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Molecular phylogeny of the large South American genus Eriosyce (Notocacteae, Cactaceae): Generic delimitation and proposed changes in infrageneric and species ranks

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Molecular phylogeny of the large South American genus Eriosyce
(Notocacteae, Cactaceae): Generic delimitation and proposed
changes in infrageneric and species ranks
Pablo C. Guerrero,
1,2
Helmut E. Walter,
3
Mary T.K. Arroyo,
2
Carol M. Peña,
1,4
Italo Tamburrino,
2
Marta De Benedictis & Isabel Larridon
1 5,6
1Departamento de Botánica, Facultad de Ciencias Naturales & Oceanográficas, Universidad de Concepción, Casilla 160C, 4030000
Concepción, Chile
2Instituto de Ecología y Biodiversidad, Universidad de Chile, Casilla 653, 7750000 Santiago, Chile
3The EXSIS Project: Cactaceae Ex-Situ & In-Situ Conservation, 31860 Emmerthal, Germany
4Departamento de Ciencia y Tecnología Vegetal, Universidad de Concepción, Campus Los Ángeles, Juan Antonio Coloma 0201,
4440000 Los Ángeles, Chile
5Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AE, U.K.
6Department of Biology, Research Group Spermatophytes, Ghent University, K.L. Ledeganckstraat 35, 9000 Gent, Belgium
Address for correspondence: Pablo C. Guerrero, pabloguerrero@udec.cl
DOI https://doi.org/10.1002/tax.12066
Abstract Eriosyce is one of most species-rich genera within Notocacteae (Cactaceae) harboring a variety of stem and flower morphol-
ogies, and fruits with basal abscission. The lack of a well-sampled molecular phylogeny contributes to the current taxonomic instability
of the genus, where its circumscription and infrageneric classification has been questioned. Specimens of Eriosyce (63 taxa) plus 19
outgroups were analyzed through sequencing three plastid noncoding introns (rpl32-trnL,trnL-trnF,trnH-psbA), one plastid gene
(ycf1), and one nuclear gene (PHYC). Individual markers and concatenated matrices were analyzed using maximum likelihood and
Bayesian approaches. Phylogenetic analyses strongly support the monophyly of Eriosyce s.l. Furthermore, seven clades within Eriosyce
s.l. were defined based on supported branches, although one of them was weakly supported. Our results suggest that some past taxo-
nomic proposals have low phylogenetic support and should no longer be used, e.g., based on their scattered positions in the phylogenetic
reconstruction, several infraspecific taxa appear unrelated to the typical form of the species in which they had been placed. We present a
phylogeny-informed infrageneric classification of the genus Eriosyce, and new combinations are proposed to update the nomenclature
of species and sections.
Keywords Andes; Atacama; Cactoideae; Caryophyllales; central Chile; succulent plants; taxonomy
Supporting Information may be found online in the Supporting Information section at the end of the article.
INTRODUCTION
Southern South America is one of the worlds main centers
of cactus diversity (Hernández-Hernández & al., 2011). It is
unique in harboring representatives of all four presently ac-
cepted subfamilies of the Cactaceae and is possibly the place
of origin of the Pereskioideae (Edwards & al., 2005),
Opuntioideae (Wallace & Gibson, 2002), the early-branching
Cactoideae lineage Blossfeldia Werderm. (Edwards & al.,
2005) and the monogeneric subfamily Maihuenioideae (Fearn,
1996). This center of cactus diversity embraces several major
arid biomes in South America as represented by the Brazilian
and eastern Paraguayan Cerrados, the Brazilian Atlantic
Forests, the Tropical Andes of Bolivia, southeast Peru and
northwestern Argentina as well as the Chilean Winter
Rainfall-Valdivian Forest (Myers & al., 2000; Arroyo & al.,
2005). Furthermore, Cactoideae contain several species
threatened with extinction in the near future due to a massive re-
duction in their distributions or population abundance
(Goettsch & al., 2015), driven by elevated rates of land conver-
sion, extraction as a biological resource, and residential and
commercial development.
The Andean region of Chile, Argentina and southern
Bolivia has been postulated to be the most probable area of ori-
gin of the Cactaceae, as well as of the species-rich subfamilies
Opuntioideae and Cactoideae (Hernández-Hernández & al.,
2014). Moreover, the tribe Notocacteae within Cactoideae is
one of the oldest cactus lineages endemic to the southern South
American biomes, estimated to have diverged 16.0 to 14.8 mil-
lion years ago (Arakaki & al., 2011), or ~12 million years ago
(Hernández-Hernández & al., 2014). Based on floral characters,
Buxbaum (1958) erected the tribe to accommodate a number of
columnar and globular genera endemic to these biomes. Until
recently, the Notocacteae have been one of the most
Article history: Received: 4 Aug 2017 | returned for (first) revision: 12 Dec 2017 | (last) revision received: 25 Jan 2018 | accepted: 5 Feb 2019
Associate Editor: Cassio van den Berg © 2019 International Association for Plant Taxonomy
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heterogeneous groups of South American cacti, often having
been considered as something of a dustbin for isolated genera
whose relationships were unclear (Bárcenas & al., 2011).
Barthlott & Hunt (1993) and Anderson (2001) reduced the num-
ber of genera in Buxbaums tribe, but retained columnar genera
such as Austrocactus Britton & Rose, Eulychnia Phil. and
Corryocactus Britton & Rose, as well as the globular genera
Blossfeldia,Frailea Britton & Rose and Copiapoa Britton &
Rose. With the advent of molecular techniques, Nyffeler
(2002) concluded that this assemblage might be polyphyletic,
a finding that reduced the tribe to the (more or less globular)
strongly supported monophyletic core Notocacteae, compris-
ing Eriosyce Phil. s.l., Parodia Speg. s.l. and Neowerdermannia
Frič. (Hunt & al., 2006, 2013). Hunt & al. (2006) still upheld the
inclusion of Blossfeldia and the orphan genera Frailea and
Copiapoa (respectively Blossfeldia and Frailea, Hunt & al.,
2013), but DNA sequence data-based studies by Bárcenas &
al. (2011), Hernández-Hernández & al. (2011) and Arakaki &
al. (2011) supported Nyffelers (2002) findings. In todays
widely accepted circumscription (Nyffeler & Eggli, 2010), the
Notocacteae comprise the two widespread and species-rich gen-
era Eriosyce s.l. and Parodia s.l., the bitypic genus Neo-
werdermannia, plus the two monotypic and range-restricted
genera Rimacactus Mottram and Yavia R.Kiesling & Piltz.
The members of Notocacteae are characterized by their small
to medium-sized, mainly unbranched, (sub)globular to short co-
lumnar, mostly ribbed stems and their small to medium-sized,
colorful diurnal flowers.
One of the most disputed genera in the Notocacteae is
Eriosyce. In its broad circumscription, it is a widespread,
species-rich and morphologically diverse taxon (Fig. 1).
Eriosyce mainly occurs on the western slopes of the Andes
at altitudes between sea-level and 2800 m between 13° S
and 37° S, but also occurs at high altitudes on the eastern
side of Andes in Argentina from 24° S to 36° S (Fig. 2). Its
main center of diversity lies in northern Chile between
26° S and 30° S, at altitudes between sea-level and 1500 m
(Kattermann, 1994; Guerrero & al., 2011a; Barthlott, & al.,
2015).
The genus Eriosyce has had a chequered taxonomical
history since the naturalist Rodolfo A. Philippi separated
Echinocactus sandillon Gay from Echinocactus Link & Otto,
proposing the new genus Eriosyce to accommodate
E. sandillon (Gay) Phil. in 1872 (Table 1). In their revision
of the family, Britton & Rose (1922) recognized Eriosyce
as a monotypic genus and proposed the new genus Neo-
porteria Britton & Rose for seven endemic Chilean species
with funnel-form and bristly flowers, and referred seven
more species from Argentina, Peru and Chile to
Malacocarpus Salm-Dyck. In 1929, Berger proposed the
new genus Pyrrhocactus A.Berger for species with reddish
flowers growing on the western (Cactus curvispinus Bertero,
Echinocactus tuberisulcatus Jacobi) and on the eastern slopes
of the Andes (e.g., Echinocactus strausianus K.Schum.,
E. catamarcensis Speg.). Yet, some years later, Backeberg
accepted Pyrrhocactus only for the Argentinean species and
referred the Chilean species to his new genera Horridocactus
Backeb. (Backeberg, 1938) and Neochilenia (Dölz, 1942).
However, Ritter (1959) did not accept Backebergs concept
of Pyrrhocactus and included Horridocactus and Neochilenia
within Pyrrhocactus. In 1934, Backeberg proposed another
genus (i.e., Islaya Backeb.) for plants with woolly apices
from northern Chile to southern Peru, and Itô (1957) erected
the genus Thelocephala Y.Itô for a group of small Chilean
species with tuberculate stems.
Backebergs (and Ritters) narrow generic concepts were
widely adopted between the sixties and eighties of the last cen-
tury, and by later authors (e.g., Zuloaga & al., 2007; Kiesling
& al., 2008; Saldivia & Faúndez-Yancas, 2011; Duarte & al.,
2014). Donald & Rowley (1966) proposed to unite Bergers
Pyrrhocactus, BackebergsIslaya,Horridocactus and Neo-
chilenia and ItôsThelocephala under the oldest name Neo-
porteria, but excluded PhilippisEriosyce. This broad
concept was adopted by various authors, e.g., Hunt & Taylor
(1986) and Hoffmann (1989). Later, Kattermann (1994) argued
that Eriosyce must be only part of a larger and closely related
group including taxa from central to northern Chile, northwest-
ern Argentina and southwestern Peru, and consequently pro-
posed to amplify Eriosyce by merging it with Neoporteria s.l.
He further proposed to divide the genus into two sections
and six subsections corresponding to the former segregate
genera and to greatly reduce the number of species in Neo-
porteria s.l. from 66 to 25 and Eriosyce s.str. from 7 to 2
(Table 1). Anderson (2001, 2005), Hoffmann & Walter
(2004) and Hunt & al. (2006, 2013) adopted the broad generic
concept, but somewhat modified the classification at specific
and infrageneric (using subgenusor groupinstead of sec-
tions and subsections) levels. Furthermore, morphological cla-
distics analyses gave support to the broad circumscription of
Eriosyce (Wallace in Kattermann, 1994; Nyffeler & Eggli,
1997), but since vegetative and reproductive morphological
characters in Cactaceae have been documented as convergent,
these results must be contrasted with molecular data
(Schlumpberger & Renner, 2012).
Studies based on DNA sequences did not support the
broad concept of Eriosyce s.l. A phylogenetic analysis of
the family (Nyffeler, 2002) using two plastid markers and in-
cluding only four Eriosyce species suggested the non-
monophyly of the genus. Similarly, Nyffeler & Egglis
(2010) results based on the trnL-trnF marker and including
14 Eriosyce species suggested that there is no support for
an expanded concept of Eriosyce. They also suggested that
E. laui Lüthy is more closely related to Neowerdermannia
and Yavia than to Eriosyce, thus supporting Mottrams
(2001) proposal to place it in the new monotypic genus
Rimacactus. The results of Bárcenas & al. (2011) based on
the trnK-matK marker and including four species, Arakaki
& al. (2011) based on the trnK-matK and PHYC markers
and including four species, and Hernández-Hernández & al.
(2011) based on the trnK-matK,matK,trnL-trnF,rpl16,
and ppc markers and including five species, all suggested
that Eriosyce s.l. is not monophyletic. Although the above
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Guerrero & al. Eriosyce s.l. systematics and taxonomy
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68 (3) June 2019: 557573
Fig. 1.
