, 37
000 4400 S, 72
040 1500 W, 232 m,
7 Dec 2010, Johnson and Zavala 10–105 (BRY, SI).
CALYCERA PULVINATA J. Rémy, Ann. Sci. Nat., Bot. sér. 3, 6:
352, 1846. Anomocarpus pulvinatus (J. Rémy) Miers, Ann.
Mag. Nat. Hist. ser. 3, 6(35): 354. 1860 [Nov 1860].—
TYPE: BOLIVIA. Carangas: Grand plateau des Andes,
date not indicated, A. D’Orbigny s. n. (holotype: P not
seen; isotypes: BM 000947749 not seen, photo of BM at
SI!, MO not seen).
Hemicryptophytic, glabrous, rosulate herbs, about 3–4 cm
tall, with vertical rhizomes and adventitious roots. Leaves
spathulate, base attenuate into a rectangular petiole, blade
oblong, margin crenate-mucronate; apex obtuse. Peduncles
scapiform very short. Central cephalioid surrounded by
several smaller ones crowded at the center of the plant.
Involucre indistinct. Floral bracts wide, lanceolate, apex
entire or 3-toothed, free, although clearly different from
the normal leaves, the outermost ones not arranged into
a distinct involucre. Flowers pentamerous. Calyx dimor-
phic: 5 equal sepals, 1 mm, acicular, triangular section,
partially lignified (the central terminal flower and termi-
nal flower of outer cymose group); 5 equal sepals, reduced,
triangular section, flat (solitary flowers and terminal flower of
inner cymose groups). Hypanthium 1.5 mm, green, tubulose.
Corolla white, half lobed; tube 1 mm, campanulate; lobes
0.75 mm, lanceolate, scimitar-shaped, apex obtuse, unci-
nate. Staminal tube 0.2 mm. Filaments 0.3 mm. Anthers
1 mm, lanceolate, thecae basally obtuse. Nectaries 0.2 mm.
Cypselas clearly dimorphic: armed, smooth, ridged cypselas,
and unarmed, rounded, smooth cypselas, according to the
flower position and sepal dimorphism described above.
Figure 12.
Distribution—Bolivia and northwestern Argentina, where
this species reaches the high Calchaquı́es tops, above 4,000 m
in elevation. DeVore (1994) cited the species also from the
high Andean peaks from southern Peru.
Notes—Calycera pulvinata could be confused with C. herbacea
var. sinuata (see notes at C. herbacea var. sinuata).
Representative Specimens Examined—ARGENTINA. Catamarca:
Andalgalá, Rı́o Potrero superior, 3,900 m, 28 Feb 1951, Sleumer
1908 (LIL); Tinogasta, Reales Blancos, 4,000 m, 2 Feb 1930, Schreiter
6114 (LIL). Jujuy: Dr. Manuel Belgrano, Refugio Militar, 24
020 1300 S,
65
420 5800 W, 4,610 m, 26 Jan 2012, Zanotti and Suescún 258 (SI);
Tumbaya, de Purmamarca al Abra de Lipán, 4,000 m, 11 Mar 1982,
Kiesling et al. 3534 (SI). Salta: San Carlos, Cerros del Cajón, 4,500 m,
28 Feb 1914, Rodrı́guez 1315 (SI). Tucumán: Tafı́ del Valle, Cumbres
Calchaquı́es, Huaca Huasi, 26
400 S, 65
440 W, 4,300 m, 13 Mar 1984,
Gómez-Sosa and Múlgura 173 (SI); valles de “El Pelado”, 4,000 m,
17 Mar 1912, Rodrı́guez 438 (LIL, SI); Cerro Muñoz, 4,000 m, 23 Feb
1905, Lillo 4189 (SI).
BOLIVIA. La Paz: Pacajes, Rosario, 6 Jan 1921, Shepard 233 (LIL).
2014] DENHAM ET AL.: MORPHOLOGY AND TAXONOMIC REVISION OF CALYCERA

Fig. 11. Calycera leucanthema. A. Habit. B–C. Leaves. D. Inflorescence. E–F. Flower with non spiniform sepals, corolla partially removed
in F. G. Diagrams of flowers illustrating proportions and shape of hypanthium and corolla. H–J. Cypsela polymorphism. From Johnson &
Zavala 10–105 (SI).
