A New Species of Parodia (Cactaceae, Notocacteae) from

Rio Grande do Sul, Brazil

Marlon C. Machado and Reto Nyffeler
Institut für Systematische Botanik, Universität Zürich, Zollikerstrasse 107, CH-8008 Zürich,
Switzerland. marlon.machado@systbot.uzh.ch

Urs Eggli
Sukkulenten-Sammlung Zürich, Mythenquai 88, CH-8002 Zürich, Switzerland.
urs.eggli@zuerich.ch

João F. Larocca e Silva

Universidade do Vale do Rio dos Sinos (Unisinos), Av. Unisinos 950, Bairro Cristo Rei,
93022-000, São Leopoldo, Rio Grande do Sul, Brazil. jlarocca@unisinos.br

ABSTRACT . A new species, Parodia gaucha M.
Machado & Larocca (Cactaceae, Notocacteae), from
Encruzilhada do Sul, Rio Grande do Sul, Brazil, is
described and illustrated. The new species is
morphologically very similar to P. muricata (Otto ex
Pfeiffer) Hofacker but can be distinguished from the
latter by the presence of a conical projection below
each areole, the higher number of shallower and
narrower ribs, more closely spaced areoles, higher
number of spines, different color of the spines, and
seeds with minutely striate testa-cell surfaces.
RESUMO . Descreve-se e ilustra-se uma nova espécie,
Parodia gaucha M. Machado & Larocca (Cactaceae,
Notocacteae), proveniente de Encruzilhada do Sul,
Rio Grande do Sul, Brasil. A nova espécie é
morfologicamente muito similar a P. muricata (Otto
ex Pfeiffer) Hofacker, mas pode ser distinguida desta
última pela presença de uma projeção cônica abaixo
de cada aréola, por apresentar um número mais
elevado de costelas que são menos profundas e de
menor largura, menor espaçamento entre aréolas,
maior número de espinhos, por possuir espinhos de
diferente coloração e sementes cujas células da testa
possuem uma superfı́cie finamente estriada.
Key words: Brazil, Cactaceae, campos, IUCN Red
List, Notocacteae, pampas, Parodia, Rio Grande do
Sul.
Parodia Spegazzini, as circumscribed by the
Cactaceae Working Party of the International Orga-
nization for Succulent Plant Study (Hunt & Taylor,
1986, 1990), comprises about 60 to 70 species of
mostly globular or occasionally subcolumnar cacti
(Barthlott & Hunt, 1993; Hunt, 1999, 2006; Ander-
NOVON 18: 214–219. PUBLISHED ON 22 MAY 2008.
son, 2001) distributed through the eastern part of
central South America. The species occur in two
centers of diversity (Buxbaum, 1964, 1966–1967,
1967a, b, 1975; Endler & Buxbaum, 1974): (1) the
lowland pampas regions of southern Brazil (particu-
larly in the state of Rio Grande do Sul), Uruguay,
eastern Paraguay, and northeastern Argentina; and (2)
on the eastern slopes of the Andes in northwestern
Argentina and eastern Bolivia. Many species have
limited distribution ranges and are often threatened by
grazing, agriculture, and habitat modification.
The stems of Parodia generally have tuberculate
ribs, but a few taxa, e.g., P. haselbergii (Rümpler) F.
H. Brandt and P. microsperma (F. A. C. Weber)
Spegazzini, differ in having distinct tubercles ar-
ranged in helical lines. The flowers of most Parodia
species are large and funnel- or occasionally bell-
shaped, brightly colored, and pollinated by solitary
bees (Schlindwein, 1995). The flowers arise near the
stem apex, and the outside of the floral tube and the
pericarpel is covered with bristles and hairs. The
genus is characterized by having seeds with a straight
or occasionally very slightly curved embryo and a
large hilum-micropylar region (Barthlott & Voit, 1979;
Barthlott & Hunt, 2000). Many of the seeds possess a
prominent, either pad-, dome-, or balloon-shaped
strophiole associated with the hilum-micropylar
region.
A number of well-circumscribed subgroups can be
recognized in Parodia, some of which were recognized
as distinct genera in the past. One such subgroup is
the group of species related to P. ottonis (Lehmann) N.
P. Taylor. This group consists of species placed in the
formerly separate genus Notocactus (K. Schumann)
Frič (now subsumed under Parodia) and comprises
doi: 10.3417/2007088
Volume 18, Number 2
2008
the species included in Notocactus ser. Fricianae
Havlı́ček (nom. inval., ICBN Art. 36.1 [McNeill et al.,
2006]) and Notocactus ser. Paucispini (Frič &
Kreuzinger) Havlı́ček: P. carambeiensis (Buining &
Brederoo) Hofacker, P. linkii (Lehmann) R. Kiesling,
P. muricata (Otto ex Pfeiffer) Hofacker, P. ottonis, P.
oxycostata (Buining & Brederoo) Hofacker, P. stock-
ingeri (Prestlé) Hofacker & P. J. Braun, and P.
tenuicylindrica (F. Ritter) D. R. Hunt. This group of
species currently lacks a formal infraspecific taxon
name under Parodia, and thus is referred here with
the informal name P. ottonis group. Plants of the P.
ottonis group share the morphological synapomorphy
of producing offsets on short stolons, which often
initially develop underground. Recent molecular