Diversity and morphology of some of the Eriosyce species included in the molecular analyses. A, Eriosyce aurata;B, E. marksiana var.
lissocarpa;C, E. curvispina;D, E. krausii;E, E. subgibbosa;F, E. senilis subsp. elquiensis;G, E. crispa;H, E. odieri subsp. malleolata
(= E. malleolata); I, E. napina subsp. duripulpa (= E. duripulpa). Photos AH by P.C. Guerrero; I by H. Villalobos.
Guerrero & al. Eriosyce s.l. systematics and taxonomy
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TAXON 68 (3) June 2019: 557573
mentioned molecular studies unanimously failed to support
the broad concept of Eriosyce, the narrow sampling of
Eriosyce and/or the use of few molecular markers resulted
in unresolved intergeneric relationships within the selected
members of tribe Notocacteae (Nyffeler, 2002; Nyffeler &
Eggli, 2010; Bárcenas & al., 2011). Up to this date,
Kattermanns broad concept of Eriosyce is disputed (e.g.,
Zuloaga & al., 2007; Duarte & al., 2014; Hernández-
Ledesma & al., 2015) not only for lumping six segregate
genera into one, but also for substantially reducing the num-
ber of species in these genera to 33, plus 39 infraspecific
taxathe latter being mostly former species lowered in rank.
Unfortunately, none of these proposed infraspecific taxa were
subjected to the cladistic analysis of 58 morphological char-
acters by Wallace in Kattermann (1994) upon which
Kattermanns classification hypothesis was based.
Given the above described complex taxonomic back-
ground and sometimes conflicting phylogenetic results, the
aims of this study are (1) to develop a species-level phyloge-
netic hypothesis based on four plastid and one nuclear marker
and a dense sampling of the ingroup, in order to test the mono-
phyly of Eriosyce s.l.; (2) to investigate whether previous
infrageneric taxonomic treatments are supported by the
molecular data including evaluation of the placement of infra-
specific taxa; and (3) to present a phylogeny-informed
infrageneric classification for the genus Eriosyce.
MATERIALS AND METHODS
Plant material and taxon sampling.
Most of our
plant material came from living plants, i.e., from plants
cultivated from seed in the EXSIS collection and currently
maintained at Universidad de Concepción, and less
frequently from field-collected specimens. This procedure
helped avoid extraction of mature individuals from their
habitats as several taxa are categorized as threatened with a
significant number of these species endangered or critically
endangered (Duarte & al., 2014; Larridon & al., 2014;
Goettsch & al., 2015). For some taxa, sequences published in
GenBank were used (Nyffeler, 2002; Guerrero & al., 2011a;
Franck & al., 2012; Larridon & al., 2015). Voucher
information, locality data, collection numbers and GenBank
accession numbers are given in Appendix 1.
Our sampling was mainly based on Kattermanns classi-
fication and nomenclature (1994, 2001), supplemented by al-
terations in Hoffmann & Walter (2004) and Hunt & al.
(2006, 2013), as well as relevant new taxa described after
2001 that were not included in Hunt & al. (2006, 2013).
We analyzed 63 ingroup taxa (90% of the accepted taxa), in-
cluding all of Kattermanns 33 species (1994, 2001), nearly
all of his subspecies, many of his varieties and two species
published under Thelocephala (Ritter, 1980) that were treated
as synonyms in Kattermann (1994) and Hunt & al. (2006,
2013). The outgroup included 19 species from the core
Notocacteae (sensu Nyffeler & Eggli, 2010), various genera
formerly associated with Notocacteae or still included in
the tribe by Hunt & al. (2006, 2013) (see Table 1), and
two Trichocereeae. Phylogenetic trees were rooted with
Echinopsis chiloensis subsp. litoralis (Johow) M.Lowry
assigned to tribe Trichocereeae. The taxonomic determination
of specimens was realized following relevant literature (e.g.,
Ritter, 1980; Kattermann, 1994; Hoffmann & Walter, 2004;
Hunt & al. 2006, 2013).
DNA extraction, amplification and sequencing.
Fresh samples of cortical tissue of cactus stem were
pulverized to a fine powder using an automatic homogenizer.
Genomic DNA was extracted with the DNeasy Plant Kit
(Qiagen, Valencia, California, U.S.A.). Three noncoding
chloroplast markers (rpl32-trnL,trnH-psbA,trnL-trnF), one
plastid gene (ycf1) and a nuclear one (PHYC) were amplified
using published primers and procedures: rpl32-trnL
(UAG)
(Shaw & al., 2007); trnH-psbA (Sang & al., 1997); trnL-
trnF (Nyffeler, 2002); ycf1 (Franck & al., 2012), and PHYC
(Helsen & al., 2009). These loci were selected based on
their phylogenetic usefulness in solving systematic re-
lationships in Cactaceae (Nyffeler & Eggli, 2010; Arakaki
& al., 2011; Bárcenas & al., 2011; Guerrero & al., 2011a;
Hernández-Hernández & al., 2011; Larridon & al., 2015).
Fig. 2.
Taxonomic richness distribution of Eriosyce s.l. at the species
and infraspecific ranks. Occurrence data were compiled from literature
(Ritter, 1980; Kattermann, 1994; Guerrero & al., 2011a,b; Duarte &
al., 2014), herbaria (CONC, SGO) and our own field observations.
Guerrero & al. Eriosyce s.l. systematics and taxonomy
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TAXON 68 (3) June 2019: 557573
PCR reactions were performed using Eppendorf
Mastercycler gradient (Westbury, New York, U.S.A.) in a
25 μl total volume using 22.5 μl SapphireAmp PCR Master
Mix, 0.5 μl 10 mmol of each primer, 0.5 μlH
2
O and 1 μl
DNA. PCR cycling times were as follows: for rpl32-trnL de-
naturation was performed at 80°C for 5 min followed by 30
cycles of 95°C for 1 min, a primer annealing procedure at
56°C for 1 min characterized by a ramp of 0.3°C/s and then
a primer extension at 69°C for 5 min; the final extension step
lasted 5 min at 65°C; trnL-trnF denaturation occurred at
94°C for 5 min followed by 35 cycles of 94°C for 45 s then
52°C for 1 min and then final elongation steps at 72°C for
1 min 30 s and 72°C for 7 min; for trnH-psbA denaturation
was performed at 80°C for 2 min 30 s followed by 35 cycles
of 94°C for 15 s, primer annealing at 55°C for 15 s, primer
extension at 72°C for 30 s followed by a final primer exten-
sion at 72°C for 3 min; for ycf1, denaturation happened at
96°C for 1 min 30 s followed by 35 cycles of 95°C for
30 s, primer annealing at 52°C for 20 s, primer extension at
72°C for 45 s followed by a final primer extension at 72°C
for 3 min; denaturation of PHYC occured at 94°C for 1 min
45 s followed by 35 cycles of 94°C for 25 s, primer annealing
at 62°C for 25 s and first extension at 72°C for 25 s conclud-
ing with the final step at 72°C for 1 min 45 s. PCR products
were checked on 1% agarose gels and then sent to Macrogen
(Seoul, Korea) for sequencing in both directions.
Phylogenetic analyses.
Sequences were assembled and
edited in the program CLC Main workbench v.7.7.2 (Qiagen).
Sequences for each marker were automatically aligned using
MUSCLE implemented in MEGA v.7.0.18 (Kumar & al.,
2016) and then checked manually. Each marker was at first
analyzed separately and then concatenated. A microsatellite
region in the ycf1 dataset was excluded (450 bp), because the
alignment was ambiguous for that region and many gaps had
to be introduced. The taxa with missing data were different for
each studied region (see details in Table 2). PartitionFinder
v.2.1.1 (Lanfear & al., 2017) with the -raxmlcommand line
option (Stamatakis, 2006) was used to search for the best
partition strategy and best models of molecular evolution for
the molecular dataset. These analyses used the potential
partitions that were defined a priori such as each codon
positions of the ycf1 and PHYC genes and the noncoding
markers. The best models were GTRG for rpl32-trnL and
trnH-psbA and GTRINVGAMMA for the rest of the markers.
Maximum likelihood (ML) analyses of the concatenated
matrix were performed using the program RAxML v.8.1.11
(Stamatakis, 2014) included in raxmlGUI v.1.3.1 (Silvestro &
Michalak, 2012). The search for an optimal ML tree ran com-
bined with a rapid bootstrap analysis of 10,000 replicates.
Bayesian inference (BI) was conducted in the program
MrBayes v.3.2.6 (Ronquist & al., 2012), using unlinked rate
heterogeneity, base frequencies, and substitution rates across
Table 1. Historical overview of the taxonomic classification of Eriosyce.
Philippi
(1872)
Britton & Rose
(1922)
Donald & Rowley
(1966)
Ritter
(1959, 1980)
Kattermann
(1994, 2001)
Hoffmann & Walter
(2004)*
Kiesling & al.
(2008)
Hunt & al.
(2006)
Genus:
Eriosyce
(monotypic)
Genus:
Eriosyce
(1 species)
(1 taxon)
Genus:
Neoporteria
(7 species)
(7 taxa)
Genus:
Eriosyce
Genus:
Neoporteria
(66 species)
Subgenus:
Neoporteria
(9 species)
Subgenus:
Pyrrhocactus
(56 species)
Series:
A (39 species)
B (11 species)
C (1 species)
D (4 species)
Unplaced (1
species)
Subgenus
unplaced:
(1 species)
Genus:
Eriosyce
(7 species)
(9 taxa)
Genus:
Neoporteria
(14 species)
(24 taxa)
Genus:
Islaya
(1 species)
(1 taxon)
Genus:
Telocephala
(16 species)
(17 taxa)
Genus:
Pyrrhocactus
(58 species)
(83 taxa)
Genus:
Eriosyce
(33 species)
Section:
Eriosyce
Subsection:
Eriosyce
(2 species)
(4 taxa)
Pyrrhocactus
(5 species)
(7 taxa)
Islaya
(1 species)
(1 taxon)
Section:
Neoporteria
(25 species)
Subsection:
Chileosyce
(5 species)
(5 taxa)
Horridocactus
(15 species)
(55 taxa)
Neoporteria
(5 species)
(19 taxa)
Genus:
Eriosyce
(83 taxa)
Subgenus:
Eriosyce s.str.
(2 species)
(5 taxa)
Neoporteria
(4 species)
(14 taxa)
Horridocactus
(15 species)
(44 taxa)
Thelocephala
(3 species)
(15 taxa)
Islaya
(3species)
(3 taxa)
(1 nomen nudum)
Genus:
Eriosyce
(5 species)
(7 taxa)
Genus:
Neoporteria
(6 species)
(6 species)
Genus:
Islaya
(4 species)
(4 taxa)
Genus:
Thelocephala
(7 species)
(7 taxa)
Genus:
Pyrrhocactus
(27 species)
(29 taxa)
Genus:
Eriosyce
(58 taxa)
Groups:
Eriosyce
(2 species)
(2 taxa)
Pyrrhocactus
(5 species)
(7 taxa)
Neoporteria
(6 species)
(12 taxa)
Horridocactus
(13 species)
(23 taxa)
Thelocephala
(3 species)
(10 taxa)
Islaya
(3 species)
(6 taxa)
*refers to a publication including only Chilean taxa.
Guerrero & al. Eriosyce s.l. systematics and taxonomy
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TAXON 68 (3) June 2019: 557573
partitions. Bayesian searches were run for 20 million genera-
tions across four independent runs with four chains each, sam-
pling every 1000 generations. Convergence was monitored
using standard deviation of split frequencies; when this value
stabilized below 0.001, it was considered a strong indication
of convergence. The associated likelihood values, effective
sample size (ESS) values, and burn-in values of the different
runs were verified with the program Tracer v.1.6 (Rambaut
& al., 2014). Both procedures for convergence monitoring re-
vealed that the first 2000 trees should be discarded, therefore,
the sumtfunction considered 18,000 posterior distribution
trees for building the majority-rule consensus tree. Phyloge-
netic trees obtained from ML and BI retrieved overall similar
topologies, though ML search showed less node support than
BI. Trees were visualized and edited using the software Figtree
v.1.4.3 (available at http://tree.bio.ed.ac.uk/software/figtree/)
and Adobe Photoshop CS5. We ran an extra Bayesian analysis
excluding Yavia cryptocarpa R.Kiesling & Piltz, since this ge-
nus is missing in the ycf1 dataset.