 SYSTEMATIC BOTANY
Fig. 12. Calycera pulvinata. A. Habit. B. Cypsela, hemisection. C. Cypsela. D. Cephaloid. E. Diagrams of flowers illustrating proportions and shape
of hypanthium and corolla. A–D. Modified from Weddel (1857)[1858].
CALYCERA SESSILIFLORA Phil., Linnaea 28: 706. 1856.—TYPE:
CHILE. Prope Santiago, date not indicated, R. A. Philippi
737 (lectotype, designated here: K 000588762 not seen,
photo of K at SI!; isolectotypes: BAF not seen, photo of
BAF at SI!, CONC 29656!, GOET 000508 not seen,
photo of GOET at SI!, P not seen, S 05–2022 not seen,
photo of S at SI!, SGO 057255 not seen).
Anomocarpus axillaris Miers, Ann. Mag. Nat. Hist. ser. 3,
6(35): 352. 1860. [Nov 1860], syn. nov. Calycera sessiliflora
Phil. var. axillaris (Miers) Reiche, Anales Univ. Chile
106: 1045. 1900.—TYPE: CHILE. Valparaı́so, date not
indicated, H. Cuming 664 (holotype: K 000588751 not
seen, photo of K at SI!).
Therophytic, glabrous, caulescent herbs, about 15 cm tall.
Leaves spathulate, base attenuate into a short petiole, blade
lanceolate, margin entire or toothed-mucronate, apex acute,
mucronate. Peduncle short, slender. Cephalioids erect or
nodding, associated in a secondary cymose inflorescence,
flowers few (ca. 10), cymose groups absent. Involucre
bracts clearly differenciated from distal uppermost leaves
with a campanulate fused, slightly accrescent base and
triangular lobes, the lobes with apex mucronate. Recep-
tacle punctiform, bracts absent. Flowers pentamerous. Calyx
dimorphic: 3–5 unequal sepals, up 2 mm, acicular, with
distally triangular and basally flat section, non-lignified at
anthesis (central terminal flower), 5 equal, short triangular,
flat sepals, 0.5 mm, non-lignified at anthesis (solitary flowers).
Hypanthium 0.5 mm, green, tubulose. Corolla white, half
[Volume 39
lobed; tube 0.5 mm, slightly urceolate; lobes 1.5 mm, lan-
ceolate, flat, slightly recurved, apex obtuse, uncinate.
Staminal tube reduced, 0.1 mm. Filaments 0.3–0.4 mm.
Anthers 0.6 mm, oblong, thecae basally obtuse. Nectaries
0.1 mm. Cypselas prismatic, with 5 wrinkled ridges and
vestigial, short triangular sepals (unarmed cypselas), or
3–5 smooth ridges and spiniform recurved sepals (armed
cypselas), according to the flower position and sepal dimor-
phism described above. Figure 13.
Distribution—This species is endemic to central Chile;
it occurs from the IV Región de Coquimbo to the Región
Metropolitana, between 700 and 1,900 m in elevation.
Notes—Calycera sessiliflora may be misidentified with
C. leucanthema (see notes for the latter species).
Calycera sessiliflora var. axillaris does not differ from Calycera
sessiliflora var. sessiliflora. In both original descriptions by Miers
(1860–1869) and the later descriptions by Reiche (1900, 1902)
no consistent diagnostic characters could be found; neither
could differences be found between the holotypes.
We consider it convenient to designate as lectotype the
Philippi’s (K) syntype because of its larger amount of mate-
rial, numerous flowers and cypselas, and better preservation.
The other syntype, collected by P. Germain (Germain s. n., K,
SGO), has few cypselas and is poorly preserved.
Anomocarpus subsessiliflorus Miers, Ann. Mag. Nat. Hist.
ser. 3, 6(35): 352. 1860. [Nov 1860], is a nomen superfluum.