systematic studies (Nyffeler, 1999; Machado et al.,
in prep.) have confirmed the monophyly of the P.
ottonis group.
In 2005 and 2006 the first author traveled
extensively in the state of Rio Grande do Sul, Brazil,
sampling populations of the Parodia ottonis group in
order to infer species limits in this group and
investigate infra- and interspecific relationships by
studying morphological and molecular variation
patterns for species delimitation analyses. During
the course of one of the field excursions, a population
of a distinctive taxon belonging to the P. ottonis group
was found that represents a morphologically distinct
entity; it is described here as a new species.
Parodia gaucha M. Machado & Larocca, sp. nov.
TYPE: Brazil. Rio Grande do Sul: Mun.
Encruzilhada do Sul, 90 m, 27 Oct. 2006, M.
Machado 865 (holotype, ZSS; isotypes, HASU,
HUEFS, MO). Figures 1, 2.
Haec species Parodiae muricatae (Otto ex Pfeiffer)
Hofacker affinis, sed projecturis infra quamque areolam
conicis, costis numerosioribus angustioribus minus profun-
dis, areolis densius dispositis, spinis numerosioribus color-
ibus diversis, cellulis testae seminis tenuiter striatis differt.
Plant body clavate to short-cylindric, 9–12 3 5–
8 cm, producing offsets from areoles located at the
lower portion of the body; offsets first subterranean,
produced on short stolons ca. 0.1 cm diam. and to
2 cm long; epidermis bright green; cortical tissues not
mucilaginous, yellowish green in the upper portion of
the body but becoming orange toward the base; ribs 18
to 22, straight, well-defined but slightly tuberculate,
with shallow sinuses between successive tubercles,
0.2–0.4 cm tall 3 0.6–1.0 cm thick, edges serrate due
to a conical projection (triangular in lateral view)
below each areole; areoles located at the sinuses
between consecutive tubercles, rounded, 0.2–0.25 cm
diam., densely set, 0.3–0.4 cm apart, with pale
Machado et al. 215
Parodia gaucha (Cactaceae) from Brazil
yellowish wool when young, later glabrescent; spines
spreading, acicular, straight, almost bristly, 0.5–0.8 3
0.01–0.03 cm; central spines 6, golden yellow to
reddish orange, hardly distinguished from the radials;
radial spines 18 to 26, pale yellow; flowering areoles
producing longer and curved spines to 1.5 cm. Flower
buds with a dense cover of pale brownish hairs;
flowers subapical, diurnal, opening for one to several
days about 5 hours after sunrise and closing 4 hours
later, shortly funnelform, to 4 3 5 cm; floral tube to
ca. 1 cm, internal walls colored purplish red on the
lower 1/3, yellow above; external walls yellowish
green, with triangular bract scales subtending areoles
that bear pale brownish to cream-yellow hairs and
long yellowish bristles to 2 cm; perianth multiseriate,
perianth segments pale yellow, lanceolate; stamens
numerous (hundreds), spreading, arranged along the
inner walls of the floral tube, sensitive and closing
around the style when touched, lowermost stamens
with filaments purplish red on the lower 1/3 to lower
1/2, yellow above; filaments to 0.7 cm; style yellow,
ca. 1.6 3 0.2 cm; stigma lobes exserted, 10 to 12,
dark red. Fruit 1.0–1.4 3 0.8–1.0 cm, greenish
yellow, pericarpel with areoles bearing long yellowish
brown hairs and a few curly bristles, dehiscing by one
or more longitudinal slits, pulp white; dry perianth
remnants either persistent on the fruit or falling off;
seeds 1.0–1.2 3 0.8–0.9 mm, helmet-shaped, dark
reddish brown to black, shiny; hilum-micropylar
region 0.8–0.9 mm, pear-shaped, lateral to the seed
axis, rim of hilum-micropylar region sharply edged
against the testa, without bulging; testa cells convex,
mostly pentagonal or hexagonal, ca. 50–100 3 30–
80 mm, elongated along the seed axis; cell boundaries
straight, sunken; cell surface with a minutely striate
cuticular folding pattern.
Distribution and habitat. Parodia gaucha is at
present known solely from its type locality in the
municipality of Encruzilhada do Sul, Brazil, where it
grows among rock outcrops in open grassland areas on
hillsides and hilltops. The area consists of more or
less rounded hills rising to ca. 300 m in altitude. The
vegetation in the region is a composite of shrubby
savanna-like vegetation on the lower parts of the hills,
gallery forests in the valleys, and the grassland
vegetation known as campos or pampas on the upper
parts of the hills. In habitat P. gaucha grows in
sympatry with other species of Cactaceae: P. ottonis,
Frailea pygmaea (Spegazzini) Britton & Rose, Cereus
hildmannianus K. Schumann, Opuntia Miller sp.,
Rhipsalis Gartner spp., and Lepismium Pfeiffer spp.
IUCN Red List category. The population of
Parodia gaucha is small, consisting of a few dozen
individuals, the majority of which were located on a
216 Novon