The placement of infraspecific taxa was inferred through a
phyletic approach, considering the proximity of the taxa
forming species complexes in our molecular phylogenetic hy-
pothesis. In general, we opted to split a taxonomic complex
when our phylogenetic analyses indicated that two taxa (which
were previously placed in a species complex such as Eriosyce
subgibbosa (Haw.) Katt.) are not sister taxa in the phylogenetic
hypothesis.
Morphology and distribution.
Plants of all taxa
included in the molecular analyses were studied by the
authors in situ during field expeditions and in herbaria
(CONC, SGO) as well as ex situ in living collections in
Chile. Revisions of specimens were here used to discuss the
morphological trends in the obtained phylogenetic hypothesis.
Distribution data was obtained from different sources: field ex-
cursions, literature (e.g., Eggli & al., 1995; Guerrero & al.,
2011b; Duarte & al., 2014), and the Chilean herbaria CONC
and SGO. These locality data are not included here, following
CITES norms for species listed under significant threat due to
illegal collecting (Goettsch & al., 2015).
RESULTS
The five sequence matrices included 5054 bp of aligned
sequence data for 82 taxa, of which 1169 were variable for
the complete matrix and 504 for the ingroup only
(Table 2). Of the aligned concatenated matrix (suppl.
Appendix S1), the plastid non-coding marker rpl32-trnL
contributed with 1353 bp (locus with 27% of variable sites),
trnL-trnF with 1084 bp (locus with 19% of variable sites)
and trnH-psbA with 439 bp (locus with 18% of variable
sites), while the plastid gene ycf1 added 1143 bp (locus with
35% of variable sites) and the nuclear gene PHYC contrib-
uted 1033 bp (locus with 13% of variable sites). Individual
markers retrieved overall less-supported trees compared to
the results obtained from the analysis of the concatenated
matrix, and differences among them are attributable to low
support in specific nodes (suppl. Figs. S1S6). When nodes
presented posterior probability (PP) >0.90, Eriosyce was
systematically grouped in a monophyletic lineage.
The analyses of the concatenated matrix recovered a molec-
ular phylogenetic hypothesis with high statistical support for the
evolutionary relationships between and within outgroup and
ingroup taxa (Fig. 3). Phylogenetic analyses grouped Eriosyce
species in a monophyletic clade within Notocacteae (Fig. 3).
Within Eriosyce, seven clades are recognizable. The strongly
supported first branching clade (Clade I: PP = 1.0, BS [bootstrap
support] = 100) comprises five taxa from Chile, Argentina and
Peru including E. aurata (Pfeiff.) Backeb. (Fig. 3). Next
branching is a strongly supported monotypic clade (Clade II:
PP = 1.0, BS = 100; Fig. 3). Subsequently branching is Clade
III (PP = 1.0, BS = 100) with four Argentinean endemic species
(Fig. 3). Then, a group of 12 taxa from central and northern
central Chile forms a well-supported clade (Clade IV: PP =
1.0, BS = 96), and the next branching clade (Clade V: PP =
1.0, BS = 81) harbors three species from northern central
Chile (Fig. 3). Subsequently branching is a clade (Clade VI:
PP = 1.0, BS = 97) comprising 15 taxa from southern to north-
ern central Chile. Finally, a less-supported species-rich clade
(Clade VII: PP = 0.88, BS = 47) is composed of 22 taxa from
northern Chile (Fig. 3).
DISCUSSION
Systematic relationships in Notocacteae and generic
delimitation.
Our phylogenetic analyses included 11
outgroup genera with 19 species. In the BI phylogenetic tree,
Blossfeldia and the two sampled genera of tribe Trichocereeae
(i.e., Echinopsis and Haageocereus) were placed outside of a
Table 2. Statistics for the 82-sample DNA sequence alignments.
Locus Total length
Ingroup, variable
characters
Total variable
characters
Parsimony-infor-
mative characters
Ingroup
coverage (%)
Outgroup
coverage (%)
rpl32-trnL 1353 157 362 173 97 84
trnL-trnF 1084 112 200 90 100 68
trnH-psbA 439 31 77 26 94 53
ycf1 1143 119 398 290 90 89
PHYC 1033 85 132 47 87 26
Concatenated matrix 5054 504 1169 626
Guerrero & al. Eriosyce s.l. systematics and taxonomy
562
TAXON 68 (3) June 2019: 557573
Fig. 3.
Molecular phylogenetic cladogram of Eriosyce based on five loci (four plastid markers and one nuclear marker) for 82 taxa. Bayesian posterior
probabilities are shown at the nodes, and reflected in the color of the branches (from dark blue = 1 to red = 0.5). Roman numerals show major clades
within the phylogeny. The vertical line describes major geographic distribution of taxa, i.e., green (Argentina), orange (Peru), blue (central Chile) and
black (northern Chile). The species Eriosyce islayensis is believed to be extinct in the wild in Chile (Cáceres & al., 2017).
Guerrero & al. Eriosyce s.l. systematics and taxonomy
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TAXON 68 (3) June 2019: 557573
well-supported clade harboring genera that have been
associated with the southern South American tribe
Notocacteae by different authors (e.g., Buxbaum, 1958, 1975;
Anderson, 2001; Hunt & al., 2006). The species of the genus
Frailea formed a monophyletic clade sister to the remainder
of the tree. Our results do not support the placement of
Rimacactus laui within Eriosyce s.l., which is in accordance
with Mottrams erection of the monotypic genus and results
presented by Nyffeler & Eggli (2010). Indeed, R. laui is
placed in a strongly supported clade as sister to Neo-
werdermannia plus Yavia.
Generic and infrageneric classification.
Molecular
studies suggested that Eriosyce s.l. is paraphyletic, as
E. aurata and E. islayensis (C.F.Först.) Katt. clustered with
Neowerdermannia vorwerkii Fričand not with the other
Eriosyce taxa (Hernández-Hernández & al., 2011) or,
respectively, as E. aurata and E. islayensis were placed
within a polytomous clade together with species of Parodia
s.l. and N. vorwerkii and not in the same clade together with
E. napina and E. subgibbosa (Bárcenas & al., 2011). Our
more densely sampled analyses (90% of the accepted taxa,
suppl. Table S1) do not support these earlier results, but
rather suggest that Eriosyce s.l. is monophyletic when
Rimacactus laui is excluded from the genus. Although we
use an exhaustive sampling for Eriosyce s.l., we
acknowledge that our outgroup dataset has some missing data
that may potentially affect our results. However, the generic
delimitation of Eriosyce is strongly supported by the data
since the core Notocacteae taxa were well represented in our
study for all markers used (most data gaps are found in taxa
more distantly related to Eriosyce such as Frailea spp. and
Echinopsis). Although data is missing for some Parodia
species sampled, this did not appear to influence the
phylogenetic reconstruction since these species clustered in a
strongly supported monophyletic group with the Parodia
species for which all markers were sequenced.
Our analyses show that earlier infrageneric classifications,
i.e., the sectionsand subsectionsof Eriosyce proposed by
Kattermann (1994), the groupsproposed by Hunt & al.
(2006, 2013), and the subgenerasuggested by Hoffmann &
Walter (2004) based on the six former segregate genera Islaya,
Pyrrhocactus,Neoporteria,Horridocactus,Neochilenia and
Thelocephala, are not monophyletic. Our analyses retrieved
six well-supported monophyletic clades within Eriosyce
(= Eriosyce sensu Kattermann minus R. laui). The first
branching Clade I harbors five taxa (Fig. 3) with a mainly
northerly distribution from both sides of the Andes. Only
E. aurata extends its southern distribution as far as 33° S
throughout the Andes range. Eriosyce umadeave (Werderm.)
Katt., a species from the eastern slopes of the Andes, was re-
trieved sister to the rest of Clade I consisting of two well-
supported clades composed of two species of Eriosyce s.str.
and two taxa of E. islayensis.Eriosyce umadeave was formerly
placed within subsection Pyrrhocactus (Kattermann, 1994), re-
spectively the Pyrrhocactus group(Hunt & al., 2006). How-
ever, according to our results, it is more closely related to the
west Andean than to the east Andean members of the genus.
The gross morphology of this narrow endemic species is simi-
lar to that of E. rodentiophila F.Ritter and both are distributed
at the same latitude. The members of this clade share the pres-
ence of large stem areoles with white dense wool, woolly and
bristly floral areoles and a fibrous root system.
Interestingly, the next branching Clade II consists of only
two taxa, i.e., the typical variety of Eriosyce marksiana
(F.Ritter) Katt. and E. marksiana var. lissocarpa (F.Ritter)
Katt. (Fig. 3). The two taxa are endemic to southern central
Chile, occurring in the foothills of the Andes from 100 to
1000 m. The stem diameters of both taxa are large and they
share a fibrous root system, markedly bell-shaped flowers,
and floral and fruit areoles lacking spines and bristles. Ritter
(1960a,b) placed these taxa within Pyrrhocactus, but
Backeberg (1962) included them in Horridocactus because of
the nearly naked flowers and fruits, and following Backeberg,
Kattermann (1994) placed the species E. marksiana within sec-
tion Neoporteria subsection Horridocactus. However, accord-
ing to our results E. marksiana is placed in its own clade as
sister to Clades III to VII (Fig. 3). Despite the shared charac-
ters, E. marksiana var. lissocarpa should be considered as a
different taxon since it presents some important differences
compared to E. marksiana, for example, has thin and longer
spines, wider perianth segments, and the old plants have more
elongated stems (Kattermann, 1994).
The subsequent lineage (Clade III, Fig. 3) harbors four
species exclusively distributed on the eastern slopes of the
Andes suggesting that the mountain range acted as a strong
isolating barrier. Our results refute Ritters (1980) proposal
to unite the west- and east Andean taxa under the genus
Pyrrhocactus, but partially support Backebergs (1962) pro-
posal to consider these species as a distinct group. Yet, his
proposal to consider them as a different genus was not sup-
ported by our study. The four taxa in this clade are charac-
terized by a basal partial circumscissile slit in their fruits
and by their low rib numbers.
Several species distributed in southern and northern central
Chile form Clade IV (Fig. 3). This clade comprises 12 taxa and is
nearly congruent with Kattermanns subsection Horridocactus.
With the exception of Eriosyce napina subsp. napina, subsp.
duripulpa (F.Ritter) Katt. and subsp. lembckei Katt., that occur
in the southern coastal desert, all of the taxa in this clade are
distributed in the sclerophyllous woodlands and spiniferous or
desert matorrals of central Chile. In our phylogenetic hypothe-
sis, E. heinrichiana subsp. intermedia was placed between
the subclade composed by E. curvispina (Bertero ex Colla)
Katt. plus E. curvispina subsp. tuberisulcata (Jacobi) Katt. and
the rest of the taxa in this clade, and not sister to the
typical E. heinrichiana subsp. heinrichiana, a taxon placed
within Clade VI, thus refuting the broad concept of
E. heinrichiana (Kattermann, 1994; Anderson, 2001, 2005;
Hoffmann & Walter, 2004; Hunt & al., 2006, 2013). The major-
ity of authors placed E. napina and its subspecies within Thelo-
cephala, but our results support Kattermanns (1994) proposal
to place it within his subsection Horridocactus. In our
Guerrero & al. Eriosyce s.l. systematics and taxonomy
564
TAXON 68 (3) June 2019: 557573
phylogenetic tree, E. napina subsp. duripulpa was placed sister
to the closely related sister pair E.napina subsp. napina and
E.napina subsp. lembckei in a strongly supported subclade.