Representative Specimens Examined—CHILE. IV Región de Coquimbo:
Limarı́, Tulahuen, 1,200 m, 10 Oct 1948, Jiles-P. 1011 (LIL); Choapa,
Quebrada de Torca. Auco, 31
300 S, 71
090 W, 700 m, 8 Oct 1993, Arancio
2014] DENHAM ET AL.: MORPHOLOGY AND TAXONOMIC REVISION OF CALYCERA

Fig. 13. Calycera sessiliflora. A–B. Habit. C. Cephaloids. D–E. Cypsela dimorphism. F. Diagrams of flowers illustrating proportions and shape
of hypanthium and corolla. From Jiles-P 1011 (LIL).
 SYSTEMATIC BOTANY [Volume 39

92852 (CONC). V Región de Valparaı́so: San Felipe de Aconcagua,
Los Andes, Inter Mendoza et Santa Rosa, 1868–69, R. A. Philippi s. n.
(SI 12672); Marga Marga, cuesta La Dormida vicinity, off Ruta F10G,
cerro Las Vizcachas, 33
040 2300 S, 71
010 4900 W, 1,365–1,550 m, 11 Dec
2010, Johnson and Zavala 10–153 (BRY, SI). Región Metropolitana:
Chacabuco, Cordillera de Tiltil, Nov 1865, collector not indicated
(SI 20196); Santiago, Santuario de la Naturaleza Yerba Loca, cordón
entre estero Manzanito y estero Yerba Loca, 33
200 S, 70
210 W, 1,530 m,
20 Oct 1999, Arroyo and Humaña 993603 (CONC); Comuna de La Reina.
Quebrada de Ramón, 33
260 S, 70
300 W, 1,600 m, 10 Sep 2000, Tomé-R.
64 (CONC).
CALYCERA SYMPAGANTHERA (Ruiz and Pav.) Kuntze, Revis. Gen.
Pl. 3[3]: 127. 1898 [28 Sep 1898]. Scabiosa sympaganthera
Ruiz and Pav., Fl. Peruv. [Ruiz and Pavón] 1: 49. 1798.—
TYPE: CHILE. “Chile Flora. Decembri in arenosis.
Syngenesia”, 1786, H. Ruiz and J. Pavón s. n. (lectotype,

Fig. 14. Calycera sympaganthera. A. Habit. B. Cephaloid. C–D. Cypsela dimorphism. E. Diagrams of flowers illustrating proportions and shape of
hypanthium and corolla. From R. A. Philippi s. n. (SI 12665).
2014] DENHAM ET AL.: MORPHOLOGY AND TAXONOMIC REVISION OF CALYCERA
designated here: MA not seen, photo of MA at SI!;
isolectotype: MA not seen, photo 29275 by Field Museum
of Natural History in SI!).
Chamaephytic, villous herbs, about 15–20 cm tall, with
erect and prostrate stems. Leaves spathulate, attenuate into
a rectangular pubescent petiole, blade lanceolate, margin
villous, slashed, lobes triangular, mucronate, apex obtuse.
Peduncles indistinct from the uppermost internodes of the
stem. Involucre bracts almost free, lanceolate, basally broad,
margin entire and apex mucronate, clearly differenciated
from distal uppermost leaves with a reduced, dish-shaped
fused base. Receptacle punctiform, bracts linear, spathulate
and rectangular. Flowers pentamerous. Calyx dimorphic:
4–5 unequal sepals, up to 6 mm, long triangular, naviculate,
non-lignified at anthesis (terminal central flower, terminal
flower of outer cymose groups); 5 equal sepals, 1–3 mm,
short triangular (solitary flowers, terminal flower of inner
cymose groups, and second order flowers of cymose
groups). Hypanthium 2 – 2.5 mm, green, tubulose. Corolla
white, half lobed; tube 1.5 mm, broad; lobes 2.5 mm,
oblong, slightly recurved, flat, apex obtuse, uncinate.
Staminal tube 0.3–0.4 mm. Filaments 0.3–0.4 mm. Anthers
1.2 mm, oblong, thecae basally sagittate, with caudicles.