Figure 1. Parodia gaucha M. Machado & Larocca. —A. Habit; notice the underground offsets produced on short stolons (the
dotted line indicates the soil line). —B. View of a rib with spines removed from the areoles, showing the conical projection
below each areole. —C. Longitudinal section of a flower. —D. Typical areole with six central spines and 26 radial spines; the
central spines hardly differentiable from the radials. —E. Detail of the fruits. Drawn from the holotype M. Machado 865 (ZSS).

single rocky wall at the margin of a country road. An
extensive search of the surrounding area revealed that
suitable rock outcrops among the grassland held very
few plants, the majority of which displayed signs of being
trampled and grazed by cattle and sheep. Due to the
apparent rarity, restricted distribution, and observed
threats, it seems prudent to include P. gaucha in the
Critically Endangered category of the IUCN Red List of
endangered plant species according to the following
criteria (IUCN, 2001): CR A2a B1ab(iii,v) C2a(ii) D.
Due to the rarity of Parodia gaucha at its single
known locality, no paratypes have been collected.
Etymology. The specific epithet is derived from
the vernacular term ‘‘gaúcho’’ (feminine form: ‘‘gaú-
cha’’), which in Brazil designates the natives from the
Volume 18, Number 2
2008
Figure 2. Seeds of Parodia gaucha M. Machado & Larocca. —A. Lateral view. —B. Hilum-micropylar region. —C. Testa-
cell surfaces, displaying minutely striated cuticular folding patterns. —D. Close-up of the striations of the testa-cell surface.
From the holotype M. Machado 865 (ZSS).
state of Rio Grande do Sul; this epithet is in accordance
with ICBN Art. 23.2 (McNeill et al., 2006).
Systematic relationships. Parodia gaucha belongs
to the P. ottonis group by virtue of producing offsets
on short stolons and having fruits with a thick and
fleshy pericarpel that dehisces by longitudinal slits. It
is morphologically similar to the other many-ribbed
species of this group, P. muricata, P. stockingeri, and
P. tenuicylindrica. The species that most closely
resembles P. gaucha is P. muricata, but the latter can
be distinguished from the former by the following suite
of characters: absence of a conical projection below
each areole; lower number of ribs (only to 18), which
are wider and higher than the ribs of P. gaucha;
areoles that are less closely spaced; smaller number of
spines (only to 18 including central and radial spines);
spines of different color; and seeds with smooth testa-
cell surfaces (Table 1). Parodia stockingeri shares
with P. gaucha the presence of a conical projection
below each areole, but it is a smaller plant with a
Machado et al. 217
Parodia gaucha (Cactaceae) from Brazil
lower number of ribs and spines, and it has seeds
with a smooth testa-cell surface (Table 1). Parodia
tenuicylindrica differs from all other species of the P.
ottonis group by producing hooked central spines; it
differs from P. gaucha due to its smaller size, absence
of a conical projection below each areole, the lower
number of ribs and spines, and different color of the
spines (Table 1).
Parodia gaucha is geographically well separated
from the other similar species of the P. ottonis group.
Parodia muricata occurs in southwestern Rio Grande
do Sul, Brazil, where it is known from a number of
localities around the municipality of Jaguari. The
nearest populations of P. muricata are separated from
P. gaucha by over 250 km. Parodia stockingeri also
occurs in southwestern Rio Grande do Sul, Brazil,
where it is known from a handful of localities
southeast of the village of Unistalda, municipality of
Santiago. It is separated from P. gaucha by over
320 km. Parodia tenuicylindrica occurs in southwest-
ern Rio Grande do Sul, Brazil, and northwestern
218 Novon