Thus, the very broad concept of E. napina (e.g., Hoffmann &
Walter, 2004; Hunt & al., 2006; Walter & Mächler, 2006) is
not supported by our results, as E. napina subsp. fankhauseri,
subsp. tenebrica and subsp. riparia were placed in Clade V.
The strongly supported sister pair E. curvispina subsp.
curvispina and E.curvispina subsp. tuberisulcata was retrieved
as sister to the rest of the taxa of Clade IV. A close relationship
between these two subspecies is suggested by their low genetic
divergence (Fig. 3). Yet, the monophyly of the E. curvispina
species complex was not supported since E. curvispina subsp.
armata was retrieved closely related to the E. aspillagae group.
Also we could not include in our analysis (due to DNA ex-
traction problems) E. curvispina var. mutabilis (F.Ritter)
Katt. and E. curvispina var. robusta (F.Ritter) Katt.; thus,
more research is necessary to fully resolve the relationships
within the E. curvispina complex. The relation between the
sister pair E. aspillagae subsp. aspillagae and E. aspillagae
subsp. maechlerorum also remains unresolved (P0.59, BS =
26) and needs further study. All members of Clade IV share
tubular nectaries, elongated to isodiametric ovaries and, with
the exception of E. napina subsp. napina, subsp. lembckei
and subsp. duripulpa, their floral and fruit areoles bear scant
short wool.
The subsequently branching Clade V (Fig. 3) is a small,
well-defined clade comprising three taxa (i.e., Eriosyce napina
subsp. tenebrica (F.Ritter) Ferryman, E. napina subsp. riparia
Mächler & Helmut Walter and E. napina subsp. fankhauseri
(F.Ritter) Mächler & Helmut Walter). Kattermann (1994)
erected subsection Chileosyce for small-bodied plants with
wind-dispersed woolly and bristly fruits and ribs dissolved into
tubercles arranged in parastichies. He included E. tenebrica
(F.Ritter) Katt., E. esmeraldana (F.Ritter) Katt., E. aerocarpa
(F.Ritter) Katt., E. krausii and R. laui (= E. laui). In our study,
we involved three more taxa also bearing these morphological
character states, i.e., E. napina subsp. riparia and two taxa
previously assigned to Thelocephala by Ritter (1980), i.e.,
E. napina subsp. fankhauseri and E. odieri subsp. malleolata
(F.Ritter) A.E.Hoffm. & Helmut Walter. Based on the above-
mentioned morphological characters, the above cited taxa were
considered to be members of Thelocephala or Chileosyce in
previous studies (Ritter, 1980; Hoffmann, 1989; Hoffmann &
Walter, 2004; Hunt & al., 2006, 2013). But all these grouping
concepts are not monophyletic according to our findings, as
E. aerocarpa, E. esmeraldana, and E. odieri subsp. malleolata
were placed in Clade VII. The above-mentioned morphological
characters appear to have evolved independently several times
in different lineages inhabiting similar habitats.
Clade VI (Fig. 3) comprises 15 taxa and is similar to
Kattermanns subsection Neoporteria. Its members usually
share tubular flowers adapted to humming-bird pollination,
but one species (i.e., Eriosyce chilensis (Hildm. ex K.Schum.)
Katt.) was formerly considered a member of Neochilenia by
Backeberg (1951) or of Pyrrhocactus by Ritter (1980) for its
funnel-form flowers. The discrepancy between the two differ-
ent floral types was interpreted as a reversal from a
hummingbird- to bee-pollination syndrome (Kattermann,
1994; Walter, 2008; Guerrero & al., 2011b). In our phyloge-
netic hypothesis, E. chilensis is placed in a polytomy with
E. heinrichiana and the rest of the taxa in this clade. Its rela-
tionship to the other members of the clade remains still unclear
(see Walter, 2008) and needs further study in order to enlighten
the evolution of flower morphology and the speciation of this
narrow endemic. Furthermore, our results suggest that
Kattermanns subsection Neoporteria is non-natural: on the
one hand, E. sociabilis (F.Ritter) Katt. was placed within his
subsection Neoporteria; according to our analyses, however,
it is not part of Neoporteria but part of the clade harboring taxa
from more northerly regions (Clade VII; see also Guerrero &
al., 2011b). On the other hand, the two sympatrically growing
species E. heinrichiana and E. simulans from Kattermanns
subsection Horridocactus were placed within the Neoporteria
clade in our analyses, although their flowers are funnel-form
and bee-pollinated. This result might be considered as another
example for the finding that closely related species may have
different pollination syndromes, as floral characters and polli-
nation syndromes in cacti are evolutionary labile (Nyffeler &
Eggli, 2010; Schlumpberger & Renner, 2012). The three north-
ernmost distributed taxa in the clade, E. villosa (Monv.) Katt.,
E. subgibbosa subsp. vallenarensis (F.Ritter) Katt. and
E. subgibbosa subsp. wagenknechtii (F.Ritter) Katt. form a
well-supported clade in our analyses, a finding that confirms
the results of Guerrero & al. (2011a) and Walters assumption
(2008) and refutes Kattermanns (1994) broad concept of
E. subgibbosa. Only recently, however, he accepted
E. subgibbosa subsp. vallenarensis and subsp. wagenknechtii
as species in their own right (Kattermann, 2018). The rela-
tionships between the other four infraspecific taxa of
E. subgibbosa sensu Kattermann are still not completely re-
solved in our study (see Guerrero & al., 2011a), as they all ap-
pear in a terminal polytomy together with the typical E. senilis
(Backeb.) Katt and E. senilis subsp. coimasensis (F.Ritter)
Katt. Yet, the two sympatrically occurring taxa E. subgibbosa
subsp. nigrihorrida (Backeb. ex A.W.Hill) Katt. and
E. subgibbosa var. litoralis (F.Ritter) Katt. are placed together
in a strongly supported subclade, suggesting a close relation-
ship. Finally, our analyses placed E. senilis subsp. elquiensis
Katt., a taxon growing disjunctly from E. senilis south of and
in the Rio Elqui Valley, sister to the terminal subclade.
The species-rich Clade VII (Fig. 3) harbors taxa from the
northern coastal desert. This zone is characterized by hyperarid
conditions (Luebert & Pliscoff, 2006) that have triggered pop-
ulation fragmentation and recent colonization of the biota due
to phylogenetic niche conservatism (Pinto & Kirberg, 2009;
Guerrero & al., 2013). All the members of Clade VII have tap-
roots and, with the exception of the closely related sister pair
Eriosyce confinis (F.Ritter) Katt. and E. kunzei (C.F.Först.)
Katt., the two southernmost species in this clade, they all share
basally widened nectaries and compressed ovaries. Clade VII
is an assemblage of 22 closely related taxa previously placed
Guerrero & al. Eriosyce s.l. systematics and taxonomy
565
TAXON 68 (3) June 2019: 557573
in different subsections (14 taxa in subsection Horridocactus,
4 taxa in subsection Chileosyce, 3 taxa in subsection Islaya
and 1 taxon in subsection Neoporteria) by Kattermann
(1994).
Former hypotheses of the infrageneric classification of
Eriosyce were based on morphological characters alone, and
were shown to be non-natural by our results. Convergent evo-
lution of species living in similar habitats and potential hybrid-
ization might be the reason why morphology-based
classifications are not always compatible with molecular phy-
logenetic hypotheses (Ritz & al., 2007; Guerrero & al.,
2011b; Schlumpberger & Renner, 2012).
Placement of infraspecific taxa.
Our analyses also
resolved various controversial taxonomic problems at species
and infraspecific levels. The present circumscription of some
large species complexes (Eriosyce subgibbosa,E. napina,
E. odieri,E. curvispina,E. crispa,E. senilis,E. taltalensis
(Hutchison) Katt., E. heinrichiana,E. eriosyzoides (F.Ritter)
Ferryman) proposed in a lumping tendency by various authors
was not supported by our results. In the sense of Kattermann
(1994), E. subgibbosa includes seven infraspecific taxa, only
one of which (E.subgibbosa subsp. clavata) was grouped
within the same but weakly supported and unresolved
subclade as the typical E. subgibbosa (Fig. 3). Eriosyce
subgibbosa var. litoralis is placed in a well-supported
subclade together with the sympatrically occurring
E. subgibbosa subsp. nigrihorrida. Together with E. villosa,
the two northernmost subspecies E. subgibbosa subsp.
vallenarensis and E. subgibbosa subsp. wagenknechtii were
placed as sister to the other taxa of clade VI and not with
E. subgibbosa (Walter, 2008; Guerrero & al., 2011b). Finally,
E. subgibbosa subsp. subgibbosa var. castanea (F.Ritter) Katt.
does also not appear within the same clade as E. subgibbosa.
Furthermore, only two of the taxa formerly assigned to
E. napina by Kattermann (1994, 2001), Anderson (2001),
Hoffmann & Walter (2004), Hunt & al. (2006, 2013) and
Walter & Mächler (2006) were placed in the same clade with
the typical subspecies. Also, none of the taxa assigned to
E. odieri by Kattermann (1994), Anderson (2001, 2005),
Hoffmann & Walter (2004) and Hunt & al. (2006, 2013) were
placed together with E. odieri in our trees: E. odieri subsp.
krausii was placed within Clade VII. Similarly, E. odieri
subsp. malleolata, formerly considered a synonym of
E. krausii, respectively E. odieri subsp. krausii (Kattermann,
1994; Anderson, 2001, 2005; Hunt, & al. 2006, 2013), is not
closely related to E. odieri but to E. esmeraldana. Finally,
E. odieri subsp. fulva (F.Ritter) Katt. and E. odieri subsp.
glabrescens (F.Ritter) Katt. were placed within different
subclades of clade VII, but not close to E. odieri. The
proposed separation (Ferryman, 2003) of E. taltalensis subsp.
paucicostata (F.Ritter) Katt. and E. calderana (F.Ritter)
Ferryman (a synonym of E. taltalensis sensu Kattermann,
1994) from the large E. taltalensis complex (Kattermann,
1994) was supported by our data, as E. taltalensis subsp.
taltalensis was placed in a different subclade together with the
closely related E. taltalensis subsp. pygmaea (F.Ritter)
Ferryman and E. sociabilis and not together with E. taltalensis
subsp. paucicostata or E. calderana. It should be mentioned,
however, that our sampling did not include the other two
infraspecific taxa assigned to E. taltalensis sensu Kattermann
(i.e., E. taltalensis var. echinus (F.Ritter) Katt., E. taltalensis
var. floccosa (F.Ritter) Katt.) due to DNA amplification
problems. Also, there was no support for the inclusion of
E. crispa subsp. atroviridis (F.Ritter) Katt. within E. crispa
(Kattermann, 1994). The broad concept of E. curvispina was
not supported by our data, as E. curvispina subsp. armata
(F.Ritter) Ferryman did not cluster with E. curvispina but with
E. aspillagae (F.Ritter) Katt. Furthermore, the inclusion of
E. marksiana and E. marksiana var. lissocarpa into
E. curvispina (Hoffmann, 1989; Hoffmann & Walter, 2004;
Hunt & al., 2006, 2013) was also clearly refuted by our
results. The wide concept of E. senilis (Kattermann, 1994;
Anderson, 2001, 2005; Hoffmann & Walter, 2004; Hunt & al.,
2006, 2013) was also not supported by our data, as none of its
subspecies clustered with E. senilis subsp. senilis. The
supposition that E. andreaeana Katt. is part of E. strausiana
(K.Schum.) Katt. (Hunt & al., 2006) is not supported by our
data, as the latter is more closely related to E. villicumensis
(Rausch) Katt. than to E. andreaeana. Finally, our results
suggest that the two subspecies of E. heinrichiana
(Kattermann, 1994) are not at all closely related, as
E. heinrichiana subsp. intermedia (F.Ritter) Katt. was placed
within Clade IV, while E. heinrichiana was placed in Clade
VI. The taxonomic status of the two lately described species
E. caligophila Pinto (2005) and E. spectabilis Katt. & al.
(Kattermann & al., 2011) was confirmed by our analyses: the
former is sister to the terminal clade, while the latter is sister to
a sister pair comprising E. aerocarpa and E. odieri subsp.
glabrescens.