Nectaries 0.1 mm. Cypselas prismatic, with 5 wrinkled
ridges, and 5 vestigial, short, triangular sepals (unarmed
cypselas), or with 3–5 smooth ridges and 2–5 spiniform
recurved sepals (armed cypselas), according to the flower
position and sepal dimorphism described above, usually
without intermediate forms. Figure 14.
Distribution—Calycerta sympaganthera is endemic to Chile;
it grows in the Cordillera de Nahuelbuta.
Notes—Despite having a drawing with references by Ruiz
and Pavon, the specimen photographed by the Field Museum
shows only two immature inflorescences. The specimen here
designated as lectotype has the original label handwritten
by Ruiz and Pavon, with fruits, two immature inflores-
cences, and two mature inflorescences.
Calycera balsamitaefolia (Juss.) Rich., Mem. Mus. Hist. Nat. 6:
38. 1820. Boopis balsamitaefolia Juss., Ann. Mus. Par. 2: 350. 1803,
and Calycera sympaganthera (Ruiz and Pav.) Martic. and
Quezada, Gayana Bot., 44(1–4): 40. 1987 (publ. 1988), are
nomina superflua that apply to this species.
The name Calycera coronata is not effectively published
under article 30.5 (McNeill et al. 2012), although DeVore
(1994) included a description and illustration of the taxon
in her Thesis.
Representative Specimens Examined—CHILE. IX Región de la
Araucanı́a: Malleco, Vega de Chanleo, Cordillera de Nahuelbuta,
Jan 1877, R. A. Philippi s. n. (SI 12665); Angol, Parque Nacional
Nahuelbuta, ruta Cerro Anai, 37
470 2800 S, 73
000 0400 W, 1,315 m, 20 Jan
2008, Zapata and Sede 376 (SI); Parque Nacional Nahuelbuta, Sector
Piedra del Águila, 37
470 S, 72
590 W, 1,300 m, 9 Feb 1991, De Vore and
Baeza 1611 (CONC); Parque Nacional Nahuelbuta, near to main
camping area, 37
480 2600 S, 73
000 6000 W, 1,232 m, 8 Feb 2003, Gardner
et al. DCI n
589 (CONC).
Excluded Species—
BOOPIS GRACILIS Phil., Linnaea 28: 707. 1856.—TYPE: CHILE.
“In collibus ejusdem departamenti Linares ocurrit”, 1856,
P. Germain s. n. (lectotype designated here: W 1889–
0008587 not seen, photo of W at SI!; isolectotypes:
CONC 29655!, K 000588750 not seen, photo of K at
SI!, SI 12639!, W 0021370 not seen, photo of W at SI!).
Calycera boopidea Hicken, Reunión Nac. Soc. Argent. Ci. Nat.
1: 251. 1919, syn. nov.—TYPE: ARGENTINA. Neuquén:
Confluencia Limay, 20–31 Oct 1897, collector not indi-
cated (holotype: SI 350!; isotype: LP!)
Notes—Calycera boopidea Hicken is characterized by a
caulescent therophyte life form; oblong sessile leaves, blade
pinnatifid to dentate; involucre dish-shaped; receptacle
punctiform; flowers pentamerous; sepals roundish emar-
ginate, hypanthium filiform, 1/14 the corolla length; corolla
funnel-shaped, tube enlarged parcially lobed, lobes flat,
recurved, apex straight obtuse; staminal tube enlarged as
long as the corolline tube; anthers oblong, base obtuse;
cypselas wrinkled. All of these characters match with those
of Boopis gracilis.
The voucher W1889 – 0008587 was chosen as lectotype
of Boopis gracilis since it is the one with the greatest
amount of vegetative and reproductive material in a good
state of preservation.
Acknowledgments. We thank Leigh Johnson for many valuable
comments, the curators of CONC, LIL, LP, and SI for specimen loans,
and Francisco Rojas for the illustration. This research was supported
by the “Cristóbal M. Hicken” fellowship (Academia Nacional de Ciencias
Exactas, Fı́sicas y Naturales, Argentina), and the project, “Speciation
in Patagonia: establishing sustainable international collaborations
in evolution, ecology and conservation biology,” NSF OISE-0530267.
We thank Consejo Nacional de Investigaciones Cientı́ficas y Técnicas
from Argentina.
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