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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

August 2019
Taxon 68(3)

DOI:10.1002/tax.12066

Projects: Instantaneous speciation in a sympatric cactus mediated by hybridization and the establishment of reproductive barriers in coastal central Chile.
Biodiversity and Conservation of Chilean cacti

Authors:

Pablo Guerrero

University of Concepción

Mary Kalin

University of Chile

Carol Peña

University of Concepción

Show all 7 authorsHide

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 A–H by P.C. Guerrero; I by H. Villalobos.

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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.

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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).

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Statistics for the 82-sample DNA sequence alignments.

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Historical overview of the taxonomic classification of Eriosyce.

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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,

Mary T.K. Arroyo,

Carol M. Peña,

1,4

Italo Tamburrino,

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 world’s 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

TAXON Guerrero & al. •Eriosyce s.l. systematics and taxonomy

SYSTEMATICS AND PHYLOGENY

68 (3) •June 2019: 557–573

557

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 Buxbaum’s 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 Nyffeler’s (2002) findings. In today’s

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 Backeberg’s 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.

Backeberg’s (and Ritter’s) 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 Berger’s

Pyrrhocactus, Backeberg’sIslaya,Horridocactus and Neo-

chilenia and Itô’sThelocephala under the oldest name Neo-

porteria, but excluded Philippi’sEriosyce. 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 “subgenus”or “group”instead 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 & Eggli’s

(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 Mottram’s

(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: 557–573

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 A–H 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: 557–573

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,

Kattermann’s 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

taxa—the 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

Kattermann’s 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 Kattermann’s 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 Kattermann’s 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: 557–573

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

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 “-raxml”command 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)

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: 557–573

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 “sumt”function 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. S1–S6). 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

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TAXON 68 (3) •June 2019: 557–573

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: 557–573

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 Mottram’s 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 “sections”and “subsections”of Eriosyce proposed by

Kattermann (1994), the “groups”proposed by Hunt & al.

(2006, 2013), and the “subgenera”suggested 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 Ritter’s (1980) proposal

to unite the west- and east Andean taxa under the genus

Pyrrhocactus, but partially support Backeberg’s (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 Kattermann’s 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 Kattermann’s (1994) proposal

to place it within his subsection Horridocactus. In our

Guerrero & al. •Eriosyce s.l. systematics and taxonomy

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TAXON 68 (3) •June 2019: 557–573

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

Kattermann’s 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

Kattermann’s 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 Kattermann’s

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 Walter’s assumption

(2008) and refutes Kattermann’s (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: 557–573

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: 557–573

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°30′lat-

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. –Monville’s 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 jussieui”should be treated as of uncertain

application. Ritter’sPyrrhocactus 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: 557–573

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: 1041–1042. 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 “lectotype”designated 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: 557–573

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 “Chile”as the type

locality, but mentioned that the plants are sometimes covered

by a light layer of snow in winter. Kattermann (1994) chose

material from Balala—Provincia 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 Förster’sEchinocactus 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 Ritter’s well-argued proposal to

refer Förster’s 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., Ritter’s (1980) and

Backeberg’s (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: 557–573

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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TAXON 68 (3) •June 2019: 557–573

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

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TAXON 68 (3) •June 2019: 557–573

Towards a unified taxonomic catalogue for the Chilean cacti: assembling molecular systematics and classical taxonomy

Article

Full-text available

Jun 2022

The competition between floristic catalogues and the nomenclatural issues of the treated taxa, is a problem for the botanical knowledge of countries. Consequently, it seems to be necessary to merge former taxonomical proposals into a unified list based on phylogenetic hypotheses, the rules of nomenclature and dichotomous keys to the Chilean subfamilies, tribes and genera. With this approach we here propose an updated catalogue of the Chilean cacti. It would be necessary to merge the various taxonomic proposals into a unified list based on both phylogenetic hypotheses and the rules of nomenclature. With this approach, we here propose to updated the catalogue of Chilean cactus. A neotype was designated for Echinocactus jussieui. In addition, we present a dichotomous taxonomic key to the Chilean subfamilies, tribes, and genera.