TAXONOMIC TREATMENT
The results of this study have taxonomic implications for
the classification of Eriosyce at infrageneric and (infra)specific
levels. We studied 63 species, subspecies and varieties (86%)
of the 73 here accepted Eriosyce taxa (see also suppl. Table
S1). Following the principle that only monophyletic clades
above species level should be named (Stuessy, 2009), the
seven monophyletic clades retrieved by our analyses form the
basis of our proposed infrageneric classification (see also
suppl. Table S1).
Eriosyce Phil. in Anales Univ. Chile 41: 721. 1872 Type:
E. sandillon (Gay) Phil. (Echinocactus sandillon Gay).
Clade I: Eriosyce sect. Eriosyce
Clade II: Eriosyce sect. Campanulatae P.C.Guerrero &
Helmut Walter, sect. nov. Type: E. marksiana (F.Ritter)
Katt. (Pyrrhocactus marksianus F.Ritter).
Guerrero & al. Eriosyce s.l. systematics and taxonomy
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TAXON 68 (3) June 2019: 557573
Diagnosis. Flowers markedly campanulate; flower and
fruit areoles nearly naked; root system fibrous.
Clade III: Eriosyce sect. Pyrrhocactus (A.Berger) P.C.Guer-
rero & Helmut Walter, comb. & stat. nov. Pyrrhocactus
A.Berger, Kakteen: 215, 345. 1929 Type (designated by
Kattermann in Eriosyce (Cactac.) Gen. Revis. & Ampl.
(Succ. Pl. Res. 1): 134. 1994): E. strausiana (K.Schum.)
Katt.
Clade IV: Eriosyce sect. Horridocactus (Backeb.) P.C.Guer-
rero & Helmut Walter, comb. & stat. nov. Horrido-
cactus Backeb. in Blätt. Kakteenf. 1938(6): [21]. 1938
Type: E. curvispina (Bertero ex Colla) Katt. (Cactus
curvispinus Bertero ex Colla).
Eriosyce armata (F.Ritter) P.C.Guerrero & Helmut Walter,
comb. nov. Pyrrhocactus armatus F.Ritter in Succulenta
(Netherlands) 39: 49. 1960 Horridocactus armatus
(F.Ritter) Backeb., Cactaceae 6: 3792. 1962 Neoporteria
tuberisulcata var. armata (F.Ritter) Donald & G.D.
Rowley in Cact. Succ. J. Gr. Brit. 28: 58. 1966 Neo-
porteria armata (F.Ritter) Krainz, Städt. Sukku-
lentensamml. Zürich, Kat. Kult. Stehenden Art., ed. 2:
86. 1967 Neoporteria horrida var. armata (F.Ritter)
A.E.Hoffm., Cact. Fl. Silv. Chile: 190. 1989 Eriosyce
curvispina var. armata (F.Ritter) Katt., Eriosyce (Cactac.)
Gen. Revis. & Ampl. (Succ. Pl. Res. 1): 117. 1994
E. curvispina subsp. armata (F.Ritter) Katt. in Cactaceae
Syst. Init. 12: 14. 2001 Holotype: [Chile]: on tops of
mountains S of El Paico, SW of Santiago, May 1955,
Ritter 449 (U barcode 0249336!).
Eriosyce duripulpa (F.Ritter) P.C.Guerrero & Helmut Wal-
ter, comb. nov. Chileorebutia duripulpa F.Ritter in
Taxon 12(3): 123. 1963 Neochilenia duripulpa
(F.Ritter) Backeb., Descr. Cact. Nov. 3: 9. 1963 The-
locephala duripulpa (F.Ritter) F.Ritter, Kakteen
Südamerika 3: 1010. 1980 Neoporteria napina var.
duripulpa (F.Ritter) A.E.Hoffm., Cact. Fl. Silv. Chile:
224. 1989 Eriosyce napina var. duripulpa (F.Ritter)
Katt., Eriosyce (Cactac.) Gen. Revis. & Ampl. (Succ.
Pl. Res. 1): 118. 1994 Eriosyce napina subsp.
duripulpa (F.Ritter) Katt. in Cact. Syst. Init. 12: 14.
2001 Holotype: Atacama, North Chile, Oct 1962,
Ritter 1056 (U barcode 0007624!).
=Eriosyce napina subsp. challensis I.Schaub & Keim, Cactus
Co. 9(2): 111. 2005 Thelocephala challensis (I.Schaub
& Keim) Faúndez & Saldivia in Gayana, Bot. 68(2):
317. 2011 Holotype: Chile, Coquimbo, 20 Aug 2004,
Schaub & Keim s.n. (SGO No. 151590!).
Eriosyce spinosior (F.Ritter) P.C.Guerrero & Helmut Walter,
comb. & stat. nov. Pyrrhocactus jussieui var. spinosior
F.Ritter, Kakteen Südamerika 3: 952. 1980 Holotype:
Chile, 12 km E of Vicuña, Dec 1955, Ritter 252b (U
barcode 0249327!).
=Echinocactus jussieui Monv. ex Salm-Dyck, Cact. Hort.
Dyck: 34. 1850 Neoporteria jussieui (Monv. ex Salm-
Dyck) Britton & Rose, Cactaceae 3: 96. 1922 Neo-
chilenia jussieui (Monv. ex Salm-Dyck) Backeb. in
Repert. Spec. Nov. Regni Veg. 51: 60. 1942
Pyrrhocactus jussieui (Monv. ex Salm-Dyck) F.Ritter,
Kakteen Südamerika 3: 951. 1980. Type: not designated.
= Echinocactus fobeanus Mieckley, Monatsschr. Kakteenk.
17: 187. 1907 Neoporteria jussieui var. fobeana
(Mickley) Donald & G.D.Rowley in Cact. Succ. J. Gr.
Brit. 28: 57. 1966 Type: not designated.
=Pyrrhocactus dimorphus F.Ritter in Succulenta (Netherlands)
1962: 3. 1962 Neochilenia dimorpha (F.Ritter) Backeb.,
Descr. Cact. Nov. 3: 9. 1963 Neoporteria dimorpha
(F.Ritter) Donald & G.D.Rowley in Cact. Succ. J. Gr. Brit.
28: 56. 1966 Neoporteria jussieui var. dimorpha
(F.Ritter) A.E.Hoffm., Cact. Fl. Silv. Chile: 200. 1989
Holotype: Rocky hills near Coquimbo, Chile, Dec 1957,
Ritter 707 (U barcode 0249329!).
=Pyrrhocactus setosiflorus F.Ritter in Succulenta (Nether-
lands) 1962: 70. 1962 Neochilenia setosiflora (F.Ritter)
Backeb., Descr. Cact. Nov. 3: 10. 1963 Neoporteria
setosiflora (F.Ritter) Donald & G.D.Rowley in Cact Succ.
J. Gr. Bit. 28: 58. 1966 Neoporteria jussieui var.
setosiflora (F.Ritter) A.E.Hoffm., Cact. Fl. Silv. Chile:
200. 1989 Holotype: coast of Mid-Chile, ca. 31°30lat-
itude, Dec 1957, Ritter 708 (U barcode 0249258!).
=Pyrrhocactus setosiflorus var. intermedius F.Ritter in
Succulenta (Netherlands) 1962: 70. 1962 Neochilenia
setosiflora var. intermedia (F.Ritter) Backeb., Descr. Cact.
Nov. 3: 10. 1963 Neoporteria setosiflora var. intermedia
(F.Ritter) Donald & G.D.Rowley in Cact. Succ. J. Gr. Brit.
28: 58. 1966 Eriosyce heinrichiana var. setosiflora
(F.Ritter) Katt., Eriosyce (Cactac.) Gen. Revis. & Ampl.
(Succ. Pl. Res. 1): 118. 1994 Eriosyce heinrichiana
subsp. intermedia (F.Ritter) Katt., Eriosyce (Cactac.) Gen.
Revis. & Ampl. (Succ. Pl. Res. 1): 118. 1994 Holotype:
hill ca. 60 km S of Coquimbo E of the Panamericana,
Chile, Dec 1957, Ritter 708a (U barcode 0249259!).
=Pyrrhocactus jussiieui var. australis F.Ritter, Kakteen
Südamerika 3: 952. 1980 Isotype: Las perdices, Chile,
Mar 1963, Ritter 252d (ZSS barcode ZSS-013327 n.v.).
Note. Monvilles description of Echinocactus jussieui is
poor and published without an illustration. Moreover, the lo-
cality is given as Chile; thus the application of this name to
plants from the Elqui Valley is speculative, and for that reason
the name Eriosyce jussieuishould be treated as of uncertain
application. RittersPyrrhocactus jussieui var. spinosior, how-
ever, is well described and illustrated and is here used as a
basionym for the plants from around the Elqui Valley.
Clade V: Eriosyce sect. Diaguita P.C.Guerrero & Helmut Wal-
ter, sect. nov. Type: E. tenebrica (F.Ritter) Katt. (
Guerrero & al. Eriosyce s.l. systematics and taxonomy
567
TAXON 68 (3) June 2019: 557573
E. napina subsp. tenebrica (F.Ritter) Ferryman Thelo-
cephala tenebrica F.Ritter).
Diagnosis. Flowers funnelform; flower and fruit with
wool and conspicuous porrect bristles; root system taproot.
Eriosyce fankhauseri (F.Ritter) P.C.Guerrero & Helmut Wal-
ter, comb. nov. Thelocephala fankhauseri F.Ritter,
Kakteen Südamerika 3: 1002. 1980 Neoporteria napina
var. fankhauseri (F.Ritter) A.E.Hoffm., Cact. Fl. Silv.
Chile: 224. 1989 Eriosyce napina subsp. fankhauseri
(F.Ritter) W.Mächler & Helmut Walter in Cactus World
24(3): 141. 2006 Eriosyce napina var. fankhauseri
(F.Ritter) Romulski, Kaktusy Inne 4(3): 113. 2007 Holo-
type: Chile, Atacama, mountains NW of Domeyko, 1969,
Ritter 1451 (U barcode 0249079!).
Eriosyce riparia (Mächler & Helmut Walter) P.C.Guerrero &
Helmut Walter, comb. nov. Eriosyce napina subsp.
riparia Mächler & Helmut Walter in Cactus World
24(3): 142. 2006 E. napina var. riparia (Mächler &
Helmut Walter) Romulski in Kaktusy Inne 4(3): 131.
2007 Thelocephala riparia (Mächler & Helmut Walter)
Faúndez & Saldivia in Gayana, Bot. 68(2): 317. 2011
Holotype: Chile, Prov. Elqui, east of Trapiche, 3 Nov
2004, Helmut Walter 487 (SGO No. 152410!).
Clade VI: Eriosyce sect. Neoporteria (Britton & Rose) Katt.,
Eriosyce (Cactac.) Gen. Revis. & Ampl. (Succ. Pl. Res. 1):
117. 1994 Neoporteria Briton & Rose, Cactaceae 3: 94.
1922 Type: E. subgibbosa (Haw.) Katt. (Echinocactus
subgibbosus Haw.).
Eriosyce castanea (F.Ritter) P.C.Guerrero & Helmut Walter,
comb. nov. Neoporteria castanea F.Ritter in Taxon
12(1): 34. 1963 Neoporteria subgibbosa var. castanea
(F.Ritter) Ferryman in Preston-Mafham, Cact. Ill. Dict.:
148. 1991 Eriosyce subgibbosa var. castanea (F.Ritter)
Katt., Eriosyce (Cactac.) Gen. Revis. & Ampl. (Succ. Pl.
Res. 1): 119. 1994 Holotype: Chile, Talca, Villa Prat,
Apr 1954, Ritter 236 (U barcode 0108728!).
=Neoporteria castanea var. tunensis F.Ritter in Taxon 12(1):
34. 1963 Holotype: Tuna, near San Fernando, Chile,
May 1955, Ritter 236a (U barcode 0108729!).
Eriosyce coimasensis (F.Ritter) P.C.Guerrero & Helmut
Walter, comb. nov. Neoporteria coimasensis F.Ritter
in Taxon 12(1): 34. 1963 Neoporteria nidus var.
coimasensis (F.Ritter) A.E.Hoffm., Cact. Silv. Fl.
Chile: 164. 1989 Eriosyce senilis subsp. coimasensis
(F.Ritter) Katt., Eriosyce (Cactac.) Gen. Revis. & Ampl.
(Succ. Pl. Res. 1): 119. 1994 Holotype: Chile, Aconca-
gua, Las Coimas, nr. 4, Apr 1955, Ritter 473 (U barcode
0108733!).
=Neoporteria robusta F.Ritter in Taxon 12(1): 34. 1963
Neoporteria coimasensis var. robusta (F.Ritter) F.Ritter,
Kakteen Südamerika 3: 10411042. 1980 Neoporteria
subgibbosa var. robusta (F.Ritter) A.E.Hoffm., Cact. Fl.
Silv. Chile: 168. 1989 Holotype: Chile, Las Coimas,
Jun 1955, Ritter 473b (U barcode 0108734 n.v.).
Eriosyce elquiensis (Katt.) P.C.Guerrero & Helmut Walter,
comb. & stat. nov. Eriosyce senilis subsp. elquiensis
Katt, Eriosyce (Cactac.) Gen. Revis. & Ampl. (Succ. Pl.
Res. 1): 117. 1994 Holotype: Chile, El Tambo, 1 Oct
1984, Kattermann 462 (DES n.v.).
Eriosyce litoralis (F.Ritter) P.C.Guerrero & Helmut Walter,
comb. nov. Neoporteria litoralis F.Ritter in Succulenta
(Netherlands) 38: 28. 1959 Neoporteria subgibbosa
var. litoralis (F.Ritter) A.E.Hoffm., Cact. Fl. Silv. Chile:
168. 1989 Eriosyce subgibbosa var. litoralis (F.Ritter)
Katt., Eriosyce (Cactac.) Gen. Revis. & Ampl. (Succ. Pl.
Res. 1): 119. 1994 Isotype: Chile, coastal rocks of
Coquimbo, Jan 1954, Ritter 219 (U barcode 0108744!).
Eriosyce nigrihorrida (Backeb. ex A.W.Hill) P.C.Guerrero &
Helmut Walter, comb. nov. Chilenia nigrihorrida
Backeb. ex A.W.Hill in Index Kew. Suppl. 9: 62. 1938
Neotype (designated here): Chile, Región de Coquimbo,
Totoralillo, 17 Aug 2017, Guerrero & Rosas 1322 (CONC
No. 187359!; isoneotypes: CONC No. 187360!, SGO No.
169645!).
Note. The lectotypedesignated by Kattermann
(Eriosyce (Cactac.) Gen. Revis. & Ampl. (Succ. Pl. Res. 1):
167. 1994) is based on a photograph of a cultivated specimen
published by Backeberg & Knuth (in Kaktus-ABC: 301.
1936, Chile, Coquimbo); however, this photograph cannot be
chosen as lectotype (Staples & Prado, 2018).
Clade VII: maintained unnamed until further study.
Eriosyce atroviridis (F.Ritter) P.C.Guerrero & Helmut Walter,
comb. nov. Pyrrhocactus atroviridis F.Ritter in
Succulenta (Netherlands) 1960: 89. 1960 Horridocactus
atroviridis (F.Ritter) Backeb., Cactaceae 6: 3793. 1962
Neoporteria tuberisulcata var. atroviridis (F.Ritter)
Donald & G.D.Rowley in Cact. Succ. J. Gr. Brit. 28: 58.
1966 Neoporteria atroviridis (F.Ritter) Ferryman in
Preston-Mafham, Cact. Ill. Dict.: 140. 1991 Eriosyce
crispa subsp. atroviridis (F.Ritter) Katt., Eriosyce
(Cactac.) Gen. Revis. & Ampl. (Succ. Pl. Res. 1): 118:
1994 Eriosyce eriosyzoides subsp. atroviridis (F.Ritter)
Ferryman in Cact. Syst. Init. 16: 11. 2003 Holotype:
Chile, Vallenar, Jan 1956, Ritter 475 (U barcode
0249249!).
=Pyrrhocactus eriosyzoides var. domeykoensis F.Ritter,
Kakteen Südamerika 3: 938. 1980 Holotype: Chile,
Atacama, Domeyko, 1961, Ritter 484a (U barcode
0249324!).
Eriosyce malleolata (F.Ritter) P.C.Guerrero & Helmut Walter,
comb. nov. Chileorebutia malleolata F.Ritter in Taxon
Guerrero & al. Eriosyce s.l. systematics and taxonomy
568
TAXON 68 (3) June 2019: 557573
12(3): 123. 1963 Neoporteria reichei var. malleolata
(F.Ritter) Donald & G.D.Rowley in Cact. Succ. J. Gr.
Brit. 28: 57. 1966 Thelocephala malleolata (F.Ritter)
F.Ritter, Kakteen Südamerika 3: 1020. 1980 Neo-
porteria esmeraldana var. malleolata (F.Ritter) A.E.
Hoffm., Cact. Fl. Silv. Chile: 222. 1989 Neoporteria
odieri var. malleolata (F.Ritter) Ferryman in Preston
Mafham, Cact. Ill. Dict.: 146. 1991 Eriosyce odieri
subsp. malleolata (F.Ritter) A.E.Hoffm. & Helmut Walter,
Cact. Fl. Silv. Chile: 258. 2004 Holotype: Chile, North of
Chañaral, 1956, Ritter 517 (U barcode 0007625!).
=Chileorebutia malleolata var. solitaria F.Ritter in Taxon 12:
123. 1963 Neochilenia malleolata var. solitaria
(F.Ritter) Backeb., Descr. Cact. Nov. 3: 9. 1963 Thelo-
cephala malleolata var. solitaria (F.Ritter) F.Ritter,
Kakteen Südamerika 3: 1020. 1980 Holotype: Chile,
South of Chañaral, Feb 1956, Ritter 517a (U barcode
0007643!).
=Eriosyce odieri var. weisseri A.E.Hoffmann & Helmut Wal-
ter, Cact. Fl. Silv. Chile: 258. 2004 Thelocephala
weisseri (A.E.Hoffm. & Helmut Walter) Faúndez &
Saldivia in Gayana, Bot. 68(2): 317. 2011 Holotype:
Chile, Antofagasta, Cifuncho, 1987, Hoffmann s.n. (SGO
No. 152412!).
Eriosyce fulva (F.Ritter) P.C.Guerrero & Helmut Walter,
comb. nov. Chileorebutia fulva F.Ritter in Cactus
(Paris) 15(66): 10. 1960 Thelocephala fulva (F.Ritter)
F.Ritter, Kakteen Südamerika 3: 1011. 1980 Holotype:
Chile, Atacama, Totoral, 1956, Ritter 500 (U barcode
0249077!).
Eriosyce glabrescens (F.Ritter) P.C.Guerrero & Helmut Wal-
ter, comb. nov. Chileorebutia glabrescens F.Ritter in
Cactus (Paris) 15(66): 8. 1960 Thelocephala glabre-
scens (F.Ritter) F.Ritter, Kakteen Südamerika 3: 1003.
1980 Eriosyce odieri subsp. glabrescens (F.Ritter).
Katt., Eriosyce (Cactac.) Gen. Revis. & Ampl. (Succ. Pl.
Res. 1): 118. 1994 Eriosyce napina subsp. glabrescens
(F.Ritter) Ferryman in Cact. Syst. Init. 16: 11. 2003
Eriosyce napina var. glabrescens (F.Ritter) Romulski in
Kaktusy Inne 4(3): 131. 2007 Holotype: Chile, Arica,
Sep 1959, Ritter 710 (U barcode 0007626!).
=Eriosyce napina subsp. llanensis Schaub & Keim in Cactus
Co 15(1). 2011 Holotype: Chile, P.N. Llanos del Challe,
18 Nov 2010, Schaub & Keim s.n. (SGO No. 160015!).
Eriosyce kunzei (C.F.Först.) Katt., Eriosyce (Cactac.) Gen.
Revis. & Ampl. (Succ. Pl. Res. 1): 117. 1994 Echino-
cactus kunzei C.F.Först., Handb. Cacteenk.: 293. 1846
Neoporteria kunzei (C.F.Först.) Backeb. in Backeb. &
Knuth, Kaktus-ABC: 260. 1936 Pyrrhocactus kunzei
(C.F.Först.) Borg, Cacti: 262. 1937 Chilenia kunzei
(C.F.Först.) Backeb., Kakteenkunde: 82. 1939 Neo-
chilenia kunzei (C.F.Först.) Backeb. ex Dölz in Repert.
Spec. Nov. Regni Veg. 51: 60. 1942 Neotype
(designated here): Chile, Antofagasta, Copiapó, Paipote,
1955, Ritter 220 loc. 2 Paipote(SGO No. 121487!).
=Echinocactus neumannianus Cels ex Salm-Dyck, Cact. Hort.
Dyck.: 33. 1850 Type: not designated.
=Pyrrhocactus confinis F.Ritter in Succulenta (Netherlands)
1961: 4. 1961 Neochilenia confinis (F.Ritter) Backeb.,
Descr. Cact. Nov. 3: 9. 1963 Neoporteria confinis
(F.Ritter) Donald & G.D.Rowley in Cact. Succ. J. Gr. Brit.
28: 55. 1966 Neoporteria kunzei var. confinis (F.Ritter)
A.E.Hoffm., Cact. Fl. Silv. Chile: 204. 1989 Eriosyce
confinis (F.Ritter) Katt., Eriosyce (Cactac.) Gen. Revis.
& Ampl. (Succ. Pl. Res. 1): 118. 1994 Holotype:
Chile, Atacama, South of Monte Amargo, Jan 1956, Ritter
494 (U barcode 0249244).
Note. This makes Eriosyce confinis (F.Ritter) Katt. a het-
erotypic synonym of E. kunzei (C.F.Först.) Katt.
In his protologue, Förster only added Chileas the type
locality, but mentioned that the plants are sometimes covered
by a light layer of snow in winter. Kattermann (1994) chose
material from BalalaProvincia de Elqui, F. Katterman 459
Balala, in rock cliffs (DBG)for his neotypification of
Eriosyce kunzei and based his decision not to refer the plants
from around Copiapó to FörstersEchinocactus kunzei (as pro-
posed by Ritter, 1980) on the single argument that it never
snows in this area. Yet, a recent snow episode occurred in
the area around the Copiapó Valley affecting the grape-
producing zone (Vergara, 2011), thus this assumption cannot
be upheld. We thus follow Ritters well-argued proposal to
refer Försters epithet kunzei to the plants from near
Copiapó.
CONCLUSIONS
The description of new taxa based on minor morphologi-
cal variation (i.e., splitting) clashes with the contrasting view
of broadly grouping taxa at higher ranks (i.e., lumping)
(Stuessy, 2009). The extremes of both perspectives tend to
converge when more data are considered and explicitly used
to sustain taxonomic classifications. The taxonomic history of
Cactaceae, and in particular of Eriosyce s.l., experienced both
concepts: on the one hand, e.g., Ritters (1980) and
Backebergs (1966) tendency towards extensive splitting, and
on the other hand Donald & Rowley (1966), Hoffmann
(1989), Kattermann (1994), Anderson (2001, 2005), Hoffmann
& Walter (2004) and Hunt & al. (2006, 2013) had the tendency
to lump many taxa in fewer, broadly circumscribed species.
This taxonomic uncertainty has had a concrete impact on con-
servation, since the current conservation status of many
Eriosyce species listed in the IUCN Red List (Duarte & al.,
2014; Goettsch & al., 2015) might have been underestimated
as they were based on very broadly circumscribed species
(Hunt & al., 2006). Therefore, updating taxonomic checklists
of conservation importance such as CITES Appendices should
be undertaken as well-supported new classifications are pub-
lished (Hunt, 2016). With our explicitly phylogeny-based
Guerrero & al. Eriosyce s.l. systematics and taxonomy
569
TAXON 68 (3) June 2019: 557573
classification, we expect to reduce taxonomic uncertainty pro-
duced by lumping and splitting tendencies, improve the under-
standing of evolutionary processes that led to the observed
diversity of Chilean cacti, and increase the effectiveness of
conservation actions.
AUTHOR CONTRIBUTIONS
PCG designed the entire project (including this study), generated
data, ran analyses and wrote the paper. HEW and IL designed this study
and contributed to the writing of this article. MTKA contributed to the
design of this study. CMP contributed to obtaining the data, the taxon-
omy and systematics of this study. IT and MDB contributed to obtaining
the data of this study. PCG, https://orcid.org/0000-0003-1034-1899;
MTKA, https://orcid.org/0000-0001-6315-0426; CMP, https://orcid.
org/0000-0001-8631-5895; IL, https://orcid.org/0000-0003-0285-722x
ACKNOWLEDGEMENTS
The authors thank the Chilean herbaria (CONC, SGO) curators
who provided access to their plant collections revised for this study. This
study was funded by FONDECYT 3130456 and 1160583, PFB-23,
ICM-02-005, FIBN 9/2015, and research grants from the International
Association for Plant Taxonomy (IAPT), the Connecticut Cactus and
Succulent Society, and the International Organization for Succulent
Plant Study. Camila Munso, Francisca Aravena, Marcelo Rosas and
Glenda Fuentes are gratefully acknowledged for technical assistance.
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Austrocactus spiniflorus (Phil.) F.Ritter, Chile, Santiago: Baños Morales, UDEC 107, H. Walter 42, *MF166919, *MF166993, *MF167067, *MF167134, ;
Blossfeldia liliputana Werderm., Larridon & al. (2015), XX-0-BR-1992101023, LN867998.1, , LN867891.1, ,;Copiapoa hypogaea F.Ritter, Chile,
Copiapó: Chañaral, Cultivated UDEC 194, H. Walter 414, *MF166920, *MF166994, , *MF167135, *MF167207; Copiapoa solaris (F.Ritter) F.Ritter,
Chile, Antofagasta: El Cobre, Cultivated UDEC 5, H. Walter 83, *MF166921, *MF166995, *MF167068, *MF167136, *MF167208; Echinopsis chiloensis
subsp. litoralis (Johow) M.Lowry, ,, AY566656.1, , Franck & al. (2012) JX136750.1, ;Eriosyce aerocarpa (F.Ritter) Katt., Chile, Huasco: Llanos de
Challe, Cultivated UDEC 346, H. Walter 489, *MF166950, *MF167025, *MF167096, *MF167164, *MF167233; Eriosyce andreaeana Katt., Argentina, La
Rioja: Sierra Famatina, Cultivated UDEC, F. Katterman 593, *MF166922, *MF166996, *MF167069, *MF167137, *MF167209; Eriosyce aspillagae
(Söhrens) Katt., Chile, Cardenal Caro: Hacienda Tanumé, Cultivated UDEC 602, H. Walter 156, *MF166923, *MF166997, *MF167070, *MF167138,
*MF167210; Eriosyce aspillagae subsp. maechlerorum Helmut Walter, Chile, Talca: South Constitución, Cultivated UDEC 313, H. Walter 142,
*MF166924, *MF166998, *MF167071, *MF167139, *MF167211; Eriosyce crispa subsp. atroviridis (F.Ritter) Katt., Chile, Huasco: Southwest Vallenar,
Cultivated UDEC 399, H. Walter 134, *MF166931, *MF167005, *MF167077, *MF167146, *MF167217; Eriosyce aurata (Pfeiff.) Backeb., Chile, Elqui:
Portezuelo Tres Cruces., Cultivated UDEC 237, H. Walter 15, *MF166925, *MF166999, *MF167072, *MF167140, *MF167212; Eriosyce bulbocalyx
(Werderm.) Katt., Argentina, San Juan: Marayes, Cultivated UDEC 598, F. Katterman 709, *MF166926, *MF167000, *MF167073, *MF167141,
*MF167213; Eriosyce calderana (F.Ritter) Ferryman, Chile, Copiapó: North Caldera, Cultivated UDEC 136, H. Walter 393, *MF166927, *MF167001,
*MF167074, *MF167142, *MF167214; Eriosyce caligophila R.Pinto, Chile, Tocopilla: Patache, , *MF166941, *MF167016, *MF167088, *MF167155,
*MF167225; Eriosyce chilensis (Hildmann ex K. Schum.) Katt., Chile, Petorca: 3 km North Los Molles, Cultivated UDEC 484, P. Guerrero 872,
*MF166928, *MF167002, *MF167075, *MF167143, *MF167215; Eriosyce chilensis var. albidiflora (F.Ritter) Katt., Chile, Los Vilos: South Pichidangui,
Cultivated UDEC 805, P. Guerrero 873, *MF166929, *MF167003, *MF167076, *MF167144, *MF167216; Eriosyce subgibbosa subsp. clavata (Söhrens ex
K.Schum.) Helmut Walter, Chile, Elqui: Las Rojas, Cultivated UDEC 802, P. Guerrero 1070 , *MF166970, *MF167046, *MF167115, *MF167182,
*MF167252; Eriosyce crispa (F.Ritter) Katt., Chile, Huasco: Freirina, Cultivated UDEC 95, H. Walter 185, *MF166932, *MF167007, *MF167079,
*MF167148, *MF167218; Eriosyce crispa subsp. totoralensis (F.Ritter) Katt., Chile, Copiapó: Totoral, Cultivated UDEC 698, *MF167006, ,
*MF167078, *MF167147, ;Eriosyce curvispina (Bertero ex Colla) Katt., Chile, Aconcagua: Rio Aconcagua, Cultivated UDEC 667, H. Walter 911,
*MF166934, *MF167009, *MF167081, *MF167150, ;Eriosyce curvispina subsp. armata (F.Ritter) Katt., Chile, Santiago: Cerro Cantillana, Cultivated
UDEC 806, H. Walter 912, *MF166933, *MF167008, *MF167080, *MF167149, *MF167219; Eriosyce curvispina subsp. tuberisulcata (Jacobi) Katt.,
Chile, Valparaíso: Laguna Verde, Cultivated UDEC 3, H. Walter 148, *MF166935, *MF167010, *MF167082, *MF167151, ;Eriosyce engleri (F.Ritter)
Katt., Chile, Valparaíso: Roble Alto, Cultivated UDEC 760, H. Walter s.n., *MF166936, *MF167011, *MF167083, *MF167152, *MF167220; Eriosyce
esmeraldana (F.Ritter) Katt., Chile, Taltal: Esmeralda, Cultivated UDEC 212, H. Walter 249, *MF166937, *MF167012, *MF167084, *MF167153,
*MF167221; Eriosyce odieri subsp. fulva (F.Ritter) Katt., Chile, Copiapó: Totoral, Cultivated UDEC 12, H. Walter 281, *MF166955, *MF167030,
*MF167101, *MF167168, *MF167238; Eriosyce garaventae (F.Ritter) Katt., Chile, Valparaíso: Cerro La Campana, Cultivated UDEC 173, F. Katterman
249, *MF166938, *MF167013, *MF167085, , *MF167222; Eriosyce odieri subsp. glabrescens (F.Ritter) Katt., Chile, Copiapó: Totoral Bajo, Cultivated
UDEC 390, H. Walter 280, *MF166956, *MF167031, *MF167102, *MF167169, *MF167239; Eriosyce heinrichiana (Backeb.) Katt., Chile, Elqui:
Trapiche, Cultivated UDEC 671, H. Walter 171, *MF166939, *MF167014, *MF167086, *MF167154, *MF167223; Eriosyce heinrichiana subsp.
intermedia (F.Ritter) Katt., Chile, Elqui: Vicuña, *MF166940, *MF167015, *MF167087, , *MF167224; Eriosyce iquiquensis (F.Ritter) Ferryman, Chile,
Iquique: Junín, Cultivated UDEC 215, F. Katterman 338, *MF166960, *MF167035, *MF167105, *MF167173, *MF167242; Eriosyce islayensis (C.F.Först.)
Katt., Chile, Arica: Poconchile, Cultivated UDEC 135, F. Ritter 200, *MF166942, *MF167017, *MF167089, *MF167156, ;Eriosyce islayensis subsp.
omasensis (Ostolaza & Mischler) G.J.Charles, Peru: Rio Omas, Cultivated UDEC 129, F. Katterman 890, *MF166943, *MF167018, *MF167090,
*MF167157, *MF167226; Eriosyce odieri subsp. krausii (F.Ritter) Ferryman, Chile, Chañaral: North Caldera, Cultivated UDEC 150, H. Walter 116,
*MF166944, *MF167019, *MF167091, *MF167158, *MF167227; Eriosyce odieri subsp. malleolata (F.Ritter) A.E.Hoffm. & Helmut Walter, Chile,
Chañaral: Pan de Azúcar, Cultivated UDEC 55, H. Walter 569, *MF166957, *MF167032, , *MF167170, ;Eriosyce kunzei (C.F.Först.) Katt., Chile,
Appendix 1.
List of taxa sampled.
Taxon, geographic origin, collector and collector number, GenBank accession numbers (rpl32-trnL,trnL-trnF,trnH-psbA,ycf1,PHYC, respectively) for the
sequences used in the analyses. Dashes () indicate missing data. Newly generated sequences are indicated with an asterisk (*) before the GenBank accession
number. Most of tissues were obtained from specimens maintained at botanical gardens: Universidad de Concepción (UDEC) and Botanic Garden Meise (BR).
All of specimens maintained in botanical gardens were collected directly from their habitats or were gown from seeds collected from their habitats. CONC
refers to the herbarium of Universidad de Concepción.
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Continued.