Table 1. Comparison of morphological characters of species of the Parodia ottonis group.

P. gaucha P. muricata P. stockingeri P. tenuicylindrica

Body diameter (cm) to 8 to 10 to 4 to 4
Shape of body clavate to globose to clavate to clavate to cylindric
short-cylindric short-cylindric cylindric
No. of ribs 18 to 22 16 to 18 12 to 15 13 to 21
Height of rib (mm) 2–4 6–8 3–5 2–4
Width of rib (mm) 6–10 10–14 4–8 4–6
Distance between areoles (mm) 3–4 6–10 3–4 1.5–3
No. of central spines 6 3 or 4 4 2 to 4
Color of central spines yellow to orange-red reddish brown yellow to reddish brown to black
orange-red
No. of radial spines 18 to 26 10 to 14 10 to 16 10 to 16
Color of radial spines yellow glassy white cream-yellow to cream-yellow to yellow
yellow
Testa-cell surface of seeds minutely striated smooth smooth minutely striated

Uruguay. The distribution of this species in Brazil is Literature Cited

restricted to the area around and between the
municipalities of Santana do Livramento and Quaraı́
in Rio Grande do Sul, with the northern limit south of
the municipality of Alegrete; in Uruguay, the species
is distributed in the area around and between the
municipalities of Rivera and Artigas, with the
southern limit north of the municipality of Tacuar-
embó. The nearest populations of P. tenuicylindrica
are separated from P. gaucha by over 260 km.
Superficially, Parodia gaucha is reminiscent of
yellow-spined forms of P. scopa (Sprengel) N. P. Taylor,
but the latter is clearly distinguishable from P. gaucha
because it has a higher number of ribs (24 to 40) and
radial spines (36 to 40 or more), flowers with stamens
lacking a reddish color at the base of the filaments,
smaller fruits with a thin pericarpel that becomes
papery and disintegrates after drying, and seeds with
testa-cell surfaces displaying larger striate cuticular
folding patterns (Glaetzle & Prestlé, 1986). Moreover,
offsets of P. scopa are not produced on short stolons nor
developed underground, characteristics that are syna-
pomorphic for the P. ottonis group. Furthermore, P.
scopa belongs to a clade distinct from that of the P.
ottonis group (Nyffeler, 1999; Machado et al., in prep.).
Acknowledgments. The authors would like to
thank Cassio van den Berg, Universidade Estadual
de Feira de Santana, for revising and correcting the
Latin diagnosis of Parodia gaucha; Luciano Paga-
nucci de Queiroz, Universidade Estadual de Feira de
Santana, for his assistance during fieldwork of MCM
in Rio Grande do Sul; Carla de Lima for skillfully
preparing the botanical illustration of P. gaucha; and
Julien Bachelier, University of Zürich, for support in
preparing SEM pictures of the P. gaucha seeds. This
study was supported by grant 3100A0-103583 from
the Swiss National Science Foundation to RN and UE.
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