Independent Evolutionary Lineages in a Globular Cactus Species Complex Reveals Hidden Diversity in a Central Chile Biodiversity Hotspot

Article

Full-text available

Jan 2022

Unraveling the processes involved in the origin of a substantial fraction of biodiversity can be a particularly difficult task in groups of similar, and often convergent, morphologies. The genus Eriosyce (Cactaceae) might present a greater specific diversity since much of its species richness might be hidden in morphological species complexes. The aim of this study was to investigate species delimitation using the molecular data of the globose cacti “E. curvispina”, which harbor several populations of unclear evolutionary relationships. We ran phylogenetic inferences on 87 taxa of Eriosyce, including nine E. curvispina populations, and by analyzing three plastid noncoding introns, one plastid and one nuclear gene. Additionally, we developed 12 new pairs of nuclear microsatellites to evaluate the population-level genetic structure. We identified four groups that originated in independent cladogenetic events occurring at different temporal depths; these groups presented high genetic diversity, and their populations were genetically structured. These results suggest a complex evolutionary history in the origin of globular cacti, with independent speciation events occurring at different time spans. This cryptic richness is underestimated in the Mediterranean flora of central Chile, and thus unique evolutionary diversity could be overlooked in conservation and management actions.

Large flowers can be short‐lived: Insights from a high Andean cactus

Article

Full-text available

Aug 2022

Long‐lived and large flowers signify high floral maintenance costs. Species of arid/semiarid climates with large flowers are expected to have short flower life spans and pollination‐induced flower longevity in order to curb high floral water and other maintenance costs. We explored the context‐dependent large flower/short flower longevity hypothesis in Eriosyce curvispina (Cactaceae), a large‐flowered species of the semiarid central Chilean Andes. We determined breeding system, flower visitation rates, and open‐pollination fruit set and quantified floral water content. In a temperature‐controlled field manipulative experiment, we measured potential flower longevity and tested for the presence of pollination‐induced floral senescence. We measured the time span of the complete flower cycle defined as from when the flowers began to open to when they were totally closed, and the time span of fully open flowers defined as from when they were totally open until they began to close. The potential flower life span averaged 2.8 days (complete flower cycle) to 2.3 days (fully open flower). The complete flower cycle lasted 21.5 h, but flowers were fully open for only 10.1 h across days. Flower longevity in days was far shorter than reported for a large sample of species in the area. No evidence was found for pollination‐induced flower senescence as a complementary means for reducing floral maintenance costs. Eriosyce curvispina is self‐incompatible and abundantly pollinated by two megachilid bees. The level of pollen limitation (L = 0.36) was lower than the average reported for self‐incompatible angiosperms. Thus, the short flower life span in E. curvispina is not an impediment for high fruit set. Flowers contain >5 g of water of which >2 reside in >40 petaloid tepals. The amount of water is far higher than in another large‐flowered, non‐cactus species in the area, but only about 15% of that reported in the giant saguaro cactus which has larger flowers than E. curvispina. We explored the context‐dependent large flower/short flower longevity hypothesis in Eriosyce curvispina (Cactaceae), a large‐flowered species of the semi‐arid central Chilean Andes. We determined breeding system, flower visitation rates and open‐pollination fruit set and quantified floral water content. In a temperature‐controlled field manipulative experiment we measured potential flower longevity and tested for the presence of pollination‐induced floral senescence. Flower longevity in days was far shorter than reported for a large sample of species in the area. No evidence was found for pollination‐induced flower senescence as a complementary means for reducing floral maintenance costs.

Phylogenomics of the Caribbean melocacti: Cryptic species and multiple invasions

Article

Jul 2022
TAXON

The widespread Neotropical genus Melocactus of approximately 42 currently recognized species, is most diverse in eastern Brazil and the Greater Antilles, especially Cuba. Species delimitation is notoriously problematic in the group, although this is due in part to a lack of detailed systematic studies, as well as a severely cluttered nomenclatural history. To date, no comprehensive phylogenetic hypotheses have been generated for the clade, although some population genetic and morphological studies exist. We generated the largest phylogenetic dataset of Melocactus to date based on plastome data derived from a genome‐skimming approach for 26 taxa, which provided a framework for understanding species limits and relationships among Caribbean species. Our time‐calibrated phylogeny revealed a mid‐Pleistocene origin for Melocactus, and we resolved three major clades, a Cuban clade, a mostly South American clade, and a widespread Caribbean clade, which also included some South American taxa. Our topology recovered the Cuban clade as sister to the rest of the species, although this placement was poorly supported, and several other Cuban species are scattered throughout the rest of the tree. Biogeographic analyses suggested multiple dispersal events from South America leading to the current diversity on Cuba, as well as other parts of the Antilles. Based on our phylogenetic results, previous hypotheses of species numbers and relationships in the Caribbean generated solely on morphology have, in some cases, been greatly underestimated. Our study shows that plastome data are effective for resolving clades and species limits in Melocactus, although future work will need to include broader sampling and larger datasets to fully resolve relationships in this complicated group of cacti. We describe one new cryptic species for Cuba, Melocactus santiagoensis sp. nov., and provide a new combination (Melocactus lagunaensis comb. & stat. nov.), based on our phylogenetic results and morphological data and typify numerous names in the genus. The genus Melocactus is another striking example of the exceptional diversity that has been generated in the poorly studied, seasonally dry tropical forest of the Greater Antilles.

Atacamallium minutiflorum (Amaryllidaceae, Allioideae), new genus and species from the coastal desert of northern Chile

Article

Mar 2022
TAXON

Atacamallium, a new genus in Amaryllidaceae‐Allioideae‐Leucocoryneae, is introduced with a description of its type Atacamallium minutiflorum, endemic to the coastal Atacama Desert of northern Chile. Phylogenetic analyses of DNA sequences, along with morphological characters and its distribution pattern, support the new genus and place it as a putative sister to Leucocoryne, which is readily distinguished from Atacamallium by its floral morphology. Atacamallium minutiflorum differs from other species of Leucocoryneae by its overall smaller flowers, tepals fused only at the base, six stamens in a single series, and trifid stigma. A morphological description, a distribution map, an illustration, and the assessment of the conservation status of the new species, besides an updated key to the genera of Leucocoryneae, are provided. Additional data from multiple single‐copy nuclear genes are needed to clarify the relationships within the tribe.

Gymnocalycium ×applanatum (Cactaceae, Cactoideae) – the first reported nothospecies between the subgenera Gymnocalycium and Trichomosemineum

Article

Jan 2022

This study examines an assumed hybrid between Gymnocalycium capillense and G. intertextum discovered in a mixed population of the putative parents at Villa Viso in Córdoba Province, Argentina. Eleven quantitative morphological characters (including two ratios) were measured in the natural populations of the presumed parental species and the assumed hybrid. Five of these characters differed between the parental species whereas the assumed hybrid usually showed intermediate values. Eleven qualitative morphological characters were compared for all three taxa, whereby two characters of the assumed hybrid were closer to G. intertextum, one character was closer to G. capillense, six characters were intermediate (but two of them closer to one presumed parent), and three characters were unique. The seed morphology is the most important character separating the subgenera of Gymnocalycium: the assumed hybrid is most similar in this regard to the nominate subgenus, but the seeds are larger than in both assumed parents and have a very broad hilum-micropylar region and conspicuous strophiole at the margin, like in the subgenus Trichomosemineum. Genome sizes estimated by means of flow cytometry shows that G. capillense is tetraploid and G. intertextum diploid, while the assumed hybrid plants are hexaploid (i.e. allopolyploid hybrids). Sequencing and phylogenetic analysis of a low-copy nuclear PHYC gene also revealed that this genomic region of the hybrid combines sequences present in G. capillense and G. intertextum. In combination with the morphological data, these results support the hybrid nature and parentage of the studied plants. The hybrid is formally described here as G. ×applanatum and is the first documented hybrid between the subgenera Gymnocalycium and Trichomosemineum. The results show the potential of allopolyploidy in which different subgenera participate for the evolution of the genus.

Shrub canopy matrix decreases reproductive output of a sheltered plant via pollinator exclusion

Article

May 2021
BASIC APPL ECOL

Shrubs establish microenvironments under their canopies that can favor the growth of other plants. However, the shrub canopy could impede pollination by reducing the number of pollinator visits to sheltered plants, resulting in pollen limitation and decreased reproductive output. We assessed whether the presence of a nurse shrub species (Flourensia thurifera) alters the reproductive output of a sheltered cactus (Eriosyce coimasensis) via the restriction of access by the giant hummingbird (Patagona gigas) to E. coimasensis flowers. During two consecutive years (2018 – 2019), we excluded hummingbirds from individual cacti (using cages) and studied fruit set and seed production in two microhabitats: underneath shrubs and in open sites. In addition, we compared the reproductive mode of E. coimasensis in the two microhabitats. We observed that shrubs limit the reproduction of E. coimasensis, which strongly depends on P. gigas for seed production. Plants in open sites produced 80% more fruit and 76% more seeds than those growing underneath shrubs. The reproduction of caged individuals was low and similar to those growing beneath shrubs. In addition, plants underneath shrubs, but not in open sites, may suffer pollen limitation. Our results offer novel insights into plant-plant interactions and suggest potential trade-offs for sheltered cacti between the mild microclimatic conditions under the canopy, that could lead to larger plants and pollinator preclusion, which decreases the reproductive performance of sheltered plants.

Towards a monophyletic classification of Cardueae: restoration of the genus Lophiolepis (= Cirsium p.p.) and new circumscription of Epitrachys

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Nov 2022
PLANT BIOSYST

Using molecular data and representative species coverage, we confirmed the monophyly of Cirsium sect. Eriolepis and, therefore, we propose to treat it as a separate genus (Lophiolepis). Besides, based on molecular and morphological evidence we segregate Cirsium italicum into the separate genus Epitrachys, sister to a large clade including Carduus, Cirsium s.l. and several allied genera. The name of a new hybrid genus (Lophiocirsium) is also published. Overall, 129 new combinations in Lophiolepis one in Epitrachys and one in Lophiocirsium are proposed.

Extractivist policies hurt Chile’s ecosystems

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Sep 2021

Cactaceae at Caryophyllales.org- A dynamic online species-level taxonomic backbone for the family

Article

Full-text available

Aug 2021
Willdenowia

This data paper presents a largely phylogeny-based online taxonomic backbone for the Cactaceae compiled from literature and online sources using the tools of the EDIT Platform for Cybertaxonomy. The data will form a contribution of the Caryophyllales Network for the World Flora Online and serve as the base for further integration of research results from the systematic research community. The final aim is to treat all effectively published scientific names in the family. The checklist includes 150 accepted genera, 1851 accepted species, 91 hybrids, 746 infraspecific taxa (458 heterotypic, 288 with autonyms), 17,932 synonyms of accepted taxa, 16 definitely excluded names, 389 names of uncertain application, 672 unresolved names and 454 names belonging to (probably artificial) named hybrids, totalling 22,275 names. The process of compiling this database is described and further editorial rules for the compilation of the taxonomic backbone for the Caryophyllales Network are proposed. A checklist depicting the current state of the taxonomic backbone is provided as . All results are also available online on the website of the Caryophyllales Network and will be constantly updated and expanded in the future. Citation: Korotkova N., Aquino D., Arias S., Eggli U., Franck A., Gómez-Hinostrosa C., Guerrero P. C., Hernández H. M., Kohlbecker A., Köhler M., Luther K., Majure L. C., Müller A., Metzing D., Nyffeler R., Sánchez D., Schlumpberger B. & Berendsohn W. G. 2021: Cactaceae at Caryophyllales.org – a dynamic online species-level taxonomic backbone for the family. – Willdenowia 51: 251–270. Version of record first published online on 31 August 2021 ahead of inclusion in August 2021 issue. Data published through: http://caryophyllales.org/cactaceae/Checklist

MEGA7: Molecular Evolutionary Genetics Analysis Version 7.0 for Bigger Datasets

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We present the latest version of the Molecular Evolutionary Genetics Analysis (MEGA) software, which contains many sophisticated methods and tools for phylogenomics and phylomedicine. In this major upgrade, MEGA has been optimized for use on 64-bit computing systems for analyzing bigger datasets. Researchers can now explore and analyze tens of thousands of sequences in MEGA. The new version also provides an advanced wizard for building timetrees and includes a new functionality to automatically predict gene duplication events in gene family trees. The 64-bit MEGA is made available in two interfaces: graphical and command line. The graphical user interface (GUI) is a native Microsoft Windows application that can also be used on Mac OSX. The command line MEGA is available as native applications for Windows, Linux, and Mac OSX. They are intended for use in high-throughput and scripted analysis. Both versions are available from www.megasoftware.net free of charge.

Clarification is needed in the Code for the nomenclatural status of type specimen photographs

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