Copiapó: south Copiapó, Cultivated UDEC 661, H. Walter 69, *MF166945, *MF167020, *MF167092, *MF167159, *MF167228; Eriosyce limariensis
(F.Ritter) Katt., Chile, Limarí: Valle del Encanto, Cultivated UDEC 590, H. Walter 054, *MF166947, *MF167022, *MF167094, *MF167161, *MF167230;
Eriosyce marksiana (F.Ritter) Katt., Chile, Curicó: Villa Prat, Cultivated UDEC 691, H. Walter 1, *MF166948, *MF167023, *MF167095, *MF167162,
*MF167231; Eriosyce marksiana var. lissocarpa (F.Ritter) Katt., Chile, Curicó: Tinguiririca, Cultivated UDEC 727, H. Walter 2, *MF166949,
*MF167024, , *MF167163, *MF167232; Eriosyce napina (Phil.) Katt., Chile, Huasco: Huasco, Cultivated UDEC119, H. Walter 457, *MF166953,
*MF167028, *MF167099, *MF167166, *MF167236; Eriosyce napina subsp. duripulpa (F.Ritter) P.C.Guerrero & Helmut Walter, Chile, Norte de Huasco,
Cultivated UDEC 252, H. Walter s.n., *MF166951, *MF167026, *MF167097, *MF167165, *MF167234; Eriosyce napina subsp. fankhauseri (F.Ritter)
Mächler & Helmut Walter, Chile, Elqui: West of Domeyko, Cultivated UDEC 641, H. Walter 401, *MF166980, *MF167055, *MF167125, *MF167192,
*MF167260; Eriosyce napina subsp. lembckei Katt., Chile, Huasco: Freirina, Cultivated UDEC 134, H. Walter 130, *MF166952, *MF167027,
*MF167098, , *MF167235; Eriosyce napina subsp.riparia Mächler & Helmut Walter, Chile, Huasco: Trapiche, Cultivated UDEC 103, H. Walter 487,
*MF166981, *MF167056, *MF167126, *MF167193, *MF167261; Eriosyce occulta Katt., Chile, Taltal: Las Breas, Cultivated UDEC 526, H. Walter 110,
*MF166954, *MF167029, *MF167100, *MF167167, *MF167237; Eriosyce odieri (Lem. ex Salm-Dyck) Katt., Chile, Copiapó: Puerto Viejo, Cultivated
UDEC 553, H. Walter 119, *MF166958, *MF167033, *MF167103, *MF167171, *MF167240; Eriosyce taltalensis subsp. paucicostata (F.Ritter) Katt,
Chile, Antofagasta: North Paposo, Cultivated UDEC 419, H. Walter 237, *MF166976, *MF167051, *MF167121, *MF167188, ;Eriosyce recondita
(F.Ritter) Katt., Chile, Antofagasta: La Chimba, Cultivated UDEC 179, H. Walter 739, *MF166959, *MF167034, *MF167104, *MF167172, *MF167241;
Eriosyce rodentiophila F.Ritter, Chile, Chañaral: Barquito, *MF166961, *MF167036, *MF167106, , *MF167243; Eriosyce senilis (Backeb.) Katt.,
Chile, Choapa: East Salamanca, Cultivated UDEC 805, B. Vergara 182, *MF166964, *MF167039, *MF167109, *MF167176, *MF167246; Eriosyce senilis
subsp.coimasensis (F.Ritter) Katt., Chile, Aconcagua: Las Coimas, Cultivated UDEC 134, H. Walter 9, *MF166962, *MF167037, *MF167107, *MF167174,
*MF167244; Eriosyce senilis subsp.elquiensis Katt., Chile, Elqui: El Tambo, Cultivated UDEC 750, H. Walter 636, *MF166963, *MF167038, *MF167108,
*MF167175, *MF167245; Eriosyce simulans (F.Ritter) Katt., Chile, Elqui: Trapiche, Cultivated UDEC 140, H. Walter 60, *MF166965, *MF167040,
*MF167110, *MF167177, *MF167247; Eriosyce sociabilis (F.Ritter) Katt., Chile, Copiapó: Totoral Bajo, Cultivated UDEC 443, H. Walter 279,
*MF166966, *MF167041, *MF167111, *MF167178, *MF167248; Eriosyce spectabilis Katt., Helmut Walter & J.C.Acosta, Chile, Huasco: East of Carrizal,
Cultivated UDEC 564, J. Acosta 704, *MF166967, *MF167042, *MF167112, *MF167179, *MF167249; Eriosyce strausiana (K.Schum.) Katt., Argentina,
Mendoza: Potrerillo, *MF166968, *MF167043, *MF167113, *MF167180, *MF167250; Eriosyce subgibbosa (Haw.) Katt., Chile, Valparaíso, Cachagua,
Cultivated UDEC, H. Walter 012, *MF166973, *MF167048, *MF167118, *MF167185, *MF167255; Eriosyce subsp.nigrihorrida (Backeb. ex A.W.Hill)
Katt., Chile, Región de Coquimbo, Totoralillo, Guerrero & Rosas 1322 (CONC 187359), *MF166972, *MF167047, *MF167117, *MF167184,
*MF167254; Eriosyce subgibbosa var.castanea (F.Ritter) Katt., Chile, Curicó: Villa Prat, Cultivated UDEC 807, H. Walter 034, *MF166969,
*MF167044, *MF167114, *MF167181, *MF167251; Eriosyce subgibbosa subsp. wagenknechtii (F.Ritter) Katt., Chile, Elqui: Juan Soldado, Cultivated
UDEC 538, H. Walter 043, *MF166975, *MF167050, *MF167120, *MF167187, ;Eriosyce subgibbosa var. litoralis (F.Ritter) Katt., Chile, Coquimbo:
Totoralillo, Cultivated UDEC 87, H. Walter 47, *MF166971, *MF167045, *MF167116, *MF167183, *MF167253; Eriosyce taltalensis (Hutch.) Katt.,
Chile, Antofagasta: Quebrada Guanillos, H. Walter 435, *MF166978, *MF167053, *MF167123, *MF167190, *MF167258; Eriosyce taltalensis subsp.
pygmaea (F.Ritter) Ferryman, Chile, Copiapó: Pan de Azúcar, Cultivated UDEC 62, H. Walter 398, *MF166977, *MF167052, *MF167122, *MF167189,
*MF167257; Eriosyce napina subsp. tenebrica (F.Ritter) Ferryman, Chile, Huasco: West Domeyko, Cultivated UDEC 774, H. Walter 347, *MF166979,
*MF167054, *MF167124, *MF167191, *MF167259; Eriosyce umadeave (Werderm.) Katt., Argentina, Salta: Puerta Tastil, *MF166982, *MF167057,
*MF167127, *MF167194, *MF167262; Eriosyce subgibbosa subsp. vallenarensis (F.Ritter) Katt., Chile, Huasco: Maitencillo, Cultivated UDEC 382,
H. Walter 129, *MF166974, *MF167049, *MF167119, *MF167186, *MF167256; Eriosyce villicumensis (Rausch) Katt., Argentina, San Juan: Sierra
Villicum, F. Katterman 584, *MF166983, *MF167058, *MF167128, *MF167195, *MF167263; Eriosyce villosa (Monv.) Katt., Chile, Huasco: Huasco, Cul-
tivated UDEC 709, H. Walter 187, *MF166984, *MF167059, *MF167129, *MF167196, *MF167264; Eulychnia acida Phil., Chile, Limarí: Socos, Chile,
Cultivated UDEC 452, H. Walter 37, *MF166985, *MF167060, *MF167130, *MF167197, ;Frailea knippeliana Britton & Rose, ,, *MF166991, ,,
*MF167198, ;Frailea mammifera subsp. angelesiae R.Kiesling & Metzing, , Cultivated XX-0-BR-2013064339, *MF166989, ,, *MF167199, ;
Frailea pumila Britton & Rose, , Cultivated XX-0-BR-2013064743, *MF166992, ,, *MF167201, ;Frailea schilinzkyana (F.Haage ex K.Schum.) Britton
& Rose, , Cultivated XX-0-BR-2013064238, *MF166988, ,, *MF167202, ;Frailea sp.,,, *MF166990, ,, *MF167200, ;Haageocereus
chilensis F.Ritter ex D.R.Hunt, Chile, Arica y Parinacota, Cultivated UDEC 529, P. Guerrero 870,, *MF167061, , *MF167203, ;Neowerdermannia
chilensis Backeb., Arica: Puquito, Chile, Cultivated UDEC 126, F. Ritter 199, *MF166986, *MF167062, *MF167131, *MF167204, *MF167265; Parodia
comarapana Cárdenas, , Cultivated XX-0-BR-19841122, LN867999, , LN867892, LN868093, ;Parodia maassii (Heese) A.Berger, Argentina, Jujuy:
Northwest of Iturbe, Cultivated UDEC, J. Acosta373, *MF167063, *MF167132, , *MF167205, *MF167266; Parodia ottonis (Lehm.) N.P.Taylor, ,
Cultivated XX-0-BR-2013061915, LN868001, *MF167064, LN867894, LN868095, ;Parodia stuemeri Backeb., , Cultivated XX-0-BR-19841128,
LN868002, *MF167065, LN867895, LN868096, ;Rimacactus laui (Lüthy) Mottram, Chile, Tocopilla: South of Tocopilla, Cultivated UDEC 133,
P. Guerrero 871, *MF166946, *MF167021, *MF167093, *MF167160, *MF167229; Yavia cryptocarpa R.Kiesling & Piltz, Argentina, Jujuy: La Quiaca,
Cultivated UDEC, J. Acosta 353, *MF166987, *MF167066, *MF167133, *MF167206, *MF167267;
Appendix 1.
Continued.
Guerrero & al. Eriosyce s.l. systematics and taxonomy
573
TAXON 68 (3) June 2019: 557573
... Eriosyce Phil. (see a proposed key below):-The results of the molecular phylogeny by Guerrero et al. (2019b) show that Eriosyce s.l. (see also Kattermann 1994) species from Chile, Perú, and Argentina form a strongly supported monophyletic clade, but only with the exclusion of Rimacactus laui (Lüty) Mottram, a species that had been included within Eriosyce section Neoporteria subsection Chileosyce by Kattermann (1994, as Eriosyce laui Lüthy) and within the "Islaya-group" by Hunt et al. (2006, 2013. ...
... Hoffmann (1998: Lam. 56: Neoporteria eriosyzoides) shows an old plant with yellowish spines; Hunt (2006: Fig. 3) shows an old plant with yellow spines Phylogenetic analyses retrieved seven major clades in the genus (Guerrero et al. 2019b). The first branching clade (Section Eriosyce Katt.) comprises two species from Chile, one from Argentina, and two from Perú. ...
... Chileosyce Katt.), whereas Hunt et al. (2006Hunt et al. ( , 2013 and Hoffmann & Walter (2004) proposed six "groups" (= subgenera) based on the former genera Islaya Backeb., Pyrrhocactus A.Berger, Neoporteria Britton & Rose, Horridocactus Backeb., Thelocephala Ito, and Eriosyce Phil. None of these concepts were corroborated by the results of the molecular phylogeny (Guerrero et al. 2019b). Moreover, at species level, the large "species complexes" E. napina (Phil.) ...
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... This phenomenon has largely complicated the taxonomy of the family, among which South American globose cacti are among the least studied [18]. The tribe Notocacteae (Cactoideae subfamily) harbors several globose species and is regarded as one of the oldest and most narrowly distributed lineages in southern South America [19]. Within this tribe, Eriosyce sensu lato has a complex taxonomic history with a high level of uncertainty, evidenced by the long history of taxonomic changes since its description more than 100 years ago. ...
... Here, we investigate the phylogenetic relationships and molecular diversity across E. curvispina populations in order to delimit species and to understand the sequence of origin of its diversity. To achieve these objectives, we analyzed the phylogenetic relationships within Eriosyce section Horridocactus, the clade where E. curvispina is nested [19], and developed 12 microsatellites markers to further investigate the genetic structure of the species complex. ...
... They are produced from young areoles, forming a circle around the stem apex; flowers are broad, slightly funnelform, and of 3-5 cm long by 3-5 cm wide [23]. To analyze the evolutionary relationships of the E. curvispina complex, we added 33 samples to a backbone phylogenetic data matrix of the genus published elsewhere [19]. Specifically, we sequenced 18 new individuals of the section Horridocactus and 15 new individuals of the sister clade Neoporteria, all from field collected specimens. ...
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... Eriosyce curvispina (Bertero ex Colla) Katt. (Figure 1; Guerrero, Walter, et al., 2019a) is a small globose cactus endemic to central and 2.5-4.6 cm (Mean = 3.6 cm), respectively (N = 30 on 26 plants). ...
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... In general, the coastal desert is more humid and diverse than the inland zones, especially where steep slopes intercept the moisture of the clouds, forming plant communities that have been termed "lomas", "fertile belt" or "fog oases" (Johnston, 1929a;Ellenberg, 1959;Rundel & al., 1991). The coastal Atacama Desert in northern Chile harbors several monotypic endemic plant genera, such as Rimacactus Mottram (Cactaceae; Mottram, 2001;Guerrero & al., 2019), Gypothamnium Phil. and Oxyphyllum Phil. ...